oro nets : ae ax, eer tie Srey Seesutosse ee Sorbesateceeee stneees sey est usr “: 5 Pisestae eer eatres c ; Sei Sone ety 5 “ oe Reman c : : Z : : : : = 2 : = LC eee Js ob ane: zi sets te. st or - 7 ees > = =e . : = . as Tee EE SS nt iorpel ile E ‘ = eee a - east: aa Toe: * agenda oe crete aa strate fi ERAN SAC YT LEON'S OF THE ROYAL SOCIETY OF EDINBURGH. TRANSACTIONS OF THE ren y & bh SO CTET Y OF EDINBURGH. VOL. XLVI. EDINBURGH : PUBLISHED BY ROBERT GRANT & SON, 107 PRINCES STREET, AND WILLIAMS & NORGATE, 14 HENRIETTA STREET, COVENT GARDEN, LONDON. MDCCCCXI. 1 Published April 15, 1909. May 3, 1909. May 25, 1909. May 27, 1909. June 9, 1909. September 13, 1909. September 24, 1909. October 12, 1909. October 13, 1909. October 21, 1909. December 31, 1909. January 7, 1910. January 29, 1910. February 1, 1910. XV. XVI. XVII. dO MOLE, XIX. XX, XXI. ».©.Q 05 XXIII. XXIV. XXV. XXVI. XXVII. Va Published February 8, 1910. March 3, 1910. March 24, 1910. August 30, 1910. September 21, 1910. January 26, 1911. January 13, 1911. January 21, 1911. February 10, 1911. February 4, 1911. March 17, 1911. March 31, 1911. April 18, 1911. April 18, 1911. NUMBER CONTENTS. PART I. (1908-09.) I. Notes on Hydrodynamics, chiefly on Vortex Motion. By Professor 1g if. Ie VIL. Vill ANDREW Gray, F.R.S., The Glacial Deposits of Western Cornarvonshire. By T. J. Jeuu, M.D. (Edin.), M.A.(Camb.), F.G.S., Lecturer in Geology at the University of St Andrews. (With Four Plates and One Map), Scottish National Antarctic Expedition: Observations on the Anatomy of the Weddell Seal (Leptonychotes Weddellr). By Davin Hepsurn, M.D., Professor of Anatomy, University College, Cardiff (University of Wales). (With One Plate), Supplementary Report on the Hydroids of the Scottesh National Antarctic Expedition. By James Rirconiz, M.A., B.Sc, Natural History Department, The Royal Scottish Museum, . The Meteorology of the Weddell Quadrant and Adjacent Areas. By Ropert C. Mossman. (With Five Plates), . The Monsoons of the Chilian Inttoral (Preliminary Note). By R. C. Mossman, A Carboniferous Fauna from Nowaja Semlja,-collected by Dr W. 8. Bruce. By G. W. Ler, D.Sc, H.M. Geological Survey. With Notes on the Corals by R. G. CarrutHers. (With Two Plates), Scottish National Antarctic Expedition: Osteology of Antarctic Seals. By Roserr B. THomson, M.B., Ch.B., University of Edinburgh. (With One Plate), PAGE 17 57 65 103 137 143 187 Vi CONTENTS. PART II. (1909-10.) NUMBER IX. Lower Paleozoic Hyolithide from Girvan. By F. R. Cowprr Regn, M.A., F.G.S. (With Three Plates), . v X. Report upon the Anatomy and Embryology of the Penguins collected by the Scottish National Antarctic Expedition, comprising : (1) Some Features in the Anatomy of the Pengun; (2) The Embryology of the Penguin: a Study in Embryonic Regression and Progression. By D. Warrrston, M.A., M.D., F.R.C.S.E., and A. CamppELt GEpps, M.D. (With Three Plates), . XI. The Pharmacological Action of Harmaline. By Jamus A. Gunn, M.A. M.D:, DiSe., ; XII. The Pharmacological Action of Protocatechyl-Tropeine. By Professor C. R. Marswatt, ; XII. The Pharmacological Action of Tutu, the Toot Plant of New Zealand. By Professor C. R. MarsHatu, XIV. On the Lnfe-history of Hydrobius fuscipes, L. By Frank Batrour- Browne, M.A. (Oxon.), F.Z.S. (With Three Plates), XV. Strophanthus sarmentosus : its Pharmacological Action and its Use as an Arrow-Poison. By Sir Tuomas R. Fraser, M.D., F.R.SS. L. & E., Professor of Materia Medica in the University of Edinburgh, and AisTeR |’. Macxenzim, M.A., M.B., Ch.B., Carnegie Research Scholar. (With Eleven Plates), PART IIL (1910-11.) XVI. The Aborigines of Tasmania, Part I1.—The Skeleton. By Principal Sir Wittiam Turner, K.C.B., F.R.S., President of the Society. (With Two Plates), XVII. On the Fossil Osmundaceew. Part lV. By R. Kinston, LL.D., F.R.S., F.G.S., Foreign Mem. K. Mineral. Gesell. zu St Petersburg, Hon. Sec. R.S.E.; and D. T. Gwynne-Vaucuan, M.A., F.L.8., Professor of Botany, Queen’s University, Belfast. (Plates I.-IV.), PAGE 203 223 245 273 287 317 341 411 455 NUMBER XVIII. XIX. XX. XXI. XXIT. XXIII. XXIV. XXV. XXVI. CONTENTS. The Lamellibranchs of the Silurian Rocks of Girvan. By WuHeExuron Hinp, M.D., B.S, F.R.C.S., F.G.8. (With Five Plates), The Alcyonaria of the Cape of Good Hope and Natal. Alcyonacea. By J. Stuart Tomson, Ph.D., F.L.S. (With Four Plates), The Phase of the Nucleus known as Synapsis. By A. ANSTRUTHER Lawson, Ph.D., D.Sc, F.LS., F.R.S.E., Lecturer in a University of Glasgow. (With Ta Plates), . Observations on the Body Temperature of the Domestic Fowl (Gallus gallus) during Incubation. By SurHertanp Simpson, M.D., PART IV. (1910-11.) The Temperature Seiche. Part I.—Temperature Observations in the Madiisee, Pomeranias Part L1.—Hydrodynamical Theory of Temperature Oscillations in Lakes. Part [11.—-Calculation of the Period of the Temperature Seiche in the Madiisee. By EH. M. WeppERBURN, W.S. Part 1V.—-Experimental Verification of the Hydrodynamical Theory of ‘Temperature Seiches. By EH. M. Wepprersurn, W.S., and A. M. Wiiiiams, M.A., B.Sc. (With One Plate), The Jurassic Flora of Sutherland. By A. C. Sewarp, F.R.S., Professor of Botany, Cambridge. (Plates 1—X.; text-figures 1-14; sketch-maps, and views of the coast), On the Structure and Affinities of Diplolabis rémeri (Solms). By W. T. Gorpon, M.A., B.Se., B.A., Falconer Fellow of Edinburgh University, Lecturer in Paleontology, Edinburgh University. (With Four Plates), . Thermo- Electric Diagram from — 200° C. to 100° C., deduced from the observations of Professors Dewar and Fleming. By J. D. Hamitton Dickson, M.A., Peterhouse, Cambridge, The Plant Remains in the Scottish Peat Mosses. Part IV. By Francis J. Lewis, M.Sc., F.L.S., Lecturer in Geographical Botany, University of Liverpool. (With Five Plates), ; Ne! fe ; {i } Tee eee cs (hp viwt A. Si ter nn of) p af dthowy j } ; - vil PAGE 479 549 605 619 643 Gul 737 793 vill NUMBER CONTENTS. XXVIII. On an Entoproctan Polyzoon (Barentsia benedenr) new to the British Fauna, with Remarks on Related Species. By James RitcHix, M.A., B.Sc., Natural History Department, The Royal Scottish Museum. (With One Plate), XXVIII. An Investigation into the Effects of Errors in Sea a HENRY INDEX, Briaes, B.Sc., A.R.S.M., an PAGE 835 849 879 TRANSACTIONS - OF THE ROYAL SOCIETY OF EDINBURGH. VOLUME XLVII. PART I—SESSION 1908-9. CONTENTS. I. Notes on Hydrodynamics, chiefly on Vortea: Motion. By Professor ANprew Gray, F.R.S., . 1 (Issued April 15, 1909.) Il. The Glacial Deposits of Western Carnarvonshire, By T. J. Juuu, M.D. (Edin.), M.A. (Camb.), F.G.S., Lecturer in Geology at the University of St Andrews. (With Four Plates and One Map), ‘ : : 2 : : : : ; : 17 (Issued May 3, 1909.) X IIL. Scottish National Antarctic Expedition: Observations on the Anatomy of the Weddell Seal (Leptonychotes Weddelli). By Davip Hersurn, M.D., Professor of Anatomy, University College, Cardiff (University of Wales). (With One Plate), : (Issued May 25, 1909.) or ba | IV. Supplementary Report on the Hydroids ‘of the Scottish National Antarctic Expedition. By James Ritcaiz, M.A., B.Sc., Natural History Department, The Royal Scottish Museum, 65 (Issued May 27, 1909.) V. The Meteorology of the Weddell Quadrant and Adjacent Areas. By Roprert ©. Mossman. (With Five Plates), < ; : : ; ; : s Haat (5) (Issued June 9, 1909.) VI. The Monsoons of the Chilian Littoral (Preliminary Note). By R. C. Mossman, : Rye (Issued September 13, 1909.) VIL. A Carboniferous Fauna from Nowaja Semija, collected by Dr W. S. Brucz, By G. W. Luz, -D.Se., H.M. Geological Survey. With Notes on the Corals by R. G. Carruruers. (With Two Plates), : : 4 t : . : 5 s. @ 143 (Issued September 24, 1909.) | X VIII. Scottish National Antarctic Expedition: Osteology of Antarctic Seals. By Roser B. Tuomson, M.B., Ch.B., University of Edinburgh. (With One Plate), . : oer LST (Issued October 12, 1909.) EDINBURGH: : PUBLISHED BY ROBERT GRANT & SON, 107 PRINCES STREET, AND WILLIAMS & NORGATE, 14 HENRIETTA STREET, COVENT GARDEN, LONDON. MDCCCCIX. Price Nineteen Shillings and Ninepence. * x re 8 a ae, es eee by a = ri 1 1 aed Pelel aires f :\ foe Lan eae |, ~ caren ae a sae vat fe WS. a 2m . > =. —_ , . 4 ta iat é ‘ so Fak.’ . wa - - ao Maes en TRANSACTIONS. ].—Notes on Hydrodynamics, chiefly on Vortex Motion. By Professor Andrew Gray, F.R.S. (MS. received November 6, 1908. Read November 16, 1908. Issued separately April 15, 1909.) 1. In the teaching of hydrodynamics many points of difficulty arise, both for teacher and for student. The subject abounds in subtleties even in its very elements, and the advanced student frequently finds himself in a state of doubt as to fundamental questions which crop up unexpectedly in connection with various problems. The following notes contain a discussion of a few of these fundamental matters: for example, they deal with some theorems of energy which have been found difficult by students, perhaps mainly through want of perfectly explicit state- ment of their scope and purpose. Finally will be demonstrated a theorem of vortex-motion, particular cases of which have been given by various writers, but which I have not seen stated elsewhere in the same generality. This will be found to lead to Lorp Ketvin’s well-known and far-reaching theorem of circulation, and to other theorems of the vortex-motion of a perfect fluid, some of which are already known. 2. First it may be recalled that if gq be the resultant velocity of a particle of the fluid at any point P at time ¢ (or indeed any other quantity characteristic of an element of the fluid in motion), and ds the element of path actually described by the particle in the interval of time dt beginning at ¢, the acceleration of the particle is This, of course, is the usual expression from which the component accelerations parallel to the axes are deduced. But if ds’ be an element of a line drawn from P at any angle to ds, and q’ the component of velocity in the direction of ds’, the acceleration in this direction is nC at a5" This is a very useful expression, and will be of great service in what follows. By putting u, v, w for g’ we obtain easily the usual component accelerations. That the acceleration along ds’ has this value is at once evident from the fact that, at time ¢, rg TRANS. ROY. SOC. EDIN., VOL. XLVII. PART I. (NO. I). 1 2 PROFESSOR ANDREW GRAY, the velocity in the direction of ds’ at a point Q, at a distance ds in advance of P. along the line of flow, is q’+0q’/és.ds. After the lapse of the interval dt, the particle which was at P has reached Q, and its velocity (along ds’) has become, to the first order of small quantities, q’+0q'/ot.dt+0q'/os.ds. Since q=ds/dt, the particle when at P is thus gaining velocity in the direction of ds’ at rate se + : ; : : (1). 3. An elementary point of some difficulty to the student arises in this connection. The integral [as ot is to be differentiated with respect to ds’, specified as in §2. The result is d | oq aq ds _ 0q' d — — F = = 5 4 she @ Oh ds CF (2) Here we have ds/ds’=cos 0, where @ is the angle between ds’ and the terminal element of the line of integration. It is at first a little dithcult to see that ds/ds’ has this value. But the whole question is, What is the change in /0q/dt.ds produced by a small step ds’ inclined at an angle 6 to ds? Now the step ds’ may be regarded as made up of a step ds’cos@ along the stream-line, followed by one ds’sin#@ at right angles to the stream-line. The former gives the element 0q/dt. ds’ cos@ of the integral; the latter, being at right angles to the stream-line, leaves the integral unchanged ; therefore the result of the differentiation is aq/dt . cos 0, or 0q’/dt. The student is at first tempted to take ds’ as one component of a step ds along the stream-line, that is, as dscos@. He forgets that if this is done the element dq/ot. ds of the integral, which he now makes to correspond to ds’, is the result, not of the step ds’ alone, but in part also of the coexistent step ds sin 0. 4. The flow of energy in the hydrodynamic field may be discussed in two ways. We may determine the rate of change of energy within a closed surface S fixed in the fluid, or we may find the rate of change of the energy of the definite mass of fluid which at time ¢ occupies the space within the given surface 8. These are two distinct problems—since at time t+dt the fluid which at time ¢ occupied the space within the surface no longer does so precisely—and a comparison of their solutions is instructive. As we shall see presently, the solution of one problem can be derived from that of the other; and the application of the results to particular cases gives various well-known hydrodynamical theorems. 5. We take first the problem of the rate of change of the energy of the mass of fluid contained at time ¢ within the surface 8S. Let g denote, as usual, the resultant velocity of a particle of the fluid, and q its total time-rate of variation. If, as is usually the case, the external forces (per unit mass) be derivable from a potential—which we shall call V—we have sev Gop Ae Re oe do ate NOTES ON HYDRODYNAMICS, CHIEFLY ON VORTEX-MOTION. 3 Multiplying both sides by dm, an element of the fluid mass, and integrating for the whole mass of fluid within the surface, we obtain | qgdm = — | qodm— | qPao (3), if dw be the volume (dm/p) of the element of mass dm. Now usually V is independent of ¢, and therefore 0V/ot=0. Thus for gdV/ds in the first integrand on the right we can write 0V/ot+q0V/ds, which is clearly dV/dt. Thus if T denote the kinetic energy of the fluid which fills the surface -at time ¢, and KH denote the potential energy from which the external forces are derived for the same fluid mass, we have dT [dV dE = m= el dV, _dE [adam at’ i dt dt’ The equation found above can therefore be written in the form di MA fixep S(T+E)= [Pao ! : empty This gives, in an exceedingly compact form, the value of the total time-rate of variation of the sum of the kinetic and potential energies T and H. 6. The integral on the right can be transformed by partial integration, without the introduction of Cartesian co-ordinates, by proceeding as follows. The continuity of the motion from point to point in the fluid involves the possibility of dividing the fluid mass up into narrow tubular portions bounded by non-intersecting stream- lines. We shall call these tubes of flow. Their distribution is determinate for any instant (or point of time), but varies in the general case from instant to instant. Hach such tube (unless it be endless, and entirely contained within the surface) will enter the closed surface S at one element dS, of surface, and emerge at another element dS,. We shall denote for any element of one of these tubes, say an element of length ds, the average area of cross-section by «; then o« varies along the tube, that is, it is to be regarded as a definite function of s, at each instant of time. The element of volume dw will now be cds. 7. First, then, integrating the expression on the right of (4) by parts, we get as the integrated terms for a single tube — p,q.o,+ 191%), Where 71, v, are the cross-sections of the tube at entrance into and emergence from the surface 8. If q, be the com- ponent of velocity normal to the surface inward from an element dS of the surface, this sum of terms may be written (pqndS).+(pqnd8),. The aggregate result for all the tubes crossing the surface is the integral | PQAS taken over the surface S. Thus we obtain, since dw =cds, d shel Shee Siig: G{t+B)= [pads + [ph £(qo)aa 0g , g oo = dS MS, ; 5). |r + o(Z+22\a () 4 PROFESSOR ANDREW GRAY, If there be endless tubes of flow contained entirely within the surface, no change in the surface integral in this result is required. The volume integral on the right, however, must be evaluated for each such tube. The expression is the “divergence” of the fluid within a tube of flow, that is, the time-rate of increase of the volume of the fluid, per unit of volume, at the element ds. This expression is equivalent to du, ow, ow Baney Aue? the divergence expressed in terms of the component velocities with reference to rectangular axes. [It is not difficult to verify this equivalence. | Equation (5) may thus be written in the form d ow dv. ow - —= — y) —_— —_ — d.: . 6 ithe + ») [r(0u-+-mo-+moyas | | [r(S+ 2+ =) xdydz (6), where /, m, 7 are the direction-cosines of the inward drawn normal to the element of surface d8. 8. From the equation (5) [or (6)] we can deduce the value of 0 2 a | (Zeq’ + pV)d@, that is, of the time-rate of variation of the energy within the fixed surface 8. Let U denote the volume of the space within §, then this at time ¢ is the volume of the mass of fluid which we have been considering. Let also e denote the average value of the sum of the kinetic and potential energies per unit of volume; then for the whole mass of the fluid T+H=Ue. Thus d dU de ap fe oo ee Be a ae ee therefore de a dU Jeena Wipe UF a ape for U, considered as the volume of a definite mass of fluid, is subject to change. This by § 7 is the same thing as the equation i o (bp + pV)da = | pq, + i de +2 aes oq i=) — | (Log? ot pt |; [0a +pv(Z44 a5 iD ; Qi): But by § 2 d 9 : (beg? + pV) = ai’ pq? +pV)+ 12 (3 egepN): NOTES ON HYDRODYNAMICS, CHIEFLY ON VORTEX-MOTION. 9) Thus we obtain | der + pV )l@ = Jonas + ine + ¢ 2 ae J ot Os oo Os - [eg+9¥) 14222) a0 é - [a2 Gor + pV)ae. Remembering that {éq/és + (q/c)0o/és} dw =0(qo)/es. ds, and integrating the second line on the right by parts as in § 7, we obtain ie a bzed + pV )d@ = i (207° + pV + p)gndS + |o(S+ 12 a0 . (8). The integral on the left of (8) is the rate at which the sum of the kinetic and potential energies within the surface is increasing ; and the integrals on the right show that this rate is equal to the rate of flow of energy into the space across the surface, together with the rate at which work is done by pressure in consequence of expansion of the fluid within the space and of passage of fluid across the surface. 9. The value of the divergence found above for a tube of flow leads to a corre- sponding mode of expressing the equation of continuity of the fluid. If p be the density of the fluid at the element of volume da, then, since the increase of volume in time dt is (dq/és+q/o .dc/ds)dt per unit volume, the mass for this at density p is p(oq/ds + q/c . do/ds)dt. But since the mass of the element is not changed, for it preserves its identity, this expression must have the value —dp/dt.dt. Thus we obtain the equation of continuity in the form Ba a) or ; ‘ : : (9). dp dp og gor ae ae 5) 79 If we put © for the divergence, this equation can be written dp @=0 . & Vo} are (10) 10. As an example of these results we apply them to the steady motion of the fluid within a portion of a tube of flow bounded by cross-sections perpendicular to the stream-lines. In this case the surface-integral on the right is limited to the ends of the tube. Let dS,, dS, be the areas of the ends, and q,, g, be the velocities there along the tube. Then the surface integral becomes (4ai+ Vi +2) pad, - (408+ Ve +2!)padS, 1 ‘ 2 But since the motion is steady, p,q,dS, = pyqodSq, since the former is the mass of fluid 6 PROFESSOR ANDREW GRAY, which flows into the tube, and the latter the mass which flows out, per unit of time. Denoting this constant quantity by «, we obtain Ja pg? + pV +p)q,d8 = r(dai+V, sae Va Es) : : (11). 2 Now consider the volume-integral on the right of (8). By the equation of con- tinuity given in § 9 we have d [reac = - n|r% ales where the integral on the right is to be taken along the tube here considered, in the direction of flow. Integrating by parts, we find oq ELA Bowe [oe Co =a = (2 Py Pi p ): This added to the former result gives 0g q 0a oe a er) ho ee ” dp [@ pg’ + pV + p)gndS + po +1 Vom =n( i+ Vi 22 — Vo We 2) . (13). Thus, since the left-hand side of (13) vanishes in the case of steady motion, we obtain finally for any cross-section of the tube the well-known equation ig + V+ [P=const : 5 : c (14), which is generally obtained hy another process. It is to be understood that the integral, /dp/p, is to be taken along the stream-line, from any chosen starting-point up to the cross-section to which g and V belong. [tis thus shown that $q?+ V +/dp/p is constant along a stream-line ; but its constant value, itis to be observed, may change from one stream-line to another. 11. We shall now find an expression for aD OTe | [#as+ 40 7 Be) p in the general case. This will show that when the fluid is destitute of what may be called elemental rotation the value of this expression, which is a function of the time, is the same throughout the fluid at any one instant, and becomes an absolute constant when the motion is steady. The theorem thus arrived at will be found to lead at once to Lorp Ketvin’s theorem of circulation, from which the permanence of vorticity, or non-vorticity, in any portion of a perfect fluid, can be inferred. It will be shown also how another relation can be obtained, which affords another view of the proof of the theorem of permanence. Let \ denote [aes yer v+ (2, ot p NOTES ON HYDRODYNAMICS, CHIEFLY ON VORTEX-MOTION. a where ds is in the direction of the resultant velocity g, and p is as usual a function of p. The integrals are supposed taken along the stream-line from any chosen starting-pomt to the point considered. If now we differentiate in any direction inclined at an angle 6 to the direction of ds and q, we get Ow _0q' , Og , OV , lap ag af * Vag ay * 5 a5” But we know that oq’ 0g’ oV -o ; Ot oe as” a for the first two terms make up dq’/dt, and this, by the equation of motion for the direction ds’, is —(0V/ds’ + 1/p . dp/ds’). Thus by subtraction we get Op (aq _ 6 as’ a ds ) But if ,, denote the component of ele- mental rotation of the fluid about the normal to the plane of ds and ds’, at P their point of meeting, as indicated in the figure, we can easily prove that — Que sin =o : : : : (15), so that Weer gsn ct ES 0s 12. I have not seen this theorem stated before, though various particular cases of it are known from which it might be inferred. It asserts that, at time ¢t, ~ is constant along a stream-line, and likewise along any line ds’ so drawn that ,, is zero for the plane determined by ds’ and ds. is in general a function of ¢, and the theorem shows how its variation from point to point depends on the motion of the fluid. By turning ds’, without altering 9, we can change the plane of ds and ds’ from that for which ,, sin 8 is zero to that—inclined to the former at an angle of 90°—for which Sin # is a maximum. The normal to the latter plane may or may not be the direction of the axis of resultant rotation at P (see figure). 13. Equation (15) may be regarded as an equation of motion for the direction ds’ inclined at the angle @ to ds, the direction of flow. If we take ds’ parallel to the fixed axes Ox, Oy, Oz in succession, we obtain three equations :— CG) oy ie wey sin sy = 0 Gi : ~ + 2oseg sin O57 = 0, ay (17). 2weegq sin O,2= 0 8 PROFESSOR ANDREW GRAY, If, according to the usual notation, we put a — these equations become ow oy a). + 2wy — 206=0, ay toed 208 =o] Y (18), ow ae 2vé — 2un =0 | which may be regarded as the Cartesian equations of motion of the fluid. These equations are given in Lams’s Hydrodynamics, 3rd edition, § 146, but are derived directly from the ordinary Eulerian equations of motion. 14. The proof of equation (15) deserves a little attention. We can show that the equation holds, and also make it clear that o, is an angular velocity of rotation— a point that is not always brought out in the kinematics of vortex-motion-—by pro- ceeding as follows. The average velocities along the four sides of the small parallelo- gram of the figure may be taken to be the velocities of the fluid in these directions at the middle points of the sides. These are, for the sides PQ, QR, RQ’, Q’P, aq 1 OY 5. OF 5 q+ 3-.ds, qta—ds+tards, é é aq’ = (a + yas + sys ) “- Ci + bends’). Multiplying the first and third of these by ds, and the second and fourth by ds’, and adding, we obtain - ~ “A asa ds ds’ Thus if g, be the component of velocity along the boundary of the parallelogram at any element dc, we have for the parallelogram = — | ate = @ =e dsds’. But the angular velocity of a fluid particle at dc about a point, P” say, in the plane of ds and ds’ and within the parallelogram, is q,/p, where p is the length of the perpendicular from P” on the side upon which dc is situated. Thus Je ae = (4-1) aaas. [tp ae = (Z-F) asa If , be the mean angular velocity for particles at different points of the periphery of the parallelogram, the integral on the left is evidently 2,,dsds’sin@. Thus we obtain, as stated in (15), NOTES ON HYDRODYNAMICS, CHIEFLY ON VORTEX-MOTION. 9 15. Returning now to equation (16), we write imstead of W, x+Jdq/ot. ds, so that x=i¢+V+ [2 praey Jp Equation (16) may thus be written 0 , == = Boge qsin 6 — E : (20). As in the diagram, let AB be any line drawn in the fluid, and PQ be the element ds’, which we suppose to be on this line. As before, g is the resultant velocity of the fluid at P, and q’=q cos @ is the component at P along the tangent ‘to AB at P. Now qsin@ is the velocity at P at right angles to ds’, that is, gsin@ is the velocity with which each point of the path AB is moving at P towards the right in the plane of the paper. The product qsin@ds’ ~, is therefore, the rate at which any circuit including ds’ is increasing in area, in consequence of the motion of ds’ at right angles to itself, in the plane of the paper, without alteration of length. Multiplying (20) by ds’, and integrating from A to B, we obtain X=X= - | Qu, ain Oda! — | a eels We see that the first integral on the right is twice the rate at which the surface integral of elemental rotation, taken over any surface of which AB is part of the boundary, is changing in consequence of the fact that each element ds’ is being carried at right angles to itself by the motion of the fluid. It is also, it is to be noted, the rate at which the flow of the fluid along AB is changing from this cause. 16. But the area enclosed is also changing in consequence of the motion of one end of the element ds’ relatively to the other. The surface integral is changing from this cause also, and the rate of change can be found most easily by calculating the rate of inerease of flow along AB arising from the change of length of the element. To do this, consider the new position of ds’ after the lapse of time dt. Let the plane of the initial position of ds’, and of the displacement qdt of P, be taken as a plane of reference. The other end Q of ds’ has been displaced through a distance (q+0q/as’. ds’)dt which is not in this plane, and is not parallel to the displacement qdt of P. The increase of the rate of flow of fluid along ds’ arising from the difference (4q/ds’ . ds’)dt is gaq/ds’. ds’. If this is not intuitively perceived, it can be proved as follows :— The component of the displacement (q+0q/ds’.ds')dt of Q alone ds’ is (q + 09/05". ds’)dt. cos (8+ 6), or (q cos 0 — qdO. sin 0 + 2y/2s’ . ds’ cos 0)dt. Thus the end TRANS. ROY. SOC. EDIN., VOL XLVII, PART IL (NO. 1). 2 10 PROFESSOR ANDREW GRAY, Q suffers in dt a displacement relatively to P of amount (@q/ds’ . ds’ cos 8 — qd6. sin A)dt, parallel to its original length. The additional flow due to this displacement is (q’ 0q/0s’ . ds cos 6 — qq’d@. sin 0) dt, or (at as cos? 6 — q?d6. sin 6 cos 6 )at, since q’ = q cos 8. But Q has also moved out through the distance (q+ 0q/0s’ . ds’)dt. sin (6+d0) at right angles to the original position of ds’. This is not parallel to q’ sin @dt, the displacement of P, but is so to the first order of small quantities. (This degree of approximation is, however, sufficient for the present calculation, as the velocity at Q in the direction at right angles to the plane ds, ds’ is a small quantity of the first order. Thus the flow due to the deviation from the plane ds, ds’ must be a small quantity of the second order.) We have to the degree of approximation required for the present purpose {(q + 0q/0s’. ds’) sin (0+ d0) —q sin 0} dt, or (a cos 0 d6 + of as sin @)at for the only other relative displacement that need be considered. The velocity in the direction of this displacement is approximately q sin @, and therefore the component of flow along the element due to the displacement is ( q’ sin 6 cos 640 + qetads sin? 6) dt. s Thus the whole change in the rate of flow brought about by the two displacements is oq Hj 29) — 0g 3 Iaads (sin? 6 + cos? @) Ia48. But the element ds’, carried along by the motion of the fluid, is displaced as a whole in time dt through a certain distance parallel to its own direction, and this, apart from alteration of length or direction of the element, causes an alteration of the rate of flow along it. The amount of this is easily shown to be gdq/és’.ds’. For the total growth in time dt of flow along ds’, with its given length and direction, is (aq’/at + gaq'/ds)ds'dt. This is due to the longitudinal and lateral displacements in dt taken together. But by (15), 09’/ds = 0q/0s’ + 2,, sin 8. Thus for the element ds’, with its given length and direction, the rate of increase of flow is (d9q’/at + goq/as’ + 2,.g sin 0)ds’. 17. We therefore have finally (7X - 2458 \as' = — Qtr, Sin gas’ — 24-2 as' - “Las . (22), The expression on the right is twice the total time-rate of change of the surface- integral of rotation caused by the motion of ds’, together with twice the time-rate of change of the same integral due to variation of velocity while ds’ remains at rest. Integrating along AB we get x= xX -=@igan8 | gery Sin aus’ — 2 | gids’ - | of as , Sesion. 3 AB AB AI NOTES ON HYDRODYNAMICS, CHIEFLY ON VORTEX-MOTION. iL The first two terms on the right give twice the rate of change of the surface integral of elemental rotation (for any closed path of which AB forms part) due to the motion and the lengthening of the elements of AB; the last gives twice the rate of change of this integral when the circuit is without motion. 18. If we integrate round the closed path, then under the conditions already stated as the dependence of p upon p, and the existence of the potential V, we get from the vanishing of the left-hand side | 2q% sin Ods' + 2fastas + [fe = . P : (24), j s and we see that the surface-integral of elemental rotation remains constant as the closed path moves with the fluid. It is to be noticed that for any closed path moving with the fluid [Bae — [Pee sin 6ds'’ . . . . (24’). Equation (23) with the value of x inserted is d 2 d : 1 0g , 0 ; , Ve+ | P 44° —(Va+ | oP 4q?)= - 2f qowsin Bas z 2 qas - [Bas . 5), B A AB and is equivalent to Lorp Ketvin’s theorem. The process of investigation here em- ployed, though longer than that usually adopted, has the advantage of tracing the various causes of change of the line-integral of flow along the path AB, and of throw- ing some additional light on the meaning of the theorem. 19. If we take AB parallel to the axis of x, we obtain [ 2a00sin Ods' = - / (206 — 20n)da : : (26). AB ‘AB Let us now consider a fluid in motion in virtue of vortices contained within it, and suppose that a surface S can be described within the fluid so that it is everywhere at a finite distance from the vortices. We can describe a closed path, consisting of two parts: (1) a part parallel to the axis of «, starting from a point A near the surface §, and ending at a point B, also near the surface, but remote from A; and (2) a part starting from B and returning to A, but kept so near the surface everywhere as to avoid the vortices. If this path move with the fluid, then by (24’) | Fae + | 2d Ws sin Oaate + | as + i 2quss' sin Ods' =0 . . (27). AB AB BA “BA This equation can be written ede | (206 — Qua) + | de + | Qow,y sin ds =O Om ‘AB AB "BA BA But going back to (21), we see that the last two terms on the left are Xz—X,, and as at every point of this part of the closed path »,,=0, we have _ { Ya. 2 je xe = | ie (28). BA 12 PROFESSOR ANDREW GRAY, If, further, the surface be so far from the vortices that the fluid is at rest [or, when this is not the case, the motion be steady], we know that 0g" a6 — [fas =) Hence in either of these cases X,=Xz. We have then | wd -{ (2v€ — 2wn)du =0 ; : : (29) ; “AB AB and we see that | | [ Sanayae - | [feu — 2wn)dadydz=0 . . (30), where the volume integrals are taken throughout the fluid. In the case of steady motion this gives | [ [eve - 2wn)dadydz=0 Similarly, | [ [(eue- 2ul)dudydz=0 : s ; (31). i | | (Qu — 2né)dadyde = 0 These equations are usually obtained only for the case of the fluid contained within a surface which is at every point infinitely distant from the vortex system. Unless there is some error in the reasoning here stated this restriction would appear to be unnecessary. | 20. A vortex-line is defined as a curve the tangent to which at any point is in the direction of the resultant rotation at the point. Since 0v)/ds’ is zero when ds’ is taken along a stream- line or along a vortex-line, it is zero for every direction in the plane determined at any point by the stream-line and the vortex-line through the point. It has therefore a maximum value at the point along a normal to the plane contained by these directions. 0 A tubular space bounded by vortex-lines and filled with fluid in elemental rotation is called a vortex-tube. Consider a portion of a vortex-tube between two cross-sections formed by ds, an element of a stream-line, and ds’, taken at right angles to the plane of the stream-line and a vortex-line at a point on the surface of the tube. Such a portion of a tube is shown in the diagram by P’A’CD’/ BQ. P’A’ and QB are taken at right angles to the planes of the stream- line and vortex-line at P’ and Q, and represent ds’ taken at these points. P’Q, A’B, CD’ are vortex-lines, and P’C, QD’ represent ds taken at the same points. Since ds is not necessarily at right angles to the vortex-line at P’ or Q, let PC, QD be so taken. Then in PAC DBQ we have a portion of a vortex-tube with its D NOTES ON HYDRODYNAMICS, CHIEFLY ON VORTEX-MOTION. 13 ends perpendicular to vortex-lines, and PA, QB are the directions of ds’ at P and Q. Now for the plane P’A’C we have, since P’A’ and P’C are at right angles, oy = — 20559, 0s or pe Onda os Thus, for the surface integral of rotation over P’A’C we have ow eee ee ie 9 le =—-tossdsds . : . . (32). But if be the resultant angular velocity of rotation of the fluid about the vortex- line at P’, and > the angle between P’C (ds) and P’Q (the vortex-line), @ SIN d= age’, so that 1Wat ds’ = - Luds ds.sin : . (33). s But 4dsds’sin¢ is the area of the end PAC of the portion of a vortex-tube. Calling this 7, we have Wa dt=-o0 es: We can prove that this is the same for both sections of the tube. or, consider the circuit PABQP, made up of the vortex-lines PQ, AB, and the two intercepted lines ds’, namely PA, QB. The line-integral reduces to dv-/ds’. ds’ for PA, and a similar expression, with the minus sign prefixed, for BQ. But for any closed circuit moving with the fluid the integral vanishes, and, since Y does not vary along a vortex-line, 0v//0s'.ds’ is the same for PA and AB. Thus for both ends of the vortex-tube we have the same value of wc, since we have to take ds at the two sections to correspond to the same dé. Along a vortex-tube, therefore, wc is constant; and the tube must either be endless or have its ends on the free surface of the fluid, if a free surface exist. In both cases it is really endless. 21. [February 12, 1909.] The equations of acceleration for any two directions drawn from the point P are by § 2 Ou + ou ov IL op sia once p ox ov. du oV 1 Op +g—= -——= Gen acu ays 2. | (35), where dx, dy are infinitesimal steps, and u, v the velocities, in these two directions. 14 PROFESSOR ANDREW GRAY, For simplicity we take the two directions at right angles to one another. If we differentiate the first of these with respect to y and the second with respect to x, and subtract the first result from the second, we obtain dA dw dw . ,ow dw — (0) =o —_ i. — ai * 6 ar Be He ee + me ty es (36), where &, 7, C are the components of spin at P about axes perpendicular to the three rectangular planes yz, zz, xy; wu, v, w the component velocities at P; and © the divergence. Two similar equations hold for €, 1, and can be written down by symmetry. These equations are equivalent to those given by v. HELMHOLTZ, as generalised for the case of a fluid of non-uniform density. By writing 1 dp Os See p dt the equations can be put in the more usual form, and in another, from the second expression on the right of (36). Now let ¢ be the resultant spin at P, that is, let the axis of z be taken along the direction of spin. Then at P, ==0, and we obtain, writing » for ¢, = (+5) . = eT But 0u/dx + ov/oy may be regarded as implicitly a function of ¢ which changes in value as the particle of fluid followed by the total differentiation moves, and in general will have a positive or negative finite value. If we suppose the axis of z to turn so as to be always in the direction of the axis of spin, then, generally, we shall have at each instant dw/dt=—kw, where k is a finite multiplier. Integrating then for a moving element, we obtain log w=cer+¢ or : : ‘ : (38), w= eet where 7 is the time interval of integration, and ¢ is a proper mean value of —(0u/o+0v/dy) for each successive instant of time, and c’ is a constant. If now we suppose that initially » is very small, or zero, c’ must be a very large, or infinite, negative constant. Thus », if at first imperceptible, would not, in any finite time, acquire a perceptible value. We obtain again, therefore, the theorem of the per- manence of the non-vorticity of a portion of a perfect fluid. 22. Returning now to (36), write in it bf ae Or er de alae CN! if Te, and multiply both sides of (36) by dadydz, and integrate for the space contained NOTES ON HYDRODYNAMICS, CHIEFLY ON VORTEX-MOTION. 15 within a fixed closed surface. From the first form of the right-hand side of (36) we obtain, since dé | Oy , a =()). On fy oy ee 2 [[ [Saeaude= | [ Can— rea ; : . (39), where g, is the normal velocity inwards along the normal at the element dS of the enclosing surface, and », is the angular velocity about that normal. If the surface is taken large enough to include all the vortices, ¢ and , are zero at every element of the C6 dedydz=0 . (40). OE A similar result is obtained, of course, for each of the other components. The volume-integral of the time-rate of change of each component of angular velocity of spin has thus the value indicated by the right-hand side of (39), and vanishes for any surface enclosing all the vortices. If we take the second form of the right-hand side of (36) we get Nee Fawdyde= | { equtS - [ [von cos dS — [[[0E+2auk © \aedyie o, (40); where 7 is the z-direction cosine of the inward-drawn normal to ds, and ¢ is the angle between the directions of g and ¢ at dS. By (40) this gives the equation | fe, — nwg cos p)dS = [[[E+e2 af +w 2 \dedyde ; ' (42). the result surface, and we have per k yb ate Rane aot a * - Py Fivisat with #¥ Pane ie ery ‘Pah dol re : ra acti Twa (ih ip bint cxf Edidire is i vend. ve hye R - aos a aani i. »f 1s he “i he “0s "'? Gish 2 itt i t . = ; “: ASkhiy ua} is laitia lowe. he if \°9 seo) bop, ofa ls | | : ~ | ’ i : ’ hhiys yt iP rr) Crs et eS sd i en el See i 3 hg II.—The Glacial Deposits of Western Carnarvonshire. By T. J. Jehu, M.D. (Edin.), M.A. (Camb.), F.G.8., Lecturer in Geology at the University of St Andrews. (With Four Plates and One Map.) (MS. received January 12, 1909. Read March 15, 1909. Issued separately May 3, 1909.) CONTENTS. PAGE PAGE I. InTRODUCTION . f : : 2 : 17 Ill. THe Drirt or tHE INTERIOR . F 3 38 II. Tar Coast SEctTIons F i : . 19 TV. SuMMARY AND GENERAL CONCLUSIONS . 45 I. INTRODUCTION. Since the time when Agassiz and BuckLanp made known the former presence of glaciers in the mountain valleys of North Wales, much has been written concerning the glaciation of Snowdonia, but comparatively little attention has been given to that of Western Carnarvonshire. This part of the country stretches south-westwards as the broad promontory of Lleyn between Carnarvon Bay and Cardigan Bay. Western Carnar- vonshire for the most part lay outside the paths followed by the native glaciers. None of the larger valleys of Snowdonia trend in this direction, and so the marks of recent glacia- tion are not so fresh and striking in this region as they are further east. Hitherto no one has attempted to give any detailed account of the Drift deposits over the whole of Lleyn, but various references to the glaciation of the region are found scattered in the literature dealing with the geology of Wales. Among these one of the most interesting and most accurate is that in JosHua TrtmmeEr’s Practical Geology and Mineralogy (1841), where he writes of the ‘‘ Diluvial Deposits of North Wales” (p. 398). Concerning Lleyn he says: “The promontory which divides the bays of Cardigan and Carnarvon is covered with diluvial deposits of variable thickness, but frequently exceeding 100 feet. They consist of sand, clay, and gravel, changing rapidly from one to the other, through which are dispersed broken marine shells, and, amidst much local detritus, fragments derived from the north and west. (Granite is rare, but they afford fragments of Antrim chalk, often very slightly abraded... . The south-western termination of this promontory affords several instances of the accumulation of the transported matter with shells, covering masses of angular detritus of the rock immediately subjacent, which may be supposed to have accumulated slowly under the atmosphere, after the manner of a talus, or to have been collected from the weathered surface of a bare rock by the first rush of the diluvial currents.” This extract proves that Trimmer had detected the “rock rubble” or “head” which underlies the Drift deposits at the western end of the promontory. TRANS. ROY. SOC. EDIN., VOL. XLVII. PART I. (NO. 2). 3 18 DR T. J. JEHU ON Sir AnpReEw Ramsay, in his Survey Memoir, ‘The Geology of North Wales” (2nd ed., 1881), makes only occasional references to the glaciation of Lleyn. Describing the district, he says: ‘‘ Like Anglesey it partly consists of metamorphic rocks, which are supposed to be of Cambrian age, and of Lower Silurian strata, through which numerous bosses of rock protrude of a syenitic and dioritic character. Some of these are of great extent, and in Bwlch-mawr and Moel Penllech, Yr Hifl or The Rivals, Carn Bodvean, Mynydd Nevin, Carn Madryn, Mynydd Mynytho, Mynydd-y-Rhiw, and other minor heights, they generally rise high above the plain of glacial drift that shrouds so much of the country, and thus form the loftiest hills of the district. The stratified rocks, indeed, generally lie in ground so low, and, except on coast cliffs, they are mostly so obscured by sands and marine boulder clays, that it is difficult to make out the details of this stratification” (p. 206). Again he states (p. 221) that “from the heights of Yr Eifl the promontory of Lleyn, but for the chain of Puys, looks like a mere plain.” He mentions “‘an angular breccia of post-Tertiary date resting on the slaty strata” at Porth Ceiriad and underlying the Drift deposits (p. 208). Beds of angular breccia are also noted as occurring in Aberdaron Bay (p. 211). With regard to Bardsey Island he observes that “like the mainland the country has been moulded by ice during the Glacial period, but the mammillated roches moutonnées have since been roughened by weather” (p. 212). He describes the ice-sheet which overspread Anglesey as continuing “its onward course to some unknown distance, for on the ice-ground rocks above the sea- cliff of Trwyn-y-tal, a mile north of the seaward flank of Yr Eifl, well-marked striations are seen pointing towards the Straits 43° north of east” (p. 273). In Ramsay’s Physical Geology and Geography of Great Britain (6th ed., 1894, p. 246), mention is made of the occurrence of shell-bearing deposits in the district of Lleyn. It is also stated (p. 248) that “at various levels on the low ground between Carnarvon and Criccieth there are extensive deposits of sand and gravel, well stratified, and much resembling those of Moel ‘l'ryfan, but apparently without shells.” These are overlain by boulder beds. He held the view that there had been two epochs of glaciation in Wales separated by an epoch of submergence during which the land was drowned to a depth of 1400 feet or more. Mr Carvityt Lewis in The Glacial Geology of Great Britain and Ireland, 1894 (p. 355), states that ‘‘ deep till, with many large boulders, covers the land, and makes the moorland extending from Afonwen to Carnarvon.” He also noted the peat at Ynys and Pant-glas and the “hills of till” at Brynkir; the ‘“ very large beds of coarsely stratified drift, full of large boulders and knob-like in shape,” occurring near Penygroes. ‘ All this,” he concludes, “ probably belongs to the great ice-sheet, not to the Welsh glaciers.” He gives a Glacial Map of England and Wales in which the margin of the northern ice is depicted as reaching North-Western Lleyn but leaving the southern part of the pro- montory free. In common with many other geologists, Mr CarviLtt Lewis denied that the so-called “ Intermediate Sands and Gravels” indicate a submergence of 1400 feet. He regarded the high-level shell-bearing sands, such as those of Moel Tryfan, simply as THE GLACIAL DEPOSITS OF WESTERN CARNARVONSHIRE. 19 rearranged material derived from the bottom moraine of an ice-sheet which reached this elevation. Mr T. Metiarp Reanbsg, in a paper on “The Drift Beds of the Moel Tryfan Area of the North Wales Coast,” * describes the coast sections from Dinas Dinlle to Nevin and also some sections inland. He favours the view that the sands and gravels are marine, and that they indicate a submergence of this area in Glacial times to at least 1400 feet. The base of the Lleyn promontory lies just outside the mountains of Snowdonia, and has a breadth of about 5 miles. South-westwards the breadth varies from 3 to 5 miles, becoming reduced to about a mile at the extreme end. None of the great valleys of Snowdonia trend towards Lleyn. The stratified rocks of this district are chiefly of Lower Silurian (Ordovician) age, and through these protrude a number of igneous masses which form an interrupted chain of hills along the whole extent of the promontory. Hard grits and shales of Cambrian age constitute the high ground between Porth Nigel and St Tudwal’s Road, and metamorphosed rocks which are now regarded as _pre- Cambrian extend from Porth-dinlleyn to the end of the promontory and form the Isle of Bardsey. These metamorphic rocks rise to heights of 500 feet at the extreme end of Lleyn. The lower parts of the district are covered by a mantle of Drift, and if this were entirely removed considerable parts of Lleyn would probably be submerged. II. Tae Coast SEctTIONs. The following account of the superficial deposits begins with the sections at Dinas Dinlle and the coast is followed south-westward to the end of Lleyn and thence eastwards on the south side of the promontory to Criccieth. From Dinas Dinlle south-westwards as far as Gwydir Bay the coast-line is for the greater part bounded by cliffs of varying heights which are described on the Geological Survey Map (1 inch to the mile) as consisting of stratified clay, sand, and gravel. The bed-rock is not seen anywhere along this stretch of coast. To the north of Dinas the coast is flat, consisting of alluvial land which extends towards the entrance of the Menai Straits. The character of this flat area is shown by a boring recently made for water a few yards behind the hotel at Dinas. Eighteen feet of grey sand were passed through, in which the shells of Cardium edule were abundant, as well as the shells of Scrobicularia piperata, with the two valves often still in apposi- tion. This sand is evidently a recent marine deposit. Below the sand a very tough bluish-grey boulder clay was reached, and this extends to an unknown depth. Dinas Dinile.—Vhis is a dome-shaped mound surmounted by an ancient encamp- ment. One side of the mound is being cut away by the sea, so that good sections, which have been described by Mr T. Mrtiarp Reapsg,t are exposed. The cliff varies in height up to a maximum of about 100 feet. * Proc. Liverpool Geol. Soc., vol. vii. (18938), p. 36. + Ibid., p. 42. 20 DR T. J. JEHU ON The following succession can be traced in the deposits :— 4, Soil and Blown Sand. 3. Upper Stony Clay. 2. Sands and Gravels. 1. Lower greyish-blue or purple Boulder Clay. The Upper Stony Clay and the Sands and Gravels form almost the entire cliff. The Lower Boulder Clay only appears at the base of the cliff at one or two places. This Boulder Clay is very tough and homogeneous, being comparatively stoneless. On the surface it has a somewhat reddish or purple hue, but when dug into it has more of a dark greyish-blue colour. It can at times be traced underneath the pebbly foreshore. The Sands and Gravels in this mound generally show distinct bedding, and the beds are arched in a curious way and pass under the Upper Stony Till) The gravels and sands contain many flakes and fragments of slate and shale. There are some well- marked beds of bigger water-worn boulders. The sands vary from very fine yellow to coarse grey. The Upper Stony Clay is a true boulder clay, greyish in colour where unweathered, but becoming yellowish and friable on weathering. The included boulders are of varying sizes, and are scattered irregularly throughout the mass. Many of them show well-striated faces. All are subangular or more or less rounded in form and con- sist mostly of Welsh rocks, especially greenstones. The large boulders lying on the beach below have evidently been derived to a great extent from this Upper Till. In places the till tends to become a mass of boulders or coarse gravel. Shells, in a very fragmentary condition, occur rarely in the Upper Clay. The only form recognised was a Turritella, Chalk-flints are readily detected. Among other erratics found were the Ailsa Craig micro-granite, the Goat-Fell (Arran) granite, Dalbeattie granite, schistose and serpentinous rocks, possibly from Anglesey, and a reddish felspar porphyry which may have come from no great distance. The Dinas Dinlle mound is separated by a low cliff consisting entirely of the Upper Till from another mound which occurs a few hundred yards further south in which the cliff sections attain heights of 40 to 50 feet. Here the entire cliff is made up of boulder gravel showing but little trace of stratification. Opposite the centre of the mound the matrix is sandy or gravelly, but towards the two ends it becomes more clayey in nature. | A diagrammatic section of the mound of Dinas Dinlle is given (fig. 1). For about 25 miles to the south-west of the sections just described the coast is fringed by lowlands, and the sea has thrown up a ridge of shingle just above the high-water line. There have been considerable changes along this part of the coast within recent times, for half a mile out to sea are found submerged the remains of an ancient camp known as Caer-Arianrod, which figures in some of the traditional Welsh stories —the Mabinogion. Cliffs formed by Drift deposits begin to appear again at Point Maen Dulan. At first the cliff is low, but it gradually rises, reaching a height of about 80 feet THE GLACIAL DEPOSITS OF WESTERN CARNARVONSHIRE. 2! half a mile to the north of Clynnog. It is only here and there that good clean ex- posures are displayed. Often the cliff face is obscured by slipping or by overgrowth. Good exposures are seen just to the north of Clynnog, and again opposite the village. These show that the entire cliff in this region is made up of a greyish till weathering yellow, and full of boulders of all sizes. The till encloses lenticular patches of rather fine gravel. ‘The deposit is unstratified, but the boulders often become more numerous towards the top. The beach below is strewn with big boulders which have fallen from the cliff. Shell-fragments occur rarely, but those obtained were too fragmentary for identification. This Stony Clay represents the Upper Boulder Clay, and is here of unknown thickness, for the bottom is not reached in the sections. Far-travelled boulders are not uncommon, and the following were obtained from the cliff near Clynnog :—A foliated granite identical with that forming the eastern margin of the Criffel mass in the south-west of Scotland, Dalbeattie granite, the Goat-Fell granite of Arran, the Mull of Galloway granite, chalk- ———— lak. ) Ros 2 soil and Ee i: blown sand _——_, = eee a ——— BAAS — r sli ss = == : 5 AREER DER a AT sandy clay Upper Boulder Clay sands and_ gravels Bo ny ee Fetes Upper Boulder Clay Fic. 1.—Diagram of the Cliff Section at Dinas Dinlle. flints, Carboniferous Limestone showing crinoid stems, and various schistose and serpen- tinous rocks which are probably derived from Anglesey. To the south-west of Clynnog the cliffs are again much grassed over or obscured by talus. The heights vary up to 80 or 90 feet. Where best seen the section shows— 2. Upper Stony Clay—10 to 12 feet. 1. Gravels, bedded in places—60 to 70 feet. The bottom of the Gravels is not reached in the section. A little further west fine brownish sand is seen mingled with the gravelly material, which is also finer here. At Aberafon, on the left side of the stream, near its mouth, the cliff is over 100 feet high. It is mostly grassed over, but at one place it can be seen to consist of well- stratified sands and fine gravel. There are some gritty beds, but the finer sands are stoneless. In the sands occur pieces of a carbonaceous material resembling coal. Above the gravels and sands and forming the top of the cliff is a clay full of boulders and rounded stones. Mr Metiarp RuapE* states that he saw a red clay at the base of the cliff underlying the Sands and Gravels. This was not exposed when the writer visited the spot, probably owing to the slipping of material from above and to over- growth. In the gravels of this cliff a pebble of the Ailsa Craig micro-granite was found. * Proc. Liverpool Geol. Soc., vol. vii. (1893), p. 43. 22 DR T. J. JEHU ON Just west of Aberafon the cliff face is again somewhat hidden by overgrowth. But it can be seen to be formed of laminated stoneless sands and gravel, with some of the Upper Stony Clay on top. Immediately north-east of the wooden pier the cliff attains a height of about 90 feet. The upper 20 or 30 feet are made up of the Stony yellowish Clay, and the rest of the cliff of very fine stoneless sands. As traced laterally the stone- less sands in this upper part tend to pass into gravels and boulder gravel—the sands still forming the lower part of the cliff down to its base. Along the whole stretch of coast from Point Maen Dulan to the wooden pier at the north-east end of Gwydir Bay the stiff bluish-grey Lower Boulder Clay was not seen. But in Gwydir Bay, just south-west of the pier, the Lower Boulder Clay appears again at the base of the cliff. It is very tough and homogeneous, and its dark bluish-grey colour is strongly contrasted in certain lights with the light-coloured yellow deposits above. It is exposed all along this bay, and, on account of its unctuous nature has been the cause of much land-slipping, resulting in the formation of a sort of under-cliff. This under-cliff is 10 to 15 feet in height, and above it is a gentle slope much grassed over. At the east end of the bay the Lower Boulder Clay is seen to be overlaid by gravelly material and this again by the Upper Till How much of the Lower Boulder Clay lies under the grassed slope it is difficult to ascertain. The base of this clay is not reached in the cliff sections; indeed, the foreshore is underlaid by this tough sticky clay, from which the remains of sedges or reeds are often seen protruding. Shell-frag- ments occur sparingly in the Lower Boulder Clay, but they were too small for identifica- tion. Boulders of Carboniferous Limestone, beautifully glaciated, occur plentifully in the Lower Boulder Clay, chalk-flints are not uncommon, and porphyrites from the Dalbeattie area were obtained. To the west of Gwydir Bay the coast is rock-bound. The cliffs of Trwyn-y-tal are composed of sedimentary rocks of Lower Silurian (Ordovician) age. Considerable erosion has gone on since the Glacial period. There are no traces of a raised beach platform, but the almost perpendicular post-glacial cliffs are truncated above by a rounded slope very similar to that described by Messrs Wricut and Murr on the south coast of Ireland.* Along this part of the coast, as on the south of Ireland, the slope above can be seen to be continued on stacks now separated from the mainland. To the south-west of Trwyn-y-tal there is a remarkable small dry valley which opens seawards above the cliff. It evidently marks an old overflow channel. Some drift is seen here in the cliffs, but the Lower Boulder Clay appears to be absent. Passing south, the coast is bound by the igneous mass of Yr Hifl (The Rivals), and other igneous rocks of a similar nature form headlands to within a mile of Nevin. But at Porth-y-nant small bays have been eroded in the softer sedimentary rocks which reach the coast between the harder igneous masses. In these bays Drift deposits occur and form the cliffs, but the exposures are not good. At the north-eastern end of Porth-y- * Scientific Proc. Royal Dublin Soc., vol. x. (N.S.), part ii., 1904, p. 296. THE GLACIAL DEPOSITS OF WESTERN CARNARVONSHIRE. 23 nant the stiff bluish-grey Lower Boulder Clay is exposed, in which shell-fragments can be seen. It forms the lower part of the cliff for about 10 feet above the modern beach. This Boulder Clay is followed above -by sands and gravels up to a height of about 70 feet. Again, to the south-west of Careg-y-llam, opposite Pistyll Church, deposits of a similar nature occur, but the cliffs are much grassed over. As seen from a distance, these Drift deposits form a well-marked terrace with a somewhat flattened top, border- ing the higher land. The cliffs of Porth Nevin and Porth-dinlleyn are described on the Geological Survey Map (1 inch to the mile) as being composed of sand and boulder clay. Both bays have a horseshoe or crescentic shape and open out to the north. They are ex- cavated for the most part in Drift deposits overlying green quartzose and chloritic schists, but the north-east end of Nevin Bay is carved out of the Lower Silurian (Ordovician) sedimentary rocks. As seen from a distance, the cliffs seem to be bounded above by a terrace which extends inland to the base of the hills with a very even surface. Along the greater part of this north-west coast the superficial deposits have this remarkable flat-topped aspect. The smaller bay at Porth Nevin has an extent of about a mile. The cliff attains heights of over 100 feet, but the sections are much obscured by talus formed of debris which has slipped from above, and the cliff face is often grassed over. The cliff for the greater part of its length is made up entirely of sands, gravels, and laminated silty clays. At the two extremities of the bay the bed-rock is seen at the base, and here the sands and gravels rest immediately on the solid rock. Where, as opposite the middle of the bay, the sections are deeper and the bottom of the Drift deposits not seen, the typical stiff dark greyish-blue Lower Boulder Clay appears underneath the sands and gravels. This clay contains shell-fragments. Many of the included boulders, amongst which the commonest consist of Carboniferous Limestone, are remarkably well glaciated. Where the bed-rock is visible, the sands and gravels are at some places seen to be separated from it by broken-up or shattered rock material, a foot or more deep. Fragments of marine shells are plentiful in the sands, but these are generally too imperfect for identification. This shelly sand is often stoneless, but at places it includes irregular masses, pockets, and seams of coarser gravelly material. Sometimes the gravel is so coarse that it would be more accurately described as a boulder gravel, and this occasionally swells out to form considerable portions of the cliff. The sands are stratified, the gravels sometimes show bedding, but occasionally are tumultuous, showing no definite arrangement. Contortions can here and there be observed in the sandy beds. Above the sands and gravels are buff-coloured laminated silty clays free from stones. These at places form the upper part of the cliff, but elsewhere in the same bay the cliff is capped by gravel or by a brownish-yellow weathered clayey rubble. 24 DR T. J. JEHU ON The general succession of the Drift deposits in this bay is therefore as follows :— 5. Soil. 4. Yellowish-brown weathered rubbly clay often becoming pebbly or passing into a mass of boulders. 3. Stoneless laminated silty clay. 2. Sands and Gravels. 1. Lower Boulder Clay. The silty clay (3) shows fine laminations which are arranged in ribands of lighter and darker colours, giving a striped or banded appearance as seen from the shore. The bands are sometimes wavy, and examples of faulting occur as described by Mr MELLarp Reape,” and ascribed by him to contraction and loss of volume, either by actual drying or by the draining away of their water. At the north-east end of the bay the upper part of the cliff consists of Rubble Drift. The lower part is grassed over, but appears to be sandy and gravelly. Penrhyn Nevin, the promontory separating this bay from Porth-dinlleyn, has rock at the base and Drift deposits above. Ice-scratches running N.N.E. to 8.8.W. are seen on the rocky platform. This platform runs at a height of 10 to 12 feet above the modern beach, and is now much cut into, being in process of demolition by the waves. In appearance it simulates an old shore platform. It is often clear of drift for 7 to 8 yards from its seaward margin, and has a gentle slope outwards. Portions of it can be traced on both sides of the promontory as well as round its point. Where the rock-shelf passes under the cliff of Drift it has often a hummocky and glaciated aspect, in striking contrast to the rough and jagged appearance shown by the rocks on the shore below. On the Nevin side of the promontory the Sands and Gravels are, as already mentioned, sometimes separated from the platform by a foot or so of rock rubble. On the Porth-dinlleyn side clays, somewhat laminated, rest on the rock and are followed above by sand and gravel. Porth-dinlleyn lies to the south-west of Porth Nevin. ‘his bay is wider than that at Nevin, having an extent of about 2 miles. Its western horn is formed by a mass of coarse serpentinous rock. The cliff has a very uniform height of about 100 feet. Its face is much grassed over, so that good sections are not often displayed. The deposits are very similar to those met with in Porth Nevin. They consist largely of stoneless sands and gravels capped by the Upper Stony Clay. At the north-east end of the bay stoneless, stiffish laminated clays form the lower part of the cliff. Nearer the middle of the bay these are seen to pass downwards into the typical tough Lower Boulder Clay. At a point just east of the middle a ledge of rock appears from under the Drift. The surface of this has a glaciated appearance. Beyond the ledge the Lower Boulder Clay appears at the base of the cliff with big boulders imbedded in its upper part. A little further west, beds of gravel come down to the bottom of the cliff face, and are followed above by stoneless sands. The upper portion of the cliff is here hidden by overgrowth. * (feol. Mag., 1891, p. 487. THE GLACIAL DEPOSITS OF WESTERN CARNARVONSHIRE. 25 In the stoneless sands pockets of gravel occur. ‘The sands and the finer gravels show beautiful examples of oblique bedding. These false-bedded sands and gravels can be traced for some distance in the lower part of the cliff. The sands are very fine and marine- like, and apparently form the greater part of the cliff’ Just before reaching the road that comes down to the beach the Lower Boulder Clay is again seen at the base. West of the road the cliff is still obscured by vegetation, but appears to be made up mostly of sand. At this part of the bay there has been much sliding over the unctuous Lower Boulder Clay, giving rise to an under-cliff. The under-cliff is made up of the lower stiff clay, and above it the cliff slopes away and is grassed over. The upper part consists of yellowish sandy clay full of small boulders. The Lower Boulder Clay is here very dark in colour, and presents a striking contrast to all the deposits above. Promontory of Porth-dinlleyn.—A rocky shelf can be seen on the eastern side appearing from under the Drift. It has a glaciated aspect where the deposits have been recently cleared away. The shelf or platform is well seen just before reaching the lifeboat house. At its inner (landward) end it is 15 feet above the modern beach and slopes gently seawards. The sea is gradually eating into and destroying it. Just beyond the lifeboat house the serpentinous rock shows distinct marks of glaciation— the grooves and scratches running approximately N.E. and S.W. Near the extremity of the promontory, north-east of the flag-staff, at a height of 25 feet above Ordnance datum, the rock is beautifully glaciated, showing deep and broad furrows and ice- scratchings running about N.H. and 8.W. ‘These are the finest glacial markings noted by the writer in Lleyn. The following far-travelled boulders were found in the Gravel and Sand series at Porth Nevin and Porth-dinlleyn :— Several granites from the Dalbeattie area; a granite with reddish idiomorphic felspars much resembling the Shap granite; the Ailsa Craig micro-granite. The shell- fragments in the sands were, as a rule, too broken and minute to be identified, but the following forms were obtained :—-Turritella communs (Lam.), Nucula nucleus (L.), Pecten (Chlamys) islandicus ? (Mill), Astarte sulcata (Da Costa), Cyprina islandica (L.), Cardium edule (L.). On the west side of the promontory lies the little bay which bears the name of Porth Wen. Here the Drift is again exposed in sections which are 40 to 50 feet high. The stiff bluish-grey Lower Boulder Clay forms the base of the cliff, and is succeeded above by fine yellow marine-like sands and some gravel. From this bay to Porth-ysgadan the coast is rock-bound. Some Drift caps the rocky cliffs and is usually overgrown with grass. Porth-ysgadan.—¥airly good sections of the Drift deposits are seen at this place. Opposite the middle of the bay the typical Lower Boulder Clay with included shell- fragments forms the lower part of the cliff, and the bottom of this deposit is not reached in the section. Above, the face of the cliff slopes away and is grassed over, but there are indications that sand overlies the Boulder Clay. The upper part consists of TRANS. ROY. SOC. EDIN., VOL. XLVII. PART I. (NO. 2). 4 26 DR T. J. JEHU ON yellowish weathered clay becoming sandy in places. At the two ends of the bay the cliffs are rocky, though sand is seen resting above on the rock surface at some places, while at others the rock is covered by fine rubbly material with yellowish weathered clay above. Samples of granites from the Dalbeattie area were picked up on the beach below. These had undoubtedly come from the Drift in the cliff above. Amongst the shells included in the Lower Boulder Clay the following were recog- nised :—Purpura lapillus (L.), Astarte sulcata (Da Costa), Astarte (Nicania) compressa (Mort.), Cyprina islandica ? (L.), Corbula gibba (Olivi). All along this part of the coast the rock surfaces when visible have a mammillated and smoothed appearance where they pass under the Drift deposits. Immediately to the south-west of Porth-ysgadan lies a small bay opening westwards to the sea, which appears to be nameless. The two arms of the bay are rocky, with grassy slopes above. Midway between the arms the Drift comes down to the beach in such a way as to suggest that we have here the buried channel of an old stream, or what was once an inlet of the sea. The cliff of Drift reaches a height of about 50 feet. It seems to consist entirely of the Lower Boulder Clay in which shell-fragments are again common. Towards the surface the clay is weathered and yellowish in colour, but there is no reason for separating this from the Lower Boulder Clay. Where, asin this bay, the Sands and Gravels are wanting, it is sometimes dithcult to know whether the yellow clay which usually forms the upper part of the cliff is the weathered portion of the Lower Clay or the representative of the Upper Boulder Clay. Where typically developed, however, the Upper Boulder Clay is less homogeneous, somewhat more sandy, and contains more boulders than the Lower Boulder Clay. In this small bay, at the two sides, the Lower Boulder Clay appears to be separated from the bed-rock by a little rock rubble. Porth-y-gwylan lies half a mile further to the south-west and opens westwards. It is separated on the north side from the open sea by a ridge or projection of bare rock running more or less parallel with the coast-line. The other or mainland side of the bay is also rocky in character. Between these rocky sides a mass of Drift forms the entire cliff and rests immediately on the modern beach, and it is out of this material that the bay is still being eroded. The Drift consists of tough bluish-grey Boulder Clay at the base, with included shell-fragments, followed above by a boulder gravel with a somewhat clayey matrix, and this in turn is succeeded by a yellowish clay. So we have here :— 3. Yellowish weathered Upper Boulder Clay. 2. Boulder Gravel. 1. Lower Boulder Clay. These deposits appear to have filled up what was either the channel of a stream or a narrow inlet of the sea. At the present day a small stream is seen on the north-east side of this Drift-filled depression which is cutting out for itself a new channel in the solid rock and coming down to the beach in a waterfall to escape to sea by a Trans. Roy, Soe. Edin. Vou. XLVII. Dr IT. J. Jesu on “ The Glacial Deposits of Western Carnarvonshire.” (From a photograph.) Section of Drift resting on a rock platform at Porth Colman, Fie. 2. To face p. 26. THE GLACIAL DEPOSITS OF WESTERN CARNARVONSHIRE. 27 recently eroded channel at the base of the rocky ridge which bounds the bay on its sea- ward side. Following the coast-line south-westwards we come next to Porth Uchain, a very small inlet with rocky cliffs at the sides attaining a height of about 40 feet and capped by yellowish weathered clay. Opposite the middle of the bay the Lower Boulder Clay appears and forms the lower part of the cliff; above the clay there is a slope which is grassed over. From this point to the next bay—Porth Colman—the cliffs are rocky and have a steep slope to the sea. The bay which follows is broad, and the northern portion of it is sometimes known as Porth Penllech. Here the cliff attains heights of 30 to 40 feet. In places the lower part of the cliff is rocky, but at one spot rather stiff dark grey boulder clay with many included stones is seen. This becomes yellowish and friable where weathered. It probably represents the Upper Boulder Clay, for a little further on it is seen to overlie gravel and boulder gravel. The remaining portion of the bay is usually called Porth Colman. At its southern end the base of the cliff is formed by rock an situ for heights varying up to 18 or 20 feet. Along a part of this shore a rock-shelf or platform is seen, reminding one again of the raised shore platform described by Messrs Wricut and Murr as occurring on the south coast of Ireland (fig. 2). The platform lies 10 or 12 feet above the modern beach and slopes gently upwards to about 15 feet, where it disappears under the Drift deposits. It has an exposed breadth of 6 yards or so in places. The surface of the platform is covered with lichens or partly grassed over. At its inner end it is covered by bluish-grey Boulder Clay containing shell-fragments. This is succeeded above by the Upper Yellowish Stony Clay. As we pass towards the middle of the bay the plat- form disappears and the cliffs show steep rocky faces with a capping of Drift forming a gentle slope landwards. Near the middle of the bay is a remarkable little inlet with steep cafion-like sides from which a small stream issues. A few yards up this inlet the stream is seen to descend by a waterfall over rock to the level of the beach. The following erratics were obtained from the Boulder Clay at Porth Colman :— Several granite boulders from the Dalbeattie area; a diorite very similar to that of Colvend, Dalbeattie ; boulders of hornblende porphyrites, probably from the Dalbeattie area, and chalk-flints. The following species were identified from amongst the shell-remains found in the Boulder Clay at Porth Colman:—Turritella communis (Lam.), Astarte sulcata (Da Costa), Cyprina islandica (L.), Venus (Ventricola) Casina (L.), Cardiwm echinatum (L.). Around Penrhyn Colman the coast is rocky, and at Porth-llefesig the rocky cliffs reach heights of 50 to 60 feet. Above the rock lies a grassy slope of Drift. At the northern end of the bay boulder gravel lies immediately above the rocky cliff, and this is capped by stiff clay rather full of boulders and weathering yellow near the surface. This clay contains some shell-fragments, but the only one which could be identified was Astarte sulcata, 28 DR T. J. JEHU ON Porth Whaiting.—For the greater part of its extent the cliffs are rocky, but about the middle of the bay rock rubble is seen forming the base of the cliff where rock in situ is wanting. Above the rubble is a slope of grass which covers what appears to be a clay weathering yellow. At the south-western horn of this bay ice-scratches with a N.E. to 8. W. direction were noted on the rock surface just where it disappears under the Drift. Between Porth Whiting and Porth Ferin some well-glaciated rock surfaces are displayed, the vertical face of the rock being sometimes grooved and polished in a direction parallel to the shore-line. Above the rock, fine yellow sand is at places exposed from under the grassy slope, and this is overlaid by greyish Boulder Clay weathering yellow. Further south this stiff Boulder Clay rests immediately on the rock surface, and is even sometimes overlaid by yellow marine-like sand. So it may repre- sent the Lower Boulder Clay. At Porth Ferin an artificial cutting has been made through the rock at the top of the cliff, Stiff dark blue boulder clay, which is apparently the Lower Boulder Clay, is seen overlying the rock. It is comparatively stoneless, though some well-glaciated boulders were found in it. No sections of the Drift are again seen until we come to Porth Oer. At the north- east end of this bay is solid rock, and on this, at a height of 15 to 18 feet, are beds of sand and gravel more or less conereted and much resembling raised-beach material. Just south of this a roadway runs down to the beach, and on the southern side of it the rocky side of the bay slopes down to the beach and is followed by rock rubble or ‘“‘Head” which in turn is seen to pass under typical dark greyish-blue Boulder Clay. This is in all probability the Lower Boulder Clay, and it forms the entire cliff opposite the middle of the bay to a height approaching 100 feet. Included in this clay are irregular masses of marine-like sand and here and there a little gravel. Shell-fragments occur in the Boulder Clay, and the following forms were obtained and identified :— Pectunculus glycimeris (L.), Astarte sulcata (Da Costa), Cyprina islandica (L.), Turri- tella communis (L.). At the south-west horn of the bay a bench of rock is again seen 10 to 15 feet above the modern beach. It is much like that seen at Porth Colman, and suggests the possi- bility of the presence of an old shore platform here. Stiff bluish-grey clay (Lower Boulder Clay) rests on it, but at places the clay is separated from the rock by coarse rubbly material about a foot deep. No exposures of Drift deposits are seen from Porth Oer to the end of the promontory of Lleyn. The coast is rock-bound with a steep slope, the greater part of which is grassed over. Lying to the south-west of the mainland and at a distance of 24 miles from it is the Isle of Bardsey. It is composed entirely of metamorphosed rocks, probably of pre- Cambrian age. The island has an extreme length of about 1} miles, and its greatest breadth is three-quarters ofa mile. The highest portion of the isle lies in the north-east, facing the mainland. ‘There, from an elevation of about 500 feet, the land slopes away THE GLACIAL DEPOSITS OF WESTERN CARNARVONSHIRE. 29 steeply to Bardsey Sound. The south-west side of the island consists of lowland rising gently towards the higher ground. ‘This lowland is covered by a skin of Drift. The whole island has the appearance of having been moulded by ice during the Glacial period, but, as Ramsay observes,* ‘‘the mammillated roches moutonnées have since been roughened by weather.” The island as a whole may be regarded as an example of the phenomenon known as “crag and tail,” the crag facing the north-east, from which direc- tion the ice-sheet came. At Porth Solfach, on the island, the low cliff is composed of yellowish-brown clay showing a depth of 6 to 7 feet. Though weathered near the surface, the clay gets stiffer below. Bits of shells in the clay were too fragmentary for identification. Amongst the included boulders were found chalk-tints, the Ailsa Craig micro-granite, Dalbeattie granite, a picrite, a lava closely resembling that of Borrowdale in Cumberland, Millstone Grit, Carboniferous Limestone showing crinoid stems. Some of the boulders, especially those of the limestone, were well glaciated. Again at Henllyn beach, in Bardsey, pebbles were picked up of Carboniferous Lime- stone, chalk-flints, and of a reddish-streaked rhyolite. These were probably derived from the Drift of the low cliff adjoining. Returning to the mainland, the coast is rocky as far as Aberdaron, but the cliffs are interrupted by an inlet at Porth Mendwy, where a stream comes down to the sea. Some river-like gravels and sands are seen at the sides of this inlet. At Porth Pistyll, 14 miles to the south-west of Aberdaron, the cliff is steep and rocky, but above, at a height of about 100 feet, bluish-grey boulder clay of ex- cessive toughness was noted—the deposit attaiming a thickness of 4 to 5 feet. The exposure is a very small one. This stiff clay is capped by weathered yellowish rubbly clay. Aberdaron Bay.—Situated on the southern side of the extremity of Lleyn, this bay is excavated in beds of Bala age let down by faults into the hard schistose rocks of pre- Cambrian age which bound them on either side. ‘The sides of the bay are rock-bound, but the cliff between the two arms consists entirely of Drift deposits and is described on the Geological Survey Map (1 inch to the mile) as composed of sand, clay, and gravel. The bay is over a mile broad, but the cliff is interrupted opposite the village where a stream debouches. The height reached by the cliff varies considerably up to elevations of over 100 feet. At the south-western end of the bay the rocky cliff descends with a steep slope to the beach and is followed by rough rock rubble or “ Head.” This “Head” is overlain by more rubbly material mingled with Boulder Clay, the matrix, however, becoming sandy or gritty in places. To the west of the wooden pier the “ Head” is well defined for 8 or 10 feet from the base of the cliff. It dips eastwards at a high angle and disappears under the beach level. The material above this well-defined “Head” is still rubbly, with big angular boulders of the neighbouring rock, but it becomes more and more clayey as it is traced up the cliff face, and the included boulders * Survey Memoir, “The Geology of North Wales,” 2nd ed., 1881, p. 212. 30 DR T. J. JEHU ON tend to become smaller and less angular. The material forming the upper part of the cliff at this place is very similar to the Upper Boulder Clay which is seen at other parts of the coast. Between the pier and the village the cliff is over 100 feet in height and shows steep faces. It appears to be made up almost entirely of the Lower Boulder Clay, which shows the usual characteristics and in which shell-fragments are common. ‘The face of the cliff is furrowed by running water, and at the top the weathering gives rise to pinnacles. ‘Traces of lamination are seen here and there, and masses of fine yellowish marine-like sands occur as inclusions in the otherwise stiff homogeneous clay. One of these irregular masses of sand reminds one of the great inclusion of chalk seen in the cliff sections of the Boulder Clay in the neighbourhood of Cromer. Gravelly patches also occur but are less common. To the north-east of the village the cliffs show deposits of a more varied character, but a careful examination has led the writer to conclude that the whole belongs to the horizon of the Lower Boulder Clay. The greater part of the cliff, which attains heights of about 100 feet, consists of the typical Lower Boulder Clay. This invariably forms the higher portion of the cliff, and weathers into pinnacles with steep sides. Lower down and towards the bottom of the cliff the deposit becomes more of a boulder gravel or gravel and sand. Midway between the village and the eastern end of the bay the section is as follows :— 5. Stiff greyish-blue Boulder Clay, comparatively stoneless, and containing shell- fragments—40 to 50 feet. . Boulder Gravel—10 feet. . Dark band of stiff clay —2 feet. . A talus slope probably covering sand and fine gravel—10 feet. . Stiff grey Boulder Clay—2 feet. The bottom of this is not reached in the section. Fe wow From a little distance the boulder gravels and sands can be traced as beds running across the face of the cliff. The gravel beds appear to form gentle undulations as shown in the diagram (fig. 8). Further east slipping has taken place, due to the presence of sand in the clay, and there is a tendency to form an under-cliff. At the extreme east end there is a fine example of ‘“‘ Head,” lying between the Boulder Clay and the solid rock. ‘The ‘ Head” consists of angular and flaky fragments showing rude bedding, and these fragments are derived from the neighbouring rocks. The section at this place is shown diagrammatically in the following figure (fig. 4) :— A cave has been excavated in the rubbly material by the waves. ‘The Boulder Clay overlying the “ Head ” contains pockets of boulder gravel. Imbedded in the beach at this end of the bay just under the rock cliff is a big boulder measuring 66 inches by 24 inches. It is a granite with large idiomorphic reddish felspars and closely resembles Shap granite. The felspar is orthoclase; one THE GLACIAL DEPOSITS OF WESTERN CARNARVONSHIRE, 31. of the crystals measured 24 inches in length. Several pebbles of the Ailsa Craig rock were also picked up on the beach. a—a. Boulder gravel bed. b, Laminated clay. Length of section about 450 feet. ce, Tough boulder clay weathering into pinnacles. Fia. 3.—Diagrammatic Section of the Cliff Hast of Aberdaron. Among the included stones picked out of the Boulder Clay were chalk-flints, the Ailsa Craig micro-granite, granite from the Dalbeattie area, the Goat-Fell granite of Arran, and the Loch Dee granite of the south-west of Scotland. In addition to these ae —— nt —— 4 Be rl SS Beit — yore ¥% ib 60 feet. Boulder clay ae a pebbly in places, 2 —— —— ‘ FEES bi 40 feet. Boulder clay. 5 aS / san ef IE f Fic. 4.—Section (diagrammatic) East End of Aberdaron Bay. there were boulders of Carboniferous Limestone, probably from Anglesey, of a reddish- streaked rhyolite, of jasper, and serpentine. Amongst the shell-fragments obtained the following have been recognised :— GASTEROPODA. | LAMELLIBRANCHIA, Turritella communis, Lam. Pectunculus glycimeris, L. Buccinum undatum, L. | Mytilus edulis, L. Trophon (Boreotrophon) clathratus, L. Astarte sulcata, Da Costa. Bela turricula. Astarte (Nicania) compressa, Mont. Cyprina islandica, L. Tellina (Macoma) balthica, L. | Cardium echinatum, L. | Cardium edule, L. | Mya truncata, L. A portion of a Belemnite was also found in the Boulder Clay. Porth Cadlan ( Ysgo).—The Lower Boulder Clay is not exposed in this bay. About 32 DR T. J. JEHU ON midway between its two sides a slope of Drift is seen reaching heights approaching 100 feet. It consists of yellowish weathered clay with large boulders, and contains some shell-fragments. Pebbly and sandy layers are seen here and there. At the sides of the bay this Drift is separated from the solid rock by angular rubble or “ Head.” Some of the big boulders in the cliff are well glaciated, and the beach below is strewn with blocks derived from above. Porth Nigel or Hell’s Mouth.—Situated on the southern side of the Lleyn promon- tory, this bay opens out to the south-west. Its south-eastern side is bounded by the high ground of Mynydd Cilan, which is made up of highly resisting hard gritty sand- stones and shales of Cambrian age; the north-western side partly by indurated shales of Ordovician age, but chiefly by the greenstones which extend from Mynydd-y-Rhiw to Penarfynydd. Between these sides the head of the bay is formed entirely of glacial deposits and has an extent of about 4 miles, giving one of the finest sections of the Drift to be seen in North Wales. These deposits stretch inland, covering the whole country from Llanengan to Llangian and on north-westwards by Llandegwning to Bottwnog, thence south-westwards as far as Mynydd-y-Rhiw. Erosion is still going on to a marked extent at the head of the bay. On the Geological Survey Map (1 inch to the mile) it is stated that the cliff consists of “stratified clays, sands, and gravel from 10 to 40 feet high.” This description requires considerable modification. At places the cliff attains much greater elevations. The general succession is as follows :— . Soil and Blown Sand. . Upper yellowish weathered Boulder Clay, becoming sandy and pebbly in places. . Intermediate Sands and Gravels. . Lower Boulder Clay—massive, tough, and homogeneous. (This Lower Boulder Clay descends below the level of the modern beach to an unknown depth.) Ke bo wo The cliff is highest towards the north-west end of the bay, where it attains elevations approaching 100 feet, and shows almost vertical faces which present a striking appearance. Here it consists almost entirely of the Lower Boulder Clay, which, indeed, is by far the most prominent member of the various deposits exposed in this bay. ‘The Intermediate Sands and Gravels are not seen at the western end, but are well displayed when we pass towards the eastern end. ‘Taking the deposits from below upwards, we have at the base— The Lower Boulder Clay.—This is a stiff, homogeneous clay, grey in colour on the face of the cliff but becoming darker below the surface. It presents a massive appearance, giving almost vertical faces which are traversed by fissures resembling joints running from above downwards. These cracks are widened by the action of rain and frost, and large blocks are thus wedged off and fall in masses on to the beach below. The base of the cliff is reached by the waves at high spring tides, and the clay is scooped out, leaving a smooth hollow over which the cliff above hangs. This overhanging of the cliff shows the tough nature of the clay. Here and there traces of THE GLACIAL DEPOSITS OF WESTERN CARNARVONSHIRE. 33 a kind of bedding or lamination can be seen which may possibly be due to a shearing movement, for the clay has all the appearance of having been subjected to great pressure. Boulders are fairly common but not abundant. They are distributed pell- mell throughout the clay, not in clusters or patches but singly. The majority are about the size of one’s fist, and are subangular and beautifully glaciated. Far-travelled rocks are common, and include chalk-flints, various south of Scotland granites, especially from the Dalbeattie area, the Ailsa Craig micro-granite, a granite with reddish felspars, Carboniferous Limestone, Millstone Grit, ete. Pebbles of the Ailsa Craig rock and of the other Scottish granites were also picked up on the beach and had evidently been derived from the cliff. Bits of wood were also found included in the clay. Marine shells occur scattered irregularly in the clay, generally in a very broken condition, but some complete shells were found. The forms identified are given below. The fore- shore is for the most part covered by pebbles, but at some places it consists of buttery dark blue clay evidently due to the running of the Lower Boulder Clay. The Intermediate Sands and Gravels.—These are somewhat inconstant in their occurrence, but towards the eastern end of the bay they form the greater part of the cliff. Usually they are seen to overlie the Lower Boulder Clay, but at places they rest immediately on the modern beach, where the Lower Boulder Clay is not exposed. Where the sands follow the clay in the cliff there is often much slipping. Sometimes this series consists entirely of fine siliceous sand, sometimes of fine pebbly material, and sometimes of coarse gravels. The sands and gravels often, but not at all places, show stratification. These deposits vary much in thickness and are apt to die out when traced laterally. At Porth Nigel no contortions were observed in the sandy layers. Here and there the sands become largely ferruginous. No shell-fragments were noted. The pebbles in the gravels are of rocks very similar in their origin to those found in the Lower Boulder Clay. The Upper Boulder Clay is not very conspicuous in these sections. Where it occurs it sometimes overlies the Intermediate Sands and Gravels, and sometimes rests immediately on the Lower Boulder Clay. It is yellowish in colour, and much weathered, and tends in places to become sandy and pebbly, or to pass into a rubble Drift. Ice-scratched stones can be picked out, but are not so frequently met with as in the Lower Boulder Clay. The included boulders contain erratics, but those from a great distance are not so common as in the Lower Boulder Clay. Fragments of marine shells are rare, and those obtained were very fragmentary. At the west end of the bay a little angular local detritus or ‘‘ Head” lies on the solid rock where it slopes down to the beach. There is much slipping and overgrowth here but the Lower Boulder Clay, which forms the base of the cliff of Drift, appears to overlie this “ Head.” Towards this end of the bay the cliff attains heights approaching 100 feet, and consists from top to bottom of the massive Lower Boulder Clay. Further east a somewhat sandy band runs along the face of the cliff 15 feet from the bottom. This band has a thickness of 2 to 3 feet, and when traced laterally it has more of the nature TRANS, ROY, SOC, EDIN., VOL. XLVII, PART J, (NO. 2). 5 34 DR T. J. JEHU ON of a stiff bluish clay which is laminated. ‘This can be traced continuously for a considerable distance, but eventually it comes down to the level of the beach and dis- appears. As we pass nearer the middle of the bay laminations appear near the top of the cliff. The cliff face in this region is not so steep, and considerable slipping has taken place so as to form a sort of under-cliff with a rough slope above. The top of the cliff is here formed of the Upper Boulder Clay, which is yellowish in colour and some- what sandy, and has a thickness of 12 to 20 feet. Where landslips have occurred they have given rise to a lamination or stratification in all the deposits, the beds dipping inland at a high angle. At one place about 14 miles from the west end of the bay, a bed of peat 18 inches thick was observed near the base dipping inland at an angle of about 35°. It could be traced for a horizontal distance of 20 yards, and was resting on boulder clay and overlain by sandy material and this again by boulder clay. The peat contained reeds or sedge and was full of land-shells, such as Helix. It had evidently slipped from the surface and been in turn covered by further landslips. A little to the east of the middle of the bay there is a good exposure of the Upper Boulder Clay. The cliff is here 80 to 90 feet high, and the upper 25 feet consist of rubbly clay immediately overlying the Lower Boulder Clay. This dies out when traced laterally. Further east fine yellow sand is seen near the top of the cliff. The cliff becomes lower as we travel towards Mynydd Cilan, and the Intermediate Sands and Gravels become more prominent. Where the succession is most complete we have a section as follows :— 5. Soil and Blown Sand. 4. Upper Boulder Clay, weathered and yellowish in colour, stiff in places, but often becoming sandy and pebbly or even rubbly. . Fine yellow siliceous sand becoming somewhat clayey in its lower part. co 2. Laminated greyish-blue stoneless clay becoming somewhat sandy in places, and distinctly marked off from 1. 1. The massive Lower Boulder Clay which forms the base of the cliff and the bottom of which is not reached. Near the extreme east end of the bay the gravels of the Intermediate series come down to the base of the cliff, but the Lower Boulder Clay is exposed on the foreshore below. At a short distance from this end peat appears in the upper part of the low cliff. This peat can only be traced for a horizontal distance of about 100 feet. The section here is as follows :— . Blown sand—5 feet. . Peat—3# feet. . Stiff blue peaty clay Oo —& Oo 2 feet. 2. Greyish-blue somewhat sandy clay, pebbly in places—1 foot. 1. Stiff Boulder Clay with a tendency to lamination and comparatively stoneless. (This rests on the beach and its bottom is not reached.) THE GLACIAL DEPOSITS OF WESTERN CARNARVONSHIRE. 35 Hazel-nuts were found in the peat, and the bark of birch trees. A little rubble rests on the rocky slope at the end of the bay, but its relation to the Lower Boulder Clay is obscured by slipping. Amongst the shells and fragments of shells gathered from the Lower Boulder Clay at Porth Nigel the following have been identified :— (ZASTEROPODA. | LAMELLIBRANCHIA. Turritella communis, Lam. | Pectunculus glycimeris, L. Buccinum undatum, L. Mytilus edulis, L. Tritonofusus gracilis, Da Costa. | Astarte sulcata, Da Costa. Murex (Ocinebra) erinaceus, L. | Astarte (Nicania) compressa, Mont. Purpura lapillus, L. | Astarte (T'ridonta) arctica, Gray. Trophon (Boreotrophon) clathratus, L. Cyprina islandica, L. Trophon (Boreotrophon) scalariformis, Gould. Tellina (Macoma) balthica, L. Mactra (Spisula) solida, L. Venus (Ventricola) casina, L. Cardium edule, L. Mya truncata, L. Corbula gibba® Olivi. Nuculana, sp. A cirriped was also found in the clay.* Complete specimens of Murex erinaceus and Turritella communis were obtained, and complete single values of Astarte sulcata and A. compressa. Porth Cevriad is situated at the end of the promontory which separates Porth Nigel from St Tudwal’s Road. This bay opens to the south, and is carved out of the hard gritty sandstones and shales of Cambrian age between Penrhyn-du and Mynydd Cilan. It is nearly a mile broad. The Lower Boulder Clay is not exposed in this bay. The cliff attains heights of 50 to 60 feet, and slopes away inland at the top. It consists of yellowish weathered Drift with boulders which are subangular in shape, and many of these show ice-scratches, Beds of pebbly material and sand are seen lying in or below the above, and even in places extending to near the top of the cliff. Toward the top, however, the Drift is usually rubbly in character. At the two sides where the rocky cliffs come in. the Drift is separated from the bed-rock by a mass of rubble or “ Head.” Large boulders derived from the cliff lie on the beach, and these are chiefly of local Welsh rocks, but specimens of the reddish-streaked rhyolite are fairly common. St Tudwal's Road.—The only place where the Drift is exposed is at the south- western end, where it consists of a rubbly clay, yellowish and weathered. This probably belongs to the Upper Boulder Clay horizon. The whole of this bay, with the exception of the rocky promontory and the estuary at Abersoch, is bordered by sand dunes. At the back of the sand dunes marshy land occurs at some places, as at Cors Llyferin, and this is probably underlain by Boulder Clay. The promontory on which a part of * For the determination of these and other shells mentioned in this paper the writer is indebted to Mr Hrenry Woops, M.A., Lecturer in Palzontology at the University of Cambridge. 36 DR T. J. JEHU ON Abersoch village stands is grassed over, but marine-like sands appear wherever the ground is dug into, and apparently these sands mantle the land around the village. Drift is not again exposed along the coast until we pass east of Mynydd Tir-y-ewmmwd. Llanbedrog.—Between Mynydd Tir-y-ewmmwd and Careg-y-defaid, both of which are rocky and formed of felspar porphyry, the cliff consists of Drift deposits, and good sections are exposed for a distance of over a mile. Stiff dark brownish Boulder Clay is seen on the foreshore at the west end of the bay, and at one place this clay forms the lower part of the cliff. This is the Lower Boulder Clay, and it underlies the Sand and Gravel series which forms the cliff all along the bay. The cliff has a very uniform height of a little over 30 feet. The sands and gravels are stratified, and occasionally show oblique bedding. The sands are orange-coloured and somewhat ferruginous ; they yielded no shell-fragments. So in this bay we have represented 2. The Intermediate Sands and Gravels. 1. The Lower Boulder Clay. On the beach, boulders of Carboniferous Limestone were seen and of granites from the Dalbeattie area; very common also were boulders and pebbles of the reddish- streaked rhyolite which is so often found scattered about Lleyn. From Careg-y-defaid to Pwllheli the coast is bounded by sand dunes. An old coast-line can be traced some distance inland in the shape of a cliff which is now grassed over. This old cliff is separated from the dunes by flat land some of which is marshy. Sand dunes again border the coast-line east of Pwllheli, as far as the felspar porphyry promontory of Pen-y-chain. The greenstone mass of Careg-y-rhimbill opposite Pwllheli is being rapidly de- molished by quarrying operations, but its summit presents a moutonnéed aspect. At Pwllheli, in connection with the Harbour Works, a boring was made in the estuary, and the following deposits were traversed :— 3. Mud—6 inches. 2. Ballast and sand—4 feet. 1. Stiff blue clay—8 feet. (The bottom of this clay was not reached.) Hast of Pen-y-chain, cliffs composed of Drift deposits are again met with and are being eroded by the sea. The best sections are displayed between Afonwen and Afon Dwyfor. Afonwen to Afon Dwyfor.—On the Geological Survey Map (1 inch to the mile) these cliffs are marked as consisting of “ stratified sands, clay, and gravel.” The main deposits exposed along this stretch of coast are :— 2. Upper Boulder Clay and Rubble Drift. 1. Sands and Gravels. The Lower Boulder Clay of the Lleyn promontory is not seen here. The sands are often well bedded, the gravels generally coarsely bedded. At places the sands show ———— a Trans. Roy. Soc, Edin. Vou. XLVII. Dr T. J. Jenu on “ The Glacial Deposits of Western Carnarvonshire.” Fic. 5.—Afonwen Section, near west end. Coarse gravels capped by the Upper Boulder Clay. Fic. 6.—Afonwen Section, showing Ferruginous Sands bent over and followed above by Boulder Gravel and the Upper Boulder Clay. To face p. 36. THE GLACIAL DEPOSITS OF WESTERN CARNARVONSHIRE, 37 beautiful examples of oblique bedding. The deposits in the western half of the section are very varied, and change frequently in nature when traced laterally. Near Afon- wen a bed of peat is seen under the soil and blown sand, and where the cliff descends the peat comes down on to the beach. The well-bedded sands -generally underlie the coarse gravels. Inclusions or pockets of stiff yellow clay occur in the gravels. At some places the pebbles or rounded boulders are heaped in a tumultuous fashion and show no traces of bedding (fig. 5). A curious accumulation of well-bedded ferruginous sand occurs at one place in the cliff, the beds being arched up (fig. 6). This may repre- sent an intra-glacial channel which became filled up with sand arranged in beds. When i eee Lg MAL 1 fy laf ug, Olona tie SOM SEY SEES SS aE MOLD Cy Sy, AG °° Upper Boulder Clay feet. = a Coarse gravels with some sand—18 feet. ~~ Fine-bedded sand. Sand, more clayey react at the bottom—15 _.J :—— + feet. Bedded sand. Beach. 10 feet=1 inch. Fie. 8.—Diagram of the Afonwen Section near the East End. the walls of ice disappeared the beds would tend to bend over at the sides so as to give an archlike arrangement. ‘These ferruginous beds are followed by boulder gravel and over this lies stony till full of boulders which are glaciated. About midway along the section peat again appears. Two beds can be seen separated by a thin band of very stiff bluish-grey clay with rootlets. The lower bed of peat is underlain by tough bluish clay of a similar nature. This peat was examined by Mr F. L. Lewis of Liverpool University. He states that it yielded :—Viola palustris, Carex sp. (fragments), Menyanthes trifoliata seeds (fairly numerous), frag- ments of Sphagnum leaves, Potamogeton fruits in fragments (most probably those of P. natans), also scraps of birch bark and birch wood. Further east these beds of peat seem to unite owing to the dying out of the intermediate band of stiff 38 DRT. JT. SER UCON clay (fig. 7). Where the peat appears the cliff is low and the sands and gravels disappear. Further east the cliff rises again to heights of about 50 feet, and consists from top to bottom of gravels and bedded sands, the sands forming the lower part. As we approach the Afon Dwyfor end the Upper Boulder Clay, which is here very stony in character, is seen forming the upper part of the cliff and overlying the coarse gravels, which in turn rest on the sands (fig. 8). No shell-fragments were seen in any of these deposits, and the pebbles and boulders are all apparently of Welsh origin, no foreign erratic having been noted. Sections are again exposed on both sides of Criccieth, where the cliffs consist of the Upper Boulder Clay. A fine section is shown east of the promenade at Criccieth (fig. 9). It is a typical boulder clay containing big boulders which are often beautifully striated and polished. The shore is strewn with blocks and boulders derived from the cliff. No shell-fragments were noted and no foreign erratics. The boulders consist chiefly of greenstones derived from the Snowdonian area. Ill. Tue Drirt or tHE INTERIOR. Away from the coast good sections are not often displayed. The greater part of the interior is covered by a mantle of Drift which often rounds off the lower hills, but the higher mountains rise above all the glacial deposits. Here and there, even on the lower ground, bosses of rock protrude through the Drift, and these often show rounded outlines and a more or less glaciated aspect, but the smoothness has generally been much destroyed by long exposure to the weathering agents. The region between Carnarvon and Afonwen is for most part bleak and dreary. Not only is there a deep covering of the superficial deposits over this area, but the Drift often forms mounds, ridges, and elongated hills. Many large boulders are still seen strewn over the surface, although the land has been to a great extent cleared of the loose stones which have been used in building cottages, farmhouses, and the walls and dykes which now form a network over the whole country-side. In the hollows and lower parts peat occurs, as, for instance, at Pant-glas, where it is being cut to a depth of 4 feet, and again north of Ynys. The undulating plain of Drift rises gradually from the coast-line at Dinas and reaches a level of about 400 feet at Penygroes. Between Penygroes and Clynnog Fawr hes Y Foel—a hill rismg 650 feet above sea-level. At the summit of this hill there is an outcrop of slaty rock, but with this exception the entire hill is rounded off with Drift. Sections of the superficial deposits can be seen on the banks of streams, in railway cuttings, and in gravel-pits on the surface or sides of the moraine-like hills. Borings made for water often throw light on the deeper deposits. The mounds and hills of Drift which rise above the general surface of the plain are made up of roughly bedded coarse grey sand, gravels, and boulder gravel in which some large angular and subangular boulders are frequently seen. The materials are for most part of local Trans. Roy. Soc. Edin Vor. XLVII Dr T. J. Jenv on “ The Glacial Deposits of Western Carnarvonshire.” Fic. 7.—Section east of Afonwen. Two beds of peat are seen separated by tough bluish-grey clay. As traced laterally these two beds of peat unite. Fic. 9.—Section of the Upper Boulder Clay in the cliff east of the Marine Terrace at Criccieth. To face p. 38, THE GLACIAL DEPOSITS OF WESTERN CARNARVONSHIRE. 39 origin, but erratics such as far-travelled granites and chalk-flints are occasionally found. In these hillocks and ridges the material is generally more or less rounded. The Drift of the plain rises on the flanks of the higher hills to heights of about 600 feet. But patches of glacial accumulations are met with at much greater heights. On Y Gyrn Goch, south-west of Clynnog Fawr, Drift occurs up to a height of 1200 feet above sea-level. It consists of yellowish clayey sand in which, however, no traces of marine shells were found. Mr Me.tuarp Reape notes the occurrence of “splendidly polished quartz-grains in the angular semi-decomposed gravel from near the summit of the Pass between The Rivals and Mynydd Carnguwch, about 800 feet above sea-level.”* As we pass west of a line taken as running from Clynnog Fawr to Pwllheli, the moraine-like mounds and esker-like hills and ridges tend to disappear. The gravels and sands continue, however, to cover much of the land and to form a mantle over the lower hills. The sands tend to become fine, and are more marine-like in aspect. The flats and hollows are underlain by Boulder Clay. The hills in the neighbourhood of Nevin, as, for instance, Garn Bodvean, which attains an altitude of over 900 feet above sea-level, have a distinctly moutonnéed aspect, and were during the Glacial period buried under the ice-sheet which moved from north-east to south-west. Exposures of the Drift are few in number and very poor in the western part of Lleyn, and boulders which at one time dotted the surface have for most part been cleared away. Beginning at the eastern end of the area investigated, the following notes were made of such exposures as could be found inland. Pont-Crychddwr, nearly a mile south of Llanllyfni. Mr Mrtiuarp Reape found a fragment of Eskdale granite, a Brachiopod from a Carboniferous shale, and fragments of schists probably from Anglesey in typical till exposed at a level of 400 feet above the sea on the right-hand side of the stream Afon-ddu, just above the bridge. Sections in the neighbourhood of Lianllyfni, made in driving headings in slate-quarrying operations, show boulder gravel lying upon hard buff-coloured clayey till. Again, Mr Reape found far-travelled granites, one of which was located as from Hskdale, in loamy and gravelly material seen in a section 30 to 40 feet high on the right bank of the Afonddwu, below Pont-Crychddwr, 350 feet above the sea. On the right bank of the Avon Llyfni there is a good section 35 feet in height showing sandy clay full of boulders and gravel. Near the railway station at Penygroes Mr Reape describest “a large mound of Drift, the major axis of which is one-third of a mile long, the direction being approxi- mately north and south.” The width of this mound is about 200 yards. The mound is composed largely of rounded stones, with sand and fine gravel, and stiff clay occurring in it irregularly. This can be seen in a ovavel- pit by the roadside on the way to Pont- y-Cim. Between Pont-y-Cim and Y Foel there lies a circular mound of Drift known as Craig-y-ddinas which has a height of about 40 feet above the road. It consists largely of boulder gravel and fine gravel often with a clayey matrix. The mound is surmounted * Proc. Liverpool Geol. Soc., vol. vii. (1893), p. 45. + Ibid., p. 48. 40 DRT. J. JEU" ON by an ancient encampment. The Foel itself is for the most part covered by a mantle of Drift and is grassed over. Several sand and gravel pits can be seen between Clynnog Fawr and Llanllyfni; as, for instance, just off the roadside above Tai’n-lon, and again on the side of Y Foel, where the material is gravel and in which a south of Scotland granite boulder was picked up. At Talarn a new cut for the Afon Llyfni disclosed a section of gravelly drift 440 feet above the sea. Sections of similar drift can be seen in most of the slate quarries, but according to Mr Reape its maximum thickness does not appear to exceed 20 feet in the bottom of the valley. This gravelly material has a clayey matrix. A eravel-pit of the usual character occurs at Ty-glas, about a mile to the south of Tar'n-lon. At Ynys-yr-Arch, when visited by the writer, a trench was being dug out to accom- modate a churning wheel. The section exposed showed :— 3. Soil—1 foot. 2. Gravel—4 feet. 1. Stiff bluish-grey Boulder Clay—6 feet. (The bottom of this Boulder Clay was not reached in the section.) Baron Hill, near Bwlch-y-derwydd, Pant-glas.—Just below the farmhouse on the north side boring operations in search of water were being carried out. In the shaft 18 feet were passed through when the locality was visited by the writer, and the material traversed consisted entirely of typical dark greyish Boulder Clay which was weathered yellow near the surface. Boulders were common in the deposit and were well glaciated. The bottom of this clay was not reached. At Llecheiddior Ganol, lying between Pant-glas and Ynys railway station, there occurs a gravel and sand pit. The deposits are bedded, and are coarse near the surface and fine below. The bottom of the sands is not exposed. No foreign erratics were noted. At Plas Llecheiddior, just behind the house, a section of the hillock is disclosed. Boulder rubble is seen overlying fine sand which has a depth of 4 feet in the section without reaching the bottom. Gravelly and sandy beds of the usual character were exposed in a pit at Llecheiddior Mill, but the pit is now filled up. In the neighbourhood of Brynkir are many mounds and elongated hills of gravel and coarse sand. The included stones are all more or less rounded by water action and apparently consist of locally derived or Welsh material. A rough bedding is usually shown. ‘The ridges are seen to run for most part from north to south. East of Brynkir, at, Blain-y-cae, gravel is again seen, but the deposits have become finer, passing into sand, as traced downwards from the surface. , At Graianog Farm, near Ynys station, there is a small gravel and sand pit just above the house. In this pit a chalk-flint was picked up, and also a granite with reddish felspars and black mica, which is undoubtedly a fair-travelled erratic though its parent rock has not been located. THE GLACIAL DEPOSITS OF WESTERN CARNARVONSHIRE. Al ce The place-names in this district are interesting and suggestive. xrainog” is the Welsh term for ‘‘ gravel,” and it occurs again and again in this neighbourhood, e.g. Bryn-grainog—the gravelly hillock ; Cors G'rainog—the gravelly moor; Ynys Grainog —the gravelly isle, etc. Several spots bear the name of Ynys (island), and these generally mark hillocks or mounds of gravel and sand standing above low marshy or boggy ground which is no doubt underlain by Boulder Clay. Much of this low ground is peaty, and in early historic or prehistoric times was probably under water, leaving the hillocks standing out as islands. Some of these islands may be mentioned as examples :— Ynys-wen, Ynys-y-creian, Ynys-grainog, Ynys-yr-Arch, Ynys-y-buntan, Ynys-hower; and there are several others. Most of the spots bearing such names mark hillocks of sand and gravel, but some denote rocky knolls projecting above the low ground, e.g. Ynys-galed—the hard or rocky island. Llanaelhiaiarn Gravel Pit.—This lies a quarter of a mile to the north of the village of Llanaelhiaiarn, near the mill situated just off the right-hand side of the main road from Pwllheli to Carnarvon. It is ata distance of 14 miles from the sea (Carnarvon Bay), and at a height of about 450 feet above sea-level. The section exposed is as follows :— 2. Brownish-yellow weathered clay, somewhat sandy in places, and with subangular boulders especially numerous towards the surface—é feet. 1. Stratified sands and gravels—15 feet. The beds of sand and gravel alternate, but towards the bottom of the exposure only fine gravelly material and sand are seen. The thickness attained by these gravels and sands is unknown, for their base is not reached in the section. Most of the included stones and boulders are of local origin, but specimens were also obtained of chalk-flints, and of a reddish-streaked rhyolite. This pit occurs on the side of a slope which is mostly grassed over. It is probable that the gravels and sands cover considerable areas to the north-east of this exposure. Bryn-quydin.—This place lies on the right-hand side of the road between Chwilog and Four Crosses, and near to the latter village. Here the hillock has been dug into for sand and gravel. The section is about 12 feet deep, but is much obscured by talus. Near the surface the material exposed is earthy, but below are beds of coarse gravel alternating with beds of fine gravel. The beds get finer and sandier as traced down- wards. Amongst the boulders are several examples of the reddish-streaked rhyolite which is so common in Lleyn. Just north of Four Crosses there occur pits showing deposits of a similar nature. The sand has usually a greyish colour. No marine shells are detected. West of Four Crosses, on the way to Nevin, some pits are again seen in fields just off the roadside. In these sands and gravels are again exposed, and these are in some of the pits overlain by 4 to 5 feet of bouldery rubble. Just east of Penprys, off the same road and at a height of 300 feet above sea-level, TRANS. ROY. SOC. EDIN., VOL. XLVII. PART I. (NO. 2). 6 42 DR T. J. JEHU ON there is an exposure showing 8 to 9 feet of coarsely bedded gravelly material which becomes finer when traced downwards. Amongst the pebbles were found specimens of the reddish-streaked rhyolite and one of a reddish granite which is foreign to North Wales. At Tan-y-llyn, near Glasfryn, there is a shaft for water, 30 feet deep, giving a good section. The material exposed consists entirely of tough, dark greyish-blue Boulder Clay full of boulders of all sizes, most of which are beautifully glaciated. The clay weathers to a brownish-yellow colour just at the surface. This was the deepest section met with in this region. Amongst the boulders the reddish-streaked rhyolite was common. Another shaft for water was dug at Tyddyn-Ucha about a quarter of a mile east of Tan-y-llyn. ‘The depth reached here was 9 feet, and the material traversed consisted again entirely of stiff bluish-grey Boulder Clay full of well-glaciated stones. At Tg-Cam, a quarter of a mile distant from Four Crosses, in the direction of Pwllheli, a pit was dug for a water-wheel. Stiff bluish-grey Boulder Clay was exposed to a depth of 6 feet. Below the Boulder Clay shaly rock in situ was reached. The rock was much shattered just under the clay. Numerous boulders of the reddish- streaked rhyolite were picked out of the deposit. Near Lianor, 24 miles inland from Pwllheli, and about 5 miles from Carnarvon Bay, there are brick-works in which the stiff bluish-grey Boulder Clay is exposed for a depth of about 20 feet. The bottom of the deposit is not reached in the section. The stiff clay is capped by 2 or 3 feet of yellowish weathered rubbly clay. Well-glaciated stones were common in the lower shift clay. The boulders consist chiefly of diabase, slate, etc., all of which appeared to be of Welsh origin. There were some indications of shells in an extremely fragmentary condition. The reddish-streaked rhyolite was seen imbedded in the clay. Passing west of a line from Pwllheli to Nevin the low grounds are underlaid by Boulder Clay and the smaller hills are covered by gravels and sands. Exposures are not common, but a few may be mentioned. On the hill immediately to the south of Tu-hwnt-v'r-afon Inn, Rhyd-y-clafdy, sand and gravel are exposed in many shallow pits. Near the surface the material is gravelly, with a somewhat clayey matrix, and below this we find fine yellow marine-like sand. The exposures are only 4 to 5 feet deep, but the sands must attain a considerable thickness towards the base of the hill. Several gravel pits are found between Bodvean and Cors Greirch. One of these lies a quarter of a mile west of Bodvean, and shows a section 15 feet deep. It is situated on the slope of a hill at a height of almost 200 feet above sea-level. The deposits seen are mostly sands, fine, yellow in colour, and marine-like. Some pebbly layers occur, and at the top a little yellowish clay passing into soil. A little further south is found another with an exposure 6 to 7 feet deep. ‘This is at a height of 175 feet above sea- level. It consists mostly of gravelly material getting finer and more sandy below. On the same hill slope, but at lower levels, fine yellow marine-like sands are exposed at several places. The entire hill is evidently covered by a mantle of gravel and sand. THE GLACIAL DEPOSITS OF WESTERN CARNARVONSHIRE. 43 The marshy land below, known as Cors Geirch, is underlaid by Boulder Clay. This Cors probably marks the site of a former lake. At Nanhoron there is an old sand pit in the field in front of the house. It is now partly filled up, but formerly it was open to a depth of over 10 feet. Fine marine-like sands, well-bedded, are still exposed, with gritty layers here and there. These sands are of considerable thickness, for on digging to a depth of over 10 feet their base was not reached. ‘The entire hill on which the house stands is rounded over by these sands. But to the south, where the Afon Horon runs at a lower level, the stream has exposed the lower stiff bluish-grey Boulder Clay which here, as in the coast sections, underlies the Sands and Gravels. The stream has here not cut through the Boulder Clay. Passing north to the neighbourhood of Madryn we still find similar features. In a field on rising ground some 200 yards south of the Hall, and 60 feet above it, sand and fine gravel are seen just below the surface. The sand is of the same character as that met with at Nanhoron. Again, in the park to the north of the Hall, fine yellow sand is exposed in many places and is often seen brought up from rabbit burrows. A large area is here covered by this sandy deposit. At Meillionen, near Ceidio, coarse gravelly sand is exposed at the surface and to a depth of 4 to 5 feet. In the lower ground adjoining, the stiff Boulder Clay is met with. { At Cefn-leisog, in a field just south of the farmhouse, lies a disused sand pit. Here the sand is covered by 4 to 5 feet of yellowish weathered clay full of boulders. Passing to Brynodol we find that much of the land north of the house is sandy, as seen in several poor exposures and in newly dug ditches. At Ty-Issa, Tydweihog, a boring was recently made for water. The deposits traversed were as follows :— 3. Sand—6 feet. 2. Stiff bluish-grey Boulder Clay—27 feet. 1. Coarse sand—3 feet. (The bottom of this sand was not reached.) A little higher up, at the County School, Tydweiliog, a boring was also made, and this passed through :— 2. Sandy Clay—16 feet. 1. Pure Sand—2 feet, the bottom of which was not reached. A little further down the slope of the same hill another boring was made which passed through nothing but marine-like sand to a depth of 10 feet. So we may gather from these various borings at Tydweiliog that the Drift covering the land in this neighbourhood consists of :— 3. Sandy Clay. 2. Sands. 1. Stiff bluish-grey Boulder Clay, which at one place at any rate is underlaid by or includes Sands. 44 DRT. J. JEHU ON Going westwards, at Plas-ym-mhenllech a sand pit was found in a field near the junction of the roads to Sarn and to Aberdaron. There were exposed here 5 feet of coarse sand in which were seen streaks of stiff clay. Going south again similar deposits are found at the surface in the neighbourhood of Sarn and Bottwnog. Near Llangian, just where the road turns off to Llawr-y-dref, there is a gravel pit in which 10 feet of coarse gravel are exposed. The gravels show faint indications of bedding. Another gravel pit is seen at Nezgwi-Plas, in front of the house, the gravel being coarse and disclosed to a depth of 4 or 5 feet. All about the farm are many other disused pits of a similar nature. The gravels often pass down- wards into fine sands. A pit at Neigwl Ganol shows stony yellow weathered clay at the surface and to a depth of 4 feet. In a gravel pit at Talsarn the material is more rubbly—the stones being less rounded and more flaky. Above Talsarn Bridge the Afon Horon is seen to run on the stiff bluish-grey Lower Boulder Clay. This can be traced right up the stream, which further up its course becomes a mere rill and does not appear anywhere to reach the solid rock. The area between Abersoch and Porth Nigel is largely covered with sands. These are well exposed near Cim, at a height of over 200 feet above sea-level. The sands are fine and marine-like and have a yellow colour. Rhos-lurwawn, south-west of Llangwnadl, seems to be underlain by a stony weathered Boulder Clay. As we approach the extremity of Lleyn the mantle of Drift tends to become thinner until a mere skin of yellow weathered stony clay is left. No systematic examination was made of the erratics lying on the surface of the land, but in traversing the inland region in search of sections of the Drift the following were noted :— On Y Foel, between Llanllyfni and Clynnog Fawr, a block of felsite lies perched on the slate at a height of over 600 feet above the sea-level. On Y Gyrn Goch, at a height of 800 feet, a small boulder of the Eskdale granite was found, and a little lower down another of a granite from the south-west of Scotland. Kast of Porth Nant, on the western slope of Yr Huff, a boulder of Carboniferous Limestone was picked up at a height of 450 feet. Near Tan-y-foel, between Llithfaen and Pistyll, a boulder of greyish-white granite, probably Scottish, was obtained at an elevation of over 500 feet. At the gate leading to Berthdaur-fawr, near Penllech, there stands a huge boulder measuring 5 feet by 3 feet by 1} feet. It is a very fresh-looking basic rock. A fragment was taken away and subsequently sliced and examined under the microscope. It proved to be a beautifully fresh olivine dolerite showing typical ophitic structure. It is difficult to locate its place of origin. The large size of the boulder makes one doubt whether it can have come from the west of Scotland, where similar rocks are found. It may possibly have been derived from some Tertiary dyke in Anglesey. On Penllech were found erratics consisting of granites, a quartz-porphyry with the THE GLACIAL DEPOSITS OF WESTERN CARNARVONSHIRE. Ad5 quartz showing fine pyramidal forms, the reddish-streaked rhyolite, Millstone Grit, and quartzites probably from Anglesey. The reddish-streaked rhyolite again occurred on Mynydd Annelog, near the extremity of Lleyn, at a height of 350 feet, together with a boulder of Millstone Grit. Between Llanllawen and Aberdaron many boulders of diabase are dotted about. These are probably mostly of local origin. Blocks of jasper and of serpentinous rocks also occur which are derived from the neighbourhood of Careg, not far away to the north. On Mynydd Cilan, lying between Porth Nigel and Porth Ceiriad, and at heights of about 300 feet, the following erratics were found :—Several boulders of a coarsely crystalline diabase, the reddish-streaked rhyolite, Carboniferous Limestone with crinoid stems, a coarse grit quite unlike the rock of the district, light-coloured felspar porphyry derived from the masses of the rock which are found in situ to the north-east, Millstone Grit, and a reddish agglomerate. On Mynydd Mynytho, which consists of felspar porphyry, several erratics of diabase were found, also a boulder of a dark felsitic rock at a height of 400 feet on the south side of the mountain, and near it a small boulder of the reddish-streaked rhyolite. The reddish-streaked rhyolite is perhaps the commonest of all the boulders occurring in the Drift of Lleyn. It is found on the surface, in gravel pits inland, in the coast sections, and as pebbles on the modern beach. Its parent rock cannot be very far away, and probably lies to the north-east in Anglesey or Carnarvonshire. The boulders of this rock have been submitted to several petrologists, but no one has yet located its parent rock. IV. SumMaRY AND GENERAL CONCLUSIONS. In the western part of the Lleyn peninsula the Lower Boulder Clay is seen at some places to be separated from the bed-rock by an accumulation of angular detritus having all the characteristics of a ‘“‘ Rock Rubble” or “ Head” (fig. 4). This rubble is disclosed in some of the coast sections towards the extremity of Lleyn, as, notably, at Porth Oer and at Aberdaron. It rests on the rocky slopes of the valley sides, where the valleys terminate seawards, and with these slopes it descends on either side below the level of the modern beach. This ‘‘ Head” consists of angular blocks and fragments of all sizes packed close together or lying in an earthy matrix. The material seems to be entirely of local origin, and to be derived in the main from the immediately adjoining rocks above. lt sometimes shows a rude bedding, as, for instance, at the east end of Aber- daron Bay, where it consists below of flakes and slivers of rock arranged with their long axes parallel to one another, giving a linear arrangement, and followed above by coarser material in which big angular blocks lie. During the time of the accumulation of this rock rubble the land must have stood higher relatively to the sea than it does at present. The lower ends of the valleys or inlets on the sides of which the rubble is seen are now often choked with the Drift deposits which pass below the level of the modern beach. 46 DRT. I. JERU, ON The seaward terminations of these valleys or depressions are drowned, and their lower reaches must have been eroded during pre-Glacial times, when the land stood higher and extended further out to the sea than it does at present. The rock rubble is the result of subaerial work under severe climatic conditions. It is generally admitted that ice in some form or other has aided in the formation of such deposits. ‘lhe rock fragments have all the appearance of being frost-riven, but apart from this there is not much trace of weathering. This ‘‘ Head” may therefore be regarded as angular debris of local deriva- tion moved forward or downward by the action of frost, melting snows, and thawing sub- soils. On the steeper slopes the action of gravity alone would account for the slipping down of the material. Sometimes, on the other hand, the rubble is a confused mass of broken rock showing no arrangement. As a rule this rock rubble is overlain by the Lower Boulder Clay, but there are places where the Lower Boulder Clay seems to be absent and where the rubble is immediately followed by the Upper Drift deposits. The rubble always lies directly on the bed-rock, and the fact that it is generally seen to have the Lower Boulder Clay superposed proves that it is the lowest of the superficial deposits which occur in the area. In the eastern part of the region this rock rubble is not seen unless it be represented by the shattered rock which underlies the Lower Boulder Clay at some places. The Lower Boulder Clay sometimes rests directly on the bed-rock, and where this is the case the rock surface as it passes under the clay has a markedly glaciated aspect ; some- times it is separated from the rock by a foot or so of broken and confused rock material, and sometimes, as on the rocky slopes at the seaward termination of valleys or inlets, it rests on an accumulation of rock rubble or “Head” of a considerable thickness. It is the most widespread of all the Drift deposits. This clay is very tough, compact, and homogeneous, having a massive appearance as displayed on the cliff face. It is often seen to be traversed by joints, and to break off in huge slabs along the joint faces as at Porth Nigel. The colour is dark bluish-grey, but after drying it becomes more of a light bluish-grey. Boulders are scattered sparingly through it, and are sub- angular or more or less rounded and often intensely glaciated. Distant erratics are common, and include Carboniferous Limestone, schists, etc., from Anglesey, various granites and porphyrites from the south of Scotland, and possibly some granites and lavas from the Lake District. It is interesting to note the occurrence of the Ailsa Craig riebeckite micro-granite at many places in this Boulder Clay and the frequent presence of chalk-flints which must have come from Cretaceous deposits on the floor of the Irish Sea or in the north-east of Ireland. The Boulder Clay is shelly, especially in the western portion of the peninsula; and though the shell-remains are generally very fragmentary, some complete shells were obtained and a considerable number of forms have been identified. These include some species which are Arctic and Scandinavian and do not live now in British Seas, such as 7’rophon (Boreotrophon) clathratus, Trophon (Boreotrophon) scalariformis, and Astarte (Tridonta) arctica ; some of a northern type but not confined to high latitudes, such as Cyprina islandica THE GLACIAL DEPOSITS OF WESTERN CARNARVONSHIRE. A7 Astarte compressa, and Buccinum undatum; and several species which are now common in British Seas. Thus we have in the Lower Boulder Clay a mixture of species belonging to different climates. ‘These included shells and shell-fragments are therefore evidently derived, and point to the fact that the mer de glace of which this Boulder Clay is a product travelled over a sea-floor ere it reached this region. A part of a Belemnite was also found in the Boulder Clay at Aberdaron and several pieces of wood in that at Porth Nigel. This Boulder Clay is, as a rule, strikingly homogeneous and uniform in character and has evidently been subjected to great pressure. But at places a fine lamination is seen in the clay, especially towards the top. Where this lamination occurs the clay is stoneless and sometimes sandy. It appears to be sometimes due to shearing move- ments. Wherever land-sliding has taken place lamination occurs towards the top of the Lower Boulder Clay. On the foreshore at Porth Nigel an unctuous stoneless clay is seen which seems to result from the running or sliding downwards of Boulder Clay from which the coarser material has been washed out, leaving the residue in a buttery condition with a tendency to go on sliding down the foreshore. The lamination of the stoneless clay in the upper part of the cliff at Porth Nigel may have arisen in some- what the same way, for it is seen where slipping has taken place. The best exposures of the Lower Boulder Clay are met with in the coast sections, especially towards the western end of the Lleyn peninsula, but throughout the district there are indications that it underlies the Sands and Gravels, although these often form the lower part of the cliffs further east. The Lower Boulder Clay is shown by a boring to underlie the modern alluvium which forms the flats to the north of Dinas Dinlle. It has been detected below the Sands and Gravels of the Dinas Dinlle mound, but no further exposure is seen along this coast until we reach Gwydir Bay, where it forms a conspicuous feature below the other Drift deposits and under the foreshore. From Gwydir Bay westwards it appears in most of the bays, becoming more and more prominent and forming greater parts of the cliffs of Drift as we pass towards the extremity of Lleyn. It appears to have choked up most of the depressions, channels, and inlets met with along the coasts of the western part of Lleyn, sometimes even to the brim. These drift-filled depressions are now being rapidly eroded by the sea. At the head of the bays the Boulder Clay often forms the entire cliff, and extends below the modern beach to an unknown depth; but towards the sides rock rubble is often seen lying between the Boulder Clay and the rocky slopes where these come down to the beach. At Porth Oer, Aberdaron, and Porth Nigel, cliffs formed of the Lower Boulder Clay attain heights approaching 100 feet. At Aberdaron, towards the west end of the bay, there appear to be inclusions of large irregular masses of sand in the clay, and towards the east end some bedded sands and gravel appear in the Lower Boulder Clay. The finest exposure of the Lower Boulder Clay in Western Carnarvonshire is that at Porth Nigel, where it forms high cliffs for a length of 3 to 4 miles. Following the coast on the southern side of the peninsula eastwards, the Lower Boulder Clay is 48 DRT... JEU 7ON last seen at Llanbedrog, where it forms the lower part of the cliff and the foreshore at the west end of the bay. A boring indicates its presence below the modern alluvium in the harbour at Pwllheli, but east of this it does not appear—the cliffs from Afon- wen to Criccieth being formed entirely of the Sands and Gravels and the Upper Boulder Clay. But though not exposed anywhere along this part of the coast, it probably underlies these Upper Drift deposits. Inland the Lower Boulder Clay is rarely exposed at the surface, but it may occa- sionally be seen underlying the marshy low grounds and moors or along the banks and beds of streams which have cut through the Upper Drift deposits, as, for instance, along the Afon Horon, below Nanhoron. Even where there are no exposures at the surface, borings and artificial cuttings prove it to be present under the other deposits. At Ty-Cam, Four Crosses, north of Pwllheli, it was found resting on a shattered surface of rock. Above the rocky cliffs at Porth Pistyll, south-west of Aberdaron, it is seen resting on the bed-rock at a height of over 100 feet above the sea-level. In other borings inland the bottom of this clay is not reached, but at Tydweiliog some sand was reached lying in or below the Boulder Clay. Shell-remains were not observed in the Boulder Clay away from the coast, except doubtfully in the brickfield at Llanor, north of Pwllheli. But the inland sections and exposures of this clay were too poor to render any adequate search for shell-remains or erratics possible. All that can be stated with certainty is that Boulder Clay underlies the sands and gravels in the interior as well as on the coast, and that this clay is of a dark bluish-grey colour and is very tenacious and compact ; that in some places it is full of boulders, many of which are well glaciated, and at others fairly free of boulders. The included boulders are most numerous in the eastern part of the district which approaches the mountains of Snowdonia, and these are mostly of Welsh origin. The Lower Boulder Clay has all the characteristics of a true bottom-moraine, and is undoubtedly the product of an ice-sheet which came from the north and invaded Western Carnarvonshire after having made its way over the bed of the Irish Sea and passed over Anglesey. The direction of transport of the included erratics, the configuration of the glaciated hills and the roches moutonnées, and the course of the rock-strize all indicate that the movement of the ice was roughly from north-east to south-west. Professor JAMES GEIKIB, in his classic book—The Great Ice Age—gives a map of the British Isles during the Epoch of Maximum Glaciation in which he depicts this mer de glace as overwhelming Anglesea, skirting the north-west coast of Carnarvon- shire, and passing south-westwards over the tip of the Lleyn peninsula. The late Mr CarvILL Lewis, on the other hand, in a sketch-map which is reproduced in Professor Bonney’s Ice Work (1896), p. 186, makes what he terms the Irish Sea Glacier terminate in an edge passing along and just within the north-west boundary of Lleyn westwards towards Ireland, leaving the south of the Lleyn peninsula and Cardigan Bay outside its boundary. ‘The facts recorded in this paper prove that this great ice-sheet covered the greater part of Western Carnarvonshire, passing over Lleyn into Cardigan Bay. In a previous paper communicated to this Society, the writer proved that this THE GLACIAL DEPOSITS OF WESTERN CARNARVONSHIRE. 49 ice-sheet filled Cardigan Bay and even invaded Pembrokeshire. It is impossible to define accurately the eastern boundary of this ice-sheet. The northern ice met and overcame the Welsh glaciers which moved towards Anglesey from the valleys of Snowdon ; it invaded the foot-hills, such as Moel Tryfan, lying west of the Snowdonian mass. Professor KENDALL, in Wricut’s Man and the Glacial Period (1893), p. 171, has pointed out that it was the proximity of Snowdon which really enabled foreign ice to invade Wales in the Moel Tryfan region. He observes that “a glance at the map will show that the ‘radiant point’ of the Welsh ice was situated on or near Arenig Mawr, and that the great mass of Snowdon stands quite at the periphery of the mountainous regions of North Wales, so that it would oppose its bulk to fend off the native ice-sheet and prevent it from extending seaward in that direction.” Consequently, the northern ice-sheet was able to overwhelm Western Carnarvonshire as far east as a line running from Carnarvon to Cardigan Bay somewhere in the neighbourhood of Pwllheli. The western slopes of the Snowdonian Mountains could only have given rise to small glaciers, and these would unite and become merged into the glacial mer de glace which moved south- westwards. South of Snowdonia a great glacier came down the Vale of Gwynant past Bedd-gelert, reaching Cardigan Bay at Traeth Mawr, where it was joined by another issuing from the mountains of Merioneth by the Vale of Ffestiniog. The united mass would be large enough to fend off the northern ice and to spread out as a fan towards Criccieth and Afonwen, and the volume of native ice would be increased by a smaller stream coming down the mountain valley between Moel Hebog and Carnedd Goch. The presence of the large volume of native ice in that neighbourhood explains the absence of foreign erratics and of shell-remains in the Drift deposits of the coast sections from Afonwen to Criccieth. The Sands and Gravels there seen are probably derived from a Lower Boulder Clay, although this is nowhere exposed on this part of the coast. We may assume that, if exposed, the Lower Boulder Clay in this part, like the Sands and Gravels, would show a general absence of foreign erratics. The Intermediate Gravels and Sands form a very variable series and are somewhat inconstant in their occurrence. They comprise both stratified and unstratified material, but the former predominates, especially in the coast sections. The best exposures are met with in the cliffs along the coast-line, the series being especially well developed at Dinas Dinlle, in the stretch between Clynnog and Gwydir Bay, at Nevin and Porth- dinlleyn, at the eastern end of Porth Nigel, at Llanbedrog, and east of Afonwen. The sands are, as a rule, yellowish or yellowish-brown in colour, and are marine-like in aspect ; they vary in texture, showing all grades from very fine sand to coarse gritty material. ‘lhe coarser sands pass into gravel and boulder gravel. Mr Metiarp Reaper's investigations proved that the sand contains a large propor- tion of extremely well-rounded grains often possessing an extraordinary polish and transparency. He speaks of these as “microscopic boulders,” and states that the contrast between these and the local grains is most remarkable. The sands and gravels usually show bedding, although at places confused irregular masses occur, especially of TRANS. ROY. SOC. EDIN., VOL. XLVII. PART I. (NO. 2). 7 50 DRI Ts w.. VEHU VON the boulder gravel. As a rule the sands underlie the gravels. The pebbles and boulders of the gravels have not the flattened, ovoid form which is so characteristic of the shingle on modern beaches. At some places along the margin of the north-west coast, notably at Nevin, there occur laminated silty clays usually high up in the series. These silty clays are stoneless and devoid of shell-remains and appear to have been laid down tranquilly in water of some depth. Along this part of the coast the Drift consists chiefly of gravel and sand and silty clays, and the upper surface of the Drift has a terraced aspect. Some traces of disturbance and contortions are occasionally seen in the sandy beds. The sandy series at some places lies immediately on the bed-rock, but at others and especially in the deeper depressions, it rests on the Lower Boulder Clay. The line of demarcation between the Lower Clay and the Sands and Gravels is often well marked, but at places there appears to be a gradual passage from the one to the other. The pebbles and boulders in the Gravels include distant erratics of like origin to those found in the Lower Boulder Clay. Chalk-flints are not uncommon, and specimens of the Ailsa Craig riebeckite granite were obtained at several places. Shell-remains occur especially in the coarse sands and beds of gritty material. They are commonest in the sands on the north-west coast. Neither shell-remains nor distant erratics were noted in the Sands and Gravels on the south coast east of Llanbedrog. The shells found in the sands are very broken, usually occurring as mere crumbs. A few of those obtained from the sands at Nevin and Porth-dinlleyn have been identified, but the list is too limited to yield definite conclusions as to climate. But the sands in this region are very similar to the shell-bearing sands of Moel Tryfan, and further south in Pembrokeshire,* the writer collected in similar sands a much larger number of shell-remains which were capable of identification. At both places the shells pointed to a mixture of species belonging to different climates. This, together with the fact that everywhere the shells are very broken and much rolled, indicates that they are not now in or near the positions in which they lived. In no case were Lamellibranch shells found with the two valves in apposition such as one finds in a recent and undisturbed alluvium like that which occurs to the north of Dinas Dinlle. Marine-like sands of a similar nature with gravels above are found inland in patches all over the peninsula. They lie on the slopes of, and sometimes spread over, the lower hills. Hastwards, as we approach the Snowdonian Mountains, the sands tend to become coarser and more greyish in colour and they are less marine-like. No shell-remains were noted in the sands of the interior, but the exposures met with are very poor and afford but little opportunity for a thorough search for shell- fragments. Such fragments are known to occur in the sands on Moel Tryfan and again away fromthe coast in Pembrokeshire, and so it is probable that this is the case alsoin Lleyn. Stratified sands and gravels were found at Llanaelhaiarn at a height of 450 feet above sea- level. ‘The well-known shell-bearing sands of Moel Tryfan occur at a height of 1350 feet. In Pembrokeshire marine-like sands and gravels were found at elevations of 640 feet. * Trans. Roy. Soc. Hdin., vol. xli., part i. (No. 4), 1904. THE GLACIAL DEPOSITS OF WESTERN CARNARVONSHIRE. Dill These shelly sands and gravels evidently represent the materials of a sea-bottom carried onwards and upwards by an ice-sheet and rearranged by fluvio-glacial action. That is to say, they are the remaniés of the great ice-sheet which came from the north and passed over the floor of the Irish Sea. The materials were laid down during the retreat of the ice-front, and at that time the lower parts of Lleyn must have been under water. The boulder gravel may be partly due to the action of torrential waters issuing from the melting ice-sheet and glaciers. No deposits indicating a genial inter-glacial epoch have been discovered in Western Carnarvonshire ; but this is not a matter for surprise, for such deposits, if they once existed, would stand very little chance of being preserved in a region so exposed as this. Whether the sands and gravels indicate a mere temporary retreat of the ice-sheet or the oncoming of a true inter-glacial epoch it is impossible to decide from the evidence afforded by a study of this region alone. The Upper Boulder Clay again is very sporadic in its occurrence and varies much in thickness and character. Sometimes it is a typical tumultuous unstratified till with boulders of al] shapes and sizes scattered pell-mell throughout the clayey matrix; at other places it has some of the character of rubble drift. As compared with the Lower Boulder Clay the Upper Till is less compact and homogeneous, more stony in character, and more weathered and friable. It often passes into a sandy clay. Where it becomes a rubble drift it consists of an agglomeration of subangular and more or less rounded boulders crowded thickly together in earthy debris. The included boulders are derived in the main from Welsh rocks, but many far-travelled stones are also found. Ice-scratched boulders are fairly common. The distant erratics are of similar origin to those found in the lower deposits, and include Scottish granites, the Ailsa Craig rock, and chalk-flints. Shell-fragments are again present, but these are scarcer and even less well preserved than those found in the Lower Boulder Clay. This Upper Boulder Clay is well displayed in the coast cliffs of Dinas Dinlle, and again between Point Maen Dulan and Clynnog, where it often forms the entire cliff. West of Clynnog it is not so conspicuous in the coast sections, but in many of the bays it can be seen to cap the cliffs which are formed of the Drift deposits. As a rule the Upper Clay rests on the Gravel and Sand series, but occasionally the two boulder clays occur together, the Upper Clay resting directly on the Lower, and in such cases it is sometimes difficult to separate them. In some of the smaller bays on the south-west coast of Lleyn the Lower Boulder Clay is not exposed; in the cliff sections and at the sides of these bays the Upper Boulder Clay rests immediately on the rocky slope or is separated from it by some rock rubble. East of Afonwen the Upper Boulder Clay becomes again conspicuous. There it forms the upper part of the cliff, becoming more prominent and well-defined as we approach Criccieth. To the east of the town it forms the entire cliff. In the coast sections east of Afonwen no shell-remains were noted in this clay, and the included boulders seem to be entirely of native origin. 52 DR T. J. JEHU ON Inland this Upper Boulder Clay is often absent. When it occurs it forms a thin covering only a few feet deep, and is often a mere rubble of boulders set in an earthy matrix. It has evidently suffered much from weathering and denudation. This Upper Boulder Clay may be the product of another mer de glace which followed the same course as that which produced the Lower Boulder Clay, and which was not much less extensive than its predecessor; or it may be due to the re-advance of one and the same mer de glace. Whether the two boulder-clays be the product of two separate and distinct ice-sheets separated by a genial inter-glacial epoch, or the product of one glacial epoch which was marked by an extensive oscillation of one ice-sheet, it is impossible to decide on the evidence presented in Western Carnarvonshire. The Drift deposits of this area can be accounted for on either of these theories. The ridges and mounds of gravel and coarse grey sand which run from north to south from the neighbourhood of Penygroes to that of Brynkir represent morainic material re-arranged and re-sorted by fluvio-glacial action and under the influence of the torrential waters which must have flowed over the surface at the time of the final disappearance of the ice. They occur near the eastward margin of the mer de glace which came from the north and passed over Lleyn. At the time of maximum glaciation the mountains of Lleyn must have been almost, if not entirely, buried under the ice-sheet. On Y Gyrn Goch, south-west of Clynnog, Drift was noted at a height of 1200 feet above sea-level, and a boulder of a foreign granite at 800 feet. The highest peak of Yr Hifl (The Rivals) reaches an elevation of 1849 feet, and it is possible that the summit of this mountain stood as a “nunatak above the surface of the ice. The upper part of Yr Hifl has not a glaciated aspect, but of course this may be due to subsequent weathering. It is interesting to compare the sequence of glacial deposits in the Welsh area on the eastern side of Snowdonia with that met with in Lleyn. Professor KenpaLu* gives the succession on the coast east of Little Orme’s Head as follows :— 4. Boulder Clay with northern erratics and shells. 3. Sands and Gravels with northern erratics and shells. 2. Boulder Clay with northern erratics and shells. 1. Boulder Clay with Welsh erratics and no shells. A similar succession is to be seen in the Vale of Clwyd. In these districts the Lowest Boulder Clay (1) is entirely the product of Welsh ice, which spread without hindrance seawards in the early stages of glaciation prior to the advance of the northern ice-sheet. In Lleyn this lowest Boulder Clay of the eastern area seems to be represented by the rock rubble or ‘‘Head” which underlies the Boulder Clay (2) laid down on the advance of the northern ice-sheet. So west of Snowdonia the sequence is as follows :— * WricuHt’s Man and the Glacial Period, 1893, p. 148. "y+... ° es eee ; ‘ THE GLACIAL DEPOSITS OF WESTERN CARNARVONSHIRE. 53 4. Boulder Clay with northern erratics and shells. 3. Sands and Gravels with northern erratics and shells. 2. Boulder Clay with northern erratics and shells. 1. Rock rubble or “ Head.” That the lowest Boulder Clay which is found to the east of Snowdonia, and which is the product of native ice, does not occur also in Lleyn, is explained by the fact that there are no gathering grounds for the production of glaciers of any size in that area, but the severe climatic conditions would favour the accumulation of the rock rubble or “ Head.”’ The Drift which borders the coast of Lleyn has often a markedly terraced aspect. This appearance is well seen in the neighbourhood of Clynnog, and again between Yr Kifl (The Rivals) and Porth-dinlleyn. The upper surface of the Drift in these places looks remarkably even as seen from a distance, and lies at a height of 100 to 150 feet. A similar terraced aspect was noted at Llanbedrog, but here the surface is only about 40 feet above sea-level. Where the Drift of the coast shows this more or less even surface it is found to consist for most part of the Sands and Gravels, and this may be taken to indicate a submergence of the land to an extent of 150 feet or more. But the character of the shell-remains and the form of the boulders in the boulder gravels militate against the theory that these sands and gravels are ordinary marine deposits ; and where the cliffs are rocky there are no indications that the land stood lower relative to the sea than it does at present. It is more probable that these stratified Drift deposits were accumulated between the margin of the retreating ice and the land area. The present coastal plain would at that time be under fresh water, due to melting of the ice and to the ponding of the land drainage. Along certain parts of the north-west coast of Lleyn, notably at Porth Colman, a rock- shelf or platform is seen at a height of 10 to 15 feet above the level of the present beach, which simulates an old shore platform such as that which has been described by Messrs Wricur and Murr on the south coast of Ireland.* The surface of this platform is clear of Drift for a few yards inwards from its seaward edge ; and it slopes gently upwards until it disappears under the Drift—usually the Lower Boulder Clay (fig. 2). This platform cannot be traced back to any well-marked pre-glacial cliff, nor is it overlaid anywhere by any undoubted shore-deposits. At its inner end, where it passes under the Boulder Clay, the surface often has a glaciated aspect. In early Glacial times the shore-line stood further out to sea than it does at present, and the land descended in a gentle slope to sea-level. The surface underwent severe glaciation and was buried under Drift. Since that time there has been some subsidence, and the cliff has also been cut back by marine erosion and the foreshore lowered. Where the land slopes gently to form a low cliff, the Drift has been cleared away for some distance from the edge during stormy weather and high spring tides, when the waves would be dashed against the face of the cliff to considerable heights. * Screntefic Proc. Roy. Dublin Soc., vol. x. (N.S.), part ii. (1904), p. 253. 54 DR T. J. JEHU ON On the south coast of Lleyn there are indications that during some pre-glacial period, possibly the Pliocene, the bays extended further inland than they do at present. What appears to be an ancient line of cliffs passes inland from the sides of Porth Nigel, by Llanengan and Llangian, towards Bottwnog, and back along the eastern side of Mynydd- y-Rhiw (fig. 10). The base of the old cliff is hidden under Drift, but its course follows roughly the 100 feet contour line. The Drift deposits which fill much of this ancient bay are now being rapidly eroded again. The Drift is rather higher on the coast at Porth Nigel than it is for some way inland, and so the drainage has been altered. The Afon Soch, which appears to have a natural outlet in this bay, is deflected so as to reach the sea at Abersoch (see Map). The course of an ancient pre-glacial coast-line can be traced again in the region east of Llanbedrog. At the time of the retreat of the ice-sheet glacial lakes must have been formed in Lleyn, and there are indications of overflow channels at many places, as at Trevor, in Nant Bodlas near Nanhorn, and along the valley of the Soch between Llangian and Abersoch. Some subsidence must have taken place during Glacial times, but there is no evidence to show that the land stood at.a lower level relatively to the sea than it does at present. After the passing away of glacial conditions the land area appears to have had a greater extent than it does now. Peat, with the roots of trees am situ, are occasionally exposed at low tides in Cardigan Bay. The most recent movement would therefore appear to be one of subsidence, and this view is supported by Welsh traditions regarding the loss of a land area known as Cantief Gwaclod which formerly extended over much of what is now Cardigan Bay. Some geologists hold that from the onset of the land-ice to the time of its final disappearance there was a period of continuous glaciation, during which the former sea- basins were never emptied of their ice-sheets. This view has recently been strongly advocated by Mr Lampiucn. ‘To account for the accumulation of the stratified deposits associated with the boulder clays in the West British area, somewhat extensive oscilla- tions of the mer de glace are admitted. Speaking of the Isle of Man in his Presidential Address to the Geological Section of the British Association (1906), he says that at the time of its maximum glaciation the surface of this ice-sheet stood more than 2000 feet higher than present sea-level. ‘Then followed a declining stage in glaciation, during which the ice-sheet shrank away from the hills, which were never again covered. . . . The recession of its margin was relatively accelerated in the northern part of the island, so that a broad hollow was formed there between the hills and the ice-border: in this hollow a mass of stratified drift was deposited.” This stratified material he regards as marginal ‘‘overwash” from the ice-front. ‘hough waning and shrinking for a time, he thinks the ice-sheet did not disappear entirely from the surrounding basin. “‘ Following closely upon this local deposition of stratified drift, there appears to have been a limited re-advance of the ice, which brought about the accumulation of an upper boulder clay on parts of the low ground.” He thus admits a declining stage of glaciation,which was followed after the deposition of the stratified material in a re-advance of the ice, but Vou, XLVIL. Soc. Hdin. J. SOC. Trans. Po Dr T. J. Jest on ‘* The Glacial Deposits of Western Carnarvonshire.” “OUT[-JsBOO JUoloUv Ue syTeut A[qeqoad aseq[IA ayy 07 purrur Surssed punois Suis ayy, ‘“wesueuely Jo osey]1A ayy YATM “TaStNy YAtog JO puBlur Mel A—‘OT ‘o1g cee OD P Ulta JO 1 lig To face p. = ay THE GLACIAL DEPOSITS OF WESTERN CARNARVONSHIRE. 95 only to an extent sufficient to “ close in again upon the lower flanks of the hills.” In his Survey Memoir, on “The Geology of the Isle of Man” (1903), p. 395, Mr LampLucu has adduced reasons to show that “the shrinkage of the ice-sheet covering the Isle of Man is likely to have commenced while the Welsh and Ivernian sheets were still in- creasing.” “But,” he adds, ‘‘it was not until there had been a great amelioration of climate that the island began to emerge from the waning mass” (p. 395). So this “reat amelioration of climate” must have taken place at the time in which the stratified Drift deposits were accumulating between the receding ice-sheet and the emerging land. Does not this change in the climate indicate the gradual passing away of glacial conditions at a time prior to the deposition of the Upper Boulder Clay? Mr LaMPLuGH refers again and again to the fact that a very considerable alteration in the climate must have occurred to account for the shrinkage of the ice from the island. “The shrinkage,” he observes (p. 396), ‘‘may have begun from a diminution in the amount of snowfall alone; but before it could proceed far there must have been a wide- spread change of climate and an essential difference between the conditions of this stage and of the initial stage of the period. At the beginning of the period the climate was such as to permit the permanent snow-line to descend nearly to sea-level; while, when the ice-sheet had reached its maximum, it was requisite that the climate should be warm enough to prevent snow lying permanently at over 2000 feet above sea-level before a positive lowering of the surface by melting could take place. Thus, an ice-sheet already in existence may be able for a long time to withstand a climate in which it could never originally have accumulated; and ameliorating changes may attain an advanced state before their effect on such an ice-sheet becomes marked. But when the permanent thawing of the surface once commences, it must go on with accelerated rapidity as lower levels are reached.” Again, a little further on he says: “It is clear that by the time the hill-tops had reappeared the Arctic conditions of climate had passed away, and permanent snow was no longer possible on rock-surfaces, even at the higher levels.” Ifthe Arctic conditions of climate had thus passed away by the time the hill-tops had reappeared, it is likely that the amelioration in the climate continued, and that something more than a mere temporary and short-lived recession of the ice-lobe took place. Mr LamptucH accounts for the deposition of the Upper Boulder Clay by a re-advance of the ice which followed closely again the local depositions of the stratified drift. In view of the great change of climate which is admitted, it seems more probable that at this time also the ice-plateau, though lingering for a considerable time in the Irish Sea basin, went on wasting and shrinking, passing into a condition of “dead ice,” and eventually disappearing entirely in much the same way as Mr Lamewuen thinks the final waning of ice-sheets from the British area to have taken place. Though the mass of ice may not have attained its maximum development at the same time in all parts of the West British region, and shrinkage may have started in the ice covering the Isle of Man sooner than it did further south and west, one cannot 56 DR T. J. JEHU ON THE GLACIAL DEPOSITS OF WESTERN CARNARVONSHIRE. imagine any recession of ice from the land area which was not practically contempor- aneous on all sides of the Irish Sea basin. When the ice-sheet which deposited the Lower Boulder Clay shrank away from the hills and the area now known as Lleyn emerged from its icy covering, the ice probably still lingered for a time in the basin of the Irish Sea. _ It was probably during this time of shrinkage and recession that the sands and gravels lying between the two boulder clays were accumulated. Owing to melting at the edge of the ice-sheet and to the ponding up of the land drainage, much of Western Carnarvonshire must for a time have been submerged under fresh water, and there is no need to assume any sinking of the land relatively to the sea of which there is no clear evidence in this region. The facts which militate against the supposition that these shelly sands and gravels are ordinary marine deposits laid down when the land stood lower relatively to the sea are summarised by Professor KenpaLt in Carvitt Lewis’ Glacial Geology of Great Britain and Ireland (1894), Appendix viii. Stratified Drift deposits—-shell-bearing sands and gravels—are met with lying between two boulder clays at various elevations on both sides of the Irish Sea—in Ireland as well as in West Lancashire, Cheshire and Wales. If these deposits were laid down in their present position during a retreat of the ice-sheet from the hills, it means that a shrinkage took place from the land area on both sides of the Irish Sea as well as about the Isle of Man. The recession of the ice would be accelerated by the action of land streams impinging on to ice, and wasting at the surface would become more rapid as the surface was brought lower. The shrinking mass would thus be attacked on all sides. Granting, therefore, as Mr Lampiucu does, a widespread change of climate to account for the shrinking of the ice-sheet from the Isle of Man, and recognising that a similar shrinkage must have taken place on all sides of the Irish Sea basin, the proba- bility is that the ice-plateau vanished entirely and that an inter-glacial epoch intervened between the times of the deposition of the two boulder clays. The Upper Boulder Clay would thus denote a recurrence of severe glacial conditions and the advance of another ice-sheet. It has already been stated that, as far as Lleyn is concerned, the two boulder clays maght be regarded as the product of one mer de glace subject to considerable oscillation, but a review of the whole of the Irish Sea area renders it more probable that they are the products of the ice-sheets of two glacial epochs separated by an inter-glacial epoch. (The author is indebted to the Carnegie Trust for a Research Grant towards the expenses connected with this work.) Trans. Roy. Soc. Edin. Vou, XV EE. Dr T. J. Janu on “The Glacial Deposits of Western Carnarvonshire.” ‘NAGTT fO dVW SS QNVISI AISAUVE TIIIN HLHOd Yor {59% S.TWMGAL iS’ Hlo4 wouror) YJ40d UMTYAMT UOfYY NIM NOs @ Uopobsx yplod YRC ukayurg Yyulag Se OO Ce: L434 0Or UAynp-uaDp] 7UI0e GNV1 40 LHOISH + £ SFTW 40 31V0S a nahi : , a . i COW p02, Wo a ‘ 4, cas i } . bt denovinwa amine W to sgoeC LatoalO oT“ a0 wual, . “iis j J — f: \< Pr pe ormnary A surnnkarised | Britaie. and aie hills, it means Son As well aa i. ) a* a ihre a2chon of F EoOID 7. AwDierTr? ye — id oy ‘ , a ( at, f ome nore rapid. “es ‘ f > 4 he | . ithe red On all OS) ale (De Srey hae f{ climate a ae oe NX ig . & ; } £ > ackkurnisine that ® on oa ¢ ’ ro : ah P. i ea : Ayit, ‘iM proba 4 ~ 7 _ ) | : p - Y Pe aiid Boch Uvery ened i \ > on \ { ; ta Z e 4 4 7 , cr . * . - \ a4 ; mewt conditions a . \ S J ; , ; ; z Pe te a - i ; te 3 3 } RrABrny fe, iS ai ) + a tact t fam » far as Lleyn = / + . me \ ) i ' - . 3 L& Bt SSrOM | a : a % 3 a a: gy a ‘ ‘ , » ¢ha nevwinet of one Mer de a“ > J y } asi , i» -_ ie ; : = oe ae be 1 Irish Sea arem 4 es Ms / “ e 4 : : : re at LWwo wlaciad |. et ath: bey the - ba ie ' ‘* . ¥ : po oo . 7 a i } j . if é . |. 2 d ‘ GF: Stretch (expanded) . é : 1% TRANS. ROY. SOC. EDIN., VOL. XLVII. PART I. (NO. 3). 8 58 PROFESSOR DAVID HEPBURN ON Hind flipper :— Length on outer edge 5 : : : : 13% ins, =! Fy SUC Te ne 5 ; : 3 , aA ate Stretch (expanded) . : ; : ; : 133 ,, Circumference at base : i : ‘ ; 10} ,, Its weight was 83 lbs. A portion of umbilical cord about 3 inches in length was still dangling from the umbilicus. Its fur was greyish yellow, mottled by darker spots, suggesting leopard characters. Abdominal Viscera.—On opening the abdomen, the general shape of the cavity was seen to be ovoid, being wider in front and narrowing behind to the well-marked inlet of the pelvic cavity. Regarded as a whole, the abdominal cavity presented a close resemblance to that of the porpoise,* of which an account appeared in the 7’ransactions of the Royal Society of Edinburgh, vol. xl., part ii, p. 315. Running forwards from the umbilicus, the umbilical vein, already nearly obliterated, formed a very distinct object, suspended as it was in the free edge of an extensive ventral mesentery constituting the falciform ligament of the liver. Running back- wards from the umbilicus, the urinary bladder, with its associated hypogastric arteries, was likewise suspended in a mesial ventral mesentery whose depth from the abdominal wall to the bladder measured from an inch to an inch and a half. The bladder was long and narrow, and at the umbilicus its lumen was distinct. No part of the bladder had become obliterated to form the urachus, so that the bladder represented the entire intra-abdominal extent of the allantois. The liver and stomach occupied the wide anterior end of the abdominal cavity, and in rear of them only the mass of coiled intestine was visible. PERITONEAL Foups AND REFLECTIONS. At first glance there was no sign of a great omentum, but on separating the coils of intestine from contact with the stomach, the great omentum was found crumpled up against its attachment to the greater curve of the stomach. Its greatest depth from the stomach to its free margin was 3 inches. It was devoid of visible fat, and was there- fore an extremely thin and translucent membrane. There was a well-marked gastro- hepatic or lesser omentum presenting the usual gastric and hepatic connections. The gastro-splenic omentum was also well marked, and by it the spleen was attached to the great curve of the stomach. From the dorsal aspect of the spleen the peritoneal membrane extended to the dorsal wall of the abdominal cavity without coming into relation with the left kidney, which was situated opposite to its fellow and considerably further back in the abdomen. The liver was attached to the sub-diaphragmatic surface by the usual suspensory, coronary and lateral peritoneal ligaments, to which further reference will be made in describing the liver, * HRPBURN and WATERSTON, loc. cit. OBSERVATIONS ON THE ANATOMY OF THE WEDDELL SEAL. 59 The duodenum was suspended in a dorsal mesial mesentery whose base of attachment measured nearly 5 inches, and this represented the distance between the pylorus (gastro-duodenal junction) and the duodeno-jejunal junction which was situated close behind (tailwards of) the superior mesenteric vessels and immediately to the left of the mesial plane. ‘To the right side of the duodenal mesentery there was a large peritoneal recess whose right boundary was formed by a tailed lobe of the liver extending back- wards along the dorsal wall in intimate association with the inferior vena cava. At the first glance there appeared to be no naked-eye distinction between the small intestine and the colon. At no point were teenia coli, sacculations, or appendices epiploicz visible. As measured from the duodeno-jejunal junction or flexure, 46 feet of gut were suspended from the dorsal wall in a mesentery practically corresponding to the entrance of the superior mesenteric vessels. A closer examination revealed a distinct lateral diverticulum rather more than 1 inch in length but of the same calibre as the gut, situated on that part of the gut, which was suspended in the mesentery and at a point 9 inches from the hinder end of the mesentery. ‘This diverticulum evidently represented the ccecum and the vermiform appendix in their most primitive form, and in the same condition as I have formerly described in the case of the grey seal (Halicherus grypus).* We may therefore conclude that this diverticulum marks the commencement of the colon. Consequently the first 9 inches of the colon are suspended in the same mesentery as the small intestine, exclusive of the duodenum. The remainder of the gut, 7.e. colon, was suspended in a dorsal mesial mesentery extending tailwards to the entrance of the pelvis, while through the greater part of the pelvic cavity a dorsal mesial mesentery supported the pelvic part of the colon, 7.e. the rectum. The foramen of Winslow was very distinctly defined in relation to the free right border of the gastro-hepatic omentum. Similarly the lesser sac of the peritoneum was equally distinct, although the great omentum was not in any sense a gastro-colic structure. The inlet of the pelvis measured 1 inch in the transverse diameter and 3 inches in the conjugate diameter. Its lateral boundaries were well defined by the hypogastric (umbilical) arteries, each of which was supported in a peritoneal sling or ligament attached to the dorsal wall along the pelvic inlet. The pelvic peritoneal pouch extended backwards between the gut and the bladder to a distance of 34 inches from the pubic crest, and so reached a point posterior to the bulb of the urethra, z.c. to a point which corresponded with the central point of the perineum. On the ventral abdominal wall, 1 inch to the outer side of the hypogastric arteries, there was the opening which led into the inguinal canal (see fig.). Each opening was circular and half an inch in diameter. It led into a circular peritoneal tube which extended through the abdominal wall in relation to the hinder free border of the musculus transversalis abdominis and the musculus obliquus internus abdominis. At * Hueppurn, “The Grey Seal (Halicherus grypus),” Jour. Anat. and Phys., vol. xxx. 60 PROFESSOR DAVID HEPBURN ON the plane of the musculus obliquus externus abdominis, and close to its pubic attach- ment, the peritoneal tube passed through a muscular slit corresponding to the external inguinal ring and thereafter terminated in a blind end situated close to the posterior end of the testis. The length of this tube of peritoneum was 2% inches, and its testicular part formed the tunica vaginalis testis. Each vas deferens entered the abdomen through the internal inguinal ring, being suspended from the ventral wall in a short mesentery or peritoneal sling by which each vas was carried across the abdominal aspect of the hypogastric artery to the pelvic surface of the urinary bladder about the level of the pubic crest. These two mesenteries for the vasa deferentis, formed a free transverse fold of peritoneum on the pelvic surface of the urinary bladder at the level indicated. From this level the vasa deferentia passed backwards on the pelvic surface of the bladder under cover of the peritoneum. On the pubic aspect of the bladder there was a triangular non-peritoneal surface extending for- wards from the pubic crest for 14 inches, at which point the ventral mesentery of the bladder commenced and continued to the umbilicus. ALIMENTARY ORGANS. The stomach (see fig.) presented a single chamber situated with its long diameter in the axis of the trunk. The cesophagus entered the stomach slightly to the right side of the most prominent or anterior part of the fundus. The greater convexity or curve was well defined, and measured 17 inches from the cesophagus to the pylorus. The distance between the same points along the lesser curve was 10 inches. Each of these curves was associated with the usual omenta. A very sharp bend occurred in the lesser curve, 5 inches in a straight line from the cesophagus and 3 inches from the pylorus. The sides of this bend were held in close apposition by the peritoneum, and the general appearance produced was that of a constriction in the course of the cavity of the stomach. The greatest width of the stomach on the cesophageal side of this bend was 54 inches, whereas on the pyloric side of the bend the greatest width was 4 inches, and opposite the bend the width was 25 inches. The practical result of the infolding of the stomach wall was therefore to produce two chambers communicating with each other by an aperture considerably narrower than either of the chambers. A second slight constriction was present in the pyloric section, and thus as a whole, from cesophagus to pylorus, the stomach suggested three imperfectly separated compartments or chambers. The contents of the stomach consisted of a quantity of a thick pasty substance of a somewhat light earthy colour. It was uniformly smooth, and contained no evidence of bones. In all probability it represented partially digested coagulum of milk. The pylorus was placed in the mesial plane, and was recognisable both to the eye and to the touch as a constricted ring. The duodenum extended from the pylorus in the form of a horse-shoe loop 12 to 13 inches long. It was attached dorsally in the mesial plane by a mesentery. The OBSERVATIONS ON THE ANATOMY OF THE WEDDELL SEAL. 61 proximal and distal ends of the loop were from 4 to 5 inches apart. To the right side of the duodenal mesentery there was a large peritoneal recess or pouch the mouth of which was directed anteriorly, and to which reference has already been made. The duodeno-jejunal flexure lay immediately to the left side of the vertebral column and marked the beginning of the mesentery proper and of the small intestine suspended in it. This mesentery had a very short base which practically corresponded to the entrance of the superior mesenteric vessels. It was twisted to the right and supported the jejunum and ileum, which together measured almost 46 feet in length. The ccecal diverticulum formerly referred to as representing the combined ccecum and vermiform appendix, was taken to indicate the termination of the small and the beginning of the large intestine. This primitive developmental form of the coecum and vermiform was rather more than an inch in length, while in calibre it corresponded with the gut. From this diverticulum to the end of the gut there were no other external evidences of any distinction between small and large intestine. From the ccecum the large intestine or colon pursued the first 9 inches of its course suspended in the same mesentery as the small intestine. Thereafter the colon assumed a mesial position and, as far as the pelvic inlet, 7.e. for a distance of 18 inches, it was suspended in a dorsal mesial mesentery. ‘The pelvic portion of the colon was also placed mesially, and the greater part of it was also suspended in a dorsal mesentery. From the pelvic inlet to the anal aperture the gut measured nearly 10 inches, so that the entire length of the colon from ccecum to anus was practically 3 feet. ‘Thus the total length of the gut from pylorus to anus was :— Duodenum : : : : ; : 1 foot. Small intestine . ; ' : : ; 46 feet. Colon é : d a : 5 BP gs Total i : : ; : : 50 ” The liver (see figs.) was large and deeply fissured, thereby presenting very dis- tinct lobes. It was intimately associated with the diaphragm, and occupied the anterior end of the abdominal cavity from side to side. It was provided with the usual peritoneal ligaments. The falciform and coronary ligaments presented no special features as regards their arrangement, but the left lateral ligament extended from the sharp left margin of the liver whereas the right lateral ligament was short and extended from the smooth surface of the right aspect of the right lobe. The diaphragmatic surface of the liver was smooth and convex, adapting itself to the abdominal surface of the diaphragm and presenting right and left lobes in relation to the suspensory or faleiform ligament. The right lobe was divided into mesial and lateral portions by a deep dorso-ventral fissure, and the left lobe was imperfectly divided by dorsal and ventral notches which, however, did not meet each other. On its visceral aspect the liver was much subdivided, particularly in relation to the right lobe. The right and left lobes were marked off from each other by the ligamentum teres (obliterated umbilical vein) on the ventral aspect of the hilum, and by the hepatic 62 PROFESSOR DAVID HEPBURN ON sinus on the dorsal aspect of the hilum. The right lobe presented the same mesial and lateral portions which were noted on the diaphragmatic aspect. The mesial portion was divided into dorsal and ventral parts by the hilum. To the ventral portion the gall-bladder was attached, and this part of the right lobe was connected to the left lobe by a pons hepaticee. The dorsal portion was to a large extent concealed by the large hepatic sinus. The lateral part of the right lobe was also divided into dorsal and ventral segments by the extension of the hilum across its visceral aspect. A tailed lobe extending from the dorsal segment of the mesial portion of the right lobe passed backwards on the ventral surface of the inferior vena cava, which was thus embedded in the liver substance. This tailed lobe expanded so as to conceal a consider- able length of the inferior vena cava, and into this hidden part of the cava there opened directly the hepatic veins from this particular lobe, as also the hepatic veins from the lateral part of the right lobe. The inferior vena cava itself opened into the large hepatic sinus situated close to the diaphragm and extending right and left of the suspensory ligament. This sinus received the hepatic veins from the right and left lobes of the liver, with the exception of those veins already mentioned as opening directly into the inferior vena cava. ‘I'he interior of this sinus was imperfectly divided into right and left parts by a crescentic partition which was situated to the right side of the line of attachment of the suspensory ligament, so that the part of the sinus to the left side of the crescentic fold was the larger. The inferior vena cava opened into the hepatic sinus on the right side of the imperfect partition, while on its left side it received the mouth of the ductus venosus. The hepatic sinus narrowed for the purpose of passing through the diaphragm in order to enter the right auricle of the heart, but the general size of its lumen was so much greater than that of the inferior vena cava that it is more accurate to say that the inferior vena cava opened into the sinus and the sinus joined the auricle. The conditions presented by the hepatic veins afford some interesting light upon the question of their development. Clearly the large hepatic sinus has resulted from the fusion of the two embryonic venze revehentes, although, from the position of the imperfect crescentic partition, it is evident that the left vena revehens was the larger of the two and therefore received the smaller or right vena revehens. This arrangement would therefore appear to provide a variation upon the current statement that “ the left vena revehens loses its connexion with the sinus venosus and opens into the right vena revehens” (Cunninghan’s Text-Book of Anatomy, 2nd ed., p. 935). Again, the ductus venosus is described as passing directly “ from the left vena advehens to the right hepatic vein” (vide cbid.), whereas, in the liver under consideration, the mouth of the ductus venosus opens to the left side of the crescentic fold, which appears to represent the remains of a fusion between the right and left hepatic veins. Further, to quote again from the same authority, ‘‘ The upper part of the inferior vena cava is developed as an outgrowth from the common trunk formed by the fusion of the ductus venosus with the right hepatic vein.” From the present dissection the inferior vena cava would OBSERVATIONS ON THE ANATOMY OF THE WEDDELL SEAL, 63 appear to arise from the right hepatic vein independently of the ductus venosus, more especially in view of the fact that the inferior vena cava receives direct tributaries from the caudate lobe adherent to its ventral aspect. The gall-bladder, which was empty, occupied a fossa on the visceral aspect of the mesial portion of the right lobe of the liver. It thus lay to the right side of the ligamentum teres, from which it was separated by a projecting portion of liver substance, connected to the left lobe by a pons hepatica. This portion of the liver corresponded in general position with the quadrate lobe of the human liver. The cystec duct passed towards the hilum of the liver and, having been joined by hepatic ducts corresponding in number with the three chief lobes of the liver, the common bile-duct was formed. ‘This duct pursued its course on the ventral aspect. of the foramen of Winslow and, immediately after passing this foramen, 7.e. just before reaching the duodenum, it was joined by the pancreatic duct. The duct thus resulting came in contact with the duodenum a little more than half an inch from the pylorus. It perforated the duodenal wall very obliquely, and opened on the summit of a papilla at a distance of 2 inches from the pylorus. The pancreas presented a characteristic appearance. A small part of this gland was found within the duodenal mesentery, but the greater part of the organ extended headwards behind the lesser peritoneal sac. The pancreatic duct emerged from the substance of the organ on the cephalic (anterior) side of the foramen of Winslow (not from that part within the duodenal mesentery), and, extending to the right side, it formed a junction with the common bile-duct on the cephalic side of the foramen of Winslow and fully 2 inches from the common orifice by which both ducts pour their contents into the duodenum. The spleen measured from 9 to 10 inches in length, from 1 to 6 inches in width, being narrow anteriorly and wide posteriorly. It was extremely thin, being not more than a quarter of an inch in thickness. It was situated between the stomach and the dorsal wall, being attached to the former by the gastro-splenic omentum and to the latter by a dorsal mesial mesentery. There was no intimate relation between the spleen and either of the kidneys, because these organs were removed from all immediate association by reason of their situation considerably nearer to the pelvic region. The tail end of the pancreas extended into the posterior splenic mesentery but did not come in contact with the spleen. Trans. Roy. Soc. Edin. Von, XLVII. Prof. Davin Hepsurn on “ Observations on the Anatomy of the Weddell Seal.” DORSAL. AND INF. V CAVA. ~~ SINUS FOR HEP. VS. LOBE ON VENTRAL ASPECT OF INF. V. CAVA (TURNED TO- WARDS DORSUM). i LEFT LOBE LEFT. RIGHT. J f _ / Stomach. LIG. TERES. VENTRAL. Liver of seal. View from abdominal aspect. g a > & ie) 2) z rm o o >) 7) RIGHT LAT. LIG. = * -— URINARY BLADDER. _-HYPOGASTRIC A. INT. ING. RING. ___.- URETER. ~— COLON. LIG. TERES. : Pelvic cavity of seal (seen from pelvic inlet). Liver of seal (seen from ventral aspect). ( 65 ) 1V.—Supplementary Report on the Hydroids of the Scottish National Antarctic Expedition. By James Ritchie, M.A., B.Sc., Natural History Department, The Royal Scottish Museum. Communicated by W. 8. Bruce, LL.D. (MS. received December 8, 1908. Read January 4, 1909. Issued separately May 27, 1909.) Since the report on the Hydroids collected by the Scotia was completed, almost three years ago, I have had an opportunity, thanks to the kindness of Dr W. 8. Brucs, of looking through the enormous mass of material brought together by him during his Antarctic voyages. The search, begun in the hope of finding a few minute species— which, since many are epizoic on other forms of marine life, might readily be overlooked on cursory examination—-has resulted in the addition of no less than 25 forms to the 36 already recorded. Deducting from this total of 61 forms two varieties, it appears that the Scotia collection contains in all 59 distinct species. Few, indeed, of these have been obtained in truly Antarctic latitudes—subantarctic or temperate seas having furnished the greater number—but this result is in accordance with the findings of the majority of the recent Antarctic expeditions.* Dr BirLaRp, in summarising the results of the Belgian, Swedish, and French expeditions, states that the known Hydroid fauna of Antarctic regions comprises only 32 species. But from his list he has, | think un- justly, excluded South Georgian records. His list, moreover, published before the valuable paper by Professor Hickson and Mr Gravety on the Discovery Hydroids was issued, necessarily makes no reference to the large number of forms recorded by them from Victoria Land. The following figures indicate the wealth in species of the Hydroid fauna of Antarctic seas, the Antarctic area being defined as the region lying in ‘‘higher southern latitudes than the extreme limit of floating ice, according to the most recent research” (BRucE, 1894, p. 208); that is, an area corresponding to the Antarctic Circumpolar Subregion of OrTMANN (1906). To this region belong the 14 species collected by the Belgian expedition (Hartiavus, 1904), 24 of the Swedish collection of 50 species (J ADERHOLM, 1905), the 8 species of the French expedition (BiLuarD, 1906, ), and 24 of the 25 English records (Hickson and Gravety, 1907). From the same area the Scotia obtained only 9 species, although some of the most fruitful stations, Burdwood Bank, the Falkland Islands, and Gough Island, lay just beyond its boundary. Extracting the distinct species from those lists, we find that their number amounts to 68. The Hydroid Zoophytes, then, are represented in the Antarctic circumpolar seas, * The report on the Hydroids of the German expedition has not yet been published. TRANS. ROY. SOC. EDIN., VOL. XLVII. PART I. (NO. 4). 9 66 MR JAMES RITCHIE: SUPPLEMENTARY REPORT ON according to our present knowledge, by 68 different species, 21 of which are Gymnoblasts, 47 Calyptoblasts, while of the total, 36 must, so far, be regarded as peculiar to the Antarctic region. As regards the number of individuals as distinct from variety of species, recent results indicate that the area is a thinly populated one, for comparatively few Hydroids were brought up in any one haul of trawl or dredge, a strong contrast to the abundance of tangled masses which occur in such temperate areas as the North Sea. On the whole, the Antarctic Hydroid fauna may be looked upon as a rather meagre one, comprising comparatively few species belonging to well-established genera. From the point of view of geographical distribution the collection to be described is of a miscellaneous character, for Dr Bruce collected material on his outward and homeward voyages as well as at his southern stations. Thus specimens from tropical seas— from the Cape Verde Islands, from Brazil, and from St Helena—are recorded alongside specimens from subantarctic and Antarctic localities, but so little is known regarding the Hydroid fauna of those places that it seemed better to include them here, at the same time furnishing a record of all the Hydroids brought together by the Scotia. Two species, Thyroscyphus tridentatus and Plumularia lagenfera, hitherto recorded only from the South and North Pacific respectively, have been found in the South Atlantic, while the known ranges of several other species have been considerably extended. Of new forms there have been described the trophosome and gonangium of Sertularia heterodonta from off Brazil, the trophosomes of Antenella quadriaurita from Gough Island, and of a variety of Lafoéa gracillima from deep water to the south of the South Orkneys, while the gonangia of Sertularia rathbum and Anten- nopsis scoti# have been described for the first time. Some additions have been made to the existing descriptions of rare species, and noteworthy variations have been recorded in the hope that the realisation of the con- siderable limits within which variation is possible may tend to the better understanding of reliable specific characters. I gladly seize this opportunity of setting right several unfortunate errors which occurred in the earlier report on the Scotza Hydroids. There should be substituted Hincks for H1nxs, Lamarck for LaAMArK, JOHNSTON for JOHNSTONE, and Haleciwm for Haleciun on p. 523. Aglaophenia dichotoma of the former report I now identify with A. heterodonta, Jiiderholm (imfra, p. 96). In the following list, which takes the place of that in the original report, I have included all the species collected by the Scottish expedition, those discussed in the present paper being indicated by an asterisk :— | THE HYDROIDS OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION. 67 GYMNOBLASTEA. Family PopocoryNIDA. Podocoryne carnea, Sars, 1846. Family MyRiorHELipa. * Myriothela austro-georgie, Jiéiderh., 1904, p. 69. Family HuDENDRID&. * Rudendrium annulatum (1), Norman, 1864, p. 70. Family ATRACTYLIDA. * Perigonimus repens (1), (Wright, 1858), p. 70. CALYPTOBLASTEA. Family HaLEciipsé. Halecium arboreuwm, Allman, 1888. Halecium halecinum (Linn., 1758). (=. robustum, Allman, 1888.) 3 interpolatum, Ritchie, 1907, (1). 7. beanti (Johnston, 1838). - tenellum, Hincks, 1861. Family CaMPANULARIDZ. * Clytia johnstoni (Alder, 1857), p. 71. * Kucopella crenata (?), Hartlaub, 1901, p. 73. Campanularia angulata, Hincks, 1861. Silicularia hemispherica (Allman, 1888). 4 Fe clytioides (Lamx., 1824), p. 71. . Hebella striata, Allman, 1888. x tincta, Hincks, 1861. ms var. plana, Ritchie, 1907, (1). '; sp., Ritchie, 1907, (1). Calycella syringa (Linn., 1758). * Obelia geniculata (Linn., 1758), p. 72. * Campanulina chilensis, Hartlaub, 1905, p. 74. * ,, hyalina, Clarke, 1879, p. 72. * Thyroscyphus tridentatus (Bale, 1893), p. 74. * ,, longissima (Pallas, 1766), p. 72. Family Laro#ip. Lafoéa antarctica, Hartlaub, 1905. Grammaria magellanica, Allman, 1888. » gracillima (Alder, 1857). Brucella armata, Ritchie, 1907, (1). i ;, 5 var. benthophila, n. var., p. 76. Family SERTULARID. Sertularella arborea, Kirchenpauer, 1884. * Sertularia cornicina (M‘Crady, 1859), p. 78. by contorta, Kirchenpauer, 1884. “3 - heterodonta, n. sp., p. 79.- o jiliformis, var. reticulata, Ritchie, us 59 mayert, Nutting, 1904, p. 81. 1907, (1). es i. operculata, Linn., 1758, p. 82. 7 fusiformis (2), Hincks, 1861, p. 77. a is rathbuni, Nutting, 1904, p. 83. 5 gayt (Lamx., 1821), p. 78. * Thuiaria articulata (Pallas, 1766), p. 84. es rectitheca, Ritchie, 1907, (1). (= ,, pectinata, Allman, 1888.) is tenella (Alder, 1857). Synthecium robustum, Nutting, 1904. . tricuspidata (Alder, 1856). Staurotheca reticulata, Ritchie, 1907, (1). 68 MR JAMES RITCHIE: SUPPLEMENTARY REPORT ON Family PLUMULARIDA. * Plumularia curvata, Jiderholm, 1904, p. 86. * Monostxchas quadridens (M‘Crady, 1859), p. 91. =a, magellanica, Hartlaub, 1905.) * Antenella quadriaurita, n. sp., p. 92. . echinulata, Lamarck, 1836, p. 87. * Aglaophenia allmani, Nutting, 1900, p. 93. - lagenifera, var. septifera, Torrey, 1902, * o dubia, Nutting, 1900, p. 95. p. 87. e - heterodonta, Jéderholm, 1904, p. 96. # pinnata (Linn., 1758). (= Gs dichotoma, of first Scotia Report.) ° + setacea (Ellis, 1755), p. 89. ws py minima, Nutting, 1900, p. 97. 3 unilateralis, Ritchie, 1907, (1). a es latecarinata, Allman, 1877, p. 98. Antennularia hartlaubi, Ritchie, 1907, (1). * Halicornaria longicauda, Nutting, 1900, p. 98. * Antennopsis scotix, Ritchie, 1907 (1), p. 90. The localities from which the species recorded in this paper have been obtained are so scattered that, for convenience of reference, I have brought them together in list form. Station 313, 62° 10'S., 41° 20’ W. (S. of South Orkneys). Lafoéa gracillima, var. benthophila, un. var. Scotia Bay, SoutH OrkKNEYs. Myriothela austro-georgix, Jaderholm. Burpwoop Bank, 54° 25’ S., 57° 32’ W. Campanulina chilensis, Hartlaub. Port Stanuey, FaLKuaNnD ISLANDS. Perigonimus repens (2?) (Wright). Plumularia curvata, Jaderholm, GoucH IsLAND. Obelia longissima (Pallas). Thyroscyphus tridentautus (Bale). Antenella quadriaurita, n. sp. Carer Cotony. Houtjes Bay (Saldanha Bay). Plumularia echinwlata, Lamk. Saldanha Bay, entrance to. Eudendrium annulatum (?), Norman. Obelia geniculata (Linn.). Sertularia operculata, Linn. Thuiaria articulata (Pallas). Plumularia lagenifera, var. septifera, Torrey. Antennopsis scotic, Ritchie. Aglaophenia heterodonta, Jiderholm. St HELENA. Sertularella gayi (Lamx.). Sration 81, Abrohlos Bank, Brazil, 18° 24’ S., 37° 58’ W. Sertularia cornicina (M‘Crady). 5 heterodonta, ni. sp. re rathbuni, Nutting. Monostxchas quadridens (M‘Crady). Aglaophenia allman, Nutting. ye dubia, Nutting. 4 minima, Nutting. Halicornaria longicauda, Nutting. THE HYDROIDS OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION. 69 Sr Vincent, Caps VERDE ISLANDS. Eucopella crenata (?), Hartlaub. Sertularella fusiformis (?), Hincks. Tat. 27° 54’ N., long. 33° 17’ W. Clytia johnston (Alder). Sertularia mayert, Nutting. Sration 537, 29° 54’ N., 34° 10’ W. Campanularia clytioides (Lamx.). Obelia hyalina, Clarke. Srarion 538, 32° 11’ N., 34° 10’ W. Plumularia setacea (Ellis). Aglaophenia latecarinata, Allman. Myriothela austro-georgiz, Jaderholm, 1904. Several specimens of this bizarre Hydroid have to be recorded. All came from a single neighbourhood, Scotia Bay in the South Orkneys, but the depths at which the specimens were obtained varied. Some of the examples have already been described by Professor J. ARTHUR THOMSON in a short paper in which he regards them, not without hesitation, as the separated gonostyles of some unknown giant Siphonopore (‘THomson, 1904), There can be no doubt, however, that these specimens are identical with those found by both the Swedish and the French Antarctic expeditions, and recorded by Drs JADERHOLM and BritLarD (1906, p. 4) as Myriothela austro-georgie. The length, the thickened basal portion on which the blastostyles (each bearing its male or female gonophores and a distal tentacle or two) are massed, and, most characteristic of all, the capitate tentacles scattered irregularly over the whole hydranth, even amongst the blastostyles—these features show that our examples belong to the same species as theirs. Nor can there be any doubt that J ADERHOLM was correct in regarding his specimens as belonging to the genus Myriothela, for their resemblance to the northern forms is striking,—solitary hydranths, absence of hydrocaulus, capitate tentacles scattered over the body, blastostyles grouped at the base of the hydranth, the presence of longitudinal folds of endoderm lining the inner cavity. Professor THomson remarks that some of the colonies bore solitary gonophores, while one had as many as seven on its blastostyles, and suggests the possibility of the presence of two species. Since, however, the specimens examined by JADERHOLM had generally from one to three, but sometimes as many as six female gonophores, while the male gonophores occasionally numbered even ten on a single blastostyle, the variation is so great that little stress can be laid on this as a specific character. A water-colour sketch made on the capture of one of the specimens indicates that their colour was a stronger and brighter orange than is shown by JADERHOLM’s figure. (JADERHOLM, 1905, pl. i.). Locality.—Scotia Bay, South Orkneys; dredged in 10 fathoms, April 1903; dredged in 9 to 10 fathoms, May 1903; dredged among mud and pebbles, 18th December 1903. 70 MR JAMES RITCHIE: SUPPLEMENTARY REPORT ON One specimen was found “on the surface of the water, in a hole which had been cut in the ice. The depth of the water at that place was 20 to 30 fathoms; the temperature was 29° F.” M. austro-georgie has previously been recorded from Cumberland, South Georgia, (JADERHOLM), and from Flanders Bay and Booth-Wandel [sland (Binuarp). Eudendrium annulatum (¢), Norman, 1864. Two small clumps of bushy colonies appear to belong to this species, but the weathering of our specimens, and the indefiniteness of the characters which differentiate the species of Hudendrium, render certainty impossible. The colonies are 5 cm. high, and agree with Canon Norman’s species in being bushy and beset with very numerous branchlets ; in possessing thick, rugged stems, on the surface of which, near the base, the fascicular tubes are more or less contorted ; in having branches closely covered with strongly marked rings; and in bearing hydranths with about from 16 to 18 tentacles. On the other hand the gonophores, all of which are female, are borne on tentacle- bearing hydranths and not on atrophied individuals. So many, however, are the gonophores and so closely are they packed around the hydranth, that in not a few cases it was difficult to distinguish the presence of tentacles. Since, in some species at least, the loss of the tentacles is a degenerative change keeping pace with advancing maturity, their presence in this case may be of less significance than at first one tends to regard it. These measurements were made :—The diameter of the unfascicled branches and branchlets is almost constant, about 0°18 mm. The hydranths are about twice as long as broad, the breadth being measured at the level of the bases of the tentacles (0°57 mm. long, 0°28 mm. broad). Locality.—Entrance to Saldanha Bay, Cape Colony. Depth, 25 fathoms. 21st May 1904. Eudendrium annulatum is a North Atlantic form which has been recorded from Shetland (Norman, 1864); Jan Mayen (MarkTANNER-TURNERETSCHER, 1890) ; Pas-de- Calais (BrreNcouRT, 1899); Norwegian Coast (BonNEVIE, 1899). Perigonimus repens (2?) (Wright, 1858). Scanty material which I have, not without doubt, referred to the above species was collected on the shore at Port Stanley. The stems, with a diameter of from 0°04 to 0°05 mm., arise from a stolon creeping upon an encrusting Polyzoon, and reach a height of 10 mm. ‘They bear a considerable number of branches which leave the stem at a sharp angle and carry secondary, and these sometimes twigs of tertiary degree, in such a way as to give the colonies the appearance of being dichotomously branched. The offshoots can always be distinguished, however, by the presence of a slight constriction at their bases. The stems bear distinct rings at their bases and here and there through- THE HYDROIDS OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION. 71 out their course, while the intermediate portions are more or less corrugated. A delicate chitinous envelope surrounds the ccenosare and is adorned with minute sand particles, fragments of sponge spicules, etc., this coating being continued over the lower part of the hydranth. The hydranths, which are in poor condition, appear to be rather globular in shape and have from 12 to 15 tentacles. Short-stalked gonophores occur scattered over the hydrocaulus, but they are far from mature and offer no characters of significance, I have no hesitation in identifying my specimens with those collected by PaxssLER at Port Stanley in 1895 and described by Harriavs (1905, p. 530), although our examples bear more numerous branches than his “gar nicht oder nur schwach verzweigten Hydrocauli.” And I follow Harriavs, but with considerable hesitation, in referring the colonies to the Perigonimus repens of Wricut, an almost unbranched form with a maximum height of “+ inch” (Hincks, 1868, p. 90), contenting myself merely with adding a second mark of interrogation to that which expresses HarTLavs’s doubt. Locality.—Growing on an encrusting Polyzoon, from seaweed found on the shore at Port Stanley, Falkland Islands. January 1903. Clytia johnston (Alder, 1857). Of this common European species only a few stems occur on Saragassum weed. They resemble miniature British examples of C. johnston, structurally alike in every detail, but altogether on a much smaller scale. They are even less in some measure- ments than the small variety found by Dr Bittarp (1907, p. 168) on material from the Saragassum Sea. Measurements :— Stem, length . : ‘ ‘ ; : 16 -2:1 mm. , diameter . ; ; : ‘ 0°63-0°71 ,, Hydrotheca, length . , : ; ; 0°66-0°7 -,, diameter at margin : , ; 0-41-0°48 ,, Locaiity.—Off Saragassum weed, from lat. 27° 54’ N., long. 33° 17’ W. 28th June 1904. Campanularca clytioides (Lamouroux, 1824). Several specimens of this minute species have been found creeping on Saragassum fronds. The stems are short, measuring from 0°74 mm. to 0°95 mm. in leneth and 0:1 mm. in diameter, and are marked by about eight compact rings at the base of the hydrocaulus and about six less compact rings beneath the hydrotheca. The inter- mediate portion of the stem is smooth or only slightly corrugated. The hydrothece are short (0°34 mm.), rather broad at the mouth (0°38 mm.), and taper rapidly to the base. Their walls are thick, but vary considerably in different individuals and even in the various parts of the same individual. At the margin of the shelf which divides the 72 MR JAMES RITCHIE: SUPPLEMENTARY REPORT ON cavity of the hydrotheca proper from the small globular cavity at its base is a ring of bright dots, indicating the points at which the base of the polyp was attached to the hydrothecal wall. The gonosome is not present. Locality.—On gulf weed from Station 537. Lat. 29° 54’ N., long. 34° 10’ W. 29th June 1904. Obelia geniculata (Linneeus, 1758). In addition to the Gough Island locality given in the former report, this widely distributed species occurs from another station. Gonophores are present on the colonies from habitat (0). Locality.—(a) Entrance to Saldanha Bay, Cape Colony. Depth, 25 fathoms. 21st May 1904. (b) Growing plentifully, along with Plumularia lagenifera, var. septifera, on the segments and telson of a lobster, Palinostus lalandii, from the same locality. Obelia hyalina, Clarke, 1879. Many small, 6-mm. high colonies of this species occur creeping over the fronds of Saragassum weed in association with Campanularia clytioides. The characters are as described and figured by CLarxkeE, and the dimensions of our specimens appear to agree more closely with those of the type than do the corresponding measurements of the large-sized Talisman examples described by Dr A. Brituarp (L90%eps0l70): Measureménts :— Hydrotheca, depth . 3 : : : 0-2 mm. 3 maximum diameter ; : ) Os ae Peduneles, length : : ‘ : : 0°36-0°45 mm. Stem internodes, length 5 : : 5 0°54 mm. No gonangia were present. Locality.—Creeping on gulf weed from Station 537. Lat. 29° 54’ N., long. 34° 10’ W. 29th June 1904. , Obelia longissima (Pallas, 1766). In addition to the colonies already recorded from the South Orkney Islands, a small fragment whose characters agree with those of the above species has been found at Gough Island. Measurements :— Hydrotheca, depth . 3 F i ‘ 0°38-0°45 mm. +r diameter at margin ; 5 3 0°38-0°45 ” Locality.—Off Gough Island. Depth, 25 fathoms. Bottom, rock. 28rd April 1904, THE HYDROIDS OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION. 73 Eucopella crenata (?), Hartlaub, 1901. A very few examples, whose occurrence has already been mentioned (Rircuiz, 1907, p. 488, footnote), were creeping on seaweed obtained on the shore of St Vincent, Cape Verde Islands. The absolute identification of these examples with Professor HarrLavs’s E. crenata is, inthe absence of the gonosome, impossible, and, considering differences which occur in the shape of the hydrotheca and in the ringing of the peduncle, appears to me to be a matter of doubt. Our specimens, however, are identical with those described by Dr A. Brttarp, 1907, and I follow him in assigning them (with a query) to the above species. The hydrotheca are more conical than those of the type, and their proportions differ slightly from those of BILLarp’s specimens, where the depth exceeds the breadth, for here these dimensions are identical, or the latter may even exceed the former. The margin of the hydrotheca is cut into about thirteen rounded teeth, and above the projecting septum at the base there occurs a circle of from 28 to 31 bright dots (indicated in Hartiavup’s fig. 27). The peduncles bear from 9 to 12 sharply defined rings at the base, and from 1 to 4 annulations at the summit, while the median portion is generally more or less corrugated. Measurements :— Length of peduncle. é ; : : 1:01-1°98 mm. Breadth 3 : : ; : ; 0:07-0:09 _,, Depth of hydrotheca . : ; ; ‘ 0:32-0:45 _,, Breadth 5 at margin : i : 0°34-0°40 ,, Locality.—Growing on seaweed from the shore to the N.H. of Porto Grande, St Vincent, Cape Verde Islands. 1st December 1902. Apparent evidences'of regeneration are given by some of the specimens. To mention one example. The hydrotheca is borne on a stem 1°68 mm. in length, but this stem is composed of two distinct portions, a basal section 1°04 mm. long with walls 9« in thickness, of a dark horn colour and deeply stained by eosin. ‘This portion is sharply truncated at the distal end. It is succeeded by a shorter section 0°64 mm. long, with walls only 6 in thickness, clear and transparent and faintly tinged with eosin. It is apparent that the peduncle has been truncated at a particular point, and that from this point the regenerating hydranth has built a new perisarcal protecting sheath which, as one would expect, differs in density from the older portion. It is interesting to note that the regenerated portion has, instead of continuing the smooth character of the old peduncle, assumed the character of a complete peduncle, for it bears four clean-cut rings above its point of origin, these being followed by a corrugated portion, and this by three rings below the hydrotheca. In all the cases examined the regeneration process has reproduced a complete but miniature peduncle. TRANS. ROY. SOC. EDIN., VOL. XLVII. PART I. (NO. 4). 10 74 MR JAMES RITCHIE: SUPPLEMENTARY REPORT ON Campanulina chilensis, Hartlaub, 1905. Three minute colonies epizoic on Halecowm beanw belong to this species. The ringing and branching of the stems and the shape of the hydrothecz agree with the descriptions and figures of Harriavs and of JADERHOLM (1905), but although the former says of the hydrothecz that they are “viel weniger conisch als die von C. repens, Wright” [ ! Auman, 1864], I cannot distinguish between the hydrothece of the South American and British species. C. chilensis is, however, more frequently branched than C. repens, but I doubt whether this and the other rather indefinite characters cited by Hartiavs are constant and of specific value. Scarcity of material of both the South American and British forms, however, prevents a comparison sufficiently extensive to warrant me in setting them down as belonging to the same species. Measurements :— C. chilensis. C. repens.* Stem, diameter : : : 0:06 mm. 0:08 mm. Hydrotheca, length . ' : 0:20-0:21 mm. 0:18-0:28 mm. - diameter at mouth ; 0:09 mm. 0-10 mm. Locality.—Growing upon Haleciwm bean from Station 346, Burdwood Bank. Lat. 54° 25’ S., long. 57° 32’ W. Depth, 56 fathoms. 1st December 1903. JADERHOLM has already recorded C. chilensis from Burdwood Bank, where it was growing on the carapace of a crab at 137-150 metres. Britarp (1906, p. 12), found the species in collections from Flanders Bay and Booth-Wandel Island. Harriavs’s specimens were obtained at Calbuco on Tubularia and Hudendrium. Thyroscyphus tridentatus (Bale, 1893). A few colonies have been found at a single locality. Simple, unbranched stems, 10 mm. in height, arise from a creeping hydrorhiza at intervals of some 2 or 3 mm. To the unaided eye the colonies have an erect, rigid appearance, and are seen to bear pro- minent hydrothez placed alternately on the stem. Under the microscope the stem resolves itself into a series of distinct internodes which vary considerably in size, a much shorter being occasionally wedged in between two longer individuals. That the nodes in our specimen are apparently more distinct than those in the examples recorded by Professor HarrLaus (1901, p. 369) from French Pass, to the north of the South Island of New Zealand, is of little importance, as the boundary mark between internodes is liable to considerable variation. The locality of the node is, moreover, rendered more evident in the Scotia specimens, because the proximal end of each internode is generally marked by a rude annulation. The hydrothece are arranged alternately, and the whole series lies in one plane. A hydrotheca rests upon a short process at the distal end of each internode. The hydrotheca is separated by a distinct boundary line from the internodal process, and occasionally one, two, or even three short joints intervene between them. The distal * Specimens from Plymouth in my collection. THE HYDROIDS OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION. 75 margin of such an interpolated joint is always abrupt, and the perisarc of the existing ealycle is not directly continuous with this margin, but is found to merge with that of the interpolated joint some distance within the joint (see fig. 16). From this it would appear that.the joints are not all contemporaneous, but represent the remains of former eups which, broken off by accident, have been replaced once, twice, or oftener by the regenerative power of the ccenosare. In this case, therefore, they can be of no diagnostic value. Evident knobs of chitin are present on the inner surface of the adcauline wall at the base, and of the abcauline wall at the margin. The former do not seem to be reproduced in regenerated hydrothece. The hydrothec are much more deep than broad, and are characterised by an almost straight abeauline and a strongly convex adcauline contour. They are in most cases considerably longer than the stem internodes. The margin is divided into three pro- Fic. 1.—Thyroscyphus tridentatus. x70. (a) Primary hydrotheca showing k, and ky, knobs of chitin; pp, points on hydrotheca to which the hydranth is moored by ccenosarcal strands. (6) Internode process and regenerated hydrotheca ; k, knob of chitin marking base of primary hydrotheca; w,, wall of original hydrotheca ; w,, wall of first regenerated hydrotheca ; w,, wall of second regenerated hydrotheca. nounced teeth, separated by three deep and graceful bays, and is furnished with a three- flapped operculum. No trace of a gonosome could be found. The following measurements give some indication of the variable proportions of our specimens :— Stem, length . : - ; 4 : Up to 13 mm. Internode, length : : é A 3 0°49*-0'91 mm. a breadth : ; : : : 0:13-0'22 mm. Hydrotheca, length . : 6 : : 0:60-0°63 _,, 4 breadth (maximum) ‘ : : 0:22-0:28 ,, Locality.—Gough Island. Depth, 25 fathoms. Bottom, rock. 28rd April 1904. Distribution.—The species was originally described by Bay from material obtained at Port Phillip, in the south of Australia, as Campanulamia tridentata. Subsequently it has been recorded by Professor Harriavs (1901) from French Pass, north of South Island, New Zealand. ‘These, so far as I am aware, constitute the only records of the species, so that the Scotia specimens from Gough Island, midway between Cape of Good Hope and Cape Horn, extend its known range from the South Pacific to the South Atlantic. * A regenerated internode which succeeded an old truncation of the stem. 76 MR JAMES RITCHIE: SUPPLEMENTARY REPORT ON Lafoéa gracillima (Alder, 1857), var. benthophila,* n. var. This species has to be recorded from an additional locality, to the south of the South Orkney Islands. The two specimens from this locality are, however, of a type quite distinct from the Burdwood Bank examples. They are both small and incomplete. The larger, 18 mm. high, bears two short branches; the smaller, 9 mm. high, is branch- less. Only a trace of fasciculation is exhibited by the latter, but the stem of the former is a typical rhizocaulom. ‘The hydrothece, while they closely resemble those of typical examples of L. gracillima, differ in being placed at more regular intervals on the stem, in preserving a near approach to alternation, in lying towards the stem at a much smaller angle, and in lacking a twist on the hydranthophore. Fig. 2.—Lafoéa gracillima, var. benthophila. Portion of branch with hydrothece. x 80. I do not lay much stress on this last point, however, since untwisted hydranthophores are not unfrequently present in typical colonies of the species. The most striking difference lies in the robustness of the hydrothece, the dimensions being much greater than in any other specimen I have examined. Near the base of the hydrotheca proper is a rude row of minute, refringent prominences on the internal wall, to which the base of the polyp was presumably attached. Occasionally, but only where the margin is reduplicated, two rows of dots occur. The following table indicates in millimetres the differences, in respect of size, between the present and typical specimens :— | Coat’s Land Burdwood Bank North Sea Specimen. Specimen. Specimen. | a! Hydrotheea, including hydranthophore : ; 0°87-1:01 0°63-0°64 0°57-0°76 ‘4 diameter at mouth ; F : 0-21-0:25 0°11-0°13 0-11 Diameter of a simple tube. P : ; 0:16 all 0:10 | * BévOos, the deeps ; and ¢iaos, loving. THE HYDROIDS OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION. 77 No gonangia were present. Locality—Station 313. Lat. 62° 10’S., long. 41° 20’ W. Depth, 1775 fathoms. Bottom deposit, blue mud and boulders. 18th March 1903. The distribution of the species is almost world-wide, but it has not hitherto been recorded from Antarctic Seas. Sertularella fusiformis (2) Hincks, 1861. A few minute simple stems, 7 mm. high, rising from a stolon creeping upon a sea- weed. The slightly geniculate stems are divided into internodes which vary consider- ably in length, those nearer the base being longer than the more distal, the former 1°08 mm. as against an average of 0°61 mm. for the latter. The internodes are narrow, only Fic. 3.—Sertularella fusiformis (?) Portion of stem, x 40. 0°06 mm. in diameter at the base, but they widen upwards until a ledge is formed upon which the base of the hydrotheca rests. They are separated by slanting nodes, immedi- ately above which occur one or two rings, while the remainder of the internode is more or less definitely wrinkled. The hydrothecz lie in the same plane, are alternate, and are placed one on the distal end of each internode. Rather more than half of each is free, the free portion leaning wellaway from the stem. In shape they are flask-like, bulging proximally, while towards the distal end there is formed by a sudden constriction a distinct “neck,” which is sur- mounted by the four-toothed margin. The walls of the hydrothece are smooth externally, but just within the margin the inner surface bears four distinct blunt teeth which lie midway between the marginal teeth. 78 MR JAMES RITCHIE: SUPPLEMENTARY REPORT ON Dimensions of hydrotheca :— Length ‘ ; : E : 0°45-0°52 mm. Diameter where it becomes free from internode ; 0:25-0:27 ,, Diameter of “neck” . 5 ; ; ‘ 0'14-0°16 No gonangia were present. It is with some doubt that these specimens, their gonangia lacking, have been referred to the S. fusiformis of Hincxs. The hydrothecze in our specimens appear to be more robust, to have a more decided ‘“‘ neck,” and a more pronounced inclination away from the stem, while the presence of internal teeth is not mentioned in H1ncxs’s description. The present specimens closely approach the very doubtful S. fusiformis (?2) described by Professor Ct. Hartriaus (1900, pl. 5, fig. 9) from Rovigno on the Adriatic Sea, but the ringing which is absent in his is markedly present in our examples. Locality Growing on seaweed found on the shore to the N.E. of Porto Grande, St Vincent, Cape Verde Islands. 1st December 1902. Sertulurella gayt (Lamouroux, 1821). In addition to the colony found near Gough Island, another of almost equal dimen- sions, 12 cm. high, has to be recorded from St Helena. Its minute structure is very similar to that of the Gough Island example, the rugosities on the upper surface of the hydrothecz being in some cases almost obsolete. This species, although widely distributed in the Northern Atlantic and in the Mediterranean Seas, has hitherto been recorded south of the equator only from the Cape of Good Hope (Algoa Bay), (HartLaus, 1905, p. 613). In conjunction with this occur- rence the two Scotia records, from Gough Island and St Helena, may be taken to indicate a wide distribution for the species in the South as well as in the North Atlantic. Locality.—Intertwined with an Aleyonarian, Amphilaphis regularis, from St Helena. 30th May 1904. Sertularva cormcina (M‘Crady, 1859). Scanty material, a mere half-dozen colonies, represent this species. The stems arise from a stolon creeping upon the surface of the fragment of Codiwm also invested by S. rathbum and S. heterodonta, from the former of which, indeed, they are almost indistinguishable to the unaided eye. The largest is only 6 mm. in height. The specific characters agree with those given by Nurrine (1904, p. 58), but the following variations were noted. The stem internodes are proportionally longer than in Nurtine’s specimens, for while in his examples the “height of the hydrothece is usually about equal to that portion of an internode which lies between the hydrothecal base and the node below,” in ours the latter distance considerably exceeds the former. Our hydrothecee, again, have a longer distal portion free from the stem, the result being that the proportion of the anterior adnate part to the whole length of the hydrotheca is THE HYDROIDS OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION. 79 reduced from “about two-thirds” to about one-half. As seen from the anterior aspect the hydrotheca appears to be covered in by only two opercular flaps, the free margins of which run from the tip of one lateral tooth to that of the other; but when a lateral view of the colony is obtained, so that one can look directly on the operculum, another line is seen running from the mid point of the horizontal line connecting the lateral teeth to the median superior point of the aperture. This line presumably marks the margin of two contiguous flaps of the operculum, and it would therefore seem that three, and not two, are present. The determination, however, is one of considerable difficulty owing to the delicacy of the operculum. ‘The downward projecting processes from the base of the hydrotheca are long and evident. Measurements :— Internodes, length : : ; 5 : 0°63-0'78 mm. Me breadth . ; 3 5 : 0:05-0:06 _,, Hydrotheca, length of contiguous portion ; : 0:22-0:25 _,, sp 5 free portion* . 5 ; 0°21-0°22 ,, 5 diameter near base : : : 0:08-0:09 _,, ‘. at aperture : ; : 0:08-0:09 _,, Locality.—Growing upon seaweed (Codiwm, sp.) from Station 81, Abrohlos Bank, Brazil. Lat. 18° 24’ S., long. 37° 58’ W. 36 fathoms. Bottom deposit, coral. 20th December 1902. Sertularia heterodonta,*t n. sp. Growing upon the surface of a seaweed (Codiwm, sp.) in company with S. rathbuni and S. cornicina are numerous colonies of this more minute form. The simple unbranched stems arise from a creeping stolon and cover the weed as with a coat of delicate hairs. At first glance the colonies are hard to distinguish from those of S. rathbum, but on close examination their minuteness and delicacy render them distinct even to the unaided eye. The largest are only 6 mm. in height, 3°5 to 4 mm. being much more common. The hydrorhizal tube has a diameter a little greater than that of the stem, and is strengthened in places by chitinous processes projecting down- wards from its roof or upwards from its floor. The stems are divided into fairly regular internodes separated by distinct nodes. At the base two oblique nodes occur in succession, cutting off between them a short lozenge-shaped athecate internode. The remainder of the internodes are long and slender, narrower at the proximal end and very gradually increasing in diameter upwards, until by a sudden dilation at the top they form a broad bracket upon which the hydrothece rest. The thecate internodes are separated by straight nodes, but occasionally an oblique node occurs in addition, a small athecate internode being in such a case intercalated between the longer thecate individuals. * The “free portion” is measured from the line of the stem to the tip of the lateral teeth. + €repos, other ; and éy7os, a tooth, indicating the presence of other than the usual marginal teeth. 80 MR JAMES RITCHIE: SUPPLEMENTARY REPORT ON An opposite pair of hydrothecz rests on each internode, six to seven or even ten pairs being found on one colony. ‘The hydrothece are set slightly on front of the stem, are always contingent in front for rather more than one-third of their total length, but remain separate behind. The free portion diverges abruptly at a wide angle. Viewed from the anterior aspect the sides of the adherent portion of a hydrotheca are parallel to the long axis of the stem, while those of the free portion converge towards the aperture. The contour lines, especially in the younger hydrothece, are remarkably straight and graceful. Beneath the innermost angle of the base there is a minute chitinous thickening, and another projects from the base into the interior of the hydrotheca. The latter appears to be roughly triangular in shape, the apex pointing a, Fic. 4.—Sertularia heterodonta. (a) Anterior aspect of stem. x60. (6) Lateral aspect of stem showing three-flapped operculum. x60. (c) Hydrothece with reduplicated margins, one regenerated margin bearing internal teeth. x 100. (d) Gonotheca. x 50. into the cup, the side towards the centre of the stem being thickened and concave, that remote from the centre sloping gradually till it merges with the hydrothecal floor. This process represents a small ridge bounding the posterior edge of the opening between hydrotheca and stem. ‘The margin of the hydrotheca is divided into three distinct and sharp teeth, the lateral pair the more prominent, the median and superior individual tilted slightly upwards. There are three opercular flaps, difficult to distinguish except when one is looking directly into the aperture of the hydrotheca. Just within the margin and projecting from the inner wall are three prominent internal teeth, triangular in shape, with blunt apices tilted towards the aperture. These alternate with the marginal teeth. In hydrothecee towards the base of the stem the superior pair appears occasionally to be undeveloped. Frequently the free portion of the tube is much elongated by the reduplication of the margin, and in some cases a new series of internal teeth is produced on the added portion (fig. 4, c). THE HYDROIDS OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION. 81 The gonangium, of which I have been able to find only one example, is borne on the front of the stem immediately beneath a hydrotheca. It is oval in shape, tapering below to form a distinct stalk, and truncated above by a wide aperture. This is bounded by an insignificant thickened neck punctuated by small refringent chitinous swellings on the inner surface. The walls of the gonangium are smooth. Measurements :— Internodes, length 5 : : . : 0°46-0°62 mm. i diameter near base : : : 0:03-0:04 ,, Hydrotheca, length of contiguous portion 3 5 0-10-0'14 ,, s “3 free portion* . : : 0:14-0°15_,, 2 diameter at base . : : : 0:06-0:07 _,, Py - » aperture ; ; : 0:04 mm. Gonangium, length .. ; ; 5 : O74 ,, maximum diameter : : : 0-41 ,, ” One colony shows a pair of hydrothece at an early stage of development, ere yet the aperture, or the base, or the internal teeth had been formed. The stolon has so regulated itself to the papillated surface of the Codiwm upon which it is growing that the outline of its under surface is made up of a series of crescentic bays. The three-toothed margin, the three-flapped operculum, the prominent internal teeth, the presence of a projection from the base into the interior of the hydrotheca, together with the straight contours of the hydrothecz and the length of the internodes, are points which distinguish this species. It has affinities with S. linealis, Warren, 1908. Locality.—Growing upon seaweed from Station 81, Abrohlos Bank, Brazil. Lat. 18° 24’ §., long. 37° 58’ W. Depth, 36 fathoms. Bottom deposit, coral. 20th December 1902. Sertularia mayerr, Nutting, 1904. Small colonies, 5 mm. high—less than half the height of the type specimens—occur creeping on Saragassum weed. Their characters agree with Nurrine’s description and figures, and are identical with those of specimens collected by Mr CrossLanp at the Cape Verde Islands (Rircuiz, 1907, p. 505), although, owing to their smaller size and the consequent restriction of the number of hydrotheca-pairs (in the largest specimen mounted for microscopical examination there are only six pairs), the variation between the proximal and distal pairs is not so marked as in the larger Cape Verde specimens. Even here, however, the distinction between the distal long, narrow, closely forked, gradually separating pairs, and the proximal short, dumpy, wide-spreading pairs is sufficiently distinct and characteristic. It is indicated by the measurements below. In the majority of the calycles two lateral teeth and a smaller median and superior tooth, which succeeds an indentation in the wall of the hydrotheca and curves slightly upwards, are very evident. * From the line of the stem to the tip of the lateral teeth. TRANS. ROY. SOC. EDIN., VOL. XLVII. PART TI. (NO. 4). 11 82 MR JAMES RITCHIE: SUPPLEMENTARY REPORT ON Measurements (@ and b are distinct colonies) :— Basal. Distal. b b a, . a. . “42; 0°49 mm. | 0°56; 0°66 mm. Length of internode 0-4 ss hydrotheca . : : | O29: -0°30) >, 0°38; 0°45 ,, _ Breadth of hydrotheca-pair from tip to tip : | 0-48; 0-46 ,, 0°56; 0°52 ,, 4. is 5, at base + | 0:22 0:25). OIE 02 0ms Locality.—Creeping on gulf weed, Jat. 27° 54’ N., long. 83° 17’ W. 28th June 1904. Sertularia operculata, Linneeus, 1758. Several small fragments of this species occur intertwined with other Hydroids from the same locality. The specimens are typical in branching and in minute structure, but a considerable amount of variation occurs not only in the length of the two hydrothecal teeth, but also in their prominence relative to each other (cf. HaRTLavs, 1905, pp. 665, 666). Of the specimens figured by Professor HarrLavusB our examples most nearly approach those from West Patagonia collected by F. P. Moreno; but from those they differ in the proximity of the hydrothece, for the teeth of one may reach the level of the base of its successor. The present examples are also characterised by the exceedingly minute portion of the distal extremity of the hydrotheca, which is free, the proximal side of the aperture lying almost against the internode. The points above refer specially to the younger branches. On the older portions the hydrothecz are only sub-opposite, their length is less relatively to that of the internode, while a slightly longer distal portion is free. A few typical gonangia occur on the branches. They exhibit a tendency to asymmetry, the aperture lying towards the outer side of the axis of symmetry. The following measurements indicate the relations of the various parts :— Length of branch internode. ; ‘ : 0:33-0:50 mm. os stem internode ‘ : 4 : 0°54-0°61 ,, _ hydrotheca . ; : ; * 0:20-0:25 ,, ; teeth : , , ; : 0:06-0'12 ,, Distance from tip to tip of a hydrotheca-pair . : 0:43-0°58 _,, Locality.—Dredged at the entrance to Saldanha Bay, Cape Colony, in 25 fathoms. 21st May 1904. The distribution of this species is world-wide. In addition to its European localities it has been recorded from the coasts of North and South America, of Southern Asia, of Australia and New Zealand, and of Africa, although the records from the last continent are few. The African localities other than the Scotia record are as follows :— South Africa (Busk, 1850); Cape of Good Hope (Hugenie Expedition, JADERHOLM, 1903); Port Natal (Professor J. A. WAHLBERG, J ADERHOLM, 1903); Mauritanian Coast (BittarD, 1906); North-West of Cape Blane, Soudan (Brtnarp, 1906). THE HYDROIDS OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION. 83 Sertularva rathbum, Nutting, 1904. The specimens occur plentifully on a seaweed (Codiwm, sp.), standing erect and rigid on its surface, so that it appears as if covered with a sparse coating of delicate hairs. They are simple, altogether without branches, herein differing, but immaterially, from the specimens described by ALLMAN (1877), VERSLUYs (1899), and Nurrine, some of which bore pinnules, and they attain a height of only 10 mm. The structures of the stem are as previously described. The hydrothece are in opposite pairs, the individuals of which, in the distal part of the colony, are contingent for almost half their height, but the line of contact gradually decreases until in the proximal pair the individuals Fig, 5.—Sertularia rathbuni. (a) Anterior aspect of stem. 50. (b) Three-quarters view of hydrotheca showing three-flapped operculum. x60. (c)Gonangium. x 70. may be quite apart. Behind the stem the hydrothece are always separate. Beneath each hydrotheca, at the corner where base and inner wall meet, are two chitinous processes which project downwards and lie alongside the wall of the internode. These processes are more distinct in the older hydrothece. The margin of a hydrotheca is eut into three teeth, the lateral pair longer than the single superior tooth. There are three opercular flaps. Gonosome.—The gonangia, which have not previously been described, occur on many of the colonies. They are borne on the stem, from one to three in number, and arise immediately beneath hydrothecze towards the base of the colony. In the speci- mens which I have examined only one gonangium is apportioned to each hydrotheca- pair. ‘The gonangia are broadly ovate, marked with about six rather indefinite annula- tions. Proximally they taper into a short stalk, while distally they contract into a short wide neck, ornamented at its base by a circlet of bright spots, thickenings of the 84 MR JAMES RITCHIE: SUPPLEMENTARY REPORT ON erisare. ne contents of the gonangia were frequently missing, and in n Tl tents of the g f tly ge, and © case were they in a state fit for minute examination. Measurements :— Internodes, length : . : : 0°81-0°94 mm. 3 breadth . , : ? ; 0:10-0:12 ,, Hydrotheca, portion free : ; : : 0:18-0°20 ,, e 5, fixed 3 : ; ; 0:27-0:28 ,, o diameter at mouth : : : 0:74 mm. Gonangium, length . : ‘ 3 : 0:60-0°63 _,, maximum breadth : ; ; 0-45-0°53,, ” In general build, in possessing chitinous projections from the base of the hydrotheca, and in the structure of its gonangium, this species bears close resemblance to S. cormcina (M‘Crady) as described by Nurrine. The latter species, however, is to be distin- guished by the tubular shape of its hydrothecze, by the number of the marginal teeth and of the opercular flaps, by the narrowly oval outline of its gonangium, and by the fact that the gonangia are borne on hydrorhizal tubes at the base of the colony, and not on the stem. Locality.—Growing on a Plumularian, Halicornaria longicauda, and on seaweed (Codium, sp.), both from Station 81, Abrohlos Bank, Brazil. Lat. 18° 14’ 8., long. 37° 58’ W. Depth, 40-50 fathoms. Bottom deposit, coral. 20th December 1902. Previously recorded only from the Gulf of Mexico: ALLMAN, 1877; VERSLUYS, 1899 (Dry Tortugas); Nurrine, 1904, lat. 29° 28’ N., long. 87° 56’ W. Thuwaria articulata (Pallas, 1766), (= 7. pectinata, Allman, 1888). In the earlier report, a colony of this species was recorded under ALLMAN’s name of T. pectinata. The occurrence of an additional colony with gonangia reopened the ques- tion of nomenclature, and an examination of ALLMAN’s type was made (through the kind- ness of Mr R. Kirkpatrick, of the British Museum). The examination assures me of the identity of 7. pectinata, Allman, with 7. articulata, Pallas. The larger of our two colonies was some 6 cm. high, the height of ALLMAN’s speci- mens being also “ between two and three inches.” The colonies are simply pinnate, with a monosiphonic stem about 1 mm. in diameter at the base. The stem is partitioned into regular and well-marked internodes, each bearing three pairs of opposite hydrothece, and from between the proximal and median pairs arises a pair of opposite pinne. Proximal to the first pinna-bearing internode a few destitute of offshoots occur, and on these the number of hydrothece is not constant. Apart from these only one internode has been observed in which three pairs of hydrothece have not occurred; and it is clearly abnormal, for it lacks pinne, has but one pair of hydrothece, and is so short that the hydrothece project beyond it, free for nearly half their height. The pinne stand out from the stem at a wide angle, and they too are divided into distinct but less regular internodes, each bearing from two to five pairs of sub-opposite hydrothece. THE HYDROIDS OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION. 85 (MARKTANNER-TURNERETSCHER (1890) gives the variation as from three to ten pairs per internode. ) The hydrothecz are almost wholly immersed, and the “‘ free membranaceous extension of the wall,” too prominently figured in ALLMAN’s account, has been destroyed (as indeed it was in the 7. pectinata specimen examined), leaving a jagged edge level with the general outline of the pinna. There is evident on the abcauline wall of the hydrothece, just within the opening, a small knob of chitin, and immediately above or upon this rests the Fic. 6.—Thuiaria articulata. (a) Single internode of stem with proximal internodes of pinne. x20, (6) Gonangium. x12. (c) Internode from 7, pectinata with hydrothece slightly apart. x20. (d) Internode from 7. pectinata with congested hydrothece. x 20, base of the one-flapped operculum. On the stem internodes and on the younger pinne the hydrothecz are slightly apart from each other, but in the older pinne they are compressed and lie closely packed, the distal end of one forced against the base of its successor (cf. fig. 6, c and d). The gonangia are clustered on one face of the stem and on the corresponding faces of the pinne. They arise immediately beneath a hydrotheca, are elongate oval in shape, with a wide, circular, distal opening bordered by a distinct neck, and with a tapering proximal end. ‘Their distal half bears more or less indefinite annular rugosities. 86 MR JAMES RITCHIE: SUPPLEMENTARY REPORT ON Measurements :— Scotia Allman’s Specimens, T. pectinata, Stem internodes, length 2°25 mm 2°75 mm Hydrothece on stem, length 0:50 _,, 0°52 : - breadth : OT” 5 0:20 _~—,, Fi on pinna, length : : 0-45, O48 ;, - i breadth 3 Ohh: ,, Ors = is, Gonangia, length ‘ 3 sy ) 5 greatest diameter. 1) 5p f pop mes ALLMAN’ specimens are somewhat more robust than those collected by the Scotza. Locality.—Dredged at the entrance to Saldanha Bay, Cape Colony, at a depth of 25 fathoms. 21st May 1904. The species has been recorded from Algoa Bay and Cape of Good Hope (KircHEN- PAUER, 1884); South Africa (Busk, 1850); Simon’s Bay, Cape of Good Hope (ALLMan, 1888). Plumularia curvata, Jiderholm, 1904 (=P. magellanica, Hartlaub, 1905). Dr JApERHOLM has kindly drawn my attention to the fact that the species recorded in my earlier report on the Scotza collections as P. magellanca, Hartlaub, had been pre- viously described by him as P. curvata, which name I therefore substitute for Harr- LAUB’S synonym. A variation which does not seem to have been present in the examples examined by JADERHOLM or HarTLavus was described and figured in the earlier report (pl. i, figs. 1, la); here I wish simply to add that that variation is more general in the hydroclades than I had at first supposed. My previous description reads :—‘‘In the proximal hydroclades . . . . two processes arise below and at the opposite sides of the first hydrotheca, each of which bears a thecate internode, so that after the first hydro- theca the hydroclade possesses two diverging branches each similar to the simple distal hydroclades” (1907, p. 541). But this duplication of the hydroclade occurs not only at the first hydrotheca but sometimes at successive hydrothece as well. From beneath the first hydrotheca two diverging internodes spring, each capped by its hydrotheca ; from the bases of each of those second pairs arise, and from these again, and so on in a system comparable to the false dichotomy of the Mistletoe, until dichotomous pairs of even the fourth degree may be reached. Some of the hydroclades thus assume a complicated and much-branched appearance, quite distinct from the simple type figured by JADERHOLM (1905, pl. 14, fig. 10) and Harriavs (1905, p. 684, fig. N°). Measurenients :— Stem internode, length ; . : : 0:42-0:52 mm. 93 % breadth ; i , ‘ 011-0°13 ,, Hydroclade internode, length . - ‘ 5 ; 0:24-0:28 _,, Hydrotheca, depth ; ; 3 ; ‘ 0:10 PP 3 diameter at mouth , ; ‘ O1520: i THE HYDROIDS OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION. 87 Locality.—‘‘ Growing on a sponge, Port Stanley, Falkland Islands. 3rd February 1904.” Previous records are from Port Louis and Port Albemarle, Falkland Islands (J ADER- HOLM, 1905); Southern Tierra del Fuego, and Island Picton in the neighbouring archipelago (HarTLavs, 1905). Plumularia echinulata, Lamarck, 1836. In addition to a previously mentioned occurrence at Cape Town, a second locality, also in Cape Colony, has to be recorded, namely, Saldanha Bay. The specimens from this place, while rather smaller in size than the Cape Town examples, are similar in minute structure, except that in the former the hydrotheca-bearing internodes are rather shorter, and the hydrothece therefore more congested, than in the latter. The gonangia are elongate oval, considerably longer in proportion to their diameter than those figured by Hincks (1868, pl. lxv., fig. 2b), and possessing shorter and more regularly arranged spines. They stand out from the median aspect of the stem in a densely packed row. Locality.—Shore, Houtjes Bay, Saldanha Bay, Cape Colony. 19th May 1904. Plumularia lagenfera, var. septifera, Torrey, 1902. Whereas typical specimens of P. lagenifera are about three inches long, are some- times branched, and grow in flexuous clumps, the specimens which I have referred to ToRREY’s variety are short (only 7 mm. high), never branched, and are markedly rigid in habit. The detailed structure is that of a compressed P. lagenifera, where the inter- nodes have become shorter and comparatively stouter, while the internal septa have become more distinct. In conjunction with the general shortening it has come about that in the intermediate internodes there is generally but one septum, although our specimens differ from those described by TorrEy—where “ no intermediate internode has more than one septal ridge”—in that, in several, there are traces of a second ridge on the distal side of the nematophore, while in at least one case the second ridge is quite pronounced. Torrey is equally emphatic that “there is never more than one internode between thecate internodes,” but I have observed a case in which two successive athecate internodes occurred, the distal being much the shorter and lacking anematophore. There was no evidence that this duplication was due to abnormal growth, such as regeneration. These variations, however, only show more clearly the relationship between this form and P. lagenifera type, and confirm Torrey’s placing of it as a variety of that species. In one point the Scotia specimens differ both from the type and from the variety, for they show no trace of a nematophore on any internode “on side opposite branch [v.e. hydroclade] and immediately distal to the proximal septum.” 88 MR JAMES RITCHIE: SUPPLEMENTARY REPORT ON Between the hydroclade and the internode process on which it is set occur from one to three athecate internodes. The hydrorhizal tubes are close-set, are compressed from above downwards, and are supported by thickenings of the perisare which project into the interior of the tube. Somewhat similar thickenings I have already seen in the hydrorhiza of a species of Podocoryne (Rircute, 1907, p. 499) which was growing on a minute shell. In both cases it is possible to imagine that the thickenings may be in some way correlated with the peculiar substratum upon which the specimens are growing, for either on a small, readily tossed shell, or on the ever-moving appendages of a Crustacean, hydrorhizal tubes would be submitted to a great amount of buffeting and rough usage. Reaction to such abnormal external factors might result in abnormal a. Fie. 7.—Plumularia lagenifera, var. septifera. (a) Portion of stem and hydroclade. 100. (0) Base of stem arising from complicated hydrorhizal growth with scattered nematothece. strengthening of the walls. Should such a supposition be well founded, the thickenings in the hydrorhizal tubes could have little systematic value. Frequent nematophores, similar in structure to those on the remainder of the colony, arise from the hydrorhizal tubes at irregular intervals, but always near their borders. The differences between this form and P. lagenzfera in size, in habit, in the intensity of the internal septa, in the absence of a nematophore on each stem internode, and in possessing chitinous thickenings in the hydrorhiza, I do not consider sufficient to warrant the formation of a new species. Measurements :— Stem internode, length ; ; ‘ 5 0:26 mm. 3 breadth : ; : : Olds | Hydroclade thecate internodes, length ; : 0:24 ,, - athecate . 9 : 3 0:05 ,, Hydrotheca, depth . 5 ‘ ; : 0:066-—0:090 mm. Pe diameter at margin : : . 0:105-0'120 ,, THE HYDROIDS OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION. 89 Locality.—A few colonies growing on the telson and under-parts of the body of Palinostus lalandvi (Lamk.) from Saldanha Bay, Cape Colony. 21st May 1904. Distribution.—Plumularia lagenifera, Allman, has been recorded from various locali- ties off the coast of California by MaRKTANNER-TURNERETSCHER (1890, p. 255), Nurrine (1900, p. 65), Torrey (1902, p. 77); from the neighbourhood of Vancouver Island by ALLMAN (1885, p. 157), Nurrine (/.c.); and from the coast of Alaska by Torrey (/.c.). The variety septifera has been recorded by Torrey only from Cataline Island, California. The general distribution of the species and its variety is thus along the shores of the Northern Pacific from California northwards to Alaska. That the present specimens should have occurred on the eastern margin of the South Atlantic is indeed remarkable ; but, in so mobile a group as the Hydroids, wide distribution is of little significance, and the above record but adds another to the long list of species which spread beyond the bounds of any one ocean. Plumularia setacea (Ellis, 1755). Specimens of an exceedingly minute and delicate variety of this species occur creeping on gulf weed in company with Aglaophena latecarmata. They are only 7 or 8 mm. high, and are unbranched, although in one case an appearance of bifurcation at the base (a phenomenon recorded by BrtuarD in this species (1907, p. 210)) was given by a second colony being fixed to the first by its hydrorhiza. The hydroclade internodes, both thecate and athecate, are long and slender and contain two distinct septa, a distal and a proximal. While the rule is that a single athecate internode separates two thecate internodes, very rarely two intermediate internodes occur, in which case one or the other bears a single nematophore, the other lacking such an organ. Yet each of the internodes is complete as regards the internal septa, containing one at each end. They are distinct inter- nodes and seem to be due to spontaneous variation, for no hint could be observed that, as Bitiarp found in his specimens, rupture and subsequent regeneration had taken place. Our specimens appear to be similar to the “ distinct variety” recorded by Professor Norrine from gulf weed (1900, p. 57), although Nurrine’s specimens differ in being branched. Detailed measurements indicate that the Scotva specimens stand intermediate to the variety found by the T7ravailleur at Cape Spartel and to the south of Madeira, and the typical form whose dimensions BiLLarD records. Measurements :— Height of colony : : ; " F 7-8 mm. Length of stem internodes : ‘ d : 0°33-0°39 mm. Breadth of _,, 5 : 5 : . 0:075-0:09 ,, Length of intermediate internodes ; : : 0°14-0°21 ,, 6 thecate internodes . 5 : ; 0:31-0:40 ,, Breadth of ,, s , : : 0:42—0°48 Locality.—Creeping on gulf weed found in spawn net at Station 538. Lat. 32° 11’N., long. 34° 10’ W. 30th June 1904. TRANS. ROY, SOC, EDIN., VOL, XLVII. PART I. (NO. 4). 12 ~ 90 MR JAMES RITCHIE: SUPPLEMENTARY REPORT ON Antennopsis scotix, Ritchie, 1907. An additional colony of this species from the locality from which it was originally recorded enables me to amplify the diagnosis already given and to add a description of the gonosome. The new colony is of the same height as the larger of our earlier speci- mens, 9 cm., but it bears more branches and is altogether in better condition, although here also the hydroclades are in many places wanting. Branches are frequent, but are very irregular in position. They may bear secondary branches which are long, of uniform thickness, and are seldom branched. The stem and branches are composed of a thick bundle of tubes with transparent walls and without nodes. From short processes on the outermost of these the hydroclades spring, followimg one another on the same tube at a Fic, 8.—Antennopsis scotie, (a) Portion of hydroclade showing thecate and athecate internodes, and arrangement of nematothece. x65, (b) Gonotheca, x 45. distance of some 0°73 mm., and thus forming a close-set coat round the branches. In consequence of the delicacy of the hydrothecal margins a perfect hydrotheca is rare, but where such occurs it shows a slight widening at the mouth. An important addition has to be made to the previous description of the trophosome as regards the number of the nematophores accompanying the hydrotheca. Besides the median proximal nematophore and the lateral pair surmounting the processes which run alongside the hydrotheca, there is a supplementary lateral pair, the individuals of which, one on each side of the hydrotheca, rest on the upper surface of the lateral process almost in the corner formed between it and the internode (fig. 8). These nematophores, although of similar structure to, are considerably smaller than, the others, and are so delicate that they are frequently absent, their former position being marked only by a small opening in the lateral process. They correspond exactly to the supp nena nematophores described in Antenella quadriaurita of the present paper. The athecate internodes show more variation than in the former specimens, their length in many cases preventing the margin of a hydrotheca from reaching the level of the proximal end of the succeeding thecate internode. Only two THE HYDROIDS OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION. 91 deviations from the normal structure of the athecate internodes have been observed where, instead of one, two nematophores were present. Gonosome.—The gonangia arise laterally from the hydroclades. They are situated immediately beneath the hydrothecee on a short process from the hydroclade internode, between which and the body of the gonangium a short internode intervenes. In size a gonangium is three or four times as large as a hydrotheca; in shape it is broadly ovate, tapering proximally into a short stalk which rests upon the intervening internode mentioned above, and abruptly truncated distally, where the large terminal aperture is closed, prior to the maturity of the contents, by a one-flapped operculum attached by its abeauline edge to the wall of the gonangium. Two large nematophores are present, one on each side of the gonangium near its base. Measurements :— . Athecate internodes, length* . ; ‘ : 0°28-0°32 mm. Thecate 3 Be eM jf 3 amy : 0°36-0°38 _,, Hydrotheca, length . , . _ : 0:20-0:24 ,, i breadth at mouth. : : ; 0-21-0°24 ,, Gonangium, length. ‘ ‘ : ‘ 0:88-0°91 ,, 5 greatest breadth . ; : 5 0°53-0°57 _,, Locality.—As previously recorded—entrance to Saldanha Bay, Cape Colony. 25 fathoms. 21st May 1904. Growing on a sponge from the same locality were several simple colonies up to 7mm. high. Although there are no signs of fasciculation, nor even of true branching, the minute structure corresponds so exactly with that of A. scotie that I cannot but conclude that the small colonies represent an early stage of that species. ‘I'hey bear no gonangia. Monostechas quadridens (M‘Crady, 1859). Two minute colonies of this species were found growing on the leg of a masked crab. They differ from the typical form described by Nurtine (1900, p. 75) only in their minuteness—they are less than 1 cm. high—and in the length of the athecate intermediate internodes of their hydroclades. Although each bears two or three hydro- clades, neither of the colonies is branched ; yet circular holes at the bases of some of the hydrothecz, indicating the point where a gonangium had been attached, show that the colonies are mature. The unusual position in which the colonies were growing is probably responsible for their smallness, for an unstable foundation is frequently accompanied by a dwarfed fauna. Measurements :— Stem, diameter ; ; ¢ : : 0°14 mm. Thecate internodes, length . ‘ ‘ a 0°42-0°46 mm. Intermediate internodes, length ; ; : 0:56-0°73 ,, Hydrotheca, depth : : 5 ‘ : 0°15-0°17 _,, i diameter at opening : ; : 0:20-0:22 ” * Length measured from straight node to furthest point of oblique node. 92 MR JAMES RITCHIE: SUPPLEMENTARY REPORT ON Locality.—Growing amongst seaweed on the leg of a masked crab. Station 81, Abrohlos Bank, Brazil. Lat. 18° 24’ S., long. 37° 58’ W. Depth, 36 fathoms. Bottom doposit, coral. 20th December 1902. Antenella quadriaurita,* sp. nov. A few sparse colonies which cannot be referred to any described species of Antenella were trawled off Gough Island. The stems, the largest of which is 14 mm. high, are hairlike and stand out rigidly from a creeping stolon like a group of stiff bristles. To the unaided eye the stem groups much resemble the figure of A. gracilis given by ALLMAN (1877, pl. xxii, fig. 6), but in our specimens the stems are more delicate, and, Z ZZ Z Z Z Z F ae Ay 2 Z Y A A A g Z Z Z Z AA Yj Kic. 9.—Antenella quadriaurita. (a) Portion of stem. x25. (6) Anterior aspect of a hydrotheca, x 125. (c) Lateral aspect of a hydrotheca. x 125. owing perhaps to mere accident, they are more irregular in size and more straggling in arrangement. ‘The stem is divided into a series of alternating thecate and athecate internodes, the boundary lines between these being oblique and very distinct on the proximal side of the thecate internode, and on the distal, transverse but faintly indicated. On the stems examined the maximum number of hydrothecee was nine, but that a greater number may be borne is likely, as in both the “nine” colonies the stem was incomplete. ‘The hydrothecz are borne on every other internode. As seen in profile they are cylindrical, having almost parallel edges, but viewed from in front they seem to be conical in shape, tapering rapidly to the base. A hydrotheca is rather deeper than broad, and for more than half its height is free from the internode. Its profile is straight, and its margin is very slightly everted. Kach thecate internode bears five nematophores: one median, placed on a gently * Quattuor, four ; and auritus, eared, signifying the presence of two pairs of nematothece flanking the hydrotheca. THE HYDROIDS OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION. 93 raised portion of the internode on the near side of the hydrotheca ; a supracalycine pair, each individual of which reaches just to the margin of the hydrotheca and rests upon the end of an internodal projection stretching half-way across the hydrotheca; and an additional pair—supplementary nematophores—placed on the upper and exterior surface of the supracalycine process, almost in the angle between that process and the inter- node. ‘The latter are extremely fragile and readily detached, so that in many cases they are unrepresented but by a small pore in the wall of the supracalycine process, indicating where they had been attached. ‘The intermediate internodes bear a number of nematophores varying from two to four, but three is the most frequent number. Thus, in twenty intermediate internodes examined, eight bore two nematophores, eleven bore three, while only one had four. On the athecate basal internodes, three of which generally follow one another in close succession prior to the first hydrotheca, the nematophores also vary. Of nine such, four had four nematophores, two had five, while series of two, three, and seven nematophores were represented on one internode each. Measurements :— Thecate internode, length* . : : 5 0°46 mm. Intermediate internode, length * : f ; 0:57-0:91 mm. breadth : : : 0:07-0:087 ,, Hydrotheca, depth : : E : ; 0:21-0:22 ,, * diameter at margin . 5 : 0-17-0°21 ,, This species is closely related to Plumularia secundaria (L., 1789, p. 3854), which, if the genus Antenella is to be preserved, and it seems a useful one from the point of - view of practical convenience, must be transferred to that genus. The present species has been separated from dA. secundaria on the strength of observations made by MarRKTANNER-TURNERETSCHER (1890, p. 252), Piorer and Brpor (1900, pp. 27, 28), and Bittarp (1907, p. 207), all of whom record the presence of a single nematotheca immediately above the hydrotheca, whereas in our specimens the hydrotheca is flanked by a paar of nematothecz in addition to the usual supracalycine pair. The athecate internodes in the Scotia species are twice the length of those in A. secundaria, and the other parts differ in their relative proportions, but little stress can be laid on so indefinite and so variable a character. | am unable to distinguish A. natalensis, Warren, 1908, from A. secundaria. Locality.—Gough Island. Trawled at a depth of 100 fathoms. Bottom deposit, Bryozoa and rock. 23rd April 1904. Aglaophema allmani, Nutting, 1900. Two specimens are referred to this rare species. One is a comparatively small colony, 6°5 cm. high, still retaining some fragments of the hydrorhizal tubes. The other is a strongly fascicled branch, with a diameter of 1°5 mm. at its junction with * Measured from straight node to furthest point of oblique node. 94 MR JAMES RITCHIE: SUPPLEMENTARY REPORT ON what is probably part of the main stem, and with a height of 7°5 cm. It bears rather delicate alternate ramuli which leave it almost at right angles. While the general characters—branching, structure of hydoclades, shape and approximation of hydrothecze, position of nematophores—agree with the descriptions of — ALLMAN (1877, p. 39, pl. xxii., as A. ramosa) and of NuTrinG, variations worthy of note have been observed in the last-mentioned organs. These variations seem to be mainly age differences. In the hydrothecee on the proximal parts of the hydroclades the mesial nuematophore reaches to the level of the marginal teeth, and is adnate, all but the very tip. The supracalycine nematophores are generally cylindrical, with two apertures— one terminal, the other a large oval opening on that side of the upper surface which is Fic. 10.—Aglaophenia allmani, showing variation in hydrothece and nematothece. (a) Hydrothece from the proximal end of a hydroclade. 65, (6) Hydrothece from the distal end of a hydroclade. x 65, towards the interior of the hydrotheca. Rarely this opening is much elongated, and extends through the band of perisare which separates it from the terminal opening. In such a case a single continuous opening is formed and the nematophores might be described as “almost cylindrical.” The tip of a supracalycine nematophore reaches just to the margin of a hydrotheca. Its axis, viewed from the side, lies at an angle of about 45° with the stem. ‘lhe nematophores, in general, agree with the type so far described. In those hydrothecze which occur towards the tip of the hydroclades, and which are therefore younger, the aperture is less oblique, while the adnate part of the hydrotheca is of the same length as in the older examples. As a consequence the mesial nematophore, although shown by measurement to be constant in length throughout the colony, appears to be shorter in the newly formed hydrothece, since its tip falls con- siderably short of the margin. The supracalycine nematophores are of markedly greater length, overtopping the margin by about 0°056 mm., are quite cylindrical, and lie with their axis (viewed from the side) more nearly parallel to that of the internode. The lateral aperture is smaller and is never continuous with the terminal one, and the THE HYDROIDS OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION. 95 internodal septum which in the older hydrothecee marks the base of the nematophore is indicated in the terminal individuals only by a minute indentation. I think, with Nurrie, that the general structure of the trophosome of the colonies resembles that of Lytocarpus rather than that of Aglaophenia. Measurements :— Stem internodes, Jength ; : : 0°39-0°45 mm. 5 5 breadth ’ 5 ; ; O:22=0:24 |; Hydroclade internodes, length . ; : 0:°38-0°42__,, Hydrothece, length *. é : ; 0°35 mm. “3 diameter at margin ; ; s OPI op Mesial nematophore, length . : : O2385e. Supracalycine nematophores, length of longest side, 0°11 mm. at base of hydroclade to 0:14-0:17 mm. at distal end of hydroclade. Locality—Station 81, Abrohlos Bank, Brazil. Lat. 18° 14’ S., long. 37° 58’ W. Depth, 36 fathoms. Bottom deposit, coral. 20th December 1902. Distribution.—Only recorded from Florida Reef, in the Gulf of Mexico (by ALLMaN), and from a station in the Caribbean Sea (by Nurrine). Aglaophena dubia, Nutting, 1900. Two specimens of this species, the A. gracilis of AtLMAN’s Gulf Stream Report (ALLMAN, 1877), were collected in the same locality—one simple, reaching a height of 6 cm., the other sparsely branched and rather longer. The anterior profile of the hydrotheca is not so markedly concave as in ALLMAN’s figure, nor is the mesial nematophore quite so long relatively to the height of the hydrotheca. The hydrothecee haye nine teeth (NuTTING says “about eight”), of which one on each side lies behind the supracalycine nematophores and is more acute than the others, while the anterior tooth is usually recurved. ‘The number of the basal nematophores differs from that recorded by ALLMAN and Nortina, for not only are two present on the front of each stem-internode (one close to the hydroclade and one on the proximal portion of the internode), while another small one lies at the base of the hydroclade, close to the former of those just mentioned—but, in addition, a large cup-shaped nematophore exists on the posterior aspect of internode, immediately behind the base of the hydroclade. Other- wise the specimens agree, point for point, with previous descriptions. Measurements :— Stem internodes, length ; : : i ‘ 0-52 mm. a a breadth . ; ; : 0-31 ,, Hydroclade internodes, length , : j : 0:36 ,, es a breadth at middle . , : 0:05 ,, Hydrothece, height : : , : O'32"%, a diameter at margin ; : : : 0714 ,, Mesial nematophore, length : : : , Oli 5, * Length of hydrotheca measured along the internode from the base of the cavity to the margin. 96 MR JAMES RITCHIE: SUPPLEMENTARY REPORT ON On the branched specimen two branches, which appear to belong to the colony, since they lie in the same plane and leave the stem at the same angle as the true branches, were found, on microscopic examination, to be specimens of Halicornaria longicauda, arising from hydrorhizal tubes climbing upon the stem of the Aglaophema colony. Locality.—Station 81, Abrohlos Bank, Brazil. Lat. 18° 14’ 8, long. 37° 58’ W. Depth, 36 fathoms. Bottom deposit, coral. 20th December 1902. Aglaophenia heterodonta, Jiderholm, 1903. Dr Exor JApERHOLM has described amongst the extra-EKuropean Hydroids in the Swedish Museum specimens of A..dichotoma (M. Sars), as distinct from a form with similar habit which he has named A. heterodonta. I now regard the specimens which were described in the earlier Scotia report under the name of A. dichotoma as examples of A. heterodonta. Additional material enables me to add to JADERHOLM’s description of the general habit of the colonies. His specimens were characterised by irregularly ramified stems bearing short, upward curling twigs. Our specimens exhibit two types. The first, pre- viously described (Rrrowin, 1907,” pl. iii. fig. 2), is strictly dichotomous, although the branches may not develop equally in all parts of the colony. This type of branching is exactly similar to that of A. dichotoma. The dichotomously branched specimens were growing on a sponge, and are considerably taller (10 cm.) than JADERHOLM’S examples (3°5 cm.). The habit of the second type is distinctly reminiscent of that of, A. conferta, Kirchenpauer, 1872; that is to say, simple curved stems spring in pro- fusion from a hydrorhiza creeping upon an alga. ‘There is no sign of branching. The largest of those colonies is only 18 mm. high, but that they are fully developed is shown by their sexual maturity, for several bear corbulee with male gonophores. The minute characters of the two types of colonies are identical, and agree with those of A. heterodonta. It may be, however, that this is but a synonym of A. conferta, the only characters which seem to separate the latter being the absence of an unpaired anterior reflexed tooth (which, however, appears to be present in KiRcHEN- PAUER'S figure); the outward, instead of the inward, direction assumed by the pair of teeth nearest the stem; and the angled nature attributed to the supracalycine nematophores. However, it is only by examination of KiRCHENPAUER’S type that such a question could be decided. That the branched and unbranched colonies are found on two different types of substratum probably indicates that they are environmental modifications; the fixed, settled colony (that on the sponge) becoming luxuriant, while the drifting, unsettled colony (that growing on the alga) tends, as do so many alga-borne Hydroids, to remain dwarfed and of simple habit. The development of the corbula differs slightly from that of A. pluma as described by ALLMAN (1871, p. 59) and Nurrine (1900, p. 40), for the leaves develop less simul- THE HYDROIDS OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION. 97 taneously. While in Nurrine’s specimens all the leaves had made their appearance before even the primary pair had reached full development, here, when only five pairs are recognisable, the two first-formed pairs are already full grown; and when six pairs are visible the earliest three have reached full development, the later ones being in a state of decreasing perfection. While some of the corbule are wholly closed some remain partially open, their leaves, at least towards the tip, bearing nematophores on each side, and remaining separate from each other. In the latter case the gonophores are always male, in the former no gonophores remained; but since maleness and openness go together, as they do in so many other cases, it seems probable that in this species we have an example of sexual dimorphism of the type described by Mr H. B. Torrey and Miss Martin (1906). The depth of a hydrotheca varies from 0°27 to 0°28 mm., its diameter at the mouth from 0°17 to 0°19 mm. ; measurements agreeing with those of J ADERHOLM. Locality.—On sponges and algze from the entrance to Saldanha Bay, Cape Colony. Depth, 25 fathoms. Bottom deposit, sand and kelp. 21st May 1904. Aglaophena minima, Nutting, 1900. Two colonies (the larger 1 cm. high) are distinguished by their cylindrical, keelless. hydrothecz with short projecting mesial nematophores as belonging to this species. In addition to the septal ridge, mentioned by Nurrine, which traverses the hydrotheca- bearing internode at the level of the intrathecal ridge, another is evident opposite the base of the supracalycine nematophores. The opening of the hydrotheca is bordered by nine, instead of by eight teeth ; and in the mesial nematophore of some of the hydro- thecze a slight chitinous constriction, almost in line with the profile of the hydrotheca, is apparent. ‘The nematophores on the stem internodes are arranged as follows :—A solitary long nematophore on the front and at the proximal end of the internode ; and in the angle between the hydroclade process and the stem, a double nematophore, with two diverging processes each bearing a terminal aperture. Only a single opening, however, connects the cavity of the double nematophore with that of the colony. The hydroclade process itself bears a large simple nematophore on its anterior surface, and to this appears to be due the bifurcated appearance mentioned by Nurrtine. No gonangia were present. Measurements :— Stem internodes, length : F : 3 0°45-0°56 mm. 5 diameter : ; ; : 0:08-0°10 ,, Hydroclade internodes, length . : : : 0°39 mm. Hydrotheca, depth : ; : : : 0°34 _,, *; diameter at mouth ‘ ; ; 0:14 ” Locality.—Growing amongst seaweed on the leg of a masked crab. Station 81, TRANS. ROY. SOC. EDIN., VOL. XLVII. PART I. (NO. 4). 13 98 MR JAMES RITCHIE: SUPPLEMENTARY REPORT ON Abrohlos Bank, Brazil. Lat. 18° 24’ §., long. 37° 58’ W. Depth, 36 fathoms. Bottom deposit, coral, 20th December 1902. Previously recorded from Little Cat Island, Bahamas (Nurrinc). Aglaophenia latecarinata, Allman, 1877. This common tropical species, identical, as shown by the researches of Professor Norzine (1900, p. 96), with the A. minuta of Frwxkes (1881, p. 132), occurs among the Scotia collections in its usual habitat, creeping upon the fronds and bladders of Saragassum weed. The specimens, the largest of which are 13 mm. high, correspond with Nurrine’s description in all points but one. For while he mentions only » two nematophores at the base of each hydroclade, I have observed in all cases four nematophore apertures, the two recorded by Nurrine, and, in addition, lying immedi- ately distal to the internodal process from which the hydroclade projects, in the axil between it and the stem, a double nematophore, possessing two apertures, one directed to the right, the other to the left of the hydroclade process. The ccenosare of this nematophore connects with the general ccenosare of the colony through a single median perforation in the internode wall. A similar arrangement of nematophores occurred in the specimens examined by Bittarp (1907) and VerRsLuys (1899). As the following comparative table shows, our specimens are, in all respects, somewhat larger than those described by BrtLarp :— Scotia Specimen. BILLARD's Talisman Specimen, Length of hydrocaulus ‘ : : 6-13 mm. 5-6 mm. Breadth of 5 : : : 0-1-0°13 mm, 0-:08-0'12 mm. Length of stem internodes. : F 0:34 mm. 0°25-0°30 ,, af hydrotheca ; : : 0:31 ,, 0:27-0°30 ,, Breadth of hydrotheca at mouth (excluding keel) , : é : Olay, 0°135 mm. Breadth of keel . ; : ; 0:03 ,, q Locality.—On floating Saragassum weed, caught in spawn net. Station 538, Lat. 32° 11’ N., long. 34° 10’ W. 30th June 1904. Halicornaria longicauda, Nutting, 1900. At first glance one of the specimens which I have referred to this species appears to be a fascicled colony with alternate branches ; but closer scrutiny shows that there is present a central axis with the remains of hydroclades, and that around this are grouped tubes which in their distal portions are continued as the so-called branches, and in the proximal appear to form the hydrorhizal tubes of the compound colony. The specimen really consists of a central axis upon which several distinct Halicornarza colonies happen THE HYDROIDS OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION. 99 to be creeping. The clustered hydrorhizal tubes of these create the resemblance to fascicling, but the structure is analogous rather to the rhizocaulom often exhibited by Lafoéa dumosa than to a truly fascicled stem. The rhizoid nature of this compound stem is confirmed by another specimen (Aglaophenia dubia) from the same locality, where the two lower branches, at least so they appear to the eye, turn out to be distinct climbing colonies of H. longicauda. The largest of the colonies found by the Scot#a is 11 cm. in height. The specific characters agree with those given by Professor Nurrinc, but in our specimens the hydrothecee appear to have undergone a greater degree of tilting forward than his figures indicate, while the lateral teeth are more strongly developed. The aperture is vertical and slightly constricted, with a sharp tooth projecting upwards and outwards on either side. The intrathecal ridge is well marked, arising near the middle of the Fic, 11.—Halicornaria longicauda, Hydrothece showing variation in the length of the median nematophore. x 110. adnate portion of the mesial nematophore and extending backwards at an angle of 45° with the axis of the hydroclade. Its free edge is slightly reduplicated. The supracaly- cine nematophores are short and stout, and when viewed from the front appear clearly above the hydrotheca, although they do not reach the margin of the aperture. The mesial nematophore varies greatly in length ; considerable differences may be seen even in two succeeding hydrothece. Sometimes shorter than in the examples figured by Norrine, it may, on the other hand, extend far beyond the margin of the hydrotheca, and, curving gracefully upwards, its tip may reach the level of the upper portion of the rim. There are three cauline nematophores (not two, as Nurrine implies) clustered about the stem process on which the hydrotheca rests. Two lie on the anterior surface, one at the distal, the other at the proximal side of the stem process, while one lies on the posterior aspect of the process itself. They are triangular in shape and large, normally with two apertures, one at each of the free angles of the triangle, but occasionally showing, as a variation, a third aperture, situated medianly between the normal two. The gonosome is unknown. 100 MR JAMES RITCHIE: SUPPLEMENTARY REPORT ON Measurements :— Stem internodes, length 2 : ; : 0°29-0°35 mm. a breadth ; ; : ; 013-0716 ,, Hydroclade internodes, length . ; F : 0-25-0°29 _,, Hydrotheca, depth . i ‘ : : 0°14 7 Bs vertical diameter at margin : ‘ 0°13-0°14 ,, “ horizontal ,, 3 : ; 0:18 ss Locality.—Station 81, Abrohlos Bank, Brazil. Lat. 18° 14’ 8., long. 37° 58’ W. Fathoms, 36. Bottom deposit, coral. 20th December 1902. Distribution.—The only record given by Nurrine is from the Caribbean Sea near the Isthmus of Panama (lat. 9° 32’ N., long. 79° 55’ W. 36 fathoms). The present locality, alsoa shallow water one, is off the coast of Brazil near Porto Alegre—a consider- able southwards extension of the known range. LITERATURE. Mere mention is here made of the memoirs alluded to in this paper. For their titles reference must be made to such bibliographies as that in HartLaus’s “‘ Die Hydroiden der magalhaenischen Region und chilen- ischen Kiiste” (below 1905), or those in Nurrine’s ‘Monographs of the American Hydroids.” Aver, J., 1857. Trans. Tyneside Naturalists’ Field Club, vol. iii. pp. 93-160. Auman, J. G., 1864. Ann. Mag. Nat. Hist. (3), vol. xiii. pp. 345-380. 5 1871. A Monograph of the Gymnoblastic or Tubularian Hydroids, London, Ray Society. 5 1877. Mem. Mus. Comp. Zool. Harvard, vol. v., No. 2. - 1885. Jour. Linn. Soc. London, Zool., vol. xix., pp. 132-161. 1888. Report Scientific Results, ‘‘ Challenger,” Zool., vol. xxiii. Bate, W. M., 1894. Proc. Roy. Soc. Victoria, vol. vi. BgrENcOURT, iy 1899. ‘‘Deuxieme liste des Hydraires du Pas-de-Calais,” Miscell. Biolog., Trav. Soc. Zool., Wimereux, vol. vii. pp. 1-138. Bonnevik, K., 1899. Den Norske Nordhavs Kxpedition, 1876-1878, No. 26. Christiania. Bitrarp, A., 1906, (1). Actes Soc, Linn, de Bordeaux, vol. |xi. pp. 69-76. ‘ 1906, (2). Haxpédition Antarctique Frangaise, 1903-1905, “‘Hydroides.” Paris, 1906. a 1907. xpéditions scientifiques du “ Travailleur” et du “ Talisman,” tome viii. Bruce, W.S., 1894. ‘ Antarctic Birds” in Knowledge, Sept. 1, 1894, pp. 208-210. Busk, Gro., 1850. Rep. Brit. Assoc. Adv. Sc., 20 Meet. (London, 1851), pp. 118-120. Cuarke, 8. F., 1879. Bull. Mus. Comp. Zool. Harvard, vol. v., No. 10, p. 239. Exuis, J., 1755. An Essay toward a Natural History of the Corallines, ete. London, 1755. Fewxss, J. W., 1881. Bull. Mus. Comp. Zool. Harvard, vol. viii., No. 7, p. 127. Harriavs, C., 1900. ‘ Revision der Sertularella-Arten,” Abh. naturw. Ver. Hamburg, vol. xvi. pp. 1-143. oe 1901. Zool. Jahr., vol. xiv., Syst., pp. 349-379. 53 1904. Résultats du voyage du S.Y. “ Belgica,” Zool., “ Hydroiden.” Anvers, 1904. = 1905. ‘‘ Fauna Chilensis ”—Supplement vi., Zool. Jahr., 1905. Hickson, 8. J., and Gravety, F. H., 1907. ‘‘ Hydroid Zoophytes, National Antarctic Expedition,” Natural History, vol. iii. Hincxs, T., 1861. Ann. Mag. Nat. Hist., ser. iii. vol. viii. p. 251. . 1868. A History of the British Hydroid Zoophytes. London, 1868. JApERHOLM, E., 1903. Arkiv for zool., utg. af Kgl. Svenska Vetenskapsakad., Bd. i., pp. 259-312. { ; ( ; THE HYDROIDS OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION. 101 JADERHOLM, E., 1904. Archives de zool. expér. et générale, sér. iv. vol. ill. - 1905. Wéssenschaftliche Ergebnisse der schwedischen Stidpolar-Expedition, 1901-1903, Bd. v., Lief. 8. Stockholm. Lamarck, J. P. B, A. de, 1836. Histoire naturelle des animaux sans vertebres, 2nd edit., 1836. Lamouroux, J. V. F., 1821. Hzposition méthodique des Genres de ordre des Polypiers, ete. Paris, 1821. Ms 1824. Zoologie du voyage autour du monde exécuté sur les corvettes ?Uranie et la Physictenne, etc., Paris, 1824, pp. 603-643. Linnaus, C., 1758. Systema Nature, 10th edit. a 1789. "5 5 12th edit. M‘Crapy, J., 1859. Proc. Elliott Nat. Hist. Soc., vol. i., No. 1, p. 103. MARKTANNER-TURNERETSCHER, G., 1890. Ann. des k. k. naturh. Hofmuseums, Wien, vol. v. pp. 195-286. Normay, A. M., 1864. Ann. Mag. Nat. Hist., ser. 3, vol. xiii. pp. 82-90. Nortine, C. C., 1900. “American Hydroids,” Part I. The Plumularide, Spec. Bull. Smithsonian Inst., Washington, 1900. - 1904. Jbid., Part II. The Sertularide. Washington, 1904. Orrmann, A. E., 1896. Grundziige der marinen Tieryeographie. Jena, 1896. Pawuas, P. 8., 1766. Hlenchus Zoophytorum. Haag. Picrst, C., et Bepor, M., 1900. Résultats des Campagnes Sc., etc., Monaco, fascicule xviii. Rironie, Jas., 1907 (1). ‘‘Hydroids of the Scottish National Antarctic Expedition,” Trans. Roy. Soe. Edin., vol. xlv. pp. 519-545 ; also in Report on Sc. Res. of Voyage of S.Y. ‘ Scolia,’ vol. v. pp. 61-88, Edinburgh, 1909. oP 1907 (2). Proc. Zoo. Soc. London, 1907, pp. 488-514. Tomson, J. A., 1904. ‘Note on the Gonostyles of Two Antarctic Siphonophora,” Proc. Roy. Phys. Soc. Edin., vol. xvi. pp. 19-22. Torrey, H, B., 1902. “ University of California Publications,” Zool., vol. 1. pp. 1-104. Torrey, H. B., and Martin, A., 1906. Jbid., vol. i1. pp. 47-52. Verstvyrs, J. J., 1899. Mém. Soc. Zool. France, 1899, vol. xiii. pp. 29-58. Warren, F., 1908. ‘‘On a Collection of Hydroids, mostly from the Natal Coast,” Ann. Natal Government Mus., vol. i. part i1., pp. 269-355. Wrieat, T.8., 1858. Proc. Roy. Phys. Soc. Edin., vol. i. p. 450. TRANS. ROY. SOC. EDIN., VOL. XLVII. PART I. (NO. 4). 14 —, (. 403% 4) V.—The Meteorology of the Weddell Quadrant and Adjacent Areas. By Robert C. Mossman. (With Five Plates.) (MS. received July 6, 1908. Read July 20,1908. Issued separately June 9, 1909.) Having been struck by the evident importance of Antarctic phenomena in relation to the climatic condition of lower latitudes, I have endeavoured in the following discussion to give a complete representation of these conditions, and to trace their bearing on the climate of sub-Antarctic and temperate zones. It is only now that sufficient data are available for such a purpose. I have been singularly fortunate, not only in the opportunities which I have had for obtaining access to valuable data, but also in the possession of that “practical” knowledge which has resulted from my two years residence in the Antarctic. In any discussion of this nature it is obviously an immense advantage to have studied in the field the conditions there obtaining. ‘This I have been able to do also for the Arctic during two voyages under- taken for this specific purpose, thereby being enabled to appreciate the great diversity in the fundamental conditions affecting the meteorology of the two Polar regions. In such an inquiry as this, in which an attempt is made to break new ground, it is essential that the discussion should proceed on broad and general lines, in order that the fundamental principles involved may not be obscured by minor and negligible detail. Accordingly, I have restricted this inquiry to a discussion of the more prominent elements of climate. No portion of the Antarctic affords such an excellent field for the study of atmospheric conditions as that lying to the south and east of Cape Horn. During recent years this area has been the principal centre of South Polar exploration, no less than four expeditions having investigated the region lying between the meridian of Greenwich and 100° of west longitude. Moreover, we have in the South Orkney Station of the Argentine Meteorological Office the longest series of observations made south of 55°S., while data from lower latitudes, covering in each case many years, are also available for the numerous island and mainland stations of the Argentine and Chihan Republics. Valuable supplementary material has been furnished by the various summer voyages of WEDDELL, Ross, D’UrRvIiux, and others. The difficulties involved in this discussion are many, and will be readily recognised by those who have engaged in work of a similar nature. Among these are the comparative paucity of observations, the relatively great distance separating the spheres of work of the different expeditions, and the general want of synchronism in the periods covered by the observations. And yet, in spite of these difficulties and deficiencies, an attempt must be made to co-ordinate and co-relate the available information if an TRANS, ROY, SOC, EDIN., VOL. XLYII, PART I. (NO. 5), 15 104 MR ROBERT C. MOSSMAN ON THE METEOROLOGY OF approximation towards the general principles of the atmospheric circulation is to be obtained. The region chosen is not only the best but the sole available area in the whole circumference of the globe south of 50° 8. which offers a sufficient number of observing stations to warrant serious discussion. Maps are given (see Plates I. to IV.) showing the mean monthly and annual dis- tribution of pressure, temperature, and cloud. In their preparation I have drawn lines only for the regions furnishing sufficient data. I have also, as far as possible, en- deavoured to eliminate discordances due to the exceptional characteristics of individual months. With regard to NoRDENSKJOLD’S observations at Snow Hill and Paulet Island in 1908, and those taken by CHarcor at Wandel Island in 1904, I have been able to bring to bear on their discussion the experience derived from a residence during these years at the South Orkneys, which are, from a meteorological point of view, in the same region. In other words, I have tried to represent the conditions at these places as if they had been based on the same five years as were the South Orkney observations. In this connection ordinary methods of differentiation were not of much use, especially with regard to CHarcor’s 1904 observations, as this was a very exceptional year. It is for this reason that the Isotherms, Isonephs, and the Isobars do not always exactly correspond with the values given in the tables. Even with regard to the South Orkneys and South Georgia, I have allowed myself a little latitude in this direction. For the region of the “ Belgica’s”” drift I have taken the mean monthly temperatures, calculated by Hann,* except that I am inclined to assign to February a lower mean temperature than that given by him. With reference to the observations taken at the Chilian coast stations from 1899-1905,t the investigation is restricted, except as regards one or two elements such as rainfall, to the five years 1901--1905. The mean monthly temperature, pressure, humidity, and rainfall were for this series extracted from the general summary given in each volume. The mean cloud, wind force, and direction were computed from the tri-daily observations given “im extenso.” Owing to the labour attached to work of this nature, an analysis of the winds was made only for Dungeness and Evangelists Island. For New Year’s Island the data contained in the MS. sheets of observations were similarly computed from seven daily observations. As regards the Argentine Republic, much valuable information has been obtained from Mr Davis’ work on the “Climate of the Argentine Republic,” as well as from the elaborate data contained in the “ Annals” of the Argentine Meteorological Office. In this connection I have to express my cordial thanks to Mr Davis for permission to utilise the South Orkney and South Georgia observations in so far as they bear on the subject under discussion. I am also indebted to Dr W. S. Bruce for the loan of his valuable meteorological log kept on board the “ Balena” during his voyage to the South Shetlands and Louis Philippe Land in the summer of 1892-1898. * Antarctic Manual, p. 44. + Annuario del Servicio Meteorologico de la Direccion del Territorio Maritimo, Valparaiso, 1902-1906. THE WEDDELL QUADRANT AND ADJACENT AREAS. 105 GEOGRAPHICAL PosITION oF AREAS DISCUSSED. Weddell Quadrant—(See Plate V.).—The region extending from 90° W. to the meridian of Greenwich. This area is divided into the following subsections :— Bellingshausen Sea.—The area to the west of Graham’s Land south of 64° S., extending to the 100th meridian of west longitude. Data for this region are entirely derived from the observations taken during the drift of the “ Belgica.” Graham’s Land.—Includes Graham’s Land, Danco Land, Louis Philippe Land, and the South Shetlands, by some geographers termed the Dirk Gherritz Archipelago. Data comprise a year and nine months’ observations at Snow Hill, seven months’ observations at Paulet Island, a year’s observations at Wandel Island, and the summer observations of Forster, Ross, Bruck, and others in the vicinity of the South Shet- lands and Louis Philippe Land. Weddell Sea.—The region south of 60° S. between the meridian of Greenwich and 50° of west longitude. Data include five years’ observations at the South Orkneys, and the summer observations, extending over several months, made by WEDDELL, Ross, Biscog, and particularly by the Scottish National Antarctic Expedition, which initiated the South Orkney series. Biscoe Sea.—This region, from the meridian of Greenwich to 40° E., although outside the specific area under discussion, is sometimes referred to. Data are available from the Expeditions of Moors, Biscor, and the ‘“ Valdivia” and “ Challenger.” South Atlantic.—The area discussed is that portion west of the meridian of Greenwich lying south of 40° 8S. Data from ships are very scanty for some portions, especially in the region 40° to 60° S., between 0° and 30° W., as well as from 50° to 60° S., between longitudes of 30° and 40° W. On the other hand, observations covering four years are available from South Georgia Island in lat. 54° 30’ S., and covering seven years from the Falkland Islands in 51° 41’ 8. South America.—South of 50° 8. is represented by observations from numerous stations in Chili, Tierra del Fuego, and Patagonia, including many strictly insular situations such as Evangelists Island at the Pacific entrance to the Straits of Magellan, and Staten Island, and the neighbouring New Year’s Island a little north-east of Cape Horn. Returns from a few places on the Pacific and the Atlantic side of the continent in from 40° to 43° S. are also given. In addition to the land observations, there is a large mass of data for both oceans in the publications of the London and other meteorological offices. GENERAL CLIMATIC FEATURES. The general climatic features of the region under discussion, viz. from 40° to 70° 8., may be briefly summarised as follows. Beginning with the mainland of South America, we note that the two most northerly coast stations on the Pacific side, Ancud and Gallera, have a large rainfall, and, relatively speaking, much cloud. The rainfall 106 MR ROBERT C. MOSSMAN ON THE METEOROLOGY OF shows a distinct seasonal variation, being at a maximum in winter and a minimum in summer, while the pressure, cloud, and humidity curves are directly the reverse of this. Temperature is comparatively low for the latitude. Coming 8 degrees south we have the station of Evangelists Island: here also there is a very considerable precipita- tion, but with a maximum in summer and a minimum in winter, not, however, showing anything like the seasonal range of the northern stations. The climate is essentially insular and the summer temperature very low owing to the prevalence of the cool on- shore north-west winds, which blow with great force. Passing now to the Atlantic side, we find a great diminution of rainfall, most pronounced at inland stations such as Sandy Point (Punta Arenas) and Ushuaia. Here the temperature shows much greater amplitude than at the coastal stations, and is essentially of the continental type—- higher in summer and lower in winter—in harmony with the conditions prevailing in the interior of the Argentine provinces of Santa Cruz and Chubut. Another marked feature of this region is the low relative humidity and the small cloud amount, due to the prevailing winds having been deprived of much of their moisture by the mountainous region lying to the west. With regard to coastal and island stations on the Atlantic side, a very small difference in latitude makes a very great difference in climatic conditions. Staten Island, New Year’s Island, and Orange Bay have all a considerable rainfall, much cloud and humidity, with a low and equable temperature ; but at Dungeness, on the Atlantic side of the straits, the rainfall is very much less, the skies are clearer, the air drier, and the temperature varies much more than at the island stations. These contrasts become much sharper as we proceed northwards along the Atlantic coast, where the influence of the warm Brazil current is strikingly apparent. Passing next to the Falkland Islands, we find a strictly oceanic climate, viz. strong winds, equable temperature, and a moderate rainfall, which last, however, is probably much greater on the West Falklands than at Cape Pembroke, the station here quoted. A noticeable feature here is the large amount of sunshine recorded. Between the Falklands and South Georgia, which lies only 2 degrees to the south, but some 22 degrees to the east, a great difference is found. Here the influence of the Antarctic drift makes itself felt, the mean annual temperature being 8 degrees lower than at Cape Pembroke or at Staten Island. However, as regards sunshine and wind force, the two stations are almost identical. At the South Orkneys the influence of the Antarctic -drift is for the first time the predominant factor affecting climate. Here, in the low latitude of 61° S., the mean summer temperature is below freezing point, while in the winter readings below — 40° have been recorded. Perhaps the most striking example of the effect of the Antarctic drift on the climatic conditions in low latitudes as compared with the North Polar regions occurs during summer. At this season the isotherm of 34° between the long. of 10° E. and 50° W. nowhere protrudes further south than 58° %., and falls to about 52° to the east of the meridian of Greenwich. The most southerly extension of this isotherm in the Arctic is in latitude 74° N., long. 16° W., while to the north of Spitzbergen it lies in latitude 81° N., long. 10° E. Thus on THE WEDDELL QUADRANT AND ADJACENT AREAS. 107 this meridian of 10° E. the summer isotherm of 34° is located only 540 miles from the North Pole, but, as we have seen, quite 2300 miles from the South Pole. Such is the great effect of the Antarctic drift-ice in the lowering of temperature. With reference to the characteristics of the strictly Antarctic climate, which form the principal theme of this memoir, | may briefly summarise the conditions as follows. Stations such as the South Orkneys, Port Charcot, and in a less degree Snow Hill and the south of the Bellingshausen Sea, are strongly influenced by the ocean to the north. At these places great variations of temperature and weather occur, especially in winter, according as the influence at work is continental or oceanic. With southerly winds the skies are clear, the air dry, and the temperature low ; but with a change to the north, exactly the reverse conditions obtain. For this reason the summers are much cloudier than the winters, and it is then that the maximum precipitation occurs. The tempera- ture does not fall so low as one would expect—from — 40° to —45° F. The maximum temperature depends entirely on the situation. In the open sea or in the pack it rarely rises in summer more than 2 or 3 degrees above the freezing point, but near the land it may rise to 40° F., and at stations such as the South Orkneys and Snow Hill (which are subject to occasional visitations of fohn) the thermometer may rise even in winter to from 45° to 50° ; however, these high temperatures rarely last more than a few hours. In the matter of sunshine, great differences are found,—Port Charcot in 1904, for instance, recording 32 per cent. of the possible, with 140 sunless days, while at the South Orkneys only 13 per cent. was registered, with 189 sunless days. Tn this connection I have prepared a map (see PI. IV. fig. 5) showing the mean tem- perature of the sea surface in summer. In the preparation of this I have employed all the existing material. The bending of the isotherms to the north is clearly shown ; also the higher temperature of the sea near the South Shetlands and to the west of Graham’s Land, due to the north-east winds driving the warm surface waters before them. It may be incidentally mentioned that, as one would expect, there is a close relation between the summer sea temperature and the distribution of fog. Fogs are compara- tively rare between 40 and 50° %., west of 30° W., but between these latitudes, from 30° W. to the meridian of Greenwich, fog is very frequent, amounting on the average to 13 per cent. of the observations. The indirect effect of the southern ice is apparent even to the north of latitude 40°, a little to the west of the meridian of Greenwich, on which longitude the greatest northern extension of the polar water occurs. (See plate vii. of Captain Campsett-Hepworrn’s paper “On the Relation between Pressure Temperature and Air Circulation over the South Atlantic Ocean,” Meteorological Office, London, 1905, official No. 177.) Having described the broad climatic features, I now proceed to deal with the months in detail. In January there are two well-defined areas of low pressure; one to the west of Graham’s Land in about 64° S., the other over the Weddell Sea. Round the western low-pressure area the winds blow spirally inwards, being N.W. in lat. 60° S., long. 70° 108 MR ROBERT C. MOSSMAN ON THE METEOROLOGY OF W., and N.E. in the vicinity of and to the south of the South Shetlands. On the west side of Graham’s Land from 63° to 64° §. lat. winds are somewhat variable, and the combined values from the various summer expeditions give no great preponderance of any one wind. It is probable that this region is alternately influenced by the above two low-pressure areas. ‘The Antarctic high is considerably reduced, and is shown by the isobar of 29°30 inches which covers the extreme south of the Bellinghausen Sea, the winds being almost wholly from the north-east, east, and south-east in 70° S., 87° W. The strongest winds in this region are from the north-east, and the lightest from the west. As compared with the previous month, temperature has risen everywhere, the rise being from 2 to 3 degrees, except at the Antarctic stations, where it amounts to 1 degree only. The isotherms have assumed their typical summer form, bending considerably to the south to the west of 50° W. A noticeable feature in this connection is the high temperature and dry atmosphere which prevails over Tierra del Fuego and Patagonia to the south and west of Sandy Point. At most places, except the Antarctic and strictly oceanic teak ae temperature is now at its annual maximum. The amount of cloud has considerably increased over the region south of 60° 8., but has diminished over the mainland of South America. The isoneph of 95 now appears over the northern part of the Weddell Sea, or over that region where the ocean temperature is relatively high. Rainfall is now at its annual maximum over a region running along the 53 degrees of latitude between the meridians of 68° and 75° W., reaching a total of 12°82 inches at Evangelists Island on the Pacific side of the straits, where north-west winds blow with greater frequency than during any other month of the year, and, since they are on- shore winds, are accompanied by heavy rainfall. The temperature-difference between the east and west sides of the Straits of Magellan is also at a maximum, Dungeness being 6°°2 warmer than Evangelists Island, as a result of the excessive precipitation and overcast sky prevailing at the latter region. At Ancud, Port Gallera, and Isle Mocha rainfall is at a minimum for the year, this region being now well within the influence of the South Pacific high. In February, the most marked feature in the pressure-distribution for the month is the complete recession of the Antarctic high from the area under discussion. This fall of pressure is most pronounced in the Bellingshausen Sea, and extends, though in a much smaller degree, over to Graham’s Land and the Weddell Sea. Low-pressure areas are again shown to the east and west of Graham’s Land. The former low- pressure area probably extends a very long way to the east, with the isobar of 29°2 assuming a northerly trend east of the meridian of Greenwich. Observations over a combined period of forty-five days show a reduction equal to 0°22 inch between situations in 37° W. and 64° E. longitude on the parallel of 624. The few observations that we have south of the Antarctic circle show but little difference of pressure between 23° W. and 50° E. THE WEDDELL QUADRANT AND ADJACENT AREAS. 109 The wind observations in the Biscoe Sea indicate that the Antarctic high is situated to the south of 66° S. in about 40° E., and that the gradients are exceedingly steep for southerly winds, which blow with great force. There appears to be a well-defined low-pressure area in about 64° to 65° S. in 40° E. | Temperature, as compared with January, shows little change over the region south of 60° S., being, on the whole, very slightly higher. It reaches the maximum for the year at the South Orkneys, and is virtually the same as that of the surrounding ocean. The lowest isotherm, viz. 25°, is shown over the south of the Weddell Sea, but observations are too few for its position to be accurately defined. Except at the Falkland Islands and Sandy Point, temperature has decreased 1 or 2 degrees over the area from 40° to 50° 8. Cloud amount continues very large over the greater part of the Antarctic and sub- Antarctic regions, but over Graham’s Land (where there is a marked excess of south and south-east winds) a comparatively small amount is recorded, this being associated with a relatively dry atmosphere. In March, pressure has risen slightly in the Bellingshausen Sea, but has fallen over Graham’s Land and the South Orkneys, being at its annual minimum at most places south of 50° S., with South Georgia as the only marked exception. The characteristic features of pressure-distribution remain substantially the same as in February, but both the eastern and western lows have deepened, and as pressure has given way more in the south than in the north, a general steepening of gradients has taken place between 50° and 60° S.; causing a marked increase in gale-frequency to the south and south-east of Cape Horn, on the Chilian coast south of 50° 8., and on the east side of Graham’s Land. North-west winds are now at their annual maximum at Staten Island and the Falkland Islands, and over a considerable part of this area it is the stormiest month of the year. In the Weddell Sea the barometric gradient is comparatively slight until about 68° 8. lat., and the ‘‘Scotia” observations, comprising two March months, show a steady diminution of pressure to at least 72° S. It is probable that during this month the barometric trough in the Weddell Sea occupies its most southerly position of the year, being probably located in about 70° S. lat. on the meridian of 20° W. Hasterly gales are comparatively frequent south of 67° 30’ S., pointing to very steep gradients at times in this region. The same conditions prevail in about 71° S. and 93° W., where these winds blow with the maximum force of the year. Temperature has everywhere fallen. At Wandel Island, the South Orkneys, South Georgia, and the Chilian coast stations, which are largely under oceanic influence, the fall amounts to only 1 degree, but in the Bellingshausen Sea it reaches 5 degrees, and at Snow Hill as much as 9 degrees. The isotherms show a very steep thermal gradient between the east and west of Graham’s Land. It is evident that the influence of the land area, which is shown to be probably located to the east in 66° S. and 47° W., is making itself felt. 110 MR ROBERT C. MOSSMAN ON THE METEOROLOGY OF Cloud amount is still very large over the Weddell Sea, but over Antarctic areas, where the temperature has fallen, skies are clearing. In April, pressure has risen everywhere except in the south of the Bellingshausen Sea. The most marked rise is over Graham’s Land, over which there is an area of relatively high pressure, dividing the low-pressure systems located to the east and west respectively. In these low-pressure systems the strength of the wind reaches its annual maximum, this being especially the case in the south of the Bellingshausen Sea, where the prevailing easterly winds blow with great force. Indeed, atmospheric conditions are everywhere very disturbed, and especially so in the South Atlantic and the vicinity of Cape Horn. The only marked exceptions to this are to be found over Graham’s Land and at Dungeness, where the infrequency of gales is remarkable. There is in this month a decided tendency for the winds to veer. Thus at the South Orkneys south-west winds are at their annual maximum, and at South Georgia, Staten Island, and the Falklands the prevailing north-west winds are reduced in favour of west and south-west. On the other hand, at Dungeness, Evangelists Island, and Sandy Point northerly and north-east winds, which are infrequent for these places, have increased, and are at their annual maximum for the two first-mentioned stations. Temperature is now falling rapidly, the fall being greatest at the southern continental stations and least at oceanic stations in the south of South America. At places such as Sandy Point and Ushuaia, which are not specially under oceanic influences, the fall of temperature is also considerable. In this month the isotherms begin to assume their winter shape, and the amount of cloud shows a decided diminution north and south of 60° S. lat., where the mixture of warm and cold air currents is most apparent. At Ancud, Point Gallera, and Isle Mocha the increase in rainfall is considerable, the precipitation being four times that of January—the driest month of the year. In May, the outstanding feature of the isobaric chart is the reappearance of the Antarctic high in a pronounced form in the south of the Bellingshausen Sea and adjacent continental regions. The south-east wind, although here still the strongest, has greatly diminished in force, and this reduction is shared by winds from other directions. Areas of low pressure are shown in the South Pacific and over the Weddell Sea. North-east winds are virtually absent at Wandel Island, which appears to be more under the influence of the high- than of the low-pressure area. South of 60° S. the temperature has fallen from 6° to 8°, the fall being greater on the eastern than on the western side of Graham’s Land. At the South American and Island stations the fall is from 3 to 5 degrees, and is greatest, as in the previous month, at places removed from the direct influence of the sea. The mean temperature has fallen to zero I. a little to the south-east of Snow Hill, owing to the continental conditions there prevailing and the northerly extension of the pack. On the other hand, on the Pacific side of Graham’s Land, where the oceanic influence is considerable, much higher temperatures prevail. Cloud has very generally diminished, the cloudiest zone THE WEDDELL QUADRANT AND ADJACENT AREAS. et being located further north than in April. The isoneph of 85 now indicates approxi- mately the position of the pack edge at this season. A curious point in regard to the cloud-distribution in the region of the Straits of Magellan is shown by the circumstance that while the maximum for the year is reached at Sandy Point, the minimum is recorded at the neighbouring stations of Kvangelists Island and Dungeness. Rainfall shows a large increase at Ancud, Point Gallera, and Mocha. In June, pressure has risen very generally in the Antarctic and sub-Antarctic regions. It has, however, fallen at the Northern Chilian stations, where it is at the annual minimum, while at the same time rainfall, cloud, humidity, and wind-force are at their annual maximum. The general distribution of pressure remains substantially the same as in May. The Antarctic high is in the same position, but has increased in height: the cyclonic areas in the Weddell Sea and South Pacific are still prominent features of the map, but with pressure about 0°10 inch higher than in May. Tempera- ture has fallen about 6° at the Antarctic stations, but elsewhere only half this amount, if we except Ushuaia, where the increased cold of radiation has produced a fall of 5°. The isotherms have now assumed their normal winter position, the most marked feature being the steepening of the thermal gradient south of 60° S. Owing to the extension northward of continental conditions brought about by the freezing of the ocean, the amount of cloud has largely diminished, the cloudiest zone being indicated by the isoneph of 80, embracing and to the east of Staten Island, a similar high value being reached off the Chilian coast in 40° 8. Generally speaking, relative humidity is now at its annual maximum. A marked feature is the excessive rainfall on the Chilian coast in 40° 8., where Point Gallera has a mean fall for the month of 19°18 inches. The precipitation diminishes north and south of this latitude. Staten Island also has its maximum fall for the year, viz. 6°04 inches. At South Georgia there occurs the minimum for the year. In July, the Antarctic high is shown in about 70° S. and 80° W., with a low- pressure area to the west-north-west and another over the Weddell Sea.* Pressure has fallen over the whole area to the south of 50° §., west of the meridian of 60° W. To the east of 60° W. it has risen most at the Falkland Islands. Southerly and south-west winds are at their annual maximum over large portions of the region comprised between latitudes 52° and 65° 8. and longitudes 72° and 65° W., gales from these quarters being frequent to the south of Cape Horn. Temperature is now at the minimum for the year, except at places such as Staten Island, which are strongly under oceanic influence. At this season the greatest cold is * The long series of observations at the South Orkneys indicate a distinct winter maximum of pressure which is high for the latitude as compared with that recorded in winter by the “ Belgica” in long. 87° W., and it would appear that the Antarctic anticyclone in winter is central in about 80°S. and 40° E. That the centre is there rather than at the Pole is connected with the continental conditions induced by the freezing of the Weddell and Biscoe Seas as far north as 60° S. lat., and the relatively higher temperatures that prevail on the other side of the Pole in Victoria Land. TRANS. ROY, SOC. EDIN., VOL. XLVII. PART I. (NO. 5). 16 112 MR ROBERT C. MOSSMAN ON THE METEOROLOGY OF felt at Snow Hill, where the continental effect is most pronounced ; the South Orkneys also come well within the sphere of this influence. The lowest isotherm, viz. that of — 10°, is indicated to the east of Graham’s Land in about 66° S8., the relative mildness on the same parallel 10 degrees to the west on the Pacific side being a noticeable feature. The thermal gradient continues very steep in sub-Antarctic and Antarctic regions. The cloudiest zone is indicated in from 57° to 60° 8. by the isoneph of 80, the lowest values being 50 over the coast of Graham’s Land, and 60 in Tierra del Fuego. On the Pacific side of the Straits of Magellan rainfall is at its annual minimum. In August, cyclonic conditions prevail very generally, pressure having fallen except over the extreme south of South America. The fall is greatest—-about a tenth of an inch—over Graham’s Land and to the west. The isobaric chart shows a well-marked cyclonic area over Graham's Land, and steep gradients to the south and west of Cape Horn. Associated with this is a decided increase in storm frequency. North-east gales blow with great persistence and strength to the south of the South Shetlands. A small area of relatively high pressure is shown in the Bellingshausen Sea, and in this region, west of about 85° W. and south of 65° S., winds from all quarters blow with average strength, thus showing no indication of steep gradients. ‘Temperature south of 60° S. has risen decidedly, but in strictly oceanic situations in lower latitudes the rise is small. The isotherms present much the same aspect as in July, but very clearly indicate the greater rise of temperature in the south. Cloud shows a marked increase in high southern latitudes, especially to the east of Graham’s Land, this being explained by the greater condensation accompanying the increased prevalence and strength of warm winds from lower latitudes. In Tierra del Fuego, in places removed from oceanic influence, the least amount of cloud for the year is recorded. In September, the first spring month of the southern hemisphere, the changes in pressure are somewhat irregular. There is a noticeable fall at the South Orkneys, and a slight increase, extending from 55° to 65° S., between the meridians of 67° and 55° W. The most interesting feature of the isobaric chart is the total absence of any indication of the Antarctic high, there being two large low-pressure areas, one over the Weddell Sea, the other to the west of and partially covering Graham’s Land. Gradients over the larger part of the area south of 50° S. are comparatively uniform, except in the Cape Horn region, where they are steep. Temperature has risen everywhere. ‘The rise is greatest south of 60° S., amounting to 5° or 6°, but elsewhere (with the exception of Ushuaia and Sandy Point) it does not reach half this amount. ‘The isotherms north of from 63° 8. on the 80th meridian of W. long. to 57° S. on the 40th meridian have opened out considerably, thereby show- ; ing that the continental influence of the frozen sea is becoming restricted. The thermal gradient is still very steep between the Pacific and the Weddell Sea side of Graham’s Land. Cloud shows a comparatively uniform distribution, but has increased in the far THE WEDDELL QUADRANT AND ADJACENT AREAS. 1138 south as compared with the previous month. At Sandy Point, however, it falls to the annual minimum. In October, pressure has fallen very generally west of the 55th meridian, but at South Georgia and the South Orkneys it has increased. The east and west low- pressure areas have become merged into one, whose centre is normally located over Graham’s Land, but which, judging from the winds at Port Charcot, is probably frequently central to the W.N.W. Gradients are steep west of the 60th meridian, and gales are at a maximum on the South Chilian coast and Cape Horn region, a relatively large number being from the 8. W. Temperature is now rising rapidly south of 60° S., the increase being most pro- nounced at continental stations. The rise over Tierra del Fuego and adjacent islands is remarkably uniform. The isotherms continue to open out and the thermal gradients diminish. At Staten Island and Dungeness calms are at their annual maximum, a condition offering a marked contrast to the stormy weather on the Pacific side of the straits. The rainfall has increased on the Pacific side of the Magellan Straits, but in lat. 40 §. on the Chilian coast the precipitation is only about half that of the previous month. The changes shown in the isobaric chart for November are remarkable. Pressure has risen greatly over Graham’s Land, and to a less extent in the Bellingshausen Sea, while it has fallen appreciably at the South Orkneys, South Georgia, and Tierra del Fuego. An anticyclone is in process of formation on that portion of the Antarctic continent to the south of the Bellingshausen Sea and far south of Graham’s Land. North-east winds prevail on the Pacific side of Graham’s Land, and east and north-east in the Bellingshausen Sea, where they blow with great strength. The cyclonic area over the Weddell Sea in this month is probably central about the latitude of the Antarctic circle in 10° W., with the isobars tending to the north on proceeding east of the meridian. In connection with this relatively higher pressure to the north, one may note that during the summer voyage of the “ Valdivia” the west wind system was left behind in November in as low a latitude as 56° S. in 20° H., while east winds prevailed to 65° S. between the meridians of 20° to 60° E. On the return voyage in December the west winds were again experienced in lat. 55° 8. While the position of the bottom trough on this voyage may have been located a degree or two further north than usual, it is still highly probable that the general condition is as described. That a great and widespread change of pressure distribution is in progress is evident. Speaking broadly, the month to month changes of pressure in Graham’s Land and Tierra del Fuego are in the same direction, but in November, as will be seen, they are not only the reverse of each other, but that to a very large degree. Pressure has changed little at the Falklands, and has fallen much more on the Pacific than on the Atlantic side of the straits, the result being steep 114 MR ROBERT C. MOSSMAN ON THE METEOROLOGY OF gradients for north-west winds south of 50° S. on the Pacific side, with comparatively slight gradients and a subnormal gale-frequency on the Atlantic side. This latter condition extends south of Cape Horn. Temperature has risen uniformly at all places south of 60° S., where the increase amounts to 6. Elsewhere, within the limits of the area under discussion, the rise is remarkably uniform, being from 3° to 4°. The isotherms present few features of interest, with the exception of a relatively low temperature in from 51° to 55° 8. between the meridians of 65° and 70° W. ‘This appears to be associated with an increase of south-west winds over the above region, a concomitant of the pressure-changes now in progress. The cloudiest zone, indicated by the isoneph of 90, is now well south of the 60th parallel, and on the Pacific side reaches to the Antarctic circle. North and south of this a decrease of cloud-amount is observable, the diminution being most marked in the neighbourhood of Staten Island, where it is at its minimum for the year. In December, the most marked change in the distribution of pressure is a small but general rise over Antarctic and sub-Antarctic regions. Elsewhere pressure has fallen, but not to any appreciable extent, being most noticeable at Sandy Point, where it amounts to 0°08 inch. A well-defined area of relatively high pressure is shown over Graham’s Land (where pressure is at its annual maximum), and in the south-east of the Bellingshausen Sea. To the east and west of this anticyclonic region low- pressure areas are shown. Gradients are slight everywhere, and gale-frequency has greatly diminished. From about 30° W. to east of the meridian of Greenwich, in from 50° to 60° S., gales have increased, pointing to a steepening of gradients in this region, south of which, however, there are no data. Winds from quarters to the south of west are unusually frequent in the South Orkneys, Tierra del Fuego, Evangelists Island, and the Falklands; while at the latter station, as well as at South Georgia, south-east and east winds are at their annual maximum, pointing to the effect of the widespread changes of pressure referred to. Temperature in this, the midsummer month of the southern hemisphere, has risen most in the Antarctic, where the powerful influence of the midnight sun is asserting itself. The rise is least in the South Orkneys, where cold southerly and south-westerly winds blow with greater frequency than in any other month of the year. Cloud-amount has increased over the ocean on the Pacific side of Cape Horn south- ward up to the limit of observation, the maximum for the year being very generally reached in this area. While the characteristic features of the various months are as I have described, yet from time to time abnormal conditions prevail. I have selected December 1904 for special study because of the unusual pressure distribution and wind-circulation. Normally, there is at this season an area of relatively high pressure over Graham’s Land, with low-pressure systems to the east and west, and an increase of pressure to the north of 58°, with isobars running along parallels of latitude. But in the month under THE WEDDELL QUADRANT AND ADJACENT AREAS. ESS) consideration the isobars west of the 65th meridian (and probably farther west) lay approximately north and south, with pressure as high over Graham’s Land and Cape Horn as at the Falklands (see Plate IV. fig. 4). At the South Orkneys, pressure, although above the average, was 0°30 inch lower than at Charcot’s Winter Quarters, indicating a still lower barometer to the east. The prevailing winds over a very wide area south of 50° S. on the Pacific and 40° on the Atlantic side were from the south or south-west. That these were also the prevalent winds between the South Shetlands and the South Orkneys is evident by the low latitude in which the Argentine corvette “Uruguay,” on her voyage to the South Orkneys in 1904, met with the pack, this being encountered in lat. 58° 40’, long. 50° 40’ W. The vessel arrived at the South Orkneys on 31st December, having forced her way through 250 miles of ice. On her voyage to Gerlache Straits I had an opportunity of examining the pack. I perceived that it showed hardly a trace of having been subjected to oceanic swell, while the presence of unbroken ice fields as much as 30 miles in length, not far removed from the outer edge of the main body, in lat. 59° 5’, long. 49° to 50° W., was further proof of the absence of the normally prevailing westerly and north-westerly winds. I have referred to these facts as they bear directly on the question of the prevailing winds over the region and the trend of the isobars. That unusually quiet weather prevailed to the west of 65° W. and at Cape Horn is indicated by the total absence of strong winds at Dungeness and Evangelists Island, while the comparative rarity of gales at Charcot’s Winter Quarters is also worthy of notice. At the South Orkneys, however, south-west gales were frequent, especially after the middle of the month. On the mainland of South America the most pronounced feature was the persistence of a low-pressure area off the coast of Chili in about lat. 40° 8. This type of pressure-distribution prevailed on nineteen days of the month, but on the chart for the whole month the more distinctive features are masked by the normal conditions prevailing during the remainder of the month. ‘This is the type of pressure-distribution associated with the presence of an Antarctic high at all seasons, and these highs appear to pass away to the N.E. and over the Atlantic. In the table on the following page are given values of the principal climatic elements for ten stations south of 40° S., and, as far as the data permit, they have been compared with the normal. It will be observed that at Charcot’s Winter Quarters temperature was about the normal for the region, but that at the South Orkneys and at the island and coastal stations on the east side of the southern extremity of the American continent it was markedly below the normal, the deficit being practically the same, viz. 2°. On the other hand, it was markedly above the normal at Ushuaia, the excess increasing in a N.W. direction, and reaching 2°7° at Evangelists Island, where rainfall, cloud, and wind-force were all much below the average. This result is obviously due to the relative infrequency of the normal cool, onshore, rain-bearing winds, and the excess of land winds resulting from the abnormal distribution of pressure referred to. It will be observed that while southerly winds were in excess at the South Orkneys, a relatively 116 MR ROBERT C. MOSSMAN ON THE METEOROLOGY OF large number of N.E. and E. winds, blowing with considerable force, occurred at the Falklands and New Year's Island. Reference to the daily weather maps of the Argentine Meteorological Ottice show that these winds were associated with the passage of low-pressure systems from the Argentine coast north of 50° 8. An analysis of the C. Pembroke barometer showed that the north-west wind experienced before the passage of the low-pressure area, and the south-east wind which blew after the passage of the low-pressure area, had practically the average pressure of the month, but that with the DECEMBER 1904. MEAN PREsSURE AT 32° SEA-LEVEL AND STANDARD Gravity ; MEAN TEMPERATURE REDUCED TO SEA-LEVEL; RAINFALL, CLoupD AMOUNT, WIND DIRECTION AND FoRCcE At various Stations from 40° to 65° South. Percentage Frequency of Wind. Wind Force or Velocity. Station. Lat. | Long. W. |Pressure.|/Temp.| Rain. Glens ar ama 7 : = $ Miles per | Estimated N.|N.E.}) E. |S.E.) S. |S.W.| W.| N.W.| Calm. Teinthe Scale 0-12. | ae eS Ins. 2 Ins. Port Charcot . . | 65 03 63 26 29°679 3152 | 1:21 8:0 2} 19 2 5 | 37 | 22 1 1 11 ate South Orkneys . . | 60 44 44 39 29°374 28°8 | 0°19 9°5 0 0 0; 8 | 41] 29 | 15 7 0 15°5 nie Cape Pembroke -| 51 41 57 43 29°695 | 45°5 oo 81 4 TO eR abe EX a Wie 8 0 fe 4°3 New Year’sIsland . | 54 39 64 07 29°693 44:8 | 1°99 85 9] 16 | 11 6) 12) 22 sl 10 0 15°5 Ci Ushuaia . - » , 04 52 68 07 29°707 50°6 | 2°64 ve S0 are Salat i) Je He ‘fe 45 ame Harbertown - . | 55 00 67 00 29°704 51°4 | 1'39 aye qo ai fe a0 a6 Ao ; Dungeness . «| 62°24 68 25 29°734 50°2 | 2:38 75 6 10 6 2 0) 29) | 16 e 25 2°6 Evangelists Island * 52 24 75 06 ? 49°3 | 6°46 Ut 6 0 AN SE LO A zou 1S Hat 30 Ancud 5 41 51 73 50 29°984 57°9 | 3°22 56 12 al 2 1)| 16 2a 28 11 30 Point Gallera . 40 OL 73 44 29°969 55'8 | 3°10 bl 14 0 | 0 1 52 | 3 6 9 15 374 * The barometer readings are of doubtful value at this station. DECEMBER 1904. DEPARTURE FROM THE AVERAGE. Norr.—The heavy type indicates an excess, and the italic a defect. Percentage Frequency of Wind. Wind Force. Station. Pressure.| Temp.} Rain. | Cloud. a | Miles per Scale N. |N.E.| E. | S.E./ S. | S.W.] W. | N.W.) Calm. signi 0-12. Ins. - Ins. Port Charcot A : 348 0-0 “3 10 te ae oe ae 58 ee a4 te, South Orkneys . 096 1:8 - 0:3 6 5 8 1 19 5 7 h ae 29 a0 Cape Pembroke . C 084 21 ay 0-9 a at se a ae ae se # : New Year'’sIsland . 249 20 1°82 11 1 ll 10 h h 6 2 16 6 30 5 Ushuaia : . c “263 13 0:69 a 0 a ae 56 i a8 5 An ae a Dungeness . ; 5 “196 0-9 1:21 0-7 2 7 4 0 2 21 uf 4 13 0-8 Evangelists Island + Ke 27 3°82 0:3 5 0 3 0 5 2 0 16 15 12 Ancud . 5 5 * 019 0-6 0°18 0-2 0 3 0 1 8 8 4 uf 7 06 Point Gallera : , O43 O01 0:11 03 0 0 1 1 1 6 1 0 6 0-2 east wind pressure was 0°114 inch below the normal. Captain Hepworru has referred* to the remarkable prevalence of easterly gales in the months of October and November, between the meridians of 30° and 40° W., from 45° to 50° 8. It is very probable that these gales are due to the passage of low-pressure systems which have travelled sea- wards in an easterly direction from the Argentine coast. With respect to the bearing of the South Orkney and other Antarctic observations on the weather of South America, it is proposed to prepare charts for each month, * “The Relation between Pressure Temperature and Air Circulation over the South Atlantic,” Meteorological Office, London, 1905, Official No. 177, p. 9. THE WEDDELL QUADRANT AND ADJACENT AREAS. iI ag showing the conditions over the continent for the whole period of recent Antarctic research. In this connection the South American data will be jomed up with that furnished by the Falklands and South Georgia. Mean ATMOSPHERIC PRESSURE TEMPERATURE AND CLOUD FOR THE YEAR. The maps showing the mean annual barometric pressure, mean temperature, and mean amount of cloud may be said to represent the sum and substance of the year’s weather (see figs. 1, 2, and 3, Plate IV.). The isobaric charts indicate in the clearest manner the relatively higher pressure which prevails to the south over the Bellingshausen Sea, as compared with that over the Weddell Sea in similar latitudes, and the wind-circulation follows the pressure- distribution. The Peninsula of Graham’s Land thus divides two wind systems. To the west we have, as at Wandel Island, a marked prevalence of strong north-easterly winds, due, as we have seen, to the all but permanent cyclonic area to the westward. To the south of this is the region embraced by the “ Belgica’s” drift, in which the wind- circulation is of a distinctly monsoonal character—easterly in summer, and westerly in winter. Here the easterly winds are by far the strongest, indicating the steepness of the barometric gradient between the Polar anticyclone and the South Pacific low- pressure area. The anticyclone migrates with the season—facing the Pacific Ocean in summer, and retreating over the continent to the south of the Indian Ocean in winter. While these conditions represent the normal state, different seasons appear to diverge considerably from the average. Thus in 1903 the Weddell Sea “low pressure” was deepened, and lay further to the west than usual. As pressure was about the normal north of 50° S., a very steep gradient was set up for W.N.W. and N.W. winds. That this rapid fall of pressure prevailed to at least 65° S. is shown from NoRDENSKJOLD’S Snow Hill observations. In 1904, on the contrary, the pressure gradient was less than half that of the previous year, as shown by the simultaneous observations at the Falkland Islands and the South Orkneys. Between these two stations the mean difference in the barometric pressure for the nine months April to December was 0°456 in 1903, while in the corresponding period of 1904 the difference was only 0199 inch. The remarkable persistence of easterly winds during the winter of 1902 at the winter quarters of the Gauss in 90° E. indicates that the Antarctic anticyclone lay over the frozen continent to the south and south-west of this station (see footnote, p. 111). Pending the publication of the detailed official reports of the expeditions which have taken part in the recent International Antarctic campaign, it would be premature to draw final conclusions, but a careful analysis of all the available material enables us to recognise four regions characterised by permanent low-pressure areas. These are central in the Ross Sea in lat. 70° 8., and long. 170° W.; in the Bellingshausen Sea in 65° 8. and 90° W.; in the Weddell Sea in 66° S. and 30° to 35° W.; and in the Antarctic Ocean in 63° S., between the meridians of 80° and 105° EK. It will be seen 118 MR ROBERT C. MOSSMAN ON THE METEOROLOGY OF that, with the exception of this last-named low-pressure area, these cyclonic centres are situated in those regions where the ocean extends furthest south and where the forces exerted by the mixture of the rivers of cold desiccating air from the Antarctic continent and the humid air of the oceans are ata maximum. As bearing on this point, it may be noted that every occurrence of “Glazed Frost” seen on the voyage of the “ Scotia” in the Weddell Sea was observed in from 67° 10’ to 67° 39’ S., and that at the South Orkneys this phenomenon is at a maximum in winter, when the theatre of cyclonic activity is north of its summer position. It will be further noticed that these low-pressure regions are situated approximately in the longitudes occupied by the permanent southern high-pressure belts, and, like them, are subject to seasonal oscillations. Judging from the summer observations, the winds in and near the centre of these permanent low-pressure areas are not of much force, the ring of steep gradients being, as one would expect, where the Antarctic anticyclone is most developed with reference to the contiguous low-pressure system. In the Weddell Sea the effect of the wind-circulation is clearly shown in the direction of the sea surface currents. The observations of Weddell and the “Scotia” show that there is a strong set to the N.W. to the south of 66° S., but in from 64° to 65° S. on the meridian of 40° W. the set changes to the E. and E.N.E., which is the general direction of the surface drift in lower latitudes. Not only so, but in the region dividing these two currents a large number of weatherworn bergs were seen, which, judging from their general appearance, had been in the vicinity for a long time. This region may therefore not inappropriately be termed the ‘‘ Weddell Sea Doldrums.” At the South Orkneys the whole set of the pack and of the icebergs is to the H.N.E., and the few cases of opposite movement recorded were probably a tidal effect. The circumstance that there is a distinct seasonal change in the position of these “centres of action” is evident. Not only do the surface winds change with the season, but these changes are also clearly observable at the cirrus level; thus in summer ) the prevailing direction of the upper clouds at such places as South Georgia and the South Orkneys is from the west and north-west, but im winter the upper currents show a large movement from the south and south-west. This indicates that at the cirrus level (which in these latitudes is at a comparatively low altitude) the low-pressure area to the south-east has swung more to the north. At the “ Belgica’s” position in 70° to 71° S., long. 80° to 90° W., exactly the reverse obtains ; and it is of interest to note that the maximum rate of movement was from the north-east at both seasons. The whole question is exceedingly interesting, but it is not safe to draw conclusions pending the publication of the reports of the various expeditions. It is unnecessary to dilate at length upon the maps of mean annual temperature and cloud, as the factors affecting them have been fully discussed under the monthly headings. Attention may, however, be called to the bending of the isotherms south- ward west of 55° W., asa result of the prevailing north-east winds. On the other hand, the crowding of the isotherms on the east side of Graham's Land is very striking. Here THE WEDDELL QUADRANT AND ADJACENT AREAS. eds) the thermal gradient is exceedingly steep, as shown by the synchronous observations made by the Swedes at Snow Hill and the adjacent station of Paulet Island, some fifty miles to the north. In this region the temperature falls off at the rate of about 6° to 7 F. for every degree of latitude: the abrupt bending of the isotherms to the south about the meridian of 40° W. is clearly due to the presence of the extensive land mass indicated by Dr Brucs on his bathymetrical chart in lat. 66° S., long. 47° W. All the varied information which I have collected bearing on this point affords strong evidence of the existence of a land mass in the assigned position. In this connection | may incidentally refer to some conclusions | have arrived at with reference to the probable existence of a peninsula or archipelago east of King Hdward VII. Land. If the observations taken by Ross in the year 1842, between 65° and 67° S., and long. 154° to 160° W., represent the normal meteorology of this region in summer, there is strong evidence that King Edward VII. Land sweeps round in a N.N.E. direction, with its northern apex in lat. 69° S., between the longitudes of 150° and 155° W. To the east of this the land probably falls away in a gentle curve, joming Graham’s Land in 71° S. The whole wind circulation in the Ross Sea, as well as other meteorological concurrents, support the conclusion that we have to do with conditions somewhat similar to those obtaining in the Weddell Sea area. This calls for a peninsula situated perhaps even further west than I have indicated. The map of mean annual cloud-amount shows distinctly the distribution of this element in its relation to land and sea. Over Graham’s Land and adjacent regions the skies are relatively clear, but where the pack ice and open ocean meet there is a zone of much cloud. ‘The increased clearness of the skies on the east as compared with the west side of the continent of South America is also of interest, as is the compara- tively small cloud-amount of inland stations in Tierra del Fuego. With regard to future work, it is a matter for satisfaction that there is no abate- ment of interest in Antarctic investigation. A British expedition is even now at work ‘in Victoria Land, where a third year’s observations will materially strengthen the normals for that locality. The same considerations apply to Graham’s Land, towards which a French expedition, now on the eve of departure, is to direct its efforts. Several prospective expeditions have announced their programmes, the realisation of which is earnestly to be desired. Meanwhile, the energy of the Argentine Government, as reflected in Mr Davis’ efforts. to establish the South Orkney station on a permanent basis, remains undiminished, while further developments, which it is hoped to realise in the immediate future, will enable us in time to come to discuss in a manner more thorough than is now possible the meteorology of the Weddell Quadrant and adjacent areas. GENERAL SUMMARY OF THE TABLES. In the remarks which follow I shall endeavour, as briefly as is consistent with clearness, to summarise in a general manner the contents of the tables. While the titles give a definite indication of the elements to which they refer, further particulars are desirable for the sake of clearness. Tables I. to VII. give for various stations from TRANS. ROY. SOC. EDIN., VOL. XLVII. PART I. (NO. 5). ° 17 120 MR ROBERT C. MOSSMAN ON THE METEOROLOGY OF 40° to 70° S. the mean monthly and annual values of barometric pressure, air tempera- ture, amount of cloud, rainfall, wind velocity, relative humidity, and percentage frequency of wind. In their preparation I have used all the reliable material, but have discarded returns from stations the observations from which did not inspire one with confidence. Thus the barometric pressure is not given for Evangelists Island, as the readings are obviously in error. The data from Tierra del Fuega have been carefully sifted, especially with reference to mean barometric pressure. Particular trouble was taken to obtain good barometric means for the region around Cape Horn, which is of great importance as a general control on the isobars. With regard to temperature observations, Table II., the means given are those observed, but in plotting the values for drawing the isotherms the values were corrected to sea-level by adding 1:0° F. for every 300 feet. As only two stations exceed 150 feet, these corrections were small. No correction was made to bring the values to the mean of the 24 hours, as, except New Year's Island, which was specially treated, the means of the tri-daily observations give a very close approximation to the mean of the 24-hourly values. As regards New Year's Island, the hours of observation were 7, 8,10 a.m., and 1, 4, 7, and 10 p.M., and the mean of these was brought to the mean of the 24-hourly values by adopting the values obtained from a reduction of the thermograph records at the adjacent station of Staten Island. The monthly corrections, which are all minus, are as follows :— Jan. Feb. Mar. April. May. June. July. Aug. Sept. Oct. Nov. Dec. Year. 09 08 O7 06 05 O8 O1 O02 O04 O06 OF O08 0-45 The mean monthly and annual amount of cloud is given in Table III. Table LV. gives the mean monthly and annual rainfall. As regards Sandy Point and Dungeness, the values appear to be too low. This is due in the former instance to the rain gauge being at some distance above the ground. Owing to the frequency of drifting snow at the South Orkneys, the precipitation there cannot be given, but, on a review of the whole existing material, I am inclined to assign a much larger precipitation to this” region than that given in my report.* I think about 25 to 27 inches is the probable annual rainfall, with a maximum in summer. As Laurie Island is on the lee side of the South Orkney group with reference to the prevailing westerly winds, it is probable that the precipitation is much greater on the neighbouring Coronation Island to the west. Table V. gives the mean hourly velocity in miles for the first six, and the mean force of the wind on the Beaufort scale 0—12, for the last seven stations. Table VI. gives the mean relative humidity, deduced in most cases from readings of the dry and wet bulb thermometer, but at Wandel Island the records are from a Richard Hair hygrograph. Table VII, showing the mean monthly and annual percentage frequency of the winds, has involved much labour, as the data for the South Orkneys, South Georgia, Evangelists Island, and Dungeness were either wholly or partially unreduced. In every case the values have been reduced to eight points, and the numerical results * Scientific Results of the “ Scotia” : Meteorology, p. 274. THE WEDDELL QUADRANT AND ADJACENT AREAS. 121 expressed as percentages. With regard to the seasonal values, it may be pointed out that Swmmer comprises December, January, and February ; Autwmn, March, April, and May; Winter, June, July, and August ; and Spring, September, October, and November. In place of the surface winds, the observed drift of the lower clouds at the South Orkneys is given. The table is not wholly satisfactory, as clear weather in winter usually occurs with southerly winds, so that the values for south-west, south, and south- east are above, and northerly winds below, the values given for this season. With regard to Dungeness, there would appear to be a local south-west wind at the Atlantic entrance to the Straits of Magellan, as this wind is unusually frequent. Table VIII. gives an analysis of the movement of the upper clouds (cirrus, cirro- cumulus, and cirro-stratus) for the months and the seasons at the South Orkneys and Cape Horn region, including South Georgia. [For the latter station and for the “Belgica” only seasonal means are given. The mean direction of the upper clouds has been obtained by the application of Lambert's formula to the annual and seasonal values. The data for the South Orkneys are for the five years 1903-1907. The table for the Cape Horn region is from one year’s observations, 1882-83, at Orange Bay and Ushuaia, and two years’ observations at New Year’s Island. Another table is given in which South Georgia (where twenty months’ observations are available) is included with the above-mentioned Cape Horn data. For the “ Belgica” an analysis has been made of the hourly cloud observations published in extenso in M. Doprowotskt's paper.* I may point out that the mean annual values given by me differ slightly from those given by M. Dosrowotski on page 58 of his report, but the difference is not of any moment. ‘The cloud velocities given for the ‘“‘ Belgica” have also been deduced from an examination of the detailed hourly observations. Table IX. gives mean monthly and annual sea surface temperature at four places, viz. the South Orkneys, South Georgia, Orange Bay, and New Year’s Island. The returns from this last-named station have been extracted from the MS. daily sheets of observations for the period to which they refer. Table X. gives an abstract by months of the observations made during summer expeditions on the east coast of Graham’s Land and in the Weddell and Biscoe Seas. A comparative series of observations made at Staten Island for the same period as that covered by Dr Brucr’s 1892-93 visit to Louis Philippe Land is also included. Table XI. contains an abstract of observations made in the months of February and March, so combined as to give means extending over a considerable number of days for definite areas. Table XII. contains similar means for the Ross Sea, calculated from the values given in “Contributions to our Knowledge of the Meteorology of the Antarctic.” Table XIII. gives the percentage frequency and mean direction of the summer winds in various parts of the Weddell Quadrant and Ross Sea. The mean direction has been obtained from Lambert’s formula. * EHxpédition Antarctique Belge: Résultats du Voyage du S.Y. “ Belgica” en 1897-1898-1899 : “ Météorologie : Observations des Nuages,” par A. Doprowo1zskI, Anvers, 1903. 122 MR ROBERT C. MOSSMAN ON THE METEOROLOGY OF Table XIV. gives the mean force of the different winds for the same region on the Beaufort scale of 0-12. With regard to these Tables X. to XIV., in which the data utilised are based on a different number of daily observations, the values of the various expeditions have been weighted so as to allow for this. Thus on the “Challenger” and ‘“ Scotia” hourly observations were taken, while in Ross's Expedition and others the available data refer to observations made every four hours, and in some cases at even longer intervals. With regard to Biscox’s observations a numerical value (on the Beaufort scale) has been 02 ee 1? 66 applied to his descriptive terms, such as “strong breeze,” ‘‘ moderate breeze,” “gale,” ete.; the directions given when “magnetic” have been referred to the “ true” meridian by applying the variation for the epoch and locality of observation. Table XV., showing the force of the different winds during the drift of the “ Belgica,” has been prepared from the observations given im extenso in M. ArcTrowskI’s report,* the hours utilised being 4 a.M., 8 a.M., Noon, 4 p.m., 8 p.m., Mdt. Table XVI. gives an analysis of gale frequency at Evangelists Island and Dungeness, deduced from a detailed examination of the tri-daily observations published im extenso in the annual volumes of the Maritime Department of the Republic of Chili, 1901-1905. Table XVII. gives a condensed abstract of the mean barometric pressure, tempera- ture, etc. of the different winds at Cape Pembroke, Falkland Islands, and at the South Orkneys, for the seven months April to October 1903. Table XVIII. contains similar means based on the sixty-four days’ observations taken by Dr Bruce in the vicinity of Louis Philippe Land and the South Shetlands from December 1892 to February 1893. Table XIX. shows the departure from the mean of average values of pressure, temperature, etc. during the cruises of the “Scotia” in the Weddell Sea during the summers of 1908 and 1904, for each Wind Direction. Table XX. gives the barometric and thermal windrose for the seasons and the year during the ‘“ Belgica’s” drift, and the seasonal and annual departures from the normal. This table is based on those given by M. ArctowskI on pages 36 and 38 of his work, but the directions have been reduced to eight points. With reference to these tables of windroses (Tables XVII. to XX.), it may be observed that the values have been reduced to eight directions by halving the intermediate points between the octants. The annual and seasonal means of the climatic elements are not the averages of the monthly values, each of which is derived from a different number of observations, but are the arithmetical means obtained by dividing the gross totals by the number of observations of the wind in question. Table XXI. gives the departure from the mean of the temperature of the different winds at Staten Island and Rawson for January, April, July, and October, and the mean of these four months as representing the year. The data are taken from The Climate of the Argentine Republic, by W. G. Davis, p. 28. * Hupédition Antarctique Belge: Résultats du Voyage du S.Y.“ Belgica” : “ Météorologie,” par HENRYK ARCTOWSKI, Anvers, 1904, pp. 58 to 83. THE WEDDELL QUADRANT AND ADJACENT AREAS, TABLE I. 123 Mean Monroiy AND ANNUAL BAROMETRIC PRESSURE AT 32° SEA-LEVEL AND STANDARD GRAVITY AT Lat. 45°. 20°000+ inches, F Lat. |Long.| No. of} Years Hours of Vear, Station. 8. | W. |Years. | specified. | Observation. Jan. Feb. | Mar.| Apr. | May.|June.| July.| Aug. | Sept. | Oct. | Nov. e | pe alent Ins. | Ins. | Ins. | Ins. | Ins. | Ins. | Ins. | Ins. | Ins. | Ins. | Ins. Ins. “Belgica” .|7035/8722| 1 1808-99 hourly | 9°417| 8-964 | 9°153 | 9-004 | 9-496 | 9550 | 9-450 | 9°325) 9-298] 9:293| 9°357 29°307 Port Charcot || 6503/6326) 1 | 1904-05 9-250 | 9°156 | 8-978 | 9-476 | 9-558 | 9°389 | 9506 | 9°329| 9:350) 9-344| 9-492 29°375 Snow Hill .|6422|/5700| 13 | 1902-03 S 9°375 | 9-143 | 9°198 | 9-280 | 9-242 | 9°330| 9°180| 9-141| 9-242| 9-100| 9-379 29°247 The twoabove*| 6446/6013} 23 | 1902-05 - 9°265 | 9°195 | 9°125 | 9°346 | 9°345 | 9-349 | 9-289 | 9-204| 9:278| 9-181| 9:424 29:288 South Orkneys |6044 4439/ 5 | 1903-07 is 9-251 | 9-204 | 9-157 | 9-273 | 9-239 | 9348 | 9-395 | 9-411! 9-297 | 9:322' 9-218 29-283 South 54 31 | 36 00 1882-83 1882-83 pail arene totoal (asa) aaron o: He learee ae Georgia 1/2113 |3633 |} 4 {| ioce-o8 |2ovG.07 sv2-e}| 9°322| 9'866 | 9-371 9873 | 9-352] 9-458 / 9-400 | 9°596| 9-531) 9-413 Orange Bay .|5531/6805| 1 | 1882-83 | hourly 9-441! 9:559 | 9-282 | 9-455 | 9°578 | 9-515 | 9°455 | 9°417| 9-523) 9°350| 9°279 29°447 1875-77 ) | | Port Stanley .|5141|5742, 7 | 1882-88 4hourly |, 9°552) 9-541 | 9-508 | 9-624 9°631 9-616 | 9°699| 9°655| 9-665 | 9650 9670 29°618 1903-05 Dungeness .|5224/6825) 5 | 1901-05 8-2-8 | 9°566| 9°502 | 9°482| 9°590 9°594/ 9-511 |9°577| 9°557| 9°550| 9°561| 9°531, 29°547 Sandy Point -|5310|7054| 14 | 1889-1902) 7-2-9 9°487 | 9-470 | 9°531 | 9-546 | 9584 | 9-680 | 9°566| 9°677| 9 684| 9°619| 9°527 | 29°568 sofatnaig | | 5433 | 66 00 | 18-19 | 1876-1906] 7-2-9 9°431| 9-450 | 9-417 | 9-501 | 9°513| 9-548 | 9°521| 9°557| 9-621 | 9°539| 9-439 | 29°498 MiChabat) }| 427 |6505| 14 {| 18888 fh 729 9-72 | 9°785 | 9-809 | 9352 | 9°84 | 9-959 | 9-903} 9:951| 9:974| 9-802] 9-74 | 29°858 Aneud (Chili). | 4151|7350| 6 | 1900-05 8-2-9 | 9-987 | 10°00 | 9-972| 9-961 | 9-902 | 9-840 | 9878 | 9-987 | 9-996 | 10-026 | 10-007 29°963 Point Gallera .|4000|7344| 6 | 1900-05 8-2-9 10001 | 9992 | 9-978 9-988 | 9-925 | 9-369 | 9-908 | 10-031 | 10-033 | 10 062 | 10°028 10-012 | 29-986 * These values are the mean of Port Charcot and Snow Hill combined with the observations made in this region by Ross in December and January 1842- 43, and those made by Dr BRUOE and Dr DONALD from December 1892 to February 1893. TABLE II. Mran MONTHLY AND ANNUAL TEMPERATURE. A No. of Years Hours of Long.| Height ‘ ‘ | Station. Years. |specified.| Observation. Lat. W. ect. »/ Jan.| Feb.) Mar.| Apr.} May. | June.| July. | Aug.| Sept. | Oct. | Nov.| Dec.| Year “Belgica”. 1 | 1898-99 hourly | 7035/8722] 15 |29:8|30:0/160/10'7) 20:2} 4:1) -20-6/11-6| -1:5|17-9| 19°6 14°6 Port Charcot . 1 ‘| 1904-05 rs 6503/6326} 15 | 32:9|31:2/29:8|22-6| 13:3) 11:°3| — 26|20-4] 25-7 | 19-6 | 31:5 29-9 Snow Hill. 12 | 1902-03 2 6422|5700| 39 | 380-4|95-7/13-2| 7-2] -o-8| -3-4| — 55|-3-0| 4:0] 14:8 | 17-4 10°7 South Orkneys 5 | 1903-08 ie 6044/4439} 20 | 321) 322) 31:1/25-9| 17-7] 11-7) 9-4] 14-5| 19-0| 24-1 | 29-6 23:2 . 1882-83 | hourly 1882-83| 54 31 | 36 00 Palette es, ‘ : Real lteaeel lines : South Georgia 4 { 1905-08 | 8-2-81905-08 | 5413 | 36 33 }u 41°8 | 42°0 | 39°8 | 35°4| 32°5| 29°5 28°2| 29°9| 32°6| 34°6| 37°7 35°3 Orange Bay 1 ‘| 1882-83 hourly | 5531/6805} 39 | 46:0|48-0| 426/40°8| 39:9] 36-72/ 37-8| 37-4] (42-4)| 42-8 | 44-2 417 Port Stanley . a { ieee ge + {max.andmin, | 5141|5742| 22 | 48:9|49°5| 458) 41-4| 39:8] 36-0) 36°6|37-2| 38-6 | 43-0 | 46:2 42°6 Cape Pembroke 2 | 1903-04 4hourly | 5141|/5743| 52 | 48°6| 48:8] 466| 43-9| 39°8| 39:2| 35-4|37-6| 39-2] 40-6 | 44°8 49°6 The two above 63 As above as above 51 41 | 57 43 32 48°8 | 49°3 | 46°1 | 42°2| 39°8| 387°1| 36:2|37°3] 38°8 | 42°2| 45°7 42-6 1876-79 same | oo f ise | 7-2-9 5452/6807} 98 | 528| 50-2] 461) 421| 37:3] 321] 32-7|35°5| 40-1 | 43:9 | 47-9 42-4 (| 1904-06 Staten Island 8-11 { rion 7-2-9 5423/6347] 39 | 48'7|48-0| 439/436) 40:3) 369| 36-5] 36-5| 38-5 | 41-4 | 44:6 49°1 NewYear'sIsland.| 24 | 1904-074], 7 S10 4-M-\| 5439/6407/ 164 | 47-2| 46:3] 453/415) 362| 34°8| 34-6/36-6) 38-9 | 41-4 | 43-4 40°9 | Harbertown . 4 | 1903-06 7-2-9 5500/6700] 17 | 52:3|49:8| 45-9| 42-6| 365] 33:9| 33:0| 36:4] 40-4] 44:5 | 48-2 42°7 Sandy Point ; 15 ‘| 1888-1902] 7-2-9 5310|7054| 69 | 53°4|53-0|49-4|44-6| 40:0] 368| 342|37-0| 40-6] 44-5 | 48-2 44-4 Dungeness . .| 5 | 1901-05 8-2-9 5224/6825} 10 | 54:2) 524] 49:8|46-6| 41:2] 37:1] 35-0|368| 39-2 | 43°6| 47-8 44-6 Evangelists Island 5 | 1901-05 8-9-9 5224|7506| 174 | 48-0| 47-0| 46-4| 45-4| 42:2] 39-8] 37-8) 39-2] 40-0 | 41-4 | 43:5 431 Ancud .. 6 | 1900-05 8-2-9 4151|7350| 157 | 59°5|57°6|56-0|53-0| 50°6| 47°5| 45°4|46-4| 48-6 | 50-1| 53°3 520 | Point Gallera 7 | 18999-1905} 8-2-9 4001|7344| 125 | 58°0| 566] 55-9| 54-2] 52-7] 50-4] 481/|48:0| 49-8 | 50°3 | 53-0 52:7 Rawson . eee 1898-1500) 7-2-9 © |4317|6505| 26 | 69-9| 68-2] 63:5| 54-1] 47-7] 42:4] 43-0] 44-4] 50-9| 57-4 | 62:8 55°9 PauletIsland* .|7months| 1903 3 hourly 63 30 | 56 00 30 113} 3:8] 3:1 69} 81] 7°0| 25:4 * The means for this station are only approximate, meol siirskild hinsyn till Graham Land.” having been measured off a curve accompanying GOsTA BODMAN’S paper, “Om Klimatet i Antarktis TABLE III. Man MONTHLY AND ANNUAL AMOUNT oF CLouD 0 To 10. Stati Lat. | Long. | No. of Years oureiee ion. 8. W. | Years.| specified. aes Jan. | Feb. | Mar. | Apr. | May. ie July. | Aug. | Sept. Oct. | Nov. | = rem ‘‘ Belgica” . 7035 | 8722 1 1898-99 hourly 89 | 94 | 64 | 72 | 821 65 | 49 | 73 | 64 | 8:0 | 8-0 Port Charcot 6503 | 6326 1 1904-05 oye. O20 SiS: ere) CS) 5:9! si6-38 |) 690i ss) lars || 74 |) 8:7 ‘Snow Hill. 6422 | 5700 1% | 1902-03 Dy ie, 88 | 76 | 86 | 67 | 66 | G1 | 62} 56 | 7-2] 7:3 | 84 South Orkneys . - = 39 5 1903-07 i 94 | 92 | 9:2 | 86 | 80 |] 74) 73 | 76 | 7:8 | 88 | 91 ‘ 00 1882-83 A é ‘ eg : : : ; : , : South Georgia Boral foci } an Erica avg t| 73 | 67 | TO] 70 | 70] 68 | 64 | GO| o2 | 73 | 72 Orange Bay . 5531 | 6805 1 1882-83 hourly 86 | 79 | 84 | 81 | 76 | 7-7 | 72 | 74 | 76 | 7:7 | 85 Cape Pembroke 5141 | 5743 2 1903-04 ites 72 | 66 | 66 | 72 | 72 | 76 | 68 | 72 | 73 | 66 | 66 Sandy Point. 5310 | 7054 | 15 1888-1902 7-2-9 66 | 62 | 61 | 65 | 67 | 5-9 | 56 | 58 |] 55 | 56 | 63 | Staten Island 5423 | 6347 9 1887-95 7-2-9 P22 eDale9) WeweO! \eGrk il bell be lness3) || 7-0) || 70 Evangelists Island | 52 24 75 06 5 1901-05 8-2-9 79 Wee) 75 75 69 (83 70 7°3 Ta) 6 79 Dungeness 5224 | 6825 5 1901-05 8-2-9 65 | 64 | 57 | 58 | 54] 60 | 56] 57 | 55 | 56 | 68 Ushuaia . 5452 | 6807 | 10 1876-85 7-2-9 69 | 65 | 66 | 65 | 63 | 62 | 58 | 5-4 | 59 | 65 | 69 | Rawson. aS pal iep0o),| sie | t8eO Se \ 7-2-9 42 | 38] 43148] 47 | 50 | 49 | 5a | a7 | a4 | 42 Ancud. . .| 4151 | 7350 5 1901-05 8-2-9 Bol be 7 G:ONll OZ fec-OMllerg-galmg-Onilig:a aj gdie | 738" il 5:9) Point Gallera .| 4000 | 7344 5 1901-05 8-2-9 47 | &6 | 4.9 | 62 | 7:4 | SL | 7-7 | 6:9 | 71 | 62 | 6:2 CUCL We EG ST Oo! (Gp) SO S700 =1 RO FR BOOKER © NADH IQ GIGI SY CIS Ga GIGI OA A ROAMHERS © ANKE 124 MR ROBERT C. MOSSMAN ON THE METEOROLOGY OF I — q TABLE IV. Mran MontTHLY AND ANNUAL RAINFALL. Station. ie Tene: apecy eaten Jan. | Feb. | Mar. |April.| May. | June.| July.| Aug. | Sept. | Oct. | Nov. | Dec. | Year, Port Charcot. 65 03 | 63 26) 1 1904-05 181 | 1:28] 1°72| 0:93] 0°63] 1:09| 0-41} 1°54] 1:06 | 0°85 | 2:29 | 1:21] 14-82 South Georgia Perabo } 3 Cao 3:04/ 539 | 4:83| 3:36| 2-77] 2-:07| 2:46] 2-26| 4-19 | 3°03 | 8:17] 3-93| 40°50 Port Stanley . 51 41/57 42| 3 | 1875-2 1:97 | 2°72 | 217) 1:69] 2:01) 1°69] 1:46] 1:85] 1:18 | 1-24 | 1°34] 1:14] 20°36 ; Ushuaia. 54 52 | 68 07 | 10 1876-85 2°46) 2°46] 2°01 2°73) 2:22) 2:07] 1:41| 1:46] 1-26 | 1°56 | 1°50 | 1°95| 2309 Staten Island 54.23 | 63 47| 10 | {7886-93 1! 4-00| 5:19 | 4:15) 5:25] 5:76| 6-04] 4:99| 3:86/ 3:55 | s-03| 313 | 4-70| 53°65 : Dungeness . .]| 52 24 | 6825] 3 1903-05 2°78| 1°58 | 1°47| 1:33] 052] 1:20] 1:18] 0-47) 0°64 | 0-31) 0°49 | 1:07] 12°94 Sandy Point. .| 5310] 7054] 15 1888-1902 | 1:57| 1:26 | 1°56| 1:40] 1°45] 0°96) 1-25] 1°30] 1:02 | 0°86 | 1°15 | 140) 15°18 Evangelists Island | 5224 7506] 7 1899-1905 | 12°82) 9:51 | 12°10|10-44| 7:45] 7-82' 6:76) 7-04] 7-67 | 9°79 | 9:48 | 10°38 | 11126 Ancud . . .| 4151/7350] 6 1900-05 2°84 | 4°03 | 6°98) 8°79] 12°17 | 13°24 | 12-06 | 10°42| 9:16 | 4°79 | 4°75 | 3:40] 92°63 Point Gallera 4001/7344] 7 1899-1905 | 2:92) 4:42 | 6:48 | 10°68 | 14-72 | 19°18 | 17°18} 12°41 | 8°57 | 4°77 | 4°57 | 3°21/ 109-11 Isle Mocha 38 22) 7353) 7 | 1899-1905 | 0:91) 1°40] 4:25] 4:94] 9:01 |11:39| 10°53] 6:21) 3:47 | 2°00 | 1:45 | 1:19] 56°75 | TABLE V, MEAN Hovurty VELOCITY OR STRENGTH OF THE WIND. Instrumental Records in Miles per Hour, or Estimated Force (0-12). Stati Lat. Long. | No. of Years ae Jan.| Feb.| Mar.| Apr.|May.| Jun. July. |Aug.| Sept. | Oct. | Nov.| Dec.| Year. SOS, s. Ww. Years.| specified. Sari. laces = 2 | 2 ; r i : 2 u * % Snow Hill : 6422] 5700) 13 | 1902-03 Hourly | 13:5 | 15°7 | 25-0 | 17-0 | 19°2 | 19-2 | 21:4] 19°7] 17:5 |17-0]16-8| 9-0} 176 South Orkneys 60 44| 4439] 5 1903-07 rs 8-9 | 14:2 | 16-2 | 16-5 | 15°7 | 15-7 | 13-7 | 15°0] 15-7 | 16-0 | 14:8 | 12:3] 14°5 South Georgia . 54 31| 3600] 1 1882-83 ss 14-0 | 15:6 | 14:9 | 15°5 | 14°2 | 12°8 | 15°7 | 18-6 | 17-2 | 14:8] 17:7] 15-9] 161 Orange Bay 55 31| 6805! 1 1882-83 e 20°5 | 17:5 | 14:8 | 13°2 | 18°3 | 12-4 | 13-6! 13:2] 12°6 (12:2)| 16-6] 16-8] 14:7 Sandy Point 5310| 7054| 6 1897-1902 és 98| 9:1] 82] 65) 58] 45] 62] 82} 81) 97] 85] 95] 7:8 Staten Island . 5423| 63 47| 1? 1895? as 11°5 | 14:4] 19°1] 16°3 | 17-0! 19-4 | 19-4] 17-5 | 16°6 | 15:1 | 13°7 | 125] 16-0 “Belgica” . 7035 | 8722} 1 1898-99 Fs 2°4| 2:8] 3:0] 3:5] 26] 2-4] 16] 2:4] 2:0) 2:7] 21] 2:9) 25 Cape Pembroke 51 41] 57 43 2 1903-04 4hourly | 45] 4°93) 46] 4:2) 44] 4:2] 41] 4:3] 44) 41) 46) 4:5] 43 Dungeness 5224| 6825] 5 1901-05 8-2-9 | 3:5] 4:0] 41] 3:2] 3:3] 3-7] 36] 3:9] 3:5] 3:6] 3:2] 3-4] 3:5 Ushuaia .| 5452] oso7| 7 [{i8r et | 7-29 | 56] 41] 3:8] s4| 34] 35| 38] 52) 32] 42] 43] 43] 39 Evangelists Island .| 5224] 7506| 5 1901-05 8-2-9 | 4:4] 4:4] 46] 4:2) 4:2] 4:4] 40] 4:4] 4:5] 4:5| 4:3] 4:2] 4:3 Point Gallera . 40 00| 7344] 5 1901-05 8-2-9 | 3:4] 31] 30] 34] 3:7) 44) 3:7] 36] 33] 33] 34] 3:2] 3:5 Ancud .. 41 51| 7350) 5 1901-05 8-2-9 | 2:4] 2-4] 2:2] 2:5] 8:1] 3:5] 31] 31] 3:4] 3:2] 31] 24] 2:9 TABLE VI. Mean RELATIVE Humipity (SATURATION=100). : Lat. | Long. | , No. of Years Station. 3. W. Hours. Years.| specified. Jan,| Feb.| Mar.| Apr.|May.| Jun.'/July./ Ang.| Sept. | Oct. | Nov.| Dec.) Year. Port Charcot 65 03 | 63 26 | hourly 1 1904-05 89°6 | 77-1 | 83°5 | 861 | 85°8 | 87-8 | 92-4 | 89-9 | 86'5 | 85:9 | 85:0 | 85°6 | 86°3 South Orkneys 60 44 | 44 39 = 2 | 1903-04 848 | 85-4 | 87°5 | 86°0 | 90:0 | 89°5 | 89°3 | 87-2 | 90-9 | 89-3 | 88-7 | 87°7| 88:0 | South Georgia . 54 31 | 36 00 i 1 1882-83 71'9| 70°9 | 72°6|77°3| .. |75°9|72°6|72°3| .. | .. 176217389) . Orange Bay 55 31 | 68 05 - 1 1882-83 83:0 | 80°6 | 79°5 | 83°9 | 83-2 | 88-1 | 82-0 | 76-1 | (75-1) | 83°3 | 82°4 | 83:0 | 81°7 Ushuaia 54 52| 6307 | 7-2-9 5 1876-83 68°3 | 72°4 | '72°6 | 74°6 | 79:0 | 84:1 | 83°7 | 81:0 | '75°6 | 69-7 | 68°6 | 72°5| 75°2 Dungeness 52 24 | 68 25| 8-2-9 5 1901-05 69'8 | 71°3 | '72°9 | 77:2 | '79°1 | 80°9 | 80°6 | 80°4 | 78'8 | 73-7 | 72°6 | 72°2 | 75°8 Sandy Point .| 5810) 7054] 7-2-9 | 14 1889-1902 | 70-7 | 70°9 | 7371 | 74°8 | 79:8 | 76-7 | 81-0 | 74:6 | 81:0 | 69°7 | 67-1 | 69-8 | 75°5 Evangelists Island .| 5224] 75 06| 8-2-9 2% | 1901-03 87:1 | 85°5 | 83-0 | 84°6 | 83-0 | 85'9 | 86-4 | 85°6 | 83-9 | 86°0 | 86°3 | 88°5| 85°5 Ancud .. 41 51| 7350} 8-2-9 5 1901-05 76°5 | 80°5 | 82°2 | 84°5 | 82°9 | 86-2 | 83-6 | 84:4 | 83-2 | 79°7 | 77°5| 77°6| 816 Point Gallera . 40 01| 73 44| 8-2-9 5 1901-05 83'9 | 848 | 86-2 | 85°6 | 86°3 | 86°2 | 84-5 | 85-0 | 84:9 | 81-0 | 82°8| 83°8| 84°6 | Staten Island . 5423] 6347] 7-2-9 | 10 1886-96 76°6 | '77°5 \'79'2 | 81°6 | 84-1 | 86"7 | 86-4 | 84-1| 81-2 | 78°6| 76:3. 75:6 | 80°7 | Rawson 4317| 6505] 7-2-9 | 12 |{ 18a Shoot | 71:6 73:5 |75°8 | 76-2 | 738 | 77°1| 75°5 | 849) 78-7 | 75'S | 71°6| 724) 766 THE WEDDELL QUADRANT AND ADJACENT AREAS. 125 TasLe VII. SHOWING THE MEAN MonTHLY, ANNUAL, AND SEASONAL FREQUENCY OF WIND IN THE WEDDELL SEA AND ADJACENT REGIONS, PER CENT. 3 a Ble eh B |e ; fe) PS ee Lt || teh I ps Stations and Directions. Bless ales. ie hd g 2 5 BAP 2 5 8 8 I a) a 2) P| 2 60 a “ ° s ‘ S =) See esata es eles Soe Wa eet Sole |e |e Antartic Research Ship N. 2/10] 6 “ier 4) 4 8) 10) £0) 9 | 2) 8) Lor) 4) 138 |. 5 “ Belgica.” NB 28 late) SiS) 7 | ot jae) 16 | ON 19 | La hls uid. |,.20.)12)| 5 ee ecatao E B0 27 Gis) 5 \S'\e8 | 6 iat | 7 | 20 | 22'| 15.) 13126113 | 8 y mony? Sua Seo lG sie lee Tels loli nese) Gl teil Id \ob 08. 11) “9 4 ape de er arise See as Deh hst ae ea ad 9) SE 5. GG ane ME Tale ZaleSi) 8 A eee e S\s-w.| 5] 4/111] 6| 1/14/11/a1| 9/16)10/14/ 10/12! 8| 8 | 12 8 99°97 W. W. 2| 81) 11] 13/16 | 35 | 17 | 28| 12/20] 9|10/15|14| 7/|14| 26 on NEW 0) Wee || 04 | sel oon), 19) P74 on | 8 1% | 5 2 | 10 | 10:||- 2) 18) 14 (Mean monthly positions are given in Table XV.) Calm) 760) 2 |alO) |, 69) °F |e 2@ Wes) 18 | 0 |) 17 | 14 | 11 |) we | 8) 6S has Ne PLO WG We et 6 lebie |}, tk ese & | bil 4) 2 | eel 11| 7 French Antarctic Expedition. N.E. a 4 a : j a a ae ee =e a a 23 a 22 1 year Feb. 1904 to Jan. 1905. S.E. 5 6 0 1 3 4 6 5 4 4 4 5 4 4 8 1 : Hourly. d Ss. 5 | 8 | 16 | 40 | 52 | 49 | 63 | 29 | 23 | 80 | 20 | 37 | 82 | 24 | 17 | 36 | 47 Pees S.W. 16] 7| 9|12/12] 6| 7] 6| 9|22/12| 02/12/14] 15/11] 6 Latitu s 65° 03" = Wem eSccniaOn (ONO) Ov Or) em) vo Ti) 2) tial o} 9 Longitude 63° 26’ W. NEMA ea oe ean amon ol iénl leo a tel lol phoa | 7) 3 Calm| 8 | 40/ 24| 26/12] 9| 9/ 8/10] 6| 7/11/14] 8|/19] 21] 9 South Ork ‘ N. 4711 6 2 5 4 2 3 2 5 6 6 5 4 7 4 3 ~ ce geamae MeO Sees cele |) ol Too) Oiled) Gol BF leSlionl oh al 4 3 years 1903-05. E. 6] Bh i} a! Oo il Ol a! @f Ol yl ol oi Ase a _Hourly. Seer ee eT aT he O eS lhe Sled 6). Ful 8h 8 |.6 192] 1 Latitude 60° 44’ S. S. Clee o. elton Oo keSr ial) Web|) earl) 8 on! seclend | 12.) 91] 9 Longitude 44° 39’ W. S.W.| 28 | 21 | 23 | 40 | 34 | 31 | 36 | 31 | 32 | 25 | 17 | 24 | 29 | 25 | 24 | 32 | 38 Lower Clouds. _ W. | 39 | 33 | 38 | 33 | 22 | 38 | 42 | 32 | 46! 39 | 33 | 22 | 35 | 39 | 31 | 31 | 37 Calms from Surface Wind N.W.| 7 | 14/| 22 | 11] 19 | 14] 12 | 18 | 12] 23 | 23 | 11 | 15] 20 | 11 | 18 | 15 Observations. Cpim| 6) GleFl) Gi O17 9) Bl GB Gl Bw oA Gl G 9 ’ N 12 | 17 | 19 | 830] 17 | 18 | 11 | 14 | 20 | 81 | 25 | 19 | 19 | 25 | 16 | 22 | 13 South Georgia. Nevin lO les tele teleey 3 i Sir essletOu) G6 lr SiG | 8 i 8 | 2) 6 4 Vears 1882-83. Hourly. |E DY Of al By) wy 2 sp el os] G) 2) O) 2) 4 el] Bl 3 1905-07. 8-2-8. Sie ieee eG Wedel eS lk Sle Sil, lero: | en 1 18.127") Bo | Sal vg Latitude 54° 31’ S. S. SSO Oe Gah? |e? lO ed Gy, anh 3°) 43. | “ah ath eal 9 Longitude 86° 00’ W. in 1882-83,|S.W.| 7|/10| 6] 5/ 7/ 6/ 7/ 5| 7] 5] 8| 5] 7/ 7] 8! 6| 6 Latitude 54° 13’ S, . | 23 | 18 | 22/19 | 28 | 15 | 19 | 21 | 27 | 14 | 21 | 16 | 20 | 21 | 20 | 21 | 18 Longitude 36° 33’ W. in 1905-07. NO 26 | 29 | 41 | 35 | 41 | 44 | 46 | 39 | 29 | 22 | 25 | 26 | 34/| 25 | 30! 39 | 43 alm} ... mate mate alee aie siete ee ayers + : N. | 18 | 15 | 22 | 21 | 16 | 28 | 17 | 21 | 21} 18 | 24 | 12] 19 | 21 | 15 | 20 | a9 Cape Pembroke Lighthouse, |N-E.|15] 5] 6|11] 4/12] 7] 9]13|10|16]10|10/18]10| 7| 9 Falkland Islands. SE 4 i - ; a q ‘i 4 4 : 2 : ZOO | 2 of Wed 2 years 1903 and 1904. S. Galea ae 2h ea aS) ie Slee Gy lhe dling Vl) 2 Bel cB hoedalll a ; A ee. S.W.| 18) 16] 6) 14|14| 5] 12/11 | 11] 14 | 18 | 20] 13 | 13 | 18 | 12 | 10 ; ati rok * Z 5 ALS. W. | 20 | 21 | 27 | 23 | 24 | 18 | 24 | 26 | 29 | 22 | 16 | 21 | 23 | 22 | 21 | 24 | 93 ongitude 57° 43’ W, N.W.| 19 | 28 | 35 | 22 | 81 | 24 | 80 | 25 | 22 | 24 | 26 | 17 | 25 | 24 | 22 | 29 | 26 Calm | ... abe Sh aleaas { * Includes 14 days of March 1899 as well as March 1898. 126 MR ROBERT C. MOSSMAN ON THE TABLE VII.—continued. | ete ae Beet | Stations and Directions. 3 5 3 = = 3 re aio/€&|e&/8/5 | 3 Lr) & a < = 5 5 N. TE) | LO | eee) ARTO 2 SS N.E PAN; Ae OAM LEY ah] RZ Staten Island. E. al tt eo th ean 10 years 1886-96. SHDR | Oy RE ce eA ot aI Hours 7-2-9. S. 16 | 10 | 14 | 21 | 17 | 18 | 13 Latitude 54° 23’ S. SAW] Lome2Zai tS 6) 4s SUB es at6 Longitude 63° 47’ W. W. 15 | 18 | 17 | 16 | 20 | 16 | 14 N.W.| 25 | 23 | 29 | 22 | 24 | 20 | 23 Ciba 5) SU ei) Ol, Bl) Gl) ei Punta Arenas (Sandy Point). 6 years 1897-1902. Hourly. Latitude 53° 10’ S. Longitude 70° 54’ W. N 8 2 4 9 5 6 4 | Dungeness (Lighthouse). yy a 5 ; 5 3 3 3 : 5 years 1901-05. S.E 74 eo PP a oe | Hours 8-2-9. NS) Pali al) a HO OG Latitude 52° 24’ S. S.W. | 50 | 56 | 51 | 41 | 38 | 32 | 42 Longitude 68° 25’ W. W 10 |) 13 |) 15) 15 | 19 | Is |) 16 N.W 9 9 2 DS a a ) | Calm} 13 | 11 9 | 14 9 | 15 9 N. 8 5 9 | 16 a ial 6 N.E. 0 0 1 3 2 4 2 E. 0 0 1 4 4 5 ai S.E. 1 1 2 6 | 10 6 8 2 a 5 | 10 | 10 yale, eile || alZe : COA No |) UB |) PRN IGS) G2 |), ass |) AK |) al iota Ean W. | 26 | 27 | 26 | 15 | 20 | 12 | 12 N.W.| 41 | 28 | 31 | 24] 19 | 24 | 22 Oph I 7) hie |) GN) Bi] aie / Evangelists Island (Lighthouse). 5 years 1901-05. Hours 8-2-9. nl 11 | 12 | 25 | 17 | 24 | 25 | 18 Ushuaia, coma tal Moa ealrese Peel ay : E. Al ds. |G 8 | 10 of 4) ala Tierra del Fuego. SE 5 Wo lina hbontgultio War 43 years 1876-83. 8. DU al eee oa 24 8) Hours 7-2-9. S.W.| 20 | 17 | 15 | 24 | 18 | 21 | 18 Latitude 54° 52” S. W. | 56 | 55 | 35 | 33 | 26 | 30 | 36 Longitude 68° 07’ W. NA Well 4 8b) One FON Sales Calm pee | iperae ene METEOROLOGY OF tow AaAOIMmANrer aD i © CO | September. — ont me ore co bo November. pet SEP 18 December. | Autumn. pS ore bo bo | Winter. THE WEDDELL QUADRANT AND ADJACENT AREAS. Tasie VIII. PERCENTAGE FREQUENCY oF UPPER CLOUDS. Cirrus, Crrro-Cum., AND Crr.-STR. 127 Stati d Direction Balesel aries. | Sere |e) |. |e) aie | g ations and Directions. g/ 5 | ¢ | e|g|s é 3 3 g S 4 | : g : Does nee eels Meeeeneetincelees staal mreilud a | a | N. 2 CU GIP} UE 7 el ee! Sie el) 5) 0 GUN By DN GH N. 3 2/} 0 3 1 0; 4 7 0 3) 38 0 2 2 2 1 4 EB. 3 A |) ak Py OM) 7 @)! Oy Bi al BO} si al : S.E. 3 Ih, Sef Boal TN th Oy Oy Bi i al peas Orkneys, 1908-1907 s. |16| 8| 5/171 14/ 22/12] 12| 11] 1] 1| 2] 10/| 4] 8] 19] 16 S.W,| 12 | 23) 31 | 20 | 15 | 80) 17) 26) 82) 27) 14) 16) 22) 25) 17] 21) 22 W. 48 | 32) 33 | 32 | 21 | 24| 32 | 20) 39 57 56) 63 89| 50) 51) 29) 26 INEWiallsn | e2bil i ale2on een Sil 22 TS i STE La Moi adi) 24) 4 Number of Observations. 99 }113} 86 | 89 | 98 |120] 84 | 95}118 105! 87) 133 | 1227 | 310) 345) 273 | 299 : N. By) AU GS wb A SBE BYE Gas | IG) OL ea aE aS eee Cape Horn Region. NGPA eS ero von AON Si eOiaonl Si eOl oO) 0 10, t\..O 1) 1) 2 From observations at Orange | E. 0 ON OO 0; 9 Ol O; OF Of 0; O}; OO; OO; 8 Bay and Ushuaia during the JS.E. | 0] 0] 0} 0} 0} Oj O OO Oy Wo 0; O; Oj; oO; O year 1882-83, and at New |S. d 8; 0} 13 | 10 33/0 Gi Or Si) Od SN ee Year’s Island, December 1904 |S. 21 6/ 0/| 14 | 22) 20) O 0} 20! 18} 18] 18 14] 17) 16) 138 9 to March 1907. W. 46 | 56| 47 | 42 | 33) 48| 58 | 50] 15, 39) 48] 55/ 45] 35) 52| 41) 49 N.W.| 24 | 19) 47 | 28 | 15} 11) 0} 33) 40) 21) 34] 19) 238) 31) 21) 27) 8 Number of Observations. 39 | 26] 18 | 31 | 20 | 23] 12] 16] 20) 24) 22) 37| 288] 66|102| 69) 51 | N. 5 Cl EI ey hp aly) al] aie | ee) aly Bh) by Qi Hy) ef) IG ae N. 4 OF Ol OS 0; 0 Tale 4a) Cie cO)l 8 i PH ati By Cape Horn Region. 1 Oo} O OF OH O 0) 5 ui 2) O} Ol Oo i) Oy Oi wy : « S.E. | 0 0; O| 0} 20 0; 0 Oi Oi} Ol all o iy O Gl OF © Georgia, 1882.85 eae Ss. Paes) Oe) eo Be e102 eo) ONG ol 4) 5) 18) 12) 5) 2 ‘ Seer LOM eZ Ol el om iON on DSi li 23h esis Lae ES bs 18) | 2) Wb) rs W. 47 | 59| 39 | 45 | 21 | 40} 50 | 43} 11) 29) 39) 48) 42) 51] 35] 44) 30 : N.W.| 26 | 14) 40} 30] 8] 12] 11 | 19] 81} 36| 44) 22) 25] 21) 27) 15} 39 Number of Observations. 75 | 75) 46 | 44 | 41 | 26] 22 | 44] 26] 86] 53] 65) 592/154) 215/131; 92 | Mean Direction of the Upper Clouds. Seasonal Percentage Frequency of Upper Clouds. | South Georgia. e - “ eee South South Eels) eto 7 Belgica. . with South | Spring. | Summer.| Autumn.| Winter. | Year. OEmeyE. Georgie, Georgia. = | N. 0 : ° ° ° ° | NE. 7 ; 4 ‘ A Spring S. 86 W. | 8. 76 W. | N. 47 W. | N. 82 W. | E. 4 0 0 0 0 Summer . S.10 W. | S. 85 W. | S. 87 W.| N. 76 W. | SE. 0 1 7 0 y Autumn . S. 61 W. | S. 86 W. | S. 42 W.| N. 82 W. Ss. | 0 2 20 10 7 Winter S. 69 W. | S. 68 W. | S. 68 W.| N. 85 W. S.W. 7 23 14 30 19 Year S. 66 W. | S. 81 W. | N. 89 W. | N. 77 W. W. 32 51 Nets : N.W. 50 20 a Es a Mean Annual Direction, Ushuaia : . N. 81 W. 3 95 i New Year’s Island . N. 74 W. ”) ” ” ” ” & Orange Bay N. 68 W. ‘« Belgica.” “ Belgica,” Seasonal Percentage of Upper Clouds. Mean Rate of Movement, Upper Clouds. Scale 0-5. | Spring. | Summer. | Autumn.| Winter. | Year. N. |N.E.| E. |S.E.| S. |S.W.| W. | N.W.| Mean. | a ric == sae ee ae / N. 3 3 ‘A a 2 Spring 4 . | 2°38} 290] 1°7 | 2°71 2°5 | 255 2°2| 2:0 2-3 N.E, 10 3 Sele 3 5 Summer . 110/25) 22/1 5/11°8/1°6| 1:3 16 17 E. 3 7 4 oon 3 Autumn wf] ee [10/20/17] 15/18] 2:3] 1-7 S.E. 8 23 4 a 8 ae |. (80) |... 15) o4)23| 21/93 | | S a cas 2 Bee pe Year. 6 . | 1°8| 2:2) 1°9| 1:7 | 2:0 | 2° 1] 95 eqn aie 7 34 7|2:0|20}21]| 2:0 | 21 a, re i mt — - N.W. 25 14 18 21 20 TRANS. ROY. SOC. EDIN., VOL. XLVII. PART I. (NO. 5). 18 128 MR ROBERT C. MOSSMAN ON THE METEOROLOGY OF TABLE IX. Sea SurRFACE TEMPERATURE. / / ‘Hours Station. - —_ paar) acento Serve Jan. | Feb. | Mar. Apr. |May.| Jun. | July. | Aug. | Sept. | Oct. | Nov. | Dec. | Year. jon. | | ° , ° , ' AM. ° ° ° ° ° ° ° ° ° ° ° ° ° | South Orkneys . | 60 44 | 44.39] 5 1903-07} 8 |31°8|32:2|32:0| 30°5 | 29°4| 28°7 | 28°7 | 25'7'| 28°8 | 29:0 | 29°8 | 31°3 | 80°1 South Georgia . | 54 31| 3600); 1 1882-83} 8 |39°4|40°1/37°5 | 34:2 | 32°2|380°1) 82°8 | 34:2] 30:2 | 33°8 | 35°9 | 37°8 | 34°8 Orange Bay . |55 31) 68 05] 1 1882-88} 8 |49°7|50:2| 46:9] 45°5 | 43:0 | 42°2| 42-1] 47°0 |(41°5)| 42°4 | 46°2 | 48°7 | 45°0 ) New Year’s Is. . | 54 39 | 6407] 24 | 1904-07 |7 or 8) 47°7 | 46°8 | 46°4| 43:9 | 41°5 | 39°6 | 38°6| 89°5 | 41°1 | 42°3 | 44 4| 46°0 | 43°2 TABLE X. ABSTRACT OF OBSERVATIONS MADE DURING SUMMER, EAsT CoAsTt OF GRAHAM’s LAND, WEDDELL AND BISCOE SEAS. Mean BAROMETRIV PRESSURE AT 32° Spa-LEVEL AND STANDARD GRAVITY AT LATITUDE 45°. Days of | SBaro- | Mean Air ee Sea Mean i ition. Lat if onth. ‘va-| metric _| Surface | Mean Force Position. Expedition Long M eerie in Loe, Rempee Tempera. Cloud. | Wing, rea ene Po Inches , ~ 9 0-10 | 0-12 E. coast of Graham’s Land} Ross, 1842. . | 62 47 |5812 W.| Dee. 8 29°1138 33:1) 82°7 sate 4: e 93 s 43) eLOkG-« . | 64 22 |56138 W.| Jan. 31 29°116 30°8 30°8 =n 32 = 99 », | Bruce, 1892 Dec. 15 29°402 311 aes 9°0 2°6 a P nS Ay) ICR Bie 63 40 | 5500 W. [n 31 29°318 311 9°3 4-1 a 4 an » 1893 Feb. 18 29°205 29°6 ih 9°3 5:3 ~ A », | Donald, 1892 Dec. | 15 29408 32°4 32°0 “s a . Pp " +3 193 | 63 25 |56 00 W. {s. 31 29°321 33'2 32°0 A Bo s ne a », 1893 Feb. 18 29°242 30°6 31:3 Be Weddell Sea . ; . | Ross, 1843. . | 68 49 |386 36 W.| Feb. 28 29°308 30°7 80°7 00 3°6 i pe : . | Bruce, 1903. . | 64 00 |29 45 W.| Feb. 28 29°112 27°8 30°0 91 18 e pas , : a7 L908) . | 64 01 |4052 W.| Mar.| 81 29°0385 25°0 29°4 9°5 2°6 ~ Pita F ’ 1904. . | 70 18 |17 47 W.| Mar. | 81 29°066 25°0 29°7 8°8 2°8 aS ear 5 . | Ross, 1843 — x. . | 69:06 |14382 W.| Mar. | 11 29°122 28°9 30 2 os 4°9 Biscoe Sea. : . | Moore, 1845 . . | 64 09 | 3914 E. Feb. 27 29138 32°3 32°5 e 4:0 Syne Ss Z - ; FP eee . | 61 46 | 8514 E. Mar. | 10 28°980 33°5 32°5 fs 3°9 Sat es 5 ; . | “Challenger,” 1874 | 64 01 | 86 21 KE. Feb. | 17 28°931 30°7 32°1 9:0 3°9 * Staten Island 3 f 5 1892 | 54 23 |63 47 W.| Dec. 15 29°226 46°0 ise 95) F = “a 3 : ; A 1893 | 54 23 |6347 W.| Jan. 3l 29°432 48°4 6°8 Spo = ; s ; A 1893 | 54 23 |63 47 W.| Feb. 18 29°525 45°9 6°4 | * These observations are for the same periods as those included in the observations taken in 1892-93 by Brucn and DonaLp. TABLE XI. ABSTRACT OF METEOROLOGICAL OBSERVATIONS FOR FEBRUARY AND MARCH IN THE WEDDELL AND BIScoE SEAS, FROM OBSERVATIONS MADE ON THE ‘“‘ScoTiA,” ‘‘ CHALLENGER,” ‘‘Pacopa,” ‘‘EREBUS,” AND ‘‘ TERROR.” BAROMETRIC PRESSURE AT 32° SEA-LEVEL AND STANDARD GRAVITY. Mean Position. Days. : Mea Month. Tatifate. —_s Giiserve: Barons dees Sea sumaee : Lat. S. none tions. Pressure. P - | Temperature. | i ® one oa Inches. a z | February ; i ; ; : 60 to 65 62 25 387 23 W. 45 29°258 30°4 30°9 a é 4 - : 4 2 60 ,, 65 62 42 63 45 EB. 45 29°031 32°6 32°6 % ; ; 4 F é 65 ,, 70 68 09 22 48 W. 18 29°223 26'1 29°2 - ; , - 2 : 65 ,, 70 66 20 49 52 B. 13 29°159 30°2 30°1 Mach . . . . . «| 60.65 | 6249 | sobeWwo| w 29:099 26°8 30°0. - P 3 a . ‘ P 60 ,, 65 61 57 84 12 E. 9 28°992 33 °4 382°5 5 : : F : F 65 ,, 70 67 54 23 09 W. 28 29°14] PAA 30°0 yy he oe ot kw | WO ace eo 8. NY) ooron wy eae 29°048 23°6 29°5 ” 5 ‘ ' ; uy 4 65 ,, 74 69 55 21 45 W. 51 29°099 25°9 29°8 THE WEDDELL QUADRANT AND ADJACENT AREAS. 129 TABLE XII. ABSTRACT OF OBSERVATIONS MADE IN THE Ross SEA AND VICINITY ON BOARD THE ‘‘ EREBUS” AND ‘‘ TERROR.” SuMMER BAROMETRIC PRESSURE AT 32° SEA-LEVEL AND STANDARD GRAVITY. i i Sea Surface Position S. Mean Lat. S. Mean Long. Ona Hebi ar ee ie Tee, Sat eerie: e iy. ° ft? ° / ° , Inches. ° ° 60°00 to 62°30 60 30 143 40 W. 8 29°148 34°8 35°9 65°00 ,, 67°30 66 26 156 46 W. 42 29°289 30°8 28°8 67°30 ,, 75:00 71 16 177 09 E 55 28°958 27°8 30°0 75°00 ,, 78:00 76 45 177 37 E 34 29°039 25°9 29°7 TABLE XIII. SHOWING THE PERCENTAGE FREQUENCY AND MEAN DIRECTION OF WIND IN THE WEDDELL QUADRANT, BIScoR SEA, AND Ross SEA DURING SUMMER AND EARLY AUTUMN. Days. | | Locality. Tat. | Long. Observers. Months. | Obser- | N. |N.E. E. |S.E.| S. |S.W. | W.|N.W.) Mean. : vations. : | E. Coast of Graham’s Land . | 63 48} 5513 W. Bruce and Ross Dec.-Feb. 103 12 | 14 | 10) 12 | 18] 12 9| 13 |S. 22H. z - - 63 40| 5500 W. Bruce ‘aus 64 |13|) 15/11] 14/20] 6 | 7| 14 |S.505. Ey “a 3 63 25| 5600 W. Donald ers 64 |13/ 18 | 15] 12 |94/ 2 |13| 8 |S. 70K. ; * » . | 6422] 5613 W. Ross Jan. 31 |11| 15 |10/ 8/13] 28 |12] 38 |S.95 Ww. Weddell Sea. . A .|63 49) 3636 W. Ross Feb. 28 9} 10 | 18 | 28 | 18} 11 2) 4 |8. 4445. “6 a a a . | 70 24| 21 59 W.)| Ross and Bruce, S. of 67° 30’ March 43 16 | 20 | 19 9 5 7 8/| 16 |N.18 BE. “f 1s : 5 . | 68 01 943 W. Biscoe and Bruce Jan. and Feb. 31 Gp Ge ae | 2455 aby) a8} 2 i SE Sein: Biscoe Sea ., ; . . | 6409) 3914 E. Moore eb. 27 13 | 24 | 17 9 | 2L 7 3 6 IN. 605. aoe | : .|6336| 5010 EK, Feb. and Mar.| 37 /|10/ 22 |17| 18 |20| 9 | 4] 5 IN.81E. a 6130) 5948 B. 5, 60°-62°30/ S. - a 15 Bl Ge | el ey | Sa ee] ee Reraae | eer; 6452) 4418 B. » 623°-67° S, a ie Oy | st | Be Oil) Te Be aN) GI Veo.) . .|6608) 4116: | Biscoe 5 35 52 9) 1 || 18 |} 44| 12 | §| 4 1S 44k ees 5 6. 60106] 68.41 BH, = Mar. and Apr.| 23 |16, 52/12/ 4] 9] 2 | 5!| O 'N.508. ” ” ° 4 ; .|6401| 8621 E. “‘Challenger”’ Feb. 17 LOA LOSI ae ae) Ta Ss SS 7 aWi South Atlantic . ; . | 5934] 27 29 W. Various 56°-64° S. Dec.—Feb. 45 10| 4 3 3/18) 30 | 15/ 1% |S. 72 W. = = . . .|5830| 43 22 W. » 56°-64° S. ih FE 30 8/ 0] 0] 1] 6] 24 | 42| 24 |S.88 Ww. ae « . . .|5319| 5853 W. » 52°-56° S. es 19 7/14] 2] 1 | 2] 27 | 27! 20 |S.79W. 5 . . ./4652] 759 W. Scotia ” Apr. 30 |15; 7 | 4] 4] 4] 14 | 21| 31 |N.58W. *Staten Island 5 . .| 5423] 6347 W. ae Dec.-Feb, 64 11 0} O| 5 | 26] 16 | 20} 22 |S.e8 Ww. Ross Sea Region . é - | 60 30 | 143 40 W. Ross 60°-623° Be 8 4) 0 0 DQ | 32e 240) 12) S78 aw cal een ar 3 2 . | 66 26 | 156 46 W. », 65°-673° 42 11 | 17 | 13) 15 | 16; 10 8| 10 |S. 75K. ee se) 2 \)70 16)|\ 177 09/2) > O7R°-75° 55 6 | 8| 15 |19| 18 |17| 10 |S. 9w. 93 yy . ' -| 76 45 | 177 37 EB. »» 75°-78° 34 8 7 | 10} 22 | 22} 18 9 4 |S.10E. * These observations are for the same 64 days as those included in the observations of Drs BRUCE and DONALD given above. TABLE XIV. MEAN FORCE OF THE WIND, SCALE 0-12, IN THE WEDDELL QUADRANT, BiscorE SEA, AND Ross SEA DURING SUMMER AND Earty AUTUMN. Mean Force. me Lat No. of Position. 3. Long. Observers. Months. Obser- = vations.| w.|N.E. E. S.E.| 8. |S.W.| W.| N.W.| Mean. E, coast of Graham’s Land. | 6348} 5513 W. Ross and Bruce Dec.-Feb. 641 /2-4] 3°3 | 3:0} 31)53] 3°99 | 2:9] 3:3 3°5 on ae a 63 40} 5500 W. Bruce ne “9 483 | 3°9|) 3°3 | 3°4| 3:4 |60] 5:1 |2°4/] 32 4°0 ” ” ” 62 47/ 5312 W. Ross Dec. 41 /1°5/ 20) .. .. | 43] 4:7 | 5:5] 3°6 4:0 An = = 64 22| 5613 W. Ross Jan, 180 |46/] 3:3 2:0} 2°0/3°0| 3:3 |31]| 4:1 32 Weddell Sea ~ ; . |6349| 3636 W. Ross Feb. 164 |3°9| 2°7 | 3:4) 42)3°9] 3:9 |3°5] 3:9 3°6 5 i : : . |6617] 2803 W. © Scotia” Feb. and Mar.| 2059 |2°4| 2°5 |83)| 2:°2/1°9| 2:1 |2°9| 2-4 2°5 a 3 - ; . |6906| 1432 W. Ross Mar. 63 |3°2| 6-2 /6:3| 4°7 | 24) 1:5 | .. | 2b 49 ee - P x . | 7024| 2159 W.| Ross and “Scotia,” S. of 67° 30'S. 1026 |2°5| 35/45) 38/17) 21 | 2:7] 2:4 B Biscoe Sea . : A . |6407| 39145, Moore Feb. 151 |3°5| 3:7 | 3:4) 4°7|5°6| 4:2 | 3:7) 4°9 4:0 Pees 3s 4 6886 5010 R; Fs Feb.and Mar.| 212 |3°4| 3:5 |3-7| 43/56] 41 |40] 3:9 | 3-9 ee 61830) 5048 5 60°-622° S. i . |6401} 8621 KE. “ Challenger ” Feb, 392 |45) 2°8/3°6| 4:2 /3°7| 2:7 |3:4| 4°4 3°9 Ross Sea Region . A . | 60 30 | 143 40 W. Ross 60°-62}° an 48 |5°2) .. -.| 50/40) 59 |5°6] 5:4 52 Ooh aa : “ . | 66 26 | 156 46 W. », 65°-67°30' ar 245 |3:'7|) 3:0 |2°7| 24/33) 3:4 |3°'7| 3°5 30 tae D : 3 . | 7116] 177 09 E. », 87°30'-75° He 319 | 3°3) 3°71 4°5| 52/4°7) 4:2 |4°7| 46 4°5 Bla Ser op 6 ' . | 7645 | 177 37 E. », 75°-78° 198 |3°0| 31 374 53/47) 4:5 |3°5| 3°5 4°3 130 MR ROBERT C. MOSSMAN ON THE METEOROLOGY OF TABLE XV. MEAN Force or THE WinDs, ‘‘ Bencica,” Hours 4-8 Noon, 4-8 Mipnieut. (ScALE 0-12.) Mean Position. Month. N. N.E. E. S.E. Ss. S.W. Ww. N.W. Mean. Lat. S. Long. W. January . = “ 70 13 86 56 2°83 31 2b 21 2°6 16 13 AG 27 | February . : : 70 40 92 40 2°0 29 3°5 3°6 27 274 19 16 2°9 *March. C 2 . 71 03 93 10 27 4°7 5:2 24 1'8 22, 2°38 2°83 32 April . H F A 70 57 90 12 3°0 3°3 . 48 5:2 3°0 2°0 3:2 3°2 38 | May . : 2 F 71 04 88 58 2°9 | 3°3 3:2 3-6 24 2°3 2°6 25 2°8 | June . ; : : 71 09 85 40 1:8 2°5 2°5 2°6 19 2°6 28 2-0 2°5 July . . ‘ . 70 44 86 40 16 27 20 2:9 Ly ure 19 2°5 2°0 August ; : 4 70 28 87 20 2°9 2°0 2°4 2°4 2°6 2:3 2°5 3:0 2°6 September . . 70 07 82 44 3-1 31 2°6 21 16 12 29 30 2°5 October 7 ‘ 5 70 04 81 40 2°8 24 3°5 39 3°2 3°2 3:3 2:3 30 November . ‘ “ 70 05 82 14 2°2 3°0 3:4 30 15 1-4 2°3 2°3 2°6 December . : - 70 04 84 18 ae 2°8 2°8 2°6 isk Pe 2°5 1:0 2°6 Year . . A 5 70 35 87 22 27 31 3:2 SL 21 2°2 27 26 2°6 Spring a : ‘ 70 v5 82 13 2°8 29 3:2 27 2°2 22 2°9 2°6 27 Summer . : - 70 19 87 58 ZPD 2:9 29 28 21 24 2°2 1:35 27 Autumn. , rs 71 O1 90 47 29 38 48 4°2 23 2°2 2°8 2°8 33 Winter : . ' 70 47 86 33 2°3 2°3 2°2 a7 1°8 22 2°6 a7 2:4 * Includes March 1898 and March 1-14 1899. TasLe XVI. GALES, ForcE 8 OR UPWARDS, RECORDED AT DuNGeENess (Lat. 52° 24’ S., Lone. 68° 25’ W.) anp EVANGELISTS IsLaAND (Lar. 52° 24’ S., Lone. 75° 06’ W.), 5 Years 1901-05. All Gales recorded Percentage Frequency, Dungeness. All Gales recorded Percentage Frequency, Evangelists Island. N. |N.E.| BE. |S.5.| 8. |S.W.) W. | N.W. N. |N.E.| E. |S.E. S. |S.W.) W.| N.W. Spring . 5| 1] 0] 0} 2|69/|18| 5 | Spring 4| 0| 0| 6] 1) 10 | 20) 65 Summer 1 Hea Oa |p lp 2 1 | Summer df oti |ie Citi (ih weal 2 | 13 | 20! || 538 Autumn (Hl |) 2/77!) 7| 8 | Autumn é 5 aal yO) Or) it] ae ibe ey) 43 Winter . : 8 8 2) 2 9538) 1-20) 8) 4) Wanter. ‘ By Oy) 7 i) ak) aby alte ali 40 Year 4 | eee es asl) Year HNN ON) PrN aU 97 TBs Wy a) || Bil | Gales per 1000 days of each Wind, Dungeness. Gales per 1000 days of each Wind, Evangelists Island. | nw. [NB] EB. /8.E.| 3. |S.w.| W. | N.w.| Mean. N. N.E| BE. /s.z.| s. |s.w.| w. | N.w. | Mean. Yh ca | | | | | | ae | Spring . Pe ou.) ab 0 0| 79|192)113| 44 | 108 | Spring . | 73) 0 0| 87| 19}113/181] 281 | 157 Summer . | 28 | 82 | 0 0) 955 ep 36| 16 | 114 | Summer 152, 0 0/125) 34] 87) 92] 177 | 115 Autumn - | 85 0 | 95 103| 56/151! 35) 23 85 | Autumn F 161 | 0 0| 35/152)119)131) 273 | 148 Winter. . | 41 | 68 | 75 | 62)125)186| 99) 75 92 | Winter . | 76) 0/146] 29)121/1538)152)} 245 | 141 Year | 63 | 31 | 50 46) 88/172) 74] 38 99 | Year . : 18, 0) 86] 37) 96)118]124) 242 | 140 | Monthly Gale Frequency per Cent. = a eee eee eee eee eee Jan. | Feb. Mar. | April. | May. | June. | July. | Aug. | Sept. Oct. Nov. | Dec. | Year. i a — Dungeness 5 ; . ‘ 95’ |) 1670), 109 6°0 7°6 78 SF 10229) Oras LO 8°7 9°1 9°9 Evangelists Island - » | 14°F) LLB) 16°7-) 162) | 1258 6B ea io) sae Zalelieon alaey 8°7 | 14°0 Mean BAROMETRIC PRESSURE AT 32° SEA-LEVEL AND STANDARD GRAVITY; MEAN TEMPERATURE ; THE WEDDELL QUADRANT AND ADJACENT AREAS, TaBLE XVII. CLouD 131 AMOUNT ; AND WIND Forces, witH Various WINDs, AT Lavriz IsLAND, SOUTH ORKNEYS, AND CAPE PEMBROKE, FALKLAND ISLANDS, FOR THE SEVEN Montus APRIL TO OcToBER 1903. South Orkneys. Cape Pembroke, Falkland Islands. | Wind. Pressure. | Temperature. Cloud. Wind Force, Wind. Pressure. Temperature. Cloud. | Wind Force. baie | Inches. $ 0-10 0-12 Inches. : 0-10 0-12 =| N. 29°351 24°8 eal 2°5 N. 29°712 41:9 75 4°0 | N.E. “208 22°4 9°5 1°6 N.E. 808 41°4 85 4°9 E. *250 8°1 6°6 12 E. 804 38'6 48 29 S.E. 234 5°6 72 1°3 S.E. 725 | 349 7°0 3°6 S. 215 52 6°5 1:3 Ss. 780 372 77 4:3 S.W. 318 7°4 6'3 1°4 S.W. 684 37°6 7°5 51 W. 215 23°9 70 98) W. *649 37°9 5°8 4:2 N.W. 320 26'9 8°6 2°8 N.W. 714 40:0 59 3°9 Calm 386 11°0 One 5 Calm ae | vac ‘3 ve Mean 29°297 17°6 76 2°0 Mean 29°715 | 39°8 Oy 4°] Dr W. S. Brycr oN BOARD THE ‘“‘BALmNA,” DecrEMBER 16, 1892, To FEBRUARY 18, 1898. TABLE XVIII. MEAN BAROMETRIC PRESSURE AT 32° SEA-LEVEL AND STANDARD GRAVITY, MEAN TEMPERATURE, CLOUD, WIND FoRcE, WITH VARIOUS WINDS, FROM OBSERVATIONS MADE AT JOINVILLE LAND AND ‘‘EREBUS AND TERROR GULF” BY 63° 40’ S., Mman LonerTupE 55° 00’ W. MraAn LATITUDE Wind Direction and Number of Observations. Pressure. Temperature. Cloud Wind Force. Observations, Inches. s 0-10. 0-12. 55 29°434 31°9 8°6 3°9 62 875 31°4 9°3 3°3 46 ‘501 31°2 8°7 3°4 61 “145 29°8 91 3°4 85 Alea 27'°8 9°2 6:0 23 163 31°4 9°2 5'1 30 *326 33°2 7°'0 24 58 “306 32°4 9°0 3°2 55 "486 31°9 80 50 48 29°308 30°8 9°2 40 TABLE XIX, SHOWING THE DEPARTURE FROM THE AVERAGE OF MEAN VALUES OF THE CLIMATIC ELEMENTS IN THE WEDDELL SEA WITH DIFFERENT WINDs SovutH oF 60° S. DURING THE CRUISES OF THE “‘ScoTIA” IN 1903-04, Mean Position. 1908. Lat. 64° 23’ S. 53 days. 1904. Lat. 70° 03’ S. 38 days. Feb. and March, Long. 34° 37’ W. Feb, and March. Long. 20° 43’ W. Norx.—The heavy type indicates above the average, and the italic type below the average. Direction. Wind Pressure. Inches, 0:119 ‘089 1384 229 ‘057 026 058 ‘116 079 29°090* 0-172 080 ‘11h 121 229 “112 154 “040 299 29-018* Temp. bo OOO SRENSHKHOD © SOWWA BWBGSSHSHRSYS6 TS PHS HWS &w IS DON S i) Cloud. ig aa Humidity. ae No. of Observations. 0-10, Inches. | Per cent. 0-12. Per cent. 04 012 1 O-4 212 17 0:2 ‘001 il 0-0 242 19 0'7 001 i 0-7 119 9 0°0 009 1 01 17, 9 0's ‘020 if 0:2 126 10 O04 002 3 06 144 11 08 005 3 0:7 109 9 0:5 ‘015 3 0:3 133 10 eS} ‘O14 4 a 70 6 9°4 “123 83 2°4 | Total 1272 100 0'8 019 8 0:3 128 14 0-1 ‘010 6 0:0 102 11 0'3 ‘010 6 0:8 100 11 0:2 002 2 11 52 6 12 ‘027 10 I'4 66 i 0°8 018 15 0-2 131 14 0:5 005 0 0:0 136 15 0:5 008 5 01 142 16 1s 085 4 = 55 6 8°7 118 83 2°6 Total 912 100 * At 32° Sea-Level and Standard Gravity at Lat, 45°. 132 MR ROBERT C. MOSSMAN ON THE METEOROLOGY OF TABLE XX. THERMAL AND BAROMETRIC WINDROSE DURING THE DRIFT OF THE ‘‘ BELGICA” FOR THE SEASONS AND THE YEAR. Mean Temperature of the Winds. ft ae N.E. E S.E. s S.W. Ww. N.W. | Calm. | Mean | Spring 26'2 20°3 13°6 1°4 -— 3'3 52 10°4 DALY 5'5 12°0 | Summer ‘ : ; SL edie 31:0 28°9 24°9 25°9 26°6 29°2 26°6 29°3 | Autumn . 3 ; . | 25°0 | 24°6 18°8 5 81 84 12:2 19-1 79 15°6 Winter ‘ : ; BS lelalcb 12°6 11°8 81 —- 65 | - 9:9 22 11:5 -121 M7 Year . 23°9 | 24°6 22°2 by ys) 52 9°0 176 4:3 14°6 Departure from Annual Mean.* | N N.E E S.E. 8 s.W. Ww N.W. | Calm, | Mean Spring 2:3 438 86 143 8'°8 0:0 14 41 1:2 26 Summer . : : 5 72 7:0 8:8 13-2 19°4 20°7 17°6 116 22:3 14°7 Autumn. . i ‘ ilo 0:0 34 8:2 2°6 3:2 3:2 15 3°6 1:0 Winter P : - ml) one 12°0 10°4 | 76 12°0 15-1 6°8 61 16°4 12'9 Mean Barometric Pressure of the Winds. N. N.E. E. S.E. Ss. S.W. Ww. N.W. Calm. | Mean. | Spring 9°225 9°300 9*221 9-158 9°178 9°398 9°438 9°351 9°382 | 29°316 Summer 9°012 9°394 9°288 9°166 9°288 9-315 9°296 9°296 9:°398 | 29°277 Autumn 9°343 9:091 9°012 9-036 9°347 9°162 9'288 9°382 9:°272 | 29°194 Winter 9°461 9°347 9°233 9°180 9°300 9°339 9°611 9°489 9528 | 29°442 Year 9°288 9°288 9°205 9°142 9°221 9°312 9°461 9°406 9°410 | 29°307 Departure from Annual Mean.* N. N.E. BE. S.E. Ss. S.W. Ww. N.W. Calm. | Mean Ins. Ins. Ins. Ins. Ins. Ins. Ins. Ins. Ins. Ins, Spring ‘063 012 ‘016 016 O48 086 028 055 028 ‘009 Summer 276 106 083 024 067 ‘003 165 110 012 030 Autumn 055 197 198 106 126 ‘150 Tid O24 188 118 Winter 173 059 028 038 ‘079 027 150 083 118 185 * The heavy type indicates an excess, and the italic type a defect. TaBLe XXI. THERMAL WINDROSE, STATEN ISLAND AND RAwson. DEPARTURE FROM THE GENERAL MEAN OF THE MONTHS SPECIFIED. Sraten Isnanp. LatirupE 54° 23’S., LoneirupE 63° 47’ W. N. N.E. E. 8.E. 8. S.W. w. N.W. | ° | ° ° ° ° ° ° ° | January +1°4 -1°'8 —0°4 -1°4 -0°9 -0°2 +1°3 +22 April +1°4 | +1°'8 -0°4 - 2°0 -—3'2 -1°4 +2°0 +1°8 | July | $25 | +16 | +171 -22 | -40 | =-11 | 402 | 42% | October +2°0 +1°'8 -11 —3'8 —3:'2 +0°2 +1°4 +2°7 | Year +1°8 +0°9 -—0°2 -—2°3 —2°9 —0°5 +1°3 +2°2 Rawson. LATITUDE 48° 17’S., Lonerrupr 65° 05’ W. January +2°9 -3'4 ‘-1°4 -1°8 -3'1 +0°7 +2'2 +4°1 April +18 +0°2 +0°5 -1°3 -3°6 +0°2 —0°7 +2°7 | July . +1°8 +2°2 0°0 -1°8 -25 -4°3 -02 +4°7 October é +3°1 +13 —2°5 -4°3 -5°4 +1°'1 +3°2 +3°6 Year : | +2°3 0°0 --0'9 -2°3 —3°6 -0'5 +1:1 +3'8 —— THE WEDDELL QUADRANT AND ADJACENT AREAS. 133 Additional Note, received 6th September 1908. Notre on THE Mean MontHiy anp ANNUAL BarometTRIcC PRESSURE AT SANDY Pornt, THE FaLKLAND ISLANDS, AND IN THE CAPE Horn Rg&GIONS. The maps of mean monthly pressure given in this paper may be said to approxi- mately represent the conditions prevailing during the five years 1903-1907. For this reason the trend of the isobars over the ocean in the Cape Horn region as well as their relative height will be found in some months to differ from those given in treatises dealing with the results of maritime observations obtained from ships’ logs extending over long terms of years. With a view of giving good normals for land stations in this region, | have made every effort to obtain values for Sandy Point, the Falkland Islands, and for the Cape Horn area. For Cape Pembroke (Falkland Islands) the values of mean monthly and annual pressure which follow are based on 164 years’ observations. For the vicinity of Cape Horn, means deduced from observations taken at Staten Island, Ushuaia, Harbertown, and New Year’s Island are given. The periods embraced by the records comprise 19 years for the months of February, March, and May, 20 years for January, April, June, August, and December, 21 years for July and September, and 22 years for October and November. The monthly averages were obtained by taking the mean of all or any of the above stations, observing simultaneously. Thus for the year 1905 the monthly means are the average of the mean pressures recorded at New Year’s Island, Ushuaia, and Harbertown. By this method a much closer approximation to the true pressure was obtained than if the monthly means had depended on only one station. The mean position of the stations is 54% degrees south latitude, and 65 degrees west longitude. Their range in latitude does not exceed a few miles, but in longitude the stations differ some 45 degrees or 160 miles. This variation in longitude is, how- ever, unimportant, as the isobars in this locality normally approximate to parallels of latitude. For Sandy Point the monthly means are based on 18 years’ observations, so that the values for all three districts are very nearly of the same length, although they are not quite synchronous, being for different terms of years. In spite of this deficiency, however, owing to the paucity of land observations in this part of the globe, they afford the closest approximation that it is at present possible to obtain for the distribu- tion of pressure at the south of the continent of South America, and over the ocean to the east up to the longitudes of the Falklands in about 52°S%. latitude. The follow- ing table shows the mean monthly and annual pressure at the three stations. Mran MonrHty BAROMETRIC PRESSURE AT 32° SEA-LEVEL AND STANDARD GRAVITY. Period.| Years specified. Jan. | Feb. | Mar. | Apr. | May. | June.| July. | Aug. | Sept. | Oct. | Nov. | Dec. | Year. | Station. Sandy Point . |18 years|{ 1871572, 1889- \ | 59.485 | 20-473 | 29°525 | 29°552 | 29596 | 29°651 | 29-571 | 29-644 | 29°650 | 29:599 | 29-520 | 29-446 | 29-559 ’ 1876-79, 1881-83 Cape Horn } hi9-22,,, 1886-96, 1899, +| -437| -445| -432| -502| -495| 551) -527| -570| -625| 538] -434| -438| -499 Breton: - 1902-07 C. Pembroke or 16},, (Falklands) . 1859-68, 1875-77, 481 1882-83, 1903-05 “531 Ath *539 “538 549 "644 623 "BBO 492 “B51 134 MR ROBERT C. MOSSMAN ON THE METEOROLOGY OF As regards the Cape Horn region and Sandy Point, the above means differ but little from those given in Table IJ. At the Falkland Islands, however, the differences are considerable, amounting on the mean of the year to an excess of 0°067 inch during the short period, while in July, August, November, and December the excess amounts to more than a tenth of an inch. At all three stations it will be observed the highest mean monthly pressure occurs in September. The month of minimum pressure, on the other hand, is December at Sandy Point, January and March for the Cape Horn area, and March at the Falklands. The range in the mean monthly pressures is 0°204 inch at Sandy Point, 0°194 inch in the vicinity of Cape Horn, and 0°170 inch at the Falkland Islands, being greatest in the region most removed from the direct influence of the ocean, and least at the strictly insular station on the Falklands. It is of interest to give for the Cape Horn area the difference of mean monthly pressure for the five years 1903-1907, as compared with the above long average, the plus sign indicating that pressure during these five years was in excess, and the minus sign that it was in defect, of the long average. Jan. Feb. March. April. May. June. July. Aug. Sept. Oct. Nov. Dec. _— Year. in. in. in, in. in. in. in. in. in, in. in, in. in. +058 +066 --080 +7040 4-032 --098 +084 -012 -‘114 +4:007 -015 +014 --001 Thus during the past five years pressure in the Cape Horn region has on the mean of the year been virtually the average, but with considerable variations in the months, ranging from an excess of +°084 inch in July to a defect of —0°114 inch in September. The following table showing the highest and lowest mean monthly pressures in the vicinity of Cape Horn may be of interest, owing to the comparatively (for the region) long period covered by the combined records. Care Horn Recion Latirupe 54° 30’ S., Lonertrupr 65° W. Hicuest anp Lowrst Maan Montuiy Pressure at 32° Sea-LEVEL AND STANDARD GRAVITY. Highest. Lowest. Range. Inches. Inches. Inches. January : j 29-752 1878 29°236 1890 0'516 February. ‘ 610 1896 ‘170 1891 0:440 March . ‘ ; 840 1878 "185 1895 0°655 April . : : ‘811 1878 310 1892 0°501 May . : : JB 1904 "284 1907 0°453 June . ‘ ; 816 1888 "285 1905 0°531 | July . ; é T20 1887 287 1889 0-436 | August ; , 897 1907 342 1905 0°555 | September . : 890 1886 353 1889 0°537 October : : 876 1876 256 1892 0620 | November . ; ‘759 1876 Bl 1896 0°637 December . : (Ae. 1904 ‘210 1889 0°512 "Year . é ; 29°646 1878 29°412 1889 0:234 * The years with complete observations are 1876, 1878, 1888-1893, 1896, 1903-1907. THE WEDDELL QUADRANT AND ADJACENT AREAS. 135 Hence the highest mean monthly pressure was 29°897 inches in August 1907 and the lowest 29°122 inches in November 1896, showing a range of 0°775 inches. Looking at the last column, it will be seen that the greatest monthly range or difference between the highest and lowest mean monthly pressures is 0°655 inches in March, and the least 0°436 inches in July. REFERENCES. Antarctic Manual, London, 1901. Arctowsk!, Henryk, Lxpédition Antarctique Belge: Résultats du Voyage du S.Y. ‘“‘ Belgica” en 1897- 98-99 : “‘ Météorologie: Rapport sur les Observations Météorologiques Horaires,” Anvers, 1904. Witson-Barker, D., Commander R.N.R., “The Present Position of Ocean Meteorology,” Quar. Jour. Royal Meteorological Society, vol. xxx. pp. 105-132. Bopman, Gosta, “ Meteorologische Ergebnisse der Schwedischen sudpolar Expédition,” Petermanns Greogr. Mitteilungen, 1904, Heft v. — “Om Klimatet i Antarktis mel. sirskild hinsyn till Graham Land, U R Ymer,” Yidskrift Utgifven af Svenska Gallskapet for Antropologi och Geografi, Arg. 1904, H. 3. Brucz, W. S., ‘ Bathymetrical Survey of the South Atlantic Ocean and Weddell Sea,” Scottish Geographical Magazine, August 1905. Bucuan, A., Report on Atmospheric Circulation. Cui: Annuario del Servicio Meteorologico de la Direccion del Territorio Maritimo, 1899-1905, Valparaiso, 1902-1906. Contributions to our Knowledge of the Meteorology of the Antarctic Regions, published by the authority of the Meteorological Committee, London, 1873. “ Challenger” Reports: ‘‘ Narrative,” vol. 1i. Davis, Water G., Climate of the Argentine Republic, Buenos Aires, 1902. Anales de la Oficina Meteorologica Argentina, Tomo xi. ——— “Observations made at Laurie Island, South Orkneys, in the Year 1904,” Annals of the Argentine Meteorological Office, vol. xvi., in Spanish and English, Dosrowo1skl, A., Hupédition Antarctique Belge: Résultats du Voyage du S.Y. “ Belgica” en 1897-98-99 : “‘ Météorologie : Observations des Nuages,” Anvers, 1903. GouLb, Benvamin A., Anales de la Oficina Meteorologica Argentina, Tomo iil. Hepworth, M. W. Campse.t, C.B., Commander R.N.R., The Relation between Pressure Temperature and Air Circulation over the South Atlantic Ocean, published by authority of the Meteorological Com- - mittee, Official No. 177, London, 1905. Lepaay, J., Mission Scientifique du Cap Horn, 1882-1883, Tome ii., ‘‘ Météorologie,” Paris, 1885. Marasinl, P., Observatorio Meteorolégico del Colegio Salesiano “8S. Jose” en Punta Arenas de Magallanes (Chile): Restimen de las observaciones de quince uiios (1888-1902), Santiago de Chile, 1904. Mernarpus, Winn, Uber die Wind Verhdltnisse au der Winter station des ‘* Gauss,” Berlin, 1905. Mossman, R. C., Results of the Scottish National Antarctic Expedition, vol. ii. of Report on the Scientific Results of the Voyage of the S.Y. “ Scotia,” Edinburgh, 1907. -——— ‘Preliminary Reports on the Meteorological Results of the Scottish National Antarctic Expedition,” Scottish Geographical Magazine, vol. xx. pp. 113-120; xxi. pp. 417-429; xxii. pp. 252-272. ——— “The Recent Voyage of the ‘ Uruguay,’” Scottish Geographical Magazine, June 1905. “The South Orkneys in 1907,” Scottish Geographical Magazine, July 1908. Rey, Lieut. J., “‘ Note sur les Observations Météorologiques” (Appendix to Le Frangais au Pole Sud, by J. B. Charcot), Paris, N.D. Soutn Groreia: Die Internationale Polarforschung, 1882-1883: Die Beobachtungs-Hrgebnisse der Deutschen Stationen, Band ii.: “‘ Sud-Georgien,” Berlin, 1886. Wesster, W. H. B., Narrative of a Voyage to the Southern Atlantic Ocean in the Years 1828-29-30, 2 vols., London, 1834. WEDDELL, James, A Voyage towards the South Pole, London, 1825. TRANS. ROY. SOC. EDIN., VOL. XLVII. PART I. (NO. 5). ° 19 136 METEOROLOGY OF THE WEDDELL QUADRANT AND ADJACENT AREAS. NOTE. The maps on Plates lL. to [V. show average barometric pressures, temperatures, and amounts of cloud for the region dealt with in this paper. They are based on the values given in the foregoing tables, adjusted in some cases so as to indicate what is believed to be more nearly the true average (see page 104). The isobars, etc., on these maps were drawn by Mr E. K. Carmicuarn from those on sketch-maps supplied by the author. The general map, Plate V., is based on that accompanying a paper on the Bathymetrical Survey of the South Atlantic Ocean and Weddell Sea, by Dr W. S. Brucz, LL.D. (Scot. Geographical Mag., vol. xxi. p- 402, August 1905). The map has been revised by Dr Brucs for this paper. Vou. XLVII. Edin. Trans, Roy. Soc. “YQUOTY Towa 1oj sxeqosy] ‘aInssatd ILI}eulo, leg WRT é 1eqme09q | Sq JaquUsaAON ‘PADYIUULIMY “ST “7 190300 Ol Sa Mossman on “The Meteorology of the Weddell Quadrant and Adjacent Areas.”—Puare I. Mr Rosert C, qoreyy € 51 Areniqay 6 3M daquiaydag 6 3ty Aurnue I 314 XLVII. VoL. Trans. Roy. Soc, Edin. Mr Rosert C. Mossman on “‘ The Meteorology of the Weddell Quadrant and Adjacent Areas.”—Puate II. ‘YJUOT Yowa oJ suLtoyJosy ‘ainqetroduray, weep daqma00qd Zl Si qeqmeAo N IL Sly 4290799 Ol aly ‘ADYoNULIMD “M12 Ksenigeaq 6 Od Auenue pe 1 314 VoL. XLVII. Trans. Roy. Soc. Edin. "YQUuoy yore 10y sydouosy “pnoly jo JUNO Y wed, “TADYoUU.) “YB *7ap Jaqmessq Jaq WeA0N ZI Sl II Si ant and Adjacent Areas.”—Puars III. yuadr id ( Mr Rosert C. Mossman on “ The Meteorology of the Weddell 13903009 Ol Sq saquieydag 6 Sy Kaeniqay 3 Sl Avenues 1 3a Trans. Roy. Soc. Edin. Vou. XLVI. Mr Ropert C. Mossman on ‘“‘ The Meteorology of the Weddell Quadrant and Adjacent Areas.”— Prarn LV. --—- /---1---------------- 4 \ \ del. EF. K. Carmichael. Fie, 1.—Isobars for the Year. Fic. 2.—Isotherms for the Year. Fre. 3.—Isonephs for the Year. Fic. 4.—Isobars and prevailing Winds for December 1904. Fic. 5.—Mean Surface-temperature of the Sea in Summer. Plate V. Roy Soc. Edin? Vol. XLVIL CET \ 5 erage \ <2 oueTrY9,, Elle gsez aegnsny [woupdeaZoan qZanquypy ery SUM aU avn SS So aS OO Vv T1SUUSM AU di “WOUIPIGX MIE NON 138 MR R. C. MOSSMAN ON inches, with a maximum of 18°54 inches in June and a minimum of 2°85 inches in January. ‘To the north of this wet zone the rainfall diminishes rapidly, while to the south precipitation continues high. No reports are available for the 600 miles of coast separating Ancud from Evangelists Island at the Pacific entrance to the Straits of Magellan, where the annual rainfall is 111°44 inches, with a seasonal distribution the reverse of what obtains in the monsoonal region. At the island of Juan Fernandez, in lat. 33° 37’ S., long. 78° 50’ W., the distribution of the rainfall through the months of the year is in harmony with that prevailing on the Chilian littoral in the same latitude. The variation in the mean wind direction shows, however, little change; being south in the month of January, when this region is well within the influence of the South Pacific high-pressure area, while in the month of June, when the high-pressure belt is north of its summer position, the winds at Juan Fernandez are south-west. These wind variations are also characteristic of the ocean region to the east of this island. To the north of latitude 30° S. the monsoonal effect on the littoral becomes masked or disappears ; but even at Caldera, in 27°'S., the little rain that falls is precipitated in the rainy season of the more southern stations. At Iquique, situated in 20° S., where a shower of rain is of rare occurrence, the winters are much cloudier than the summers ; the seasonal amount of cloud on the mean of the three years 1901-03 being as follows :— IeuiguE. Croup Amount 0-10. Years 1901-03. Hours of Observation 8-2-9. Spring. Summer. Autumn. Winter. Year. 6°4 3°9 4°7 72 hh) The indirect monsoonal effect would therefore appear to extend for some distance into the northern rainless region. There can be little doubt that the influence of the mountain chain of the Andes in modifying the general circulation of the air is considerable, causing the coastal winds to conform to the shores of the littoral, and to blow parallel with the main axis of the Andean Cordilleras. The whole question clearly calls for a complete discussion of all the available Chilian data, and the co-ordination and correlation of these data with the observations from the Argentine Republic, Bolivia, Peru, and Brazil, as well as the joining up of the land values with those given in various memoirs dealing with the meteorology of the adjacent ocean regions. Among the factors bearing on the causation of the monsoonal effect may be mentioned the seasonal migration of the South Pacific anticyclone, the varying intensity of the high-pressure area in the north- west of Argentina, and the fluctuations in the southern atmospheric cyclonic centres of action in the Weddell and Bellingshausen Seas. The temperature gradient between the land and the ocean at different seasons will also have important bearings on the solution of the whole question, and also variations in the temperature of the cold polar current THE MONSOONS OF THE CHILIAN LITTORAL. 139 which flows along the west coast of South America, extending almost to Juan Fernandez. | The monsoonal effect varies greatly from year to year; some years being character- ised by weak monsoons, while in others the rain-bearing northerly air currents are greatly intensified. There appears to be a very definite interrelation between the position occupied by the Pacific and Atlantic ‘‘ highs” and the Antarctic and sub- Antarctic circulation which may help to throw light on the varying weather conditions prevailing over the South American continent. To take merely a single instance, that afforded by a comparison of the conditions from 20° to 65° S. for July of 1903 and of 1904, based on returns from a large number of stations in South America and at the Falklands, as well as the sub-Antarctic stations of the South Orkneys, Snow Hill, and Wandel Island. In July 1903 the South Pacific “ high,” as shown by the winds and pressure at Juan Fernandez, was well south of its normal position, and rainfall was relatively small at all the Chilian coastal stations within the monsoonal area. In this region southerly winds were in excess, and northerly winds, which bring the rain, in defect of the average. Accompanying this excess of winds from higher latitudes, low temperatures prevailed over Argentina and Chili, except in the extreme south of the continent. Barometric pressure was below the normal at the South Orkneys and at Snow Hill on the east side of Graham’s Land, the Weddell Sea “low” being not only deepened but lying farther west than usual. At Evangelists Island at the Pacific entrance to the Straits of Magellan the weather was mild and precipitation high, the same conditions extending to the Falklands and the South Orkneys, with westerly and north-westerly winds predominating. In July 1904, on the other hand, conditions were exactly the reverse of the month described. The South Pacific “ high” was considerably to the north of the island of Juan Fernandez, where barometric pressure was 0°38 inch lower than in the July previous, while on the coast of Chili in the monsoon zone the fall of pressure averaged 0°20 inch as compared with the July of 1903. A permanent low-pressure area lay off the coast of Chili in lat. 42° S., as shown by the Daily Weather Maps of the Argentine Meteoro- logical Office. To the north-west of this region of low pressure the prevailing winds, as at Juan Fernandez, were from the west-south-west, while at coastal stations to the north of it the winds blew from the north and north-west. All over this region deluges of rain fell (see Table V.), the zone of heavy rainfall extending well into the centre of Argentina between latitudes 38° and 45° S., and over to the Atlantic coast. With this excess of equatorial winds, mean temperature was above the average. South of 45° S. the winds were southerly, blowing in on the northern “low.” Temperature in the southern provinces of South America, at the Falklands, the South Orkneys, and Wandel Island was markedly below the normal, the winds in general circulating round the Weddell Sea “low,” which was far to the east of its position in the July previous. Barometric pressure fell from west to east south of 55° S., and was as high as 29°51 * inches at Wandel Island on the * At 32°, sea-level and standard gravity. 140 MR R. C. MOSSMAN ON mean of the month, diminishing to 29°33 inches at the South Orkneys, and still lower to the eastward. The month in its main features bore a very close resemblance to that of December 1904, which | have treated with some detail in the paper on the Weddell Sea meteorology. It will be seen from the foregoing brief synopsis how the climatic conditions over South America vary according to the position of the southern ‘‘centres of action.” It is not sufticient, however, to confine our investigation to the Southern Hemisphere, because there is an intimate relation between both hemispheres. For example, the cyclonic “centres of action” situated in the Weddell Sea and in the Icelandic region stand at certain periods of the year in a definite relation to each other. In the month of May, in which that relation appears to be most pronounced, a positive departure from the normal barometric pressure at Stykkisholm (Iceland) is a direct index of a negative deviation from the normal in the pressure over a region to the south-east of and including Cape Horn. When there is a posdtive deviation over this latter area, then the departures in the Icelandic region are negative. This, be it understood, applies only to the month of May, and is based on twenty years’ data from Cape Horn and six years’ data from the South Orkneys; but the whole inquiry is in an early stage of development, and further interesting contrasts are to be expected from a discussion of bipolar conditions. TABLE I. PERCENTAGE FREQUENCY OF WIND At Point GaLmrRA, Lar, 40° 01’ S., Lone. 78° 44’ W. 7 Years 1899-1905. Hours or OBSERVATION 8-2-9. Jan. Feb. Mar, | April. May. June. July. | Aug. Sept. Oct, Nov. Dec. Year, ——_—. | N. 20 20 30 42 47 45 32 33 28 13 19 15 29 N.E. 1 1 1 3 5 5 4 3 2 1 1 0 2 E. 0 1 2 1 7 6 6 5 3 1 0 1 3 S.E. ul 2 3 5 2 2 6 5 3 3 2 2 3 Ss. 52 51 36 18 10 9 9 9 20 34 32 52 28 S. W. 11 9 9 6 3 5 7 8 8 16 11 10 8 W. 5 2 4 4 4 4 9 8 11 10 6 6 6 N.W. 5 4 4 10 15 16 22 22 20 5 21 7 13 Calm, 5 10 11 11 il 8 by 7 5 7 8 7 8 | Tape II, PERCENTAGE FREQUENCY OF WinD aT Ancup, Lar. 41° 51’ S., Lone. 73° 50’ W. 6 Yrars 1900-05. Hours OF OBSERVATION 8-2-9, Jan. Feb. Mar, April. May. | June. July. Aug. Sept. Oct. Nov. Dec. Year. | -# 1) oe 16 re at 14 Loy as Le 8 SY Sete 11 12 N.E. 1 8 2 7 12 14 13 9 5 4 3 4 6 E. 2 1 4 4 8 8 5 3 3 2 2 2 4 S.E. 3 4 5 5 15 ual 11 6 6 4 3 2 6 Ss. 9 13 12 15 iy) 13 11 12 9 12 9 8 11 8. W. Lalo i) 8 6 4 6 5 6 5 8 if 10 7 W a 25 24 18 12 | 8 10 13 16 19 28 25 24 19 N.W. | |. 22 18 17 17 546 15 14 24 24 20 27 19 20 Calm. 19 18 18 19 14 9 18 11 11 iat 18 20 15 THE MONSOONS OF THE CHILIAN LITTORAL. TABLE III, 141 PERCENTAGE FREQUENCY OF WIND AT CHILIAN STATIONS FOR THE MONTHS OF JANUARY AND JUNE. 5 YEArs 1901-05, Hours 8-2-9. Nore.—For Angeles, owing to the large number of calms at 8 a.m. and 9 p.m., the data refer to 2 p.m, only. For Santa Maria the means are for the 4 years 1900-03. i January. June. A Lat. ong. Station. 8. Ww. Tez N.| N.E.| E. | S.E. | S.|S.W. | W.|N.W.|Calm.| N.| N.E.| E. | S.E. | S. | S.W. | W. | N.W.| Calm. Caldera c 27 03: 70 53 | 6 3 1 2 |13) 59 5 5 6 | 16 6 9 8 | 17] 23 2 i 12 Point Tortuga . | 29.56 | 71 21 | 11 3 2 4 2 26 24 6 22 9 8 22 7 i 14 14 6 19 Point Angeles . | 33 01 | 71 38 | 2 1 2 0 |13] 52 2 2 26 | 17 6 5 1 5 | 15 2 2 47 Juan Fernandez 33 37 | 78 50 | O 0 0 31 34 26 4 1 4 2 0 1 12 19 32 23 8 3 Point Carranza 35 56 | 72 38 | 12 0 0 0 | 66 2 1 1 18 | 41 0 3 7 | 14 4 it | ale, 18 Santa Maria . | 36 59 | 73 52 9 0 0 1 62 2 2 4 20 35 8 1 1 14 3 6 13 19 Mocha Island . | 38 21 | 73 58 | 11 1 Wy 3" {ies |) aly) 2 5 7 | 38} 13 0 6 | 11 9 4| 14 5 Point Galera 40 01 | 73 44 | 18 i 0 PS NB 9 3 5 5 | 44 6 7 3 9 6 5 16 4 | Ancud 5 ees 41 51 | 73 50} 8 I 2 3 CPI) at Wy Ces |) il PAN) |) ally) ab} 5 8 | 14 cy} atat 18 9 Evangelists Island BDROAn ero KOGN aS) Ol Onl dy bine 13) |h260 4. GQ (ial a a Ge ate |) GB. te) Be 5 TaBLe IV. MEAN MontTHLY RAINFALL, COAST OF CHILI. Station. Tat. Hone. Years.| Period. | Jan. | Feb. | Mar.| April. | May.| June.| July. | Aug. |Sept. | Oct. | Nov. | Dec. | Year. : ey Ins. | Ins. | Ins. | Ins. | Ins. | Ins. | Ins. | Ins. | Ins. | Ins, | Ins. | Ins. | Ins. Caldera 27 03 | 70 53 74 1899-1906 | 0°02) 0°00] 0°00 0°00 0°04] 0°05] 0-44} 019{ 0°03} 0°02) 0°02) O-u2 0°83 | Chanaral. . 29 00 | 71 36 6 1899-1904 | 0°00) 0:00} 0-00 0°06 0°34] 0°86} 1°23) 0°82] 0°02} 0°04} 0:00] 0-00 3°37 | Point Tortuga 29 56 | 71 21 8 1899-1906 | 0°00); 0°00} 0°15, 0-03 1°79| 2°28) 2:22] 1:38] 0°06] 0°01] 0:01) 0:01 794 5 Point Angeles 33°01 | 71 38: § 1899-1906 0°00) 0°02} 0°68 0-19 5°72| 6:6)! 7°76] 4°68) 0°44! 0°37) 0°00! 0°35] 26-82 Juan Fernandez 33 37 | 78 50 5 1902-1906 | 0°72) 0°86} 2°25 311 764] 7:01| 7:79) 3°34) 2°45] 1°39} 1°40] 0°38] 38°34 Point Carranza 35 56| 7238] 4 { tee one | 0-03 0:13] 2:37| 212 | 571] 9-07| 9:93] 3-30] 1-68] 1-44) 0-47] 1-07] 37°32 Santa Maria Island | 36 59 | 73 52 4h 1899-1908 | 0°21} 0°49! 1°56 | 2°25 6°55 | 10°68} 5°87! 3°50] 1°78] 0°98! 0°43] 0°50;| 34:80 Point Lavapie . | 37 08 | 73 36 3 1904-1906 2°48} 343) 368 2°67 7°42| 7:52) 7:83] 2°83] 3°83 1:43] 0°48] 0°79] 43°85 Isle Mocha (West) | 38 21 | 73 58 8 1899-1906 | O°77| 1:23] 3°43 4°87 9°07 | 10°93 | 10°08} 5°68] 3°61} 1:93) 1°33] 1713] 54:06 Point Galera . 40 01 | 73 44 8 1899-1906 2°85| 4°36] 629 10°35 | 14°93 | 18-54/16°28| 12°17) 8°44] 4:41] 3°52] 3°10] 10524 Port Ancud_ . . | 41 51 | 73 50 73 1899-1906 | 3°01} 4°01} 7°04, 9°73 | 11°36) 11°11| 9°76] 884) 8°63] 4°45] 4°09] 2°92] 84°95 Evangelists Island | 52 24 | 75 06 8 1899-196 | 13°15] 8°81!12:03 1018 7°28] 7°72) 7°00) 7:27) 7°65) 10°14! 9°38] 10°83 | 111°44 TABLE V, RAINFALL ON Coast oF CuiLi, Juty 1903 AND 1904. ‘ . Juan Bre Isle Mocha. | Point Port Evangelists Caldera. | Chanaral. Tortuga. | Angeles Wennnnden Carranza. (West). | Galera. iene, evan aS Ins. Ins. Ins. Ins. Ins. Ins. Tus. | Thing, Ins. Ins. July 1903 0:00 0°73 1°74 2°57 1°62 1:02 | 4°66 | 8°66 8°70 12°81 July 1904 0°62 0°62 5°27 yen 1217 LOE22) | 19289 | 25°18 14°64 3°98 TRANS. ROY, SOC. EDIN., VOL. XLVII. PART I. (NO. 6). 21 a lh } t \ ( 143 ) VII.—A Carboniferous Fauna from Nowaja Semlja, collected by Dr W. 8. Bruce. By G. W. Lee, D.Sc., H.M. Geological Survey. With Notes on the Corals by R. G. Carruthers. Communicated by Dr Horne, ¥.R.8. (With 2 Plates.) (MS. received May 29, 1909. Read July 5, 1909. Issued separately September 24, 1909.) INTRODUCTION. The present paper is based on the study of a suite of fossils collected in 1898 by Dr W. 8. Brucs, during a cruise with Major ANDREW Coats in the Barents Sea, on board the yacht Blencathra. The fossils were found wm stu in a cliff at the extremity of Cape Cherney, a promon- tory situated on the west coast of Southern Nowaja Semlja, in lat. 70° 49’ and long. 56° 37’. Contrary to what often obtains in the case of materials collected by explorers, they are all from the same bed, a fact which greatly increases the value of the collection, as there is thus no possibility of a mixing of forms from different horizons. The rock is a compact, brownish-grey limestone, emitting a pronounced fetid odour when struck with a hammer. Numerous thin sections and the residue left after attack by acid show that clastic minerals are small and poorly represented, and it is interesting to note that small patches of violet fluorite, of secondary origin, occasionally fill in eavities in the rock, or replace the aragonite of some of the shells. The limestone is essentially foraminiferal, with thin crinoidal layers, and Dr Brucr tells me that in the field its characteristic feature is the presence of a very large shell [Productus giganteus (Mart.)]; at one spot in that same bed he also observed a large branching body, which may possibly represent a drifted stigmarian root. Most of the fossils are in a perfect state of preservation, but many suffered during the process of developing from the matrix, owing to their small size. In nearly every ease the original shell-substance is preserved, calcite pseudomorphs occurring only in some of the Gasteropods; but even in this case the minutest details of ornamentation are reproduced. I wish here to emphasise the fact that all the fossils must be regarded as contemporaneous, there being no evidence of any having been “ remanié.” An unusual interest attaches to the fauna collected by Dr Brucs, as it proves the hitherto unrecorded presence of marine Lower Carboniferous rocks in Nowaja Semlja ; that is to say, we know now for the first time that the Lower Carboniferous sea did extend to such high latitudes in the western hemisphere; this discovery has therefore more than a merely local importance, as it entails considerable modifications in our theoretical conceptions of the Arctic Lower Carboniferous Continent. For the benefit of those who may not be conversant with Arctic geology, I will add that geologists who have a special knowledge of the Upper Paleozoic formations of these regions have pointed out in TRANS. ROY. SOC. EDIN., VOL. XLVII. PART I. (NO. 7). 22 144 DR G. W. LEE ON ’ various publications that the so-called ‘“‘ Mountain Limestone” of Spitsbergen, Beeren Kiland, Nowaja Semlja, ete., really belongs to the upper divisions of the system, and was identified as Mountain Limestone owing to the fact that the older paleontologists did not realise the importance of a closer comparison with the marine phases of the Upper Paleozoic formations. * As regards former records of Carboniferous rocks in Nowaja Semlja, very little need be said here. Professor TscHERNYSCHEW has ably summarised in his great work (loc. cit., pp. 679-681) all the data bearing on this question, the main facts being as follows:— According to him, the numerous species described by Professor TouLa from the Barents Island,f and generally believed to be Upper Carboniferous, belong to beds of Artinsk age; the other Carboniferous fossils recorded from various parts of the two main islands are too scanty to permit of determining their true horizons. However, a few fossils collected during the expedition of the Jermak in 1901 (Kreuz Bay and Maschigin Bay) indicate a horizon which is not more recent than the Middle Carboniferous of the Urals. These circumstances explain the absence of any reference by me in the descriptions below to species previously recorded from Nowaja Semlja, careful comparison having shown that this fauna is more closely related to that of the typical Lower Carboniferous districts of Russia and Western Europe. On the other hand, a region of which so little is known might have been the seat of the persistence of older types, or of the early appearance of forms known elsewhere from higher horizons only, so that in the determina- tions | have avoided confining myself to literature dealing exclusively with Lower Carboniferous paleontology. Before passing to the descriptive part of this paper, I wish to thank Dr Bruce for having kindly allowed me to study his valuable collection ; and I owe a debt of gratitude to Professor TH. T'scHERNYSCHEW, who generously furnished me with valuable information touching the geology of Arctic Russia. For help of a varied nature I must express my indebtedness to many of my colleagues, specially to Mr R. Lunn, who sacrificed much of his time in the preparation of the accompanying plates. Figs. 42, 43, 44, 46, 47, 48, and 50a are from drawings by Mr H. 8. M‘Vry, and figs. 12a, 13d, 15a, 17a, 22-22d, 24b, 84a, 35, 36, 37a, and 49 are from drawings by myself. DESCRIPTIVE Part. As the main object of the following descriptions is to bring out the faunistic and stratigraphical relations of the species, I have not dwelt at length on morphological features unless these have appeared to me to be new or of importance in systematics. * Freon, Lethea Geognostica (passim); J. G. ANDERSSON, Uber die Stratigraphie wnd Tektonik der Biren Insel, 1899, p. 16; TH. TscHerNyscuew, “ Die obercarbonischen Brachiopoden des Ural und des Timan,” Mém. Com. Géol. Russie, 1902, p. 694. + Sitaungsb. k. Akad. Wiss, Wien, Bd. 1xxi., 1. Abth., 1875. A CARBONIFEROUS FAUNA FROM NOWAJA SEMLJA. 145 For the same reason I have refrained from giving synonymice lists, the references I give being sufficient to show the interpretation adopted in any ambiguous cases, and where no reference is given it is implied that the original specific designation is accepted. Class RHIZOPODA. Order FORAMINIFERA. These organisms are very abundant and well preserved; many of them I succeeded in freeing from the matrix, in order to base the determinations on both external and internal characters. The Carboniferous Foraminifera of Russia have been so accurately described in the exhaustive monograph of V. von MOLLER that the task of identifymg the Nowaja Semlja forms was a specially easy one; consequently I have nothing of importance to add to what has been stated by the Russian paleeontologist.* Genus Enporuyra Phillips. Endothyra globula (dV Hichwald). Moéuter, Joc. cit., 1878, p. 98, pl. xiii. figs. 1-4. This species is represented by a number of specimens in various stages of growth ; one of the largest measures approximately 15 mm. in diameter and 0°8 mm. in thickness. Genus Spirityina Ehrenberg. Spirilina? sp. indet. Two of the slides contain horizontal sections of a non-septate tube coiled on itself in one plane; as the specimens consist of seven whorls—although they measure only 0°5 mm. in diameter—comparison with the species referred to Spirillina by MOLLER is difficult. Likewise I am unable to express an opinion as to their true affinities. In his review of the genus, H. B. Brapy ¢ states that Sprrilline are not known to occur in formations older than those of the Miocene period, while making no reference to MOLLER’s recogni- tion of that genus in Carboniferous strata. Genus CrisrostomumM Moller. I have not sufficient knowledge of Foraminifera to form an independent opinion re- garding the respective claims of p’'Orpicny’s Bigenerina and of MOLLER’s Cribrostomum. * V. von Mouusr, “Die spiral-gewundenen Foraminiferen des russischen Kohlenkalks,” Mém. Acad. Imp. Sciences de St Pétersbourg, 1878 ; and “‘ Die Foraminiferen des russischen Kohlenkalks,” ibid., 1879. + Challenger Reports, vol. ix., 1884, p. 629. 146 DR G. W. LEE ON H. B. Brapy* infers that the Carboniferous species placed under Cribrostomum by Mouter belong to Textularia and Bigenerina, and the latter name is adopted by Dr H. Yaset for Cribrostomum bradyi Moller. “ Oribrostomum” gracile Moller. Moxter, loc. cit., 1879, pp. 59, 60, fig. 23, pl. ili. fig. 4. The examples at hand appear quite typical, but I have not observed any showing the uniserial mode of growth, and none attain the maximum size of the Russian specimens. Genus Terraraxis Ehrenberg. Tetrataais gibba Moller. Moéuter, loc. cit., 1879, p. 73, pl. ii. fig. 4, pl. vii. fig. 3. As regards shape and size the few examples observed agree in every respect with the original definition of the species. Genus ArcHa&DIScUS Brady. Archediscus karreri Brady. Bravy, Monograph of Carboniferous and Permian Foraminifera, Pal. Soc., 1876, p. 142, pl. xi. figs. 1-6. This species is very abundant, and the specimens average 1 mm. in diameter, being thus appreciably larger than the Russian examples known to MOLLER. Genus FusutinELuaA Moller. Fusulinella struver Moller. Motter, loc. cit., 1879, p. 22, pl. ii. fig. 1, pl. v. fig. 4. Fusulinella struvei is represented by numerous specimens in various stages of growth, agreeing remarkably well with the Russian form as delineated by MOLLER. Genus Lacena Walker. Lagena? sp. indet. Small flask-shaped bodies may possibly be referable to some species of Lagena, but as they were observed in thin sections only, they had better be left unclassified, even generically. * Loc. cit., p. 364. + Journal of the College of Science, Imperial University of Tokyo, vol. xxi., article 5, 1906. A CARBONIFEROUS FAUNA FROM NOWAJA SEMLJA. 147 Order RADIOLARIA. Pounded portions of the limestone, after treatment with acid, indicated the presence of a few siliceous organisms, among which were observed unrecognisable fragments showing a regular lattice-work, and a whole radiolarian. The latter was filled with some carbonaceous or ferrugineous substance, but the lattice-work could still be detected, though not very distinctly. In thin sections the small bodies referred here to Radiolaria are more easily seen, and the following genera appear to be represented :— Genus CenospH@RA Ehrenberg. ” Cenosphera “sp.” a. Simple spheres, the lattice-work of which is too indistinct for description, probably belong here. They vary in diameter from 0°! mm. to 0°3 mm., but it is not, possible to say how many “ species” are present. Cenosphera? sp. b. I am in doubt concerning the generic affinities of a few small bodies, 0°1 mm. in diameter, consisting of an apparently simple sphere, the surface of which is covered with numerous short bristles. Genus CarposPpH#RA Haeckel. The genus is perhaps represented by small bodies measuring up to 0°2 mm. in diameter, and apparently made up of two concentric spheres. ‘The distance between the two spheres is greater than the diameter of the smaller one; there are still a few indica- tions of lattice-structure, but the connecting rays are not preserved. Genus CENELLIPSIS Haeckel. In the form which I doubtfuily refer to this genus, the test is simple and oval, and its long diameter measures 0°15 mm. Class SPONGL/L. The few spicules observed are slightly arcuate, with obtusely rounded extremities, and bear a general resemblance to those of Reneria clavata Hinde, as described by Dr G. J. Hrypr.* The largest one is approximately 0°6 long and 0°06 mm. thick. * A Monograph of the British Fossil Sponges, Pal. Soc., 1887, p. 143, pl. ix. fig. 5. 148 DR G. W. LEE ON Class ANTHOZOA. Novres ON THE CARBONIFEROUS CoRALS COLLECTED IN Nowasa SEMLJA By Dr W. S. Bruce. By R. G. CarruruErs. Dr Ler has kindly allowed me to examine the corals included in the collection of paleozoic fossils collected in Nowaja Semlja by Dr W. 8S. Bruce. For the opportunity thus afforded of studying an assemblage of unusual interest, I am sincerely grateful. These corals are all embedded in limestone, together with the rich fauna described by Dr Leg. The specimens are fragmentary, but are otherwise in a fine state of preservation. Notwithstanding the paucity of the material, five distinct genera and seven species can be recognised. ‘These are :— Aulophyllum aff. moseleyranum (Thom.). Campophyllum carinatum sp. nov. Campophyllum kungurense (Stuck. ). Carcinophyllum sp. Lophophyllum cherneyense sp. nov. Lophophyllum sp. Hexaphyllia m‘coyt (Duncan). All of the above genera occur in Carboniferous strata, and, with the exception of Campophyllum, attain their maximum development in strata of Upper Visean age, or in the immediately succeeding beds (Yoredale Rocks of England, Lower Limestone series of Scotland, or lower Mosquensis zone of Russia). The genera Aulophyllum, Carceinophyllum, and Hexaphylha are particularly distinctive, and it is also from beds belonging to this general horizon that the species nearest to the Nowaja Semlja forms have been described. The whole assemblage is clearly indicative of some horizon between the base of the Upper Carboniferous and top of the Lower Carboniferous of the Russian classification. This conclusion is in harmony with that arrived at by Dr Lex from his examination of the other groups of fossils. . Contrary to expectation, only one species can be definitely identified with a Russian form, but it is quite possible that a more complete collection might show a closer affinity to the Russian fauna than the present one does. Attention may be drawn to the revised definition of the genus Lophophyllum that is here furnished, while the discovery of a Carboniferous coral, Campophyllum carimatum, with well-developed carinze on the septa is certainly a point of interest. A CARBONIFEROUS FAUNA FROM NOWAJA SEMLJA. 149 Genus AuLopHYLLUM M. Ed. and H. The characters relied upon by Duncan and THomson in separating their genus Cyclophyllum from the older genus Aulophyllum certainly do not seem of generic value, and, in the writer’s opinion, Cyclophyllum should be discarded. Aulophyllum aff. moseleyranum Thom. The single specimen found of this genus was unfortunately broken off through the floor of the calyx, and as the lower part of the coral is missing, specific determination is impossible. The broken fragment, however, is identical in every way with the transverse section of A. moseleyianum figured by THomson,* except that the vesicles of the central area seem somewhat larger than in THomson’s species. Genus CampopHyittum M. Ed. and H. MM. Mine Epwarps and Haims selected as the type of their genus Campophyllum a coral identified by them with the Cyathophyllum flecuoswm of Goipruss; but according to Scuiurert this identification is erroneous, and their figured specimens (Polypes Fossiles, pl. viii. figs. 4 and 4a) belong to a new species. Such a conclusion, however, does not affect the validity of the genus. A recent examination of the genotype, for which I am indebted to the courtesy of the authorities at the Musée d'Histoire Naturelle, at Paris, and to M. Gravier in particular, shows that the original figures are of a very faithful nature. There is little to add to the description given in the Polypes Fossiles. It may be mentioned, how- ever. that a thin section cut from one of the figured specimens (fig. 4a) showed that at an early stage of growth the septa are amplexoid and dissepiments are developed ; the septa are without carine, and the cardinal fossula is barely discernible. The genus is intermediate in character between Cyathophyllum and Caninia, to both of which it is closely allied. In their mature growth-stages, indeed, it is impossible to distinguish Canina from Campophyllum; it has been held that the depth of the cardinal] fossula is greater in the former genus, but this factor is so variable that little reliance can be placed on it. The greatest difference between these two genera is shown in the young growth-stages. At this period, in Caninia, the septa reach to the centre of the corallum, and dissepiments have not yet appeared; whereas in a Campophyllum, from a very early stage onwards, the septa are short and amplexoid, and dissepiments apparent. In Cyathophyllum dissepiments also appear at an early stage, but the septa reach to the centre of the corallum throughout. These distinctions are not, perhaps, of a very striking character, and it may * James Tuomson, “A New Family of Rugose Corals,” Proc. Phil. Soc. Glasg., 1882, p. 33, pl. iii. fig. 12. + “Anthoz, d. Rhein. Mittel Devon,” Abhand. Geol. Specialkarte v. Preussen, Bd. viii., Heft 4, p. 42 (1889). 150 DR G. W. LEE ON be questioned whether Canina and Campophyllum are in reality generically separable. The retention of the latter genus, however, seems advisable on the whole, if only for convenience in working purposes. Campophyllum carinatum sp. nov. (Pl. I. figs. 3-6.) Two complete specimens and six fragments referable to this species were found in the collection. EXTERNAL CHARACTERS. Corallum elongated, more or less cylindrical, and often slightly twisted. Largest diameter known, 2 cm.; but most of the specimens are much smaller. Hpitheca, thin, smooth, with abundant faint lines and constrictions of growth. In the young and conical part of the corallum, faint longitudinal ribbing, with the pinnate arrangement characteristic of Rugose corals, can sometimes be seen, Calyx unknown except in vertical section. (PI. I. fig. 3.) INTERNAL CHARACTERS. (a) Transverse Sections.—Major septa of uniform thinness throughout, closely set, and never reaching to the centre of the section. The base tabular area in the centre of the corallum is of very variable width (compare figs. 3a and 4, Pl. L.), owing to the amplexoid nature of the septa. The mnor septa are long, and project a short distance past the dissepiments into the tabular zone. Both major and minor septa possess well-marked carinz#, appearing in transverse sections as minute irregular teeth-like projections along the septa. No distinct fossula has been observed. (b) Vertical Sections.—The tabule are numerous, horizontal in the centre of the corallum, but curving abruptly downwards before reaching the dissepiments, which are small, globose, and developed at an early stage. The breadth of the tabulate zone is uniform in the more mature part of the corallum, but that of the dissepimeatal zone is very variable (see figs. 3 and 5, Pl. L.). When the vertical section is cut down the centre of the corallum and passes down the surface of a septum, the true nature of the carinz can be readily observed ; they form closely set spiny ridges running upwards and inwards over the surface of the septa, in a somewhat irregular manner (see figs. 3 and 5, Pl. I.). Under a high magnifica- tion, in the centre of certain of these ridges there is a line of minute black points from which radiating fibres proceed. These black points are probably centres of calcification, and indicate that the carinal ridges are composite, being formed by the fusion of closely set spines, lineally arranged. Remarks.—There are many points of resemblance between this interesting species A CARBONIFEROUS FAUNA FROM NOWAJA SEMLJA. tot and THomson’s Campophyllum echinatum,* but in the latter there are no carine on the septa, and the tabule are much flatter. The same distinctions also separate MILNE Epwarps and Haime’s genotype from the new species. A deceptive resemblance to the inner wall of the Devonian genus Craspedophyllum is often seen in transverse sections, owing to the intersection of a tabula against which the septa terminate, or in vertical sections from the intersection of two septal edges within the tabulate area. The great interest of this new species lies in the carinee that cover the septal surfaces. Such a phenomenon is, of course, common in Devonian corals, but is of extreme rarity in Carboniferous species. A low stratigraphical position for the species does not necessarily follow. Cyathaxona rushiana Vaughan, which seems to be the only Carboniferous coral yet found to have caring, comes from horizons very high in, or above, the Visean. [An examination of C. rushiana shows that the projections on the septa seen in trans- verse sections, to which Dr VaucuHan has drawn attention (Quart. Jour. Geol. Soc., 1908, p. 460), are in reality carine. | Campophyllum kungurense (Stuckenberg). Amplexus kungurensis, A. StuckENBERG, ‘‘ Korallen u. Bryoz. d. Ural u. d. Timan,” Mém. Com. Géol. Russie, vol. x. No. 3 (1895), p. 189 of German text, pl. iii. figs. 10-10c. Only one example of this species has been found in the collection, but fortunately the specimen is complete and well preserved. All the characters are in perfect agreement with SruckENBERG’s figures and descrip- tions, and there is little to add to his diagnosis. The septa become amplexoid at a very early growth-stage, and a vertical section down the calyx confirmed STtucKENBERG’S observation that in the nature part of the corallum a single row of rather large dissepi- ments appears between the tabulee and the wall. Such a feature is not found in a true Amplexus (e.g. A. coralloides), and the coral may more fitly be placed in the genus Campophyllum. Campophyllum kungurense is practically identical with the common spineless form of the Tournaisian coral Campophyllum spinosum (Ampleaus spinosus de Kon.). An examination of several specimens of the latter species, procured from Tournai, showed that the septa and dissepiments were somewhat thinner than in C. kungurense, but no other difference was discernible. It is probable that such a distinction is not of specific value, but, pending further investigation, SrUCKENBERG’s name has been retained. His specimens were obtained from the east side of the Urals, presumably from strata of Permo-Carboniferous age; it seems probable, therefore, that we are dealing with a remarkably stable type, ranging throughout Carboniferous time with but little structural alteration. * James THomson, “A New Family of Rugose Corals,” Proc. Phil. Soc. Glasgow, 1882, pl. iv. figs. 10, 10a. In his large paper, published by the Society in the succeeding year, the same figures are referred to Campophyllum paracida M‘Coy. TRANS. ROY. SOC. EDIN., VOL. XLVII. PART I. (NO. 7). 23 152 DR G. W. LEE ON Genus CARCINOPHYLLUM Thomson. Carcinophyllum sp. (Pl. I. figs. 8-8.) Besides the figured specimen, another example, complete but of small size, was obtained. The corallum is conical and very slightly curved. The calyx is deep and the epitheca smooth, with shght accretions of growth. In the mature growth-stages, when it is separated from the septa, the epitheca is seen to be of medium thickness, but at the proximal end of the corallum appears to be of great thickness, owing to the complete fusion of the septal bases. A comparison of figs. 8 and 8b shows that, with increasing age, the complexity of the lamellz in the central area increases, an Aulophyllwm-like character being assumed. Such a fact, observable also in other species of the genus, suggests that the Aulophylla may have been evolved directly from the Carcinophylla. It is interesting, therefore, to note that Dr Vauauan finds the maximum of the genus Carcinophyllum to be at the base of his D, sub-zone, while the Aulophylla appear much later, at the top of the D, sub-zone. Such a genetic connection as is here suggested does not, of course, preclude the possibility of representatives of these two genera existing side by side. They un- doubtedly do so in this Nowaja Semljan fauna, and also in the Scottish Carboniferous Limestones; Dr Sipty has also found such an association in the D, sub-zone of the English Midlands. Genus LopHopHytium M. Edw. and H. 1845, Cyathaxonia (pars), Michelin. 1850. Lophophyllum, M. Ed. and H. 1876. Koninckophyllum, Nich. and Thom. 1883. Acrophyllum, Thom. (non Nich.) ; non Lophophyllum, Nich, and Thom. The opportunity is here taken to give a revised diagnosis of the genus Lophophyllum, two species of which occur in Dr Brucr’s collection. MICHELIN, in 1846, when describing the corals of Tournai, referred to his genus Cyathaxona a new species which he named C. tortwosa. Later, in 1850, Mine Epwarps and Harmer established the genus Lophophyllum, selecting as the type L. konincki, which also came from Tournai. _ An examination of a large number of topotypes of these two “species” shows that (1) Lophophyllum konincki is simply a young form of “ Cyathaxonia” tortuosa, which (2) cannot be referred to the genus Cyathaxonia, as in the mature growth-stages a well-marked zone of dissepiments appears between the tabulee and the wall, and such a feature is quite absent in Cyathaxonia proper (e.g. C. cornu Mich., or C. rushiana Vaughan). The genotype of Lophophyllum therefore is L, tortuoswm (Mich.), and the revised A CARBONIFEROUS FAUNA FROM NOWAJA SEMLJA. 153 diagnosis of the genus, founded on an examination of this and allied species, is appended below :— Corallum simple and turbinate. Major septa meeting in the centre of the coral in the early growth-stages. One of the septa, usually the counter septum, is strongly thickened at the inner end, giving rise to a prominent columella, which may be discontinuous. In the more mature growth-stages, the columella persists, but the septa usually retreat from the centre and become amplexoid, while dissepiments appear between the tabule and the wall. The tabula are arched upwards in the centre to a varying degree, but, unlike such genera as Dibunophyllum, there is no central zone where the tabule are more numerous or vesicular, nor is there a system of vertical lamellze distinct from the septa. All the simple forms of the genus Koninckophyilum of NicHotson and THomson, and also those corals referred by THomson to Acrophyllum,* thus fall into Lophophyllum as above defined. It is suggested that the compound forms of Koninckophyllum should also be included, as they only differ from a true Lophophyllum in their pronounced lateral budding. From Lithostrotions of the L. martini type the genus is distinguished by being almost always simple and not forming tubular colonies. Those corals referred by NicHo.son and THomson and other authors to Lophophyllum (e.g. L. proliferum and L. eruca) do not develop dissepiments at any stage of growth, and are essentially Zaphrentes having one of the septa thickened at the inner end. It may be convenient at some future time to group them as a sub-genus of Zaphrentis, but for the present such a course is not considered advisable. Lophophyllum cherneyense sp. nov.- (PI. I. figs. 1-2d.) EXTERNAL CHARACTERS. Corallum cylindrical and twisted for the greater part of its length ; conical and curved only at the proximal end. Hpitheca thin, smooth, with numerous constrictions of growth, but no longitudinal ribbing. Calyx unknown. The largest specimen measures 2°5 cm. by 4 cm., but is incomplete; eight other fragments were also found. The full length may have been 8 cm. or more. INTERNAL CHARACTERS. Transverse Sections.—Major septa thin, straight, and tapering towards the inner end; no stereoplasmic thickening. In proximal sections (Pl. I. figs. 2c, 2d) they all reach the central columella formed by a thickening of the inner ends of the cardinal and counter septa. In later growth-stages, most of the septa fall away slightly from the columella, although a few are still prolonged over the central area in an irregular manner. * THOMSON was mistaken in his conception of this genus, which was distinctly stated by NicHoLson to have no columella. Good figures of the genotype, A. oneidense (Billings) are given by LamBE in Contrib. to Canadian Pal., vol. iv. part 1, pl. xvi. figs. 1 and 2 (1899). 154 DR G. W. LEE ON Minor septa fairly well developed, but irregular and not reaching to the tabular area. The cardinal fossula is indistinct, and can only be recognised in some sections by a shortening of the cardinal septum, towards which the axial columella is directed. Vertical Sections.—The median vertical section figured (Pl. I. fig. 2) sufficiently indicates the nature of the tabule and dissepiments. The latter are developed at an early growth-stage. Remarks.—Certain sections (e.g. Pl. I. fig. 2a) of this species closely resemble that figured by THomson* as Acrophyllum sp., but in the latter the major septa are affected by a thickening of stereoplasma within the tabular area. In distal transverse sections (Pl. I. fig. 1) the central area of Lophophyllum cher- neyense approaches in structural character that of a Dibunophyllum, As regards the septa and columella, the Lophophyllum stage is strongly marked in the younger stages of the new species (PI. I. figs. 2a, 2d), while the tabulee throughout approximate to a Lophophyllum type, as opposed to Dibunophyllum, where the tabule have a distinct central area in which they are more numerous and vesicular. Probably all the Dibunophylla pass through a Lophophyllum stage, but assume their characteristic features at so early a period that as a rule they are readily separable from the later genus. Lophophyllum sp. (Pl. I. figs. 7, 7a.) Of this species, apparently new, there is only a single example in the collection, and the conical end is missing. The corallum is continuously conical and slightly curved. The calyx is deep (4 cm.), the diameter of the rim being 4°5 em. The epitheca is thin, smooth, with a few lines and constrictions of growth. The columella is a prominent feature but from the vertical section (PI. I. fig. 7a) is apparently discontinuous. As the specimen is embedded in limestone, it is not possible to say whether this is in reality the case, or is only due to a slight twist in the corallum taking the columella out of the plane of section at this point. The cardinal fossula is clearly visible in cross section (PI. I. fig. 7), owing to the sharply marked boundary of the tabular area encroaching on the dissepimental zone at this point. IncERTaZ SEDIS. Genus Hrxapuy.uia Stuckenberg. Heterophyllia, M‘Coy (pars). The genus Hexaphyllia was established by SruckenBerGt for those species of Heterophylha with six septa only. Although the multi-septate forms of M‘Coy’s genus (e.g. Heterophyllia grandis) probably pass through a hexameral stage early in life, still * “Corals Carb. Syst. Scotland,” pl. xi. figs. 14 and 14a, Proc. Phil. Soc. Glasgow, 1883. + “Anthoz. u. Bryoz. d. unter. Kohlenkalks v. Central Russlands,” Mém. Com. Géol. Russie, nouv. sér., livr. xiv. (1904), p. 72 of German text. A CARBONIFEROUS FAUNA FROM NOWAJA SEMLJA. 155 on the whole the separation of the hexameral species seems advisable. Many of the latter are of considerable size, and certainly cannot be regarded as early stages of the multi-septate forms. These peculiar genera are difficult to classify. They differ from ordinary Paleozoic corals in at least two important respects. In the first case, certain species apparently have no epitheca, for the dark median line of the septa passes completely through the wall to the exterior, instead of being interrupted by an epitheca. In the second case, in certain species a protoseptal stage with four septa at right angles to one another seems to be indicated, and such an arrangement is at variance with our knowledge of other Anthozoa. The remarkable external ornamentation of such species as Hexaphyllia mirabilis, and the general habit of growth, are also peculiar. Further investigation of these curious genera is certainly needed before their systematic position can be secured. Hexaphylliia m‘coyi (Duncan). (PI. L fig. 9.) Heterophyllia m‘coyi, Duncan, Phil. Trans., clvii. p. 645, pl. xxxi. figs. 3a, 3c. Heterophyllia m‘coyi, Tuomson, ‘“ Corals Carb. Syst. Scotland,” Proc. Phil. Soc. Glasgow, 1883, p. 118, pl. x. figs. 20, 20a. There are several fragments of this species, varying from 2 to 6 mm. in diameter. The corallum has a smooth exterier and is six-sided, with parallel walls. There is a low ridge at each angle, to which one of the six septa extends. The dark median line of each septum passes through the wall (which is very thick) to the exterior, and in weathering out gives rise to a faint longitudinal groove down the middle of each of the external ridges (‘‘ costee” of Duncan). No complete specimens have yet been recorded ; the whole corallum may have been 20 cm. or more in length, tapering with extreme slowness. In transverse sections of mature examples the septa are rather thick, and have a regular hexameral disposition. In young specimens they are cruciform, one arm of the eross being forked at each end; while six septa are therefore still present, it appears possible that in still younger stages there may be only four, arranged at right angles to one another. The tabulz are very close together ; they are bent sharply downwards near the wall, and upwards in the centre, but are almost flat in the middle. The appearance of a vertical section varies according to its distance from the centre of the coral. If the section be cut close to the wall, the tabula appear very steeply inclined upwards, but when cut down the centre of the coral they are more flattened (see Pl. I. fig. 9). Remarks.—Neither Duncan nor THomson figures any vertical section of this species, the former simply remarking that “ the endotheca | = tabulze] is very abundant.” The type was not available for examination, but another specimen, in every way identical with that figured by Duncan, and procured from a similar horizon in Ayrshire, is in the collection of Mr James Netuson of Glasgow. Mr Nettson very kindly allowed 156 DR G. W. LEE ON this specimen to be cut, so that the nature of the tabule could be ascertained, and a very necessary addition made to the diagnosis of the species. The fragments from Nowaja Semlja are in complete accordance with the Scottish specimens. Hexaphyllia m‘coyi resembles H. prismatica Stuckenberg,* but in the latter the tabulee are much farther apart. Class CRINOIDEA. Crinoid stems are very abundant, and some of them attain a large size; but as no portions of calyx were observed, any attempt at identification is out of the question. However, two forms at least appear to be represented. Class BRYOZOA. This class is very poorly represented ; the few examples at hand, belonging to the Suborders Trepostomata and Cryptostomata, are specifically indeterminable. Yet the presence of a Stenopora is worth noting, as the genus is not mentioned in SrucKEN- BERG’s recent monograph of the Lower Carboniferous Corals and Bryozoa of Russia.t Genus Srenopora Lonsdale. Stenopora sp. Although thin sections could not be made, on account of paucity of materials, the generic position of the fragments is proved by the nature of the mature end of the zozecia, the casts of which exhibit swellings corresponding to unthickened portions of the wall. : Fenestellid indet. (cf. Polypora papillata M‘Coy). A small fragment of a Fenestellid, the obverse face of which is not shown, bears a general resemblance to Polypora papillata M‘Coy. The arrangement of the fenestrules is the same, and, in common with it, it possesses a small papillated pore at the origin of most of the dissepiments, a character shared by Polypora spininodata Ulrich,t which is otherwise different. INCERTA SEDIS. I place tentatively among the Bryozoa an indeterminable fragment possibly belongs ing to the genus Cystodictya Ulrich. * Loc. supra cit., p. 72 of German text. t+ Mém. Com. Géol. Russie, 1904. | Geol. Survey of Illinois, vol. viii., 1890, pl. Ix. fig. 3. A CARBONIFEROUS FAUNA FROM NOWAJA SEMLJA. RST Class BRACHIOPODA. Judging from the materials at hand, Brachiopods are highly characteristic of the Cape Cherney limestone, where they probably exhibit a rich and varied character, to which full justice cannot be done here, on account of the fact that many of the most interesting forms observed came to grief when being developed, or were so sparsely represented as to preclude the possibility of making satisfactory serial sections. Modern workers on fossil Brachiopods have shown that within certain groups there ean occur remarkable phenomena of homceomorphy and convergence which often make it impossible to estimate the true genetic affinities of many species, unless a knowledge of their internal characters be obtained. Such instances of forms differing in the nature of their internal characters, although apparently nearly allied so far as their general appearance indicated, were met with in a few cases, especially among the Spiriferids, and will be referred to in due course. The mode of occurrence of the various species shows that peculiar conditions, not yet understood, affected the development of some forms more than others. For instance, it is difficult to explain how it is that Productus longispinus J. Sow., is represented by full-grown individuals only, as is also Productus giganteus (Mart.), whilst Productus elegans M‘Coy, is represented by a series of specimens ranging in length from a few millimetres to about 40 mm. The explanation may be that conditions which killed the immature individuals of the latter, left those of the former unaffected ; this explanation seems more satisfactory than that based on the assumption that the nature of the shell in the young stages of Productus longispinus and Productus giganteus was such as to render fossilisation impossible. The assemblage as a whole is quite normal, and there is no evidence of dwarfing or of a depauperised condition. TEREBRATULIDA. Genus DIELASMA King. Drelasma lenticulare de Koninck. (PI. I. figs. 10-10c.) De Koninox, Faune du Calcaire Carbonifére, 6™° partie, 1887, p. 17, pl. 2, figs. 1-9. Few of the numerous species of Dielasma established by pz Koninck have been unreservedly adopted in paleontological literature, but the well-defined characters of his Dielasma lenticulare justify its claims to specific distinctiveness. The Drelasma I refer to it is, like the Belgian form, lenticular and globose, with non-sinuate valves. Frontally, the junction line is perfectly straight. It is represented by a complete specimen, the dimensions of which are: length, 7 mm. ; breadth, 6 mm.; depth, 5 mm. A large pedicle-valve, measuring approximately 20 mm. in length, probably belongs to the same species. Comparison with Davinson’s figures will show that it cannot be 158 DR G. W. LEE ON confounded with any of the forms usually referred to Dielasma hastatum (J. de C. Sow.), and it does not appear to have closely allied representatives in the upper divisions of the Carboniferous system in Kastern Europe. Dielasma gillingense (Davidson). Davinson, British Carboniferous Brachiopoda, Pal. Soc., 1857-1862, p. 17, pl. iii. fig. 1. Two small incomplete specimens, characterised by their very depressed shape, probably belong to this species. SPIRIFERID&. Genus SprrRIFERINA dOrbigny. Spiriferina imsculpta (Phillips). Davipson, loc, cit., p. 42, pl. vil. figs, 48-55. The collection contains a few typical examples of this species. Spiriferina insculpta (Phillips), var. (PI. [. figs. 11-110.) Six specimens, in various stages of growth, differ in no essential degree from the British representatives of Spiriferina insculpta ; the only difference is that the strongly marked growth-lines typical of the species are here very indistinct, although the test is well preserved. Failing to obtain access to the internal characters, owing to the presence of infiltrated calcite, | am unable to say how far this surface feature should affect the systematic position of these specimens. WaacEN* and 8. Nixitint have indicated the differences between Sporiferina msculpta and the Upper Carboniferous Spiriferina ornata Waagen. Spiriferina cristata (Schlotheim), var. octoplicata (J. de C. Sowerby). Davinson, loc. cit., p. 38, pl. vil. figs. 37-47. All the external characters of this common fossil are well exhibited in the materials at hand, consisting of a specimen nearly complete and a few fragmentary ones. The area is large, very much like that depicted by Davipson, fig. 37 (loc. cit.). Spiriferina sp. indet. A very imperfect specimen, exhibiting the characteristic punctate shell-structure of the genus Spiriferina, differs from the above in having the ribs more sharply defined, that is, higher and narrower. Too little is seen to enable one to say whether this difference is to be attributed to anything more than an individual character. * Pal. Indica: Salt Range Fossils, 1883. + Mém. Com. Géol. Russie, t. v., No. 5, 1890, A CARBONIFEROUS FAUNA FROM NOWAJA SEMLJA. 159 Genus Sprrirer J. Sowerby. Spirifer cf. bisulcatus J. de C. Sowerby. The specimen is too incomplete for a satisfactory determination, but it certainly belongs to the group of Spirifer bisulcatus J. de C. Sowerby. Cf. Sporifer treradialis Phillips. One specimen, measuring 6°5 mm. in length and 3°5 mm. in breadth, should possibly be placed here, but its true affinities must remain doubtful, on account of its fragmentary condition. Genus Marriniopsis Waagen. Martimopsis? sp. (Pl. I. fig. 12.) Two pedicle-valves of a Spiriferid show a peculiar combination of characters which might determine its reference to either of the two genera Spirifer or Martinopsis, according to the view taken as to the value of these characters, considered individually. The outline is sub-rhomboidal, with length exceeding width. The larger specimen is 18 mm. long and 16 mm. wide: it is not very convex and has gently sloping sides. The area appears to be small and the beak is not much incurved. The sulcus is narrow and does not increase in width towards the anterior margin. The surface—excepting the cardinal slopes, which are smooth—is ornamented by ten broad, very flat ribs, separ- ated by narrow sulci. One rib occupies the bottom of the median sulcus. The external layer is minutely pitted, as in Martiniopsis and in most species of Martina. Inter- nally, the shell is characterised by two well-developed dental plates. The true generic affinities of this form cannot be determined, owing to the brachial valve (typically septate in Martiniopsis) being unknown; but it is interesting to note that the elongate shape, the median sulcus, and the ribbing of the shell are characters not included in Waacrn’s diagnosis of his genus Martimzopsis, under which the present form ought to be placed, on account of its pitted surface and small area. Hven in the event of the latter view being proved correct, this would not be the first record of an elongate and ribbed Martimopsis.* Genus Martinia M‘Coy. Martina sp. (cf. Spirifera planata Phillips). (PI. I. fig. 14.) A pedicle-valve, 11 mm. in length and 10 mm. in breadth, is externally very much like “ Spirifera” planata Phillips, as redescribed by Davipson (loc. cit., p. 26, pl. vii. figs. 25-36), with this difference, that the sulcus is more sharply defined and ornamented by three ribs only, instead of four or five. * Compare, for instance, Martiniopsis aschensis, TSCHERNYSCHEW, loc. cit., pl. 1., fig. 4. TRANS. ROY. SOC. EDIN., VOL. XLVII. PART I. (NO. 7). 24 160 DR G. W. LEE ON Closer comparison is not possible, as the internal characters of Puituips’ type- specimen are not known; the species described here has no dental plates, and the surface is apparently not punctate. Martina brucei sp. nov. (PI. I. figs 13-138e.) This shell is depressed, sub-rectangular, with width slightly in excess of length; the hinge-line is shorter than the greatest width of the shell. The sulcus and the mesial fold are indistinct. Both valves are very shallow and of equal depth, and the beak is not much incurved. If I am right in assuming that the two specimens described here under this name really belong to the same species, then the ornamentation varies considerably with age ; on the small one it consists of eighteen rounded ribs, whilst on the larger speci- men there are thirty such ribs, a few of which are due to bifurcation. In both, the surface shows an extremely minute pitting, and is covered with sharp, close-set lines of growth quite visible to the naked eye. The area is very narrow and there are no dental plates. The smaller specimen, in which both valves are in apposition, has the following dimensions :— Length . 3 : : , 13 mm. Width . LabhGeiud Depth . : ; ; ; Aa) a5 The larger one is a pedicle-valve 15 mm. long and 18 mm. wide. I do not know any Carboniferous Martinia which could be considered as closely allied to this species. The nature of the ribbing and the lack of distinct suleus and fold give it a general resemblance to the young stage of Martwuwa? inteyricosta (Phillips), as figured by Davipson (loc. cit., pl. ix., figs. 17, 18), from which it differs, however, in having a much narrower area and the pedicle-valve not more convex than the brachial one. There is also a curious superficial resemblance between the smaller specimen and the otherwise very different Martiniopsis baschkirica Tschernyschew,* this being a striking example of close external resemblance between two forms which, according to the modern views, are to be considered as generically distinct on account of their internal characters. Martina sp. (Pl. I. figs. 15-154.) Two pedicle-valves, belonging to a species apparently undescribed, are characterised by their rhomboidal shape, linguiform sinus, and peculiar ornamentation consisting of faint ribs so flat that the narrow intervening sulci are more conspicuous than the ribs themselves. No trace of ribbing can be seen on the cardinal slopes. The larger specimen, 24 mm. long and about 20 mm. wide, has some twenty ribs; the smaller one, 8 mm. long and 7°5 mm. wide, has only a little over half this number. . The whole surface of the shell is covered with fine longitudinal striz regularly * TSCHERNYSCHEW, loc. cit., pl. lxiii. fig. 1. A CARBONIFEROUS FAUNA FROM NOWAJA SEMLJA. 161 distributed, ten of which occupy the space of one rib. No pitting is evident. There are no dental plates. This species might be compared with Martinia rhomboidalis M‘Coy [non Girty, 1908],* in common with which it has steeply falling sides, a linguiform sulcus, and obsolete ribbing. Its elongate shape, flatter ribs, and characteristic longitudinal strize distinguish it from M‘Coy’s species, which is also appreciably smaller. The peculiar strize so conspicuous here have been described in many species of Spiriferids from various horizons, and cannot be said to be distinctive of any particular group. t Genus Reticutaria M‘Coy. @ Reticularia ombricata (J. Sowerby). A small and incomplete specimen has the ornamentation characteristic of this species, but, as I have been unable to see its internal characters, its true position must remain doubtful. Genus SQUAMULARIA Gemmelaro. Squamularia sp. a. (PI. I. figs. 18-180.) The collection contains a small Spiriferid, the general outline and ornamentation of which would determine its reference to Reticularia lineata, were it not that it lacks all traces of the septum and dental plates characteristic of the genus Retzcularia, as defined by M‘Coy. ‘Ten specimens in different stages of growth up to 12 mm. were examined, and all are remarkable for their extremely small and indefinite area. The dimensions of the specimen figured are :— Length . : 5 : : 10 mm. Width . : 5 F : 10%; Depth . ; : : ; BE Squamularia sp. b. (Pl. L., figs. 19-190.) A few examples of a Squamulara differ from specimens of the same size belonging to sp. @ in having a well-defined area and a more robust articulating process. Their ornamentation is the same as that in sp. a. The dimensions of the specimen figured are :— Length . : : 3 : 6 mm. Width = «. ; f : ‘ Gays Depth . , 2 : : 35 ,, * M‘Coy, Synopsis Carb. Limestone Fossils of Ireland, 1844, pl. xxii. fig. 11. : + Good figures of this surface structure are given in Palzxontology of New York, vol. viii. (passim), and in Professor TSCHERNYSCHEW'S Obercarbonischen Brachiopoden (passim). 162 DR G. W. LEE ON I have described these two forms separately in order to show their distinguishing features, but whether these have a specific value cannot be stated at present. Comparison with the known species of Squamularia and with many of the figures of “ Reticularia lineata” is difficult, owing to the general likeness obtaining between the various members of these two groups, so that the claims to specific distinctiveness of the two forms described here cannot be settled without a detailed examination of materials from various parts of the world. Remarks on the Classification of the Spirifercds. Considerable difficulty was experienced in classifying the species enumerated above, owing to the internal characters of some of them pointing to generic (?) affinities different from those indicated by the external features. : An elongate and ribbed Martiniopsis (2), if represented by the pedicle-valve only, is not easily distinguished from those forms of Spirtfer which Mr §. S. Buckman has recently proposed to place under M‘Coy’s genus Brachythyris, as it has not yet been conclusively proved that a small area, the presence of dental plates, anda pitted surface are characters not obtaining in this group. Martinia brucei sp. nov., is more strongly ribbed than most Martine, and bears, in its young stage, a striking resemblance to an Upper Carboniferous shell referred to Martiniopsis; and Squamularia sp., is hardly distinguish- able from forms referred to Reticularia lineata. That externally similar shells may differ in the nature of certain internal features has long been known, but paleontologists have not always agreed as to what degree of importance should be attached to these particular features, to wit, in this case, dental plates. Hatt and Cuarks* indicated these characters in some forms of their “ Glabratc” section, but without formulating a definite opinion as to their value, and the genus Reticularia was believed by them to include forms devoid of dental plates, this belief being probably based upon Waacen’s re-definition of the genus.t But neither WaacEn nor Haun and CiarKeE paid sufficient attention to the fact that M‘Coy, when establishing his genus Letecularia, emphasised in his diagnosis the presence of a septum and dental lamelle,{ a character observed in 1887 by Professor TscHERNYSCHEW in a specimen from Visé, figured for comparison with Devonian materials.§ In 1895 Dr A. Tornquisr commented upon the generic value of the term Reticularia, and restricted it to its original implication ;|| and under the name of Martinia lineata M. A. JuLtEen figured in 1895 a Reticularia in which a septum and dental plates are discernible. 1 * Paleontology of New York, vol, viii., 1898, and Introduction to the Study of the Brachiopoda, 1893. t Waacen, Pal. Indica: Salt Range Fossils, p. 588, 1887. { M‘Coy, Synopsis, p. 142, 1844. § Tscurrnyscuew, “Die Fauna des mittleren und oberen Devon am West-Abhange des Ural,” Mém. Com. Géol. Russie, 1887, pl. x. fig. 1. || A. Tornquist, Das fossilfiihrende Untercarbon am dstlichen Rossbergmassiv in den Vogesen, 1895, p. 119. A. Junin, Le Terrain Carbonifere marin de la France Centrale, 1896, pl. ii. fig. 10. A CARBONIFEROUS FAUNA FROM NOWAJA SEMIJA. 163 Dr G. H. Girty, who treats of this matter very exhaustively, proposes to restrict Reticularia to its original implication, and quotes a number of American Mississipian forms referable to this genus.* English authors who have of late years given much attention to the study of the internal organisation of Carboniferous Brachiopods also consider dental plates to be characteristic of Reticularia, but no mention is made by them of the presence of a median septum,t and it would be rather interesting to know whether there are actually such forms of “ Reticularia” with dental plates, but without median septum. Now, the interesting point already fully elucidated by Dr Girry is that there are forms having the typical ornamentation of “ Sprifer lineatus,” but quite devoid of dental and septal plates, and for these forms Dr Girty adopts GEMMELARO’S genus Squamularva, with this interesting remark, that in America it is an Upper Carboni- ferous genus, whilst the septate Reticularia is essentially Lower Carboniferous. On the other hand, such is not the case in Europe, as proved by the researches of various authors. WaacEN, who compared his Indian materials with specimens of “ eticu- laria” lineata from Visé, found the latter to be devoid of internal partitions, and Dr H. Scurin, after examining a number of specimens of “ Spuifer” lineatus from Germany, stated that this species is usually without dental plates.[ In this same work he points out that German examples of Spirifer glaber often have dental plates, and on this evidence he denies a generic value to Reticularia and Marta, a point of view shared by HE. ScHELLwien.§ The genus Squamularia having been created for the reception of Permian species, the above review was necessary in order to show that its presence is not incompatible with the Lower Carboniferous affinities of the Cape Cherney fauna, since the Lower Carboniferous rocks of the Continent are said to contain representatives of a non-septate “ Spurifer lineatus,” and such forms have also been recorded from Britain.|| Genus Ampoca@:iia Hall. ? Ambocelia wrei (Fleming). This species is possibly represented by an incomplete specimen of which I have not been able to determine the internal characters. * G. H. Girty, “The Carboniferous Formation and Fauna of Colorado,” U.S.G.S., Professional Paper No. 15, 1903, p. 31. + T. F. Srety, “On the Carboniferous Limestone in the Mendip Area,” Quart. Journ. Geol. Soc., 1906, p. 375 ; A. VauaHAN, “On the Faunal Succession in the Carboniferous Rocks at Loughshinny,” zbid., 1908, p. 469; S. S. Bucxmay, “ Brachiopod Homeomorphy,” 7zbzd., 1908, p. 31. t H. Scupin, ‘‘ Die Spiriferen Deutschlands,” Palxontographica, 1900, pp. 5 and 6. § E. ScHEeLiwien, “ Die Fauna der Trogkofelschichten,” Abh. k. k. Geol. Reichsanstalt, 1900, p. 67. || A. Vauanan, “The Carboniferous Limestone Series of the Avon Gorge,” Bristol Naturalists’ Society Proceedings, 1906, p. 159; S. S. Buckman, loc, czt., 1908, p. 33. 164 DR G. W. LEE ON RHYNCHONELLID&. The collection contains only a few and _ totally indeterminable fragments of Rhynchonellids. ATHYRIDZ. Athyris ? spp. indet. (PI. I. figs. 20, 21.) The Athyrids are represented by a few small specimens referable to two distinet forms. Their shape is very depressed, but as this character appears to be partly due to crushing, they had better be left undetermined. Genus CamMARosPIRA Hall and Clarke. Camarospira ? sp. (Pl. L. figs. 22-22d.) A small Brachiopod exhibiting remarkable internal features appears to be specially abundant in the Cape Cherney Limestone; unfortunately, none of the specimens obtained is complete enough to permit of a satisfactory determination. Externally, the shell has an athyroid aspect, but the pedicle-valve is much larger than the brachial one, and has a high and apparently incurved beak. The brachial valve is much flatter than the pedicle-valve, and its umbo is very low. The hinge-line is arcuate, and the cardinal area appears to be obscure, but the shoulders are well marked. The apical region being broken in all the specimens, the presence or absence of a foramen could not be determined. Both valves are evenly convex but for a faint depression in the anterior portion of the pedicle-valve, and a certain mesial gibbosity in the brachial valve. The shell is quite smooth, and its structure is fibrous and impunctate. Internally, the pedicle-valve is characterised by highly developed and strongly convergent dental plates uniting to a high septum and thus forming a spondylium which extends to about three-fourths of the length of the shell. The supporting septum appears to reach the frontal margin, so that longitudinal splitting is common among the specimens observed. Of the brachial valve, two specimens only were available for study. There is no evidence of a spondylium; but a septum, the dimensions of which could not be estimated, is present and exhibits a thin black median line. The nature of the latter is not easily explained, since the septum does not seem to be the result of the joining together of septal plates. The articulating process could not be satisfactorily studied; it appears, however, to be very robust for a shell of this size—the largest specimen observed being only 12 mm. long. The only Brachiopod to which this species bears some structural resemblance is the A CARBONIFEROUS FAUNA FROM NOWAJA SEMLJA. 165 Devonian Camarospira eucharis (Hall).* The features common to both are the presence of a spondylium in the pedicle-valve, and that of a septum in the brachial one; but here the spondylium extends anteriorly to about three-fourths of the length of the shell, whilst in Camarospira eucharvs it is only about one-fourth of that length, and its supporting septum is much shorter. It must be understood that the reference of this species to the genus Camarospira is for the sake of convenience only, and cannot be substantiated without confirmatory evidence from other important characters. In any case, enough is known to show that it has no affinities to any of the smooth Camarophorids. STROPHOMENID. Genus SCHUCHERTELLA Girty. Schuchertella crenistria (Phillips). Davinson, Joc. cit, pl. xxvi. A few fragmentary specimens of this species are in the collection. Dr G. H. Girry has fully explained his reasons for restricting the term Orthotetes to its original impli- cation, viz. to septate forms allied to Derbya Waagen; and for the group of Strepto- rhynchus lens White, and of Spirifere crenistria Phillips, thus left anonymous, he pro- poses the generic term Schuchertella. In the present determination it is assumed that PHILuips’ type-specimen belongs to the aseptate form as interpreted by Davipson.t Orthotetid indet. (PI. I. figs. 23-23a.) A small Orthotetid, represented by a pedicle-valve and two brachial valves, is char- acterised by its biconvex shape, with the hinge-line shorter than the greatest width of the shell. The cardinal area of the pedicle-valve is very elevated, and the beak is dis- torted; the brachial valve has no area. ‘The ornamentation consists of fine raised lines separated by broad sulci. This combination of characters gives it a superficial resemblance to Meekella (Ortho- tetina) oluvieri (de Verneuil), but sections made in both valves show the total absence of the dental and septal plates characteristic of Meekella, and further, there is no median septum asin Derbya and in Orthotetes [emend. Girty]. Access to the details of the articulating process being impossible, | am unable to say whether this form should be considered as an aberrant Schuchertella or be referred to the genus Streptorhynchus. It would be interesting to prove the occurrence of a Streptorhynchus in the Cape Cherney Limestone, since the genus is said by SCHELLWIEN to have a far greater vertical range * Hau and CrarKe, Palxontology of New York, vol. viii., pt. ii., 1893, pl. 1. figs. 46-52, p. 82. + G. H. Girry, “The Guadalupian Fauna,” U.S.G.S. Professional Paper No. 58, 1908, pp: 156-199, 166 DR G. W. LEE ON than is usually admitted, having, according to him, made its first appearance in Devonian times.* ORTHIDA, Genus RuiPrpoMELLA Oehlert. Rhipidomella machelini (L’Eveillé). (Pl. I. fig. 16.) Davinson, loc. cit., pl. xxx. figs. 6-8. So far as external characters may be trusted, the numerous specimens which I refer to Rhipidomella michelini are undistinguishable from the west of Europe form. Genus ScuizopHoriA King. Schizophoria aff. resupinata (Martin). (Pl. I. fig. 17-174.) Compare Davipson, loc, cit., pl. xxx. fig. 1. A single pedicle-valve, globose and very transverse, with a shallow mesial sulcus, has the general appearance of Schizophoria resupinata, from which it differs in the nature of its ornamentation. The ribs are stronger and separated by wider sulci than in the British form; near the margin the sulci are as broad as the ribs separating them. The shell is very thick. The pores and the pits corresponding to the spine-bases are disposed as in Schizophoria resupinata. Schizophoria sp. (Pl. IL figs. 24-24c.) This species is represented by a complete specimen measuring :— Length . : ; : ¢ 11 mm. Width =. ; ; ; : Los i Depth : ; : : : Des and by two isolated valves. The ribbing is very fine, and the valves are non-sinuate and much depressed ; the cardinal area is short and low. The shell is very thin. A puzzling feature exhibited by the brachial valve after decortication is the hexa- plicate appearance of the muscular impression; it may be, as suggested by my colleague Dr [vor Tuomas, who kindly looked at the specimen, that the posterior con- centric division is but the termination of unusual accretion of material below or about the cardinal process. The cast of the brachial valve of Orthis pulvinata Salter, shows an identical feature, due, as explained by Satrmr, to the impression of the “ hinge-teeth” — and of the cardinal process, the three happening to be of the same length in this species ; * E. ScHELLwien, loc, cit., p. 17 ; and “ Beitrage zur Systematik der Strophomeniden des Oberen Palaeozoicum,” Neues Jahrbuch, Ba. i., 1900, p. 5. A CARBONIFEROUS FAUNA FROM NOWAJA SEMLJA. 167 but in the present case, what is left of the shell in the cardinal region is more in favour of Dr THomas’ explanation.* The depressed shape of this shell gives it a greater resemblance to the form referred by SzemEeNow and Mozuer to “ Orthis striatula” than to any other Schizophoria, but in the Nowaja Semlja shell the hinge-line is shorter.t PRODUCTID. Genus CHoNETES Fischer de Waldheim. Chonetes papilionacea (Phillips). (Pl. U. fig. 25.) Davipsoy, loc. cit., pl. xxxv. figs. 3-5. The Cape Cherney form is undistinguishable from the flatter variety of the British Chonetes papilionacea. Dr T. F. Srpty has recently remarked that the term papilionacea embraces more than one form, and for the flat kind—as is the present one—he proposed the new name “compressa,” a name very expressive of the general appearance of the shell, but unfortunately already applied to another species of Chonetes.{ Chonetes (sp. plur.?) (PI. IL., figs. 26-284.) Numerous specimens of all sizes up to 12 mm. belong to one or more species, the general characters of which are as follows :—The shape is more or less semicircular, and the pedicle-valve evenly convex, but rather shallow ; the brachial valve is very concave, so that there is little space between it and the brachial one. The ribs increase by forking, but do not number more than about forty, on the larger specimens. Owing to the fragmentary condition of the material an accurate determination cannot be given, but it may be indicated here that where the shell’s width does not exceed its length, and the shell is ornamented by few ribs, it bears a certain resemblance to the Devonian Chonetes armata (BoucuaRD), as figured by DE Konincx,§ whilst the more transverse ones, with more numerous ribs, appear identical with the form provisionally described as Chonetes cf. crassistria by Dr A. VaucHan.|| A third type is represented by a specimen more globose than the above, with a com- paratively low area, as obtains, for instance, in Chonetes minuta (Goldfuss), from which it differs in the nature of its ribbing.4 * J. W. Sauter, “Note on the Fossils from the Budleigh Salterton Pebble-Bed,” Quart. Jour. Geol. Soc., 1864, p. 295, pl. xvii. fig. 8; Davinson, Brachiopoda of the Budleigh Salterton Pebble-Bed, Pal. Soc. 1881, pl. xli. fig. 11. + “Uber die oberen devonischen Schichten des Mittleren Russlands,” von P. SeMENow und V. von MétteEr, Bull. Acad. Imp. Scrences, St Pétersbourg, 1863, p. 691, pl. ii. fig. 10. ¢t WaaaeEn, Pal. Indica: Salt Range Fossils, 1884, iv. p. 630; T. F. Sipty, “On the Faunal Succession in the Carboniferous Limestone of the Midland Area,” Quart. Jour. Geol. Soc., 1908, p. 78, pl. i. fig. 7. § Monographie des Genres Productus et Chonetes, 1847, pl. xx. fig. 14. || “ Palzeontological Sequence in the Bristol Area,” Quart. Jour. Geol. Soc., 1905, p. 294, pl. xxvi. fig. 2. 4] As interpreted by De Kontnck, loc. cit., fig. 18. TRANS. ROY. SOC. EDIN., VOL. XLVII. PART I. (NO. 7). 25 168 DR G. W. LEE ON Genus Propuctus J. Sowerby. Productus giganteus (Martin). The materials are fragmentary, but enough is at hand to show that more than one form may be represented. | (1) A specimen which when complete must have measured about 15 em. in width has the type of ribbing and the furrowed surface characteristic of the British examples figured by Davinsoy, pl. xxxviil. and xxxix. (loc. cit.). (2) A specimen about 12 em. wide and very transverse, the surface of which is not furrowed, has the ribs separated by wider sulci than in the above. (8) Avery large specimen, about 15 cm. wide, is less transverse than the two others, and has finer and more close-set ribs; its surface is not furrowed. (4) Three small specimens have the Hdelburgensis type of ornamentation. These four forms are too poorly represented to enable one to say whether they belong to distinct varieties (or species?) or are but the different growth-stages of one and the same species. Productus antiquatus J. Sowerby. Mineral Conchology, pl. 317, figs. 5-6. The collection contains a large specimen, 7 cm. in width, agreeing very well with J. Sowersy’s figure. This form and the following I cite separately, but without expressing an opinion as to their relationship to the Productus semireticulatus of authors. Speaking of the latter, Professor 'I'scHERNYSCHEW has made the interesting remark that the flatness of its dorsal valve is a good criterion for distinguishing it from certain species of the group of Productus boliviensis d'Orbigny, to which it often bears some resemblance. Productus concinnus J. Sowerby. (Pl. Il. fig. 33.) Mineral Conchology, pl. 318, fig. 1. A strongly geniculated brachial valve seems referable to this species, the main characters of which are well exhibited, with this difference, that in the frontal portion the ribs are separated by wider sulci than appears to be the case in the British form; but this may be an age-character, this being a large specimen 30 mm. in width. Productus longispinus J. Sowerby. (Pl. Il. fig. 31.) Davinson, /oc, cit., pl. xxxv. figs. 5 and 12. The numerous specimens at hand belong to the slightly transverse, evenly convex type as figured by Davipson, fig. 5 (oc. cit.). One specimen only shows a slight con- vergence towards the lobate form. I have not been able to ascertain whether they possess the “ marginifera” ridge usually present in this species. A CARBONIFEROUS FAUNA FROM NOWAJA SEMLJA. 169 Productus longispinus? (var.?). (PI. Il. fig. 32.) A few specimens are characterised by their fine ribbing and very convex visceral region. They resemble Productus spinosus J. Sowerby (Davipson, loc. cit., fig. 17), but the materials are too fragmentary to enable one to make a decided statement as regards their true affinities. Productus sp. indet. A fragmentary pedicle-valve, too incomplete for a detailed description, but belonging probably to the semireticulati, is suggestive, in the general style of its sculpture, of the form referred by SemeNnow and MOo.ier to Productus carbonarius de Koninck.* It may, however, well be that this is but an appearance due to the imperfect state of preservation of the specimen. Productus ef. margaritaceus Phillips. (Pl. Il. fig. 29.) A pedicle-valve 10 mm. wide, and about 8 mm. long, is too small to permit of close comparison with the figures of Productus margaritaceus given by Puiuirps and Davipson, but on the other hand appears almost identical with the small form referred to this species by Dr A. Tornquist.t The Upper Carboniferous Productus parvulus Nikitin [non Winchell, 1863]{ has the same outline and ribbing, but differs from it in being more convex, with a broader visceral region on which are spines, here, however, very scarce and restricted to the cardinal slopes. Productus coats: sp. nov. (PI. IL figs. 35-35a.) This shell is longitudinally oval, and the hinge-line is a little shorter than the width of the shell. The pedicle-valve is gibbous, without a sinus; the ears are very small. The beak is very prominent, and its extremity overlies the hinge-line. The brachial valve is just perceptibly concave, so that there is a large space between it and the pedicle-valve. The ornamentation of the pedicle-valve consists of tubercles which are very elongate and rib-like on the mesial and frontal portions, and rounded on the ears and cardinal slopes. The ornamentation of the brachial valve is very different, consisting of rounded pits. The specimen figured is a slightly distorted pedicle-valve, measuring :— Length . ; § : ; 12 mm. Width . : i : : oe Depth . : : : ‘ ae * Loc. cit., Bull. Acad. Imp. Sciences St Petersbourg, 1863, p. 704, pl. iv. fig. B. + Das fossilfiihrende Untercarbon am éstlichen Rossbergmassiv in den Vogesen, 1895, pl. xiv. fig. 4. + Nixitin, Mém. Com. Géol. Russie, vol. v., No. 5, 1890, pl. i. figs. 13-14. 170 . DR G. W. LEE ON Another specimen, 11 mm. long, is partly broken, but shows both valves in apposition ; the maximum distance between the two valves is 5 mm. The general appearance of the shell is very much like that of Productus youngianus Davidson, a shell which differs from the present one in having a very concave brachial valve closely following the curves of the pedicle-valve ; besides, the brachial valve of Productus youngianus is ornamented by ribs, the place of which is taken here by pits. The nature of the ornamentation on the ears and cardinal slopes can also serve to dis- tinguish it from Productus youngianus. The Devonian Productus dissimilis de Koninck,* is remarkably like Productus brucei as regards the ornamentation of the pedicle-valve, but it is a less elongate shell, with a differently ornamented brachial valve. Productus sp. indet. A specifically indeterminable fragment exhibits the ribbing typical of Productus scabriculus (Martin). Productus cherneyensis sp. nov. (PI. II. figs. 37--370.) No specimens showing both valves in apposition are at hand, but the pedicle-valve described here is sufhiciently characterised to merit a distinct appellation. It is semi- circular, measuring 17 mm. in length and 19 mm. in width. The visceral portion is high and narrow ; the ears are well defined and the beak is much incurved, overlying the hinge-line. The surface is ornamented with ten broad concentric folds bearing thick, rounded tubercles separated by wide intervals. Between the folds, numerous concentric growth-lines are visible to the naked eye. The shell-substance is very thin. Two small specimens, 5 mm. long, show the same characters, the thick tubercles being very pronounced for such small shells. This species probably belongs to the group of Productus fimbriatus J. Sowerby, but it differs from most of its members in having a narrower visceral cavity, and fewer tubercles, which are moreover rounded, not elongate as in Productus fimbriatus. These characters are well shown in a species which Dr A. VauaHan provisionally describes as Productus cf. fimbriatus,t with the remark that it probably represents a link between the fimbriate and the aculeate Product:. Productus cf. fimbriatus may possibly be identical with the present one, but the specimen figured by Dr VauGHAN is more elongate than the Cape Cherney form. Productus elegans M‘Coy. (PI. II. figs. 38-380.) M‘Ooy, British Palxozoic Fossils, 1855, p. 460, pl. iii.H, fig. 4. This is the most abundantly represented Productus in the collection, and it is undistinguishable from its British representatives. One specimen, showing part of both * Monographie des Genres Productus et Chonetes, p. 147, pl. xvi. fig. 5. + Quart. Jour, Geol. Soc., 1906, p. 308, pl. xxx. fig. 6. A CARBONIFEROUS FAUNA FROM NOWAJA SEMLJA. rae valves in apposition, reaches a size exceeding that commonly attained by the species ; when complete it probably measured some 40 mm. in length. As this large specimen shows no trace of the sinus characteristic of Productus punctatus (Martin), we have here another conclusive proof of the fact that Productus elegans is not to be considered as the young stage of Marrin’s species. Productus keyserlingianus de Koninck, var. (PI. I. figs. 36—36a.) De Koninck, Monographie du Genre Productus, 1849, p. 134, pl. xiv. fig. 6. Four specimens, two of which have both valves in apposition, differ from Productus keyserlingianus as described by DE Koninck in having fewer tubercles ; they are other- wise quite like the Belgian form. The specimen figured under this name by M. Krotow is remarkable for an unusually large number of tubercles.* Productus spinulosus J. Sowerby. (Pl. II. fig. 30.) Davipson, loc. cit., pl. xxxiv. figs. 18, 19. The materials are fragmentary, but abundant; they are quite characteristic, with the exception of one or two specimens which appear to be slightly flatter than the type as refigured by Davinson. Sub-genus PROBOSCIDELLA Oehlert. Productus (Proboscidella) nysti de Koninck. (PI. II. figs. 34, 34a.) De Kownrncx, loc. cit., p. 65, pl. xiv. fig. 5. Materials indifferently preserved and belonging to this species might easily be taken for brachial valves of other Productids, but the specimen in the collection is in a fair state of preservation, consisting of a nearly complete pedicle-valve with part of the other valve still in apposition. Its proportions and ornamentation, and its short bent margin, are exactly as depicted by pe Koninox. IncERTa SeEpIs. The collection contains two fragmentary pedicle-valves of a Productoid shell, characterised by its greatly depressed shape and the extreme thinness of the test. The ornamentation consists of broad longitudinal folds bearing a few obtuse tubercles, with indications of wrinkling on the ears. * Mém. Com. Géol. Russie, vol. vi., 1888, pl. i. fig. 20. 172 DR G. W. LEE ON Class LAMELLIBRANCHIATA. Genus Patazoutima Hind. Palxolima aft. simplex (Phillips). (PI. IL. fig. 45.) Hinp, Monograph of the British Carboniferous Lamellibranchiata, Pal. Soc., 1903, vol. ii. p. 39, pl. xix. figs. 24-27. A right valve measuring 6 mm. dorso-ventrally and about 6 mm. antero-posteriorly is very much like the British form as interpreted by Dr Hinp, but its ribs are not appreciably flattened on the anterior side, and in this one respect differs from that form. From the Upper Carboniferous Palzolima retifera (SHuMaRD) it differs in having fewer ribs. Genus AvIcULOPECTEN M‘Coy. Aviculopecten clathratus (M‘Coy). Hinp, loc. cit., p. 82, pl. xv. figs. 1-7. A left valve measuring approximately 8 mm. dorso-ventrally agrees in every respect with Dr Hinp’s re-description of the species. Aviculopecten planoclathratus (M‘Coy). Hinp, loc. cit., p. 91, pl. xv. figs, 8-12. The shape and ornamentation of this specimen, which measures 10 mm. dorso-ventrally and 10 mm. antero-posteriorly, are exactly as in the British form, only the radial ornamentation is more accentuated ; but this may be accounted for by the excellent state of preservation of the surface characters in the Nowaja Semlja shell. Aviculopecten® sp. indet. A badly preserved pectiniform shell, measuring 30 mm. approximately in its two dimensions, probably belongs to this genus, but not to either of the above species. Genus Epmonpia de Koninck. Cf. Hdmondia m‘coyt Hind. Hinp, loc. cit., vol. i., 1899, p. 329, pl. xxxvi. figs. 23-29. A left valve, somewhat obscure and incomplete, belongs probably to a species of the group of Hdmondia scalaris (M‘Coy), and its affinities to Hdmondia m‘coy: Hind, are indicated by the shape of the broad concentric ridges, which point to a transversely subquadrate shell rather than to an obliquely suboval one. It measures approximately 15 mm. dorso-ventrally and 20 mm. antero-posteriorly. A CARBONIFEROUS FAUNA FROM NOWAJA SEMLJA. 173 Genus Conocarpium Bronn. Conocardium rostratum (Martin). (Pl. II. fig. 40.) Hinp, loc. cit., vol. i., 1900, p. 453, pl. li. figs. 6-9. This species is represented by two small specimens quite complete but for part of the anterior portion. The posterior rostrum of the specimen figured is in the collection, but could not conveniently be photographed along with it. This specimen measures 3°5 mm. dorso-ventrally and 3 mm. from side to side, and owing to its small size | am doubtful as to the degree of importance to be attached to the fact that the lower border is not concave, as in the form figured by Dr Hrnp, loc. cit., fig. 7, but is rather slightly convex, as in fig. 6, cbid. ; this may be only an appearance due to the incompleteness of the anterior end. Conocardium aleforme (J. de C. Sowerby). (PI. IL. figs. 41, 41a.) Hinp, Joc. cit., vol. i., 1900, p. 460, pl. liv. figs. 1-10, Conocardium aleforme is represented by seven specimens in all stages of growth up to about 10 mm. dorso-ventrally, and apparently undistinguishable from the British form, with this exception, that the larger one is slightly less inflated than SowrerBy’s type. At all events, it seems that the degree of gibbosity of the shell varies even in typical Lower Carboniferous districts—since a specimen of Conocardium aleforme figured by DE KonINCK is appreciably compressed laterally.* Class SCAPHOPODA. The collection contains a specifically indeterminable Dentaliid. Class GASTEROPODA. The Gasteropod fauna of the Cape Cherney limestone is very varied, but the greater number of the species is represented by extremely small specimens, as if some pauperising influences had arrested their normal development. In those cases where the types of the species which [ believe to be present here are actually minute forms, my identifications will probably be found to be fairly correct, but it is not without some misgiving that I have made bold to compare some of these small fossils with common west of Hurope species, although these have been founded on large specimens. Yet resemblance is often so striking—except in matter of size—that the latter course is perhaps after all the safest. * Faune du Calcaire Carbonifere de la Belgique, 1885, vol. v. pl. xviii. fig. 17. 174 DR G. W. LEE ON PLEUROTOMARIDZ. Genus Mourtonta de Koninck. Mourlona levis (M‘Coy). (PI. I. figs. 44-440.) M‘Coy, Synopsis of the Carboniferous Limestone Fossils of Ireland, 1844, p. 41, pl. v. fig. 15. This very depressed species appears to be represented by two small specimens measuring 6 mm. in diameter. ‘The slit-band is perhaps slightly wider than is the case in the type-figure. Genus Mourcuisonia d’Archiac and de Verneuil. Murchisonia aff. archiaciana de Koninek. (Pl. Il. fig. 42.) De Konincr, Faune du Calcaire Carboniféere de la Belgique, vol. iv., 1883, p. 16, pl. xxxiv. figs. 5-6, 19-20. Dr Konrncx’s species was founded on a much larger form. The fragmentary specimen which I compare to it is much smaller, the length of the four whorls com- posing it being only 6°5 mm., and the diameter of the last whorl 3 mm. The tapering is the same as in the Belgian species, and the ornamentation is similar, but there are four keels on each side of the sinual band, whilst in Murchisonia archiaciana four keels are stated to be above the band, and six or seven below it; the latter are also less pronounced. BELLEROPHONTID 2. Genus BELLEROPHON de Montfort. Bellerophon hiulcus (Martin). J. pB C. Sowersy, Mineral Conchology, 1825, pl. 470, fig. 1. Bellerophon hiulcus is represented by a few specimens, the largest of which does not exceed 12 mm. in diameter. Bellerophon aff. tenuifascia J. de C. Sowerby. J. pp C. Sowersy, Mineral Conchology, 1825, pl. 470, figs. 2-3. The ornamentation and the fine, thread-like ridge are as in the British species, from which the specimen at hand—measuring 18 mm. in diameter—differs in being less globose, the periphery being somewhat flattened, especially on the last part of the whorl. A CARBONIFEROUS FAUNA FROM NOWAJA SEMLJA, ifs EUOMPHALID ZA. Genus STRAPAROLLUS de Montfort. Straparollus dionyst de Montfort. De Koniwok, loc. cet., vol. ii1., 1881, p. 120. This species, as interpreted by DE Konrtnck, is represented by a few specimens greatly dwarfed, the largest of which is only 10 mm. in diameter. Straparollus pileopsideus (Phillips). Puiuuirs, Geology of Yorkshire, vol. ii., pl. xiii. fig. 6. The collection contains numerous specimens of a small species of Straparollus which in every detail except size seems identical with PurLiips’ species; they average only 5 mm. in diameter. PYRAMIDELLID A. Genus Loxonema Phillips. Loxonema levigatum (Kichwald). (Pl. II. fig. 46.) Lethza Rossica, 1860, p. 1117, pl. xlii. fig. 6. The specimens which I refer to this species agree very well with Loxonema levigatum, in particular in the smooth surface, the shape of the whorls, the linear suture, and the tapering of the shell. They are, however, much larger, the specimen figured, for example, measuring 4 mm. across the last whorl, a dimension which far exceeds that of EicHwaLp’s type. Loxonema sutwrale (Puiturrs) has the same general appearance, but its suture is wide and deeply excavated. If my identification proves correct, this will be the only Gasteropod in the collection actually larger than the type of the species to which it is referred. Loxonema sp. a. (Pl. IL figs. 48, 48a.) An incomplete specimen of four whorls, the total length of which is 4°5 mm., is characterised by a wide spiral angle and an ornamentation consisting, on the juvenile whorls, of numerous fine lines, equidistant and raised, which, on the last whorl, are arranged in sets of three separated by a smooth interval, the central line being a little stronger than the two bordering it. TRANS. ROY. SOC. EDIN., VOL. XLVII. PART I. (NO. 7). 26 176 DR G. W. LEE ON Loxonema ef. anglicum d’Orbigny. Puituips, Paleozoic Fossils of Cornwall, 1841, fig. 188. Four specimens, very small and fragmentary, must be placed in the group of Loxonema rugiferum (Puiuuirs), but the whorls are higher than in this species, thus pointing to closer affinities to Loxonema anglheum WVOrbigny. Another character they appear to have in common with it, is that the ribs are continuously uniform in size, showing no tendency to become evanescent in their upper part. This latter feature has already been pointed out by the Rev. G. F. Wumsorne.* The name Loxonema anglicum was given by pD’ORBIGNyt to a Devonian species which Parties had described in 1841 (loc. cit.) as a Devonian representative of his Carboniferous Melania rugifera, from which it differs in the characters indicated. The materials are too fragmentary to determine whether they really belong to the Devonian species, which has, however, already been recorded from Carboniferous rocks, the name figuring in Srruve’s list of Carboniferous fossils from the Moscow basin. Loxonema sp. b. (Pl. IL fig. 43.) The largest of three fragments is composed of four whorls measuring 3°5 mm. in length. The ornamentation consists of straight, sharp equidistant costee, more numerous on the younger whorls than on the later ones, and belongs to the type exhibited in Loxonema strigillatum de Koninck (loc. cit., 1881, pl. vi. fig. 22) and in Loxonema wishere Krotow.{ But the Nowaja Semlja shell differs from these in having the whorls less convex and a more shallow suture. In shape it resembles Loxonema semicostatum Meek and Worthen; § but in the latter the costee increase in number in the older part of the shell, whilst in this case it is the reverse. Loxonema? sp. c. (PI. IL. fig. 49.) A nearly complete specimen, measuring 5°5 mm. in length and 1$ mm. across the body-whorl, has doubtful generic affinities, since it is bucciniform, with a large body- whorl and an apparently twisted columella, as in certain species of Macrocheilina, but the ornamentation is like that of a Loxonema. Since the state of preservation is such as to preclude the possibility of determining whether the columella was callous or not, its true generic position must remain doubtful. The first two whorls are unornamented; the third and the fourth have a few broad transverse folds, whilst the ornamentation of the body-whorl is totally different, consist- ing of straight raised lines, closely set and very numerous. * A Monograph of the Devonian Fauna of the South of England, Pal. Soc., 1896, vol. iii. p. 43, + Prodrome de Paléontologie, vol. i., 1850, p. 62. {t Mém. Vom. Géol. Russie, vol. vi., 1888, pl. i. fig. i. § Geol. Survey Illinois, vol. v., pl. xxix. fig. 2. A CARBONIFEROUS FAUNA FROM NOWAJA SEMLJA. ae Its ornamentation distinguishes it from Loxonema? buccinordeum de Koninek (loc. cit., 1881, pl. vi. figs. 12-18), a species of doubtful generic affinities but of a similar shape, as is also Loxonema semicostatum de Koninck [non Meek and Worthen]. LIrToRINIDA. Genus TURBONITELLA de Koninck. Turbomtella biservalis (Phillips). (Pl. I. figs. 47, 47a.) Puituirs, Geology of Yorkshire, pl. xiii. figs. 10, 11. This species is represented by four small specimens, the largest of which is only 5°5 mm. in length. ‘The small ones have the ornamentation of Puiiurps’ Turbo semisulcatus (loc. cat., fig. 10), and the greater portion of the last whorl of the largest one has the two sets of alternating ribs as typically developed in Turbo biserralis (loc. cit., fig. 11), its beginning showing the transition of the “semzsulcatus” type to the “ biserialis” type. This is also exhibited on the end of the last whorl of the intermediate specimen. Genus PortTLockta de Koninck. Portlockia lacordavrer? (de Koninck). De Koninek, loc. cit., 1883, pl. xxv. figs. 17-19. Portlockia parallela? (Phillips). Puaruures, Geology of Yorkshire, pl. xvi. fig. 8. Each of these two species seems to be represented by a fairly complete but ex- tremely minute specimen, in all respects like the west of Hurope form, except in size. CaAPULIDA. Genus OrtHonycuta Hall. Orthonychia sp. (Pl. II. figs. 39, 39a.) Without expressing an opinion as to the value of the generic term Orthonychia, I place under it a very small (6 mm. from apex to frontal margin) capuliform shell which exhibits the features said to be characteristic ofthe genus, that is to say, the apex is incurved to a slight extent only, and shows no tendency to be spirally enrolled. Its specific characters are as follows :—The slopes are steep and meet under an acute angle forming a ridge bordered on each side by a very faint carina ; a shallow groove extends from the apex to the margin, and divides the left side into two equal portions; this groove is apparently absent on the right side; the surface is covered with undulating lines of growth. 178 DR G. W. LEE ON Comparison with other species is diflicult, since it is very small and there is nothing to indicate the stage of growth of the specimen, but in many respects it bears a general resemblance to Orthonychia unewm (Meek and Worthen),* a species which American palzeontologists consider to be a synonym of Orthonychia acutirostris (Hall).t Class CRUSTACEA. ‘TRILOBITA. Genus Puruuiesta Portlock. Philiipsia eichwaldi ? (Fischer de Waldheim). Woopwarp, Monograph of the British Carboniferous Trilobita, 1883, p. 22, pl. iv. A pygidium and a fragment of the head, seen from the under side, are possibly referable to this species. The under side of head is very much like that figured by Dr H. Woopwarp (loc. cit., fig. 8). I am indebted to Dr B. N. Preacu, F.R.S., for helping me in this determination. The following are generically indeterminable, on account of their imperfect preservation :— Sp. a. A fragment of pygidium characterised by the extreme convexity of the axis and the corrugated appearance of the somites, as obtains, for instance, in Griffithides globiceps (Phillips). Sp. 6. (Pl. IL. figs. 50, 50a.) A fragmentary pygidium, to which are attached three axial thoracic somites. Only the axis, consisting of fourteen somites, and a few of the pleural segments are preserved. The three anterior axial rings of the pygidium are ornamented, on their median portion, by a single row of strong tubercles; laterally, these tubercles are smaller and disposed in a double row. On the other rings the tubercles are in one row on the whole surface, and on every third ring the central tubercle is much more prominent than those next to it. ‘The pleural segments are less numerous but broader than the axial rings, and are separated by wide and deep sulci. Hach segment is longitudinally divided by a well-defined furrow in two sub-equal ridges, of which the anterior one is the broader ; both are ornamented by a row of closely set tubercles. The fragments of thoracic segments are quite smooth, and are well defined. The specimen is too poorly preserved to permit of any suggestion as to its * Geol. Survey Illinois, vol. v. 1878, pl xvi. fig. 1. + Geol. Survey Iowa, 1858, pl. xxiii. fig. 14. A CARBONIFEROUS FAUNA FROM NOWAJA SEMLJA. 179 relationships, but it might be mentioned that it appears to belong to an undescribed species bearing a certain resemblance to Brachymetopus owralicus (de Verneuil), as figured in Murcuison’s Russia, from which it differs, however, in the proportions obtaining between the divided parts of the lateral segments, and in the nature of the ornamentation on the three anterior axial rings. OSTRACODA. Thin sections of the limestone show that carapaces of Ostracods are common, but only a few could successfully be freed from the matrix. These are in excellent state of preservation, the finer details being well exhibited. They appear to be undistinguish- able from the following species :— . Leperditia okent (Minster). Borrdia curta (M‘Coy). Cytherella ? inflata (Minster). GENERAL REMARKS. Considering that the fossils described in this paper were found in a few fragments of limestone, and these from a single bed, the great variety of groups and species is really surprising ; representatives of most of the Lower Carboniferous invertebrate groups are met with, with the exception of those which are usually absent from limestone strata— the Cephalopods, for instance. So far as the evidence at hand goes, it seems probable that the bed yielding these fossils was deposited in comparatively clear water at some distance from land, but not necessarily at any great depth. The Protozoa, the Corals, the Crinoids, and the Brachiopods grew in the midst of conditions eminently suitable for their development ; it is therefore all the more remarkable that the Lamellibranchs and the Gasteropods should be represented by so many dwarfed individuals. At the same time, these dwarfing influences did not prevent the multiplication of species in the groups affected. As regards the stratigraphical position of the Cape Cherney limestone, the assem- blage as a whole points to a Lower Carboniferous age, whilst the presence of many typical species warrants us in assuming that the bed is probably in a series homotaxial with the Productus giganteus zone of Russia or Upper Visean of Western Europe. The collection does not contain any of the species characterising the upper divisions of the Carboniferous system, but at the same time not a few of the species cited in this paper have been recorded from horizons higher than the Visean. However, with our extended knowledge of the Upper Paleozoic formations, competent paleontologists have found it necessary to extend greatly the range attributed to some of the more common fossils; these forms have probably undergone certain evolutionary changes, but since these are often not discernible, new names have not been given to the apparently 180 DR G. W. LEE ON unmodified descendants of the forms originally described from lower horizons. Through the kindness of Professor ‘mH. TscHeRNyscHEW, I am enabled to mention a striking instance of persistence in the most familiar of Lower Carboniferous fossils: he found a specimen of Productus guganteus in the Sprrifer mosquensis limestone of the Timan, a region where the Lower Carboniferous is absent and the Mosquensis zone transgresses on the Devonian. Vice versa, Upper Carboniferous species may have appeared earlier than usually supposed, and, to mention an example taken from this country, Dr VaucuHan cites Derbya grandis Waagen, from the top of the Carboniferous limestone at Loughshinny, Ireland.* Bearing these facts in mind, one might hesitate to assign a definite age to a limited collection of Carboniferous fossils ; but in the present case the evidence is very strongly in favour of a correlation with the Guganteus zone, since the number of species typical of this zone is as large as that from any bed in a typical Visean locality, not to mention the complete absence of typical Upper Carboniferous forms. In the following table I give a list of the species found in the Cape Cherney lime- stone, with an indication of their occurrence in the Lower Carboniferous beds of Russia and Western Hurope. Their occurrence in higher horizons is also indicated, but these data have been compiled from various sources, and needless to say no Upper Carboni- ferous locality is known where so many Lower Carboniferous forms would occur together. It must also be understood that many of the species quoted from Upper Carboniferous beds are probably not strictly identical with the Lower Carboniferous forms the names of which they bear, for the reason stated above. Had they all been described instead of being simply cited in lists, I should perhaps have been justified in dispensing with many of these comparisons. As regards the relation of the fauna to its distribution in space, it seems to be very closely allied to that of the Giganteus zone of the Urals and Central Russia, which is practically the same as that of Western Europe. The list of fossils described in this paper has many points in common with those given respectively by Messrs Tscuerny- scHEW, Krorow, KRAsNOPOLSKY, etc., in various monographs treating of the Carboni- ferous of the Urals.t It is true that, of the districts surveyed by these authors, the most septentrional one, viz. the district of Tscherdyn and Ssolikamsk described by M. Krorow, lies some ten degrees south of Cape Cherney ; but Professor T’scHeRNyscHEW kindly informs me that Productus giganteus has been found much farther north, in the basin of the river Adzva, a branch of the Oussa. ‘Thus, although marine Lower Carboniferous beds’ have not yet been proved to exist in the extreme north of the Ural range (Pai-Khoi), it appears probable that the Cape Cherney limestone was deposited in a sea situated in the con- tinuation of the Uralian geosyncline. At all events, the Lower Carboniferous sea did not apparently extend farther west—over the Timan—and the Cape Cherney fauna * Quart. Jowr. Geol. Soc., 1908, p. 446. + Mém. Com. Géol. Russie, t. iii., 1889, t. vi., 1888, and t. xi., 1889. A CARBONIFEROUS FAUNA FROM NOWAJA SEMLJA. 181 shows no evidence of having inhabited an isolated basin, nor does it bear any affinities to the Lower Carboniferous fauna of North America. A still closer agreement obtains between it and the Lower Carboniferous fauna of the Moscow and Donetz basins* and of Western Europe, but this is obviously due to these regions being more accessible than the Northern Urals to exhaustive paleonto- logical studies. Comparison with Srruve’s list shows a striking resemblance to the lower portion of the Giganteus zone in the Moscow basin (‘‘ Kalkstein mit Stigmarien ”), but of course this may be due to nothing more than mere similarity in the conditions of deposition. In spite of the fact that Cape Cherney lies very far from the typical Lower Carboni- ferous districts, its fauna shows very few characters peculiar to itself. The maximum isolation appears to obtain in the Corals, but it is still possible that those forms which are new may at some future date be found in the Northern Urals.t The Brachiopods are closely allied to those of the typical Lower Carboniferous districts, and if I have ventured to establish a few new species and left others unnamed, the greater number of them are undistinguishable from their representatives as developed elsewhere. ‘The other groups are likewise represented by forms which appear to be remarkably cosmopolitan, and call for no special remarks. In the following table the sign + denotes the presence of species believed to be identical with those described in this paper, and cf. indicates that nearly allied or vicarious species are probably represented. * See Srruve’s list in “Schichtenfolge in den Carbonablagerungen im siidlichen Theil des moskauer Kohlen- beckens,” Mém. Acad. Imp. St Petersbowrg, 1886 ; and “Excursion XVI.,” by Messrs TscHERNYSCHEW and Lourovuain, in Guide des Excursions du VII™ Congres Géologique International, 1897. + See notes by Mr R. G. CARRUTHERS. [ TABLE. 182 DR G. W. LEE ON TABLE SHOWING THE FAUNA OF THE CAPE CHERNEY LIMESTONE AND ITS HoRIZONTAL AND VERTICAL DIsTRIBUTION IN Russi AND WESTERN EUROPE. * A ws Bs g a g 2 : . 2 3) sass BA 5S 8.2 8 ae 2) | BS eel OR. |S?) ape alae ee ES BESS lies B oe | er gis Oe |e eees é 6 Fe | haa aA Az 4 A ae a Endothyra globula (Kich.) + + + ef, Spirillina? sp. 5 + * Cribrostomum” gracile Mall. af + ae Tetrataxis gibba Moll. + + Archediscus karrert Brady + + + + Fusulinella struvet Moll. a + + ? Lagena sp. + Cenosphexra sp. a + cf, a tsp. b oo ef. Carposphera? sp. + ef. 1 Cenellipsis sp. + Glie Renieria clavata Hinde q + oF Aulophyllum sp... + Lophophyllum cherneyense °P nov. + sp. + . Campophyllum car imatum sp. nov. . + kungurense (Stiick.) . + =P Careinophyllum sp. + Hexaphyllia m‘coyt (Dunean) + + Crinoid remains + Stenopora sp. + el Ci. Polypora papillata M‘Coy q + + oF af * Dielasma lenticulare de Kon. + + * gillingense (Dav.) . ; : + ef. + Gi Spiriferina tnsculpta (Phill.) ° pall ce fmavaline + + - octoplicata (J. de C. Sow. 2 + + + cf + + 5 sp. ? + : laminosa M‘Coy. q + Gi Spirifer bisulcatus J. de C. Sow. q + ae 36 ef » triradialis Phill. q + ae Martiniopsis? sp... SF Martinia sp. (ef. Sp. planata Phill.) + ef. - brucei sp.nov. . + » sp. [aff. rhomboidalis M‘ Coy] + et, Reticularia imbricata (J. Sow.) t ah Squamularia sp. a + ef ef. ef. ef 53 ou) + cf cf. on cf Ambocelia uret (Flem.) q + + a Rhynchonellids + ef ef ef. ef. Athyrids + ef, cf ch ef, Camarospira? sp. . + Schuchertella crenistria (Phill. ) + + ae ci Orthotetid 4: cf. Rhipidomella michelini (L’Eveillé) . + sk an t +: Schizophoria resupinata (Mart.) var. + a5 ar ar * Tournaisian species. A CARBONIFEROUS FAUNA FROM NOWAJA SEMLJA. TABLE SHOWING THE FAUNA OF THE Care CHERNEY LIMESTONE—continued. Cape Cherney. Transition Beds of Malewka-Murajewna (‘‘ Devono-Carboniferous ” of Central Russia). 183 Lower Carboniferous. Moscow and Donetz Basins Lower Carboniferous. Urals Lower Carboniferous (Visean). Western Europe. Higher Horizons. Arctic and Eastern Russia, Europe. Schizophoria sp. Chonetes papilionacea (Phill. ) Sp. Productus giganteus (Mart.) . : Fs antiquatus J. Sow. ‘ concinnus J. Sow. - longispinus J. Sow. 9 9» evar. 2. x coatst sp. nov. £ scabriculus (Mart.) 0 cherneyensis sp. nov. 33 elegans M‘Coy 5 keyserlingianus de Kon. a spinulosus J, Sow. sp. [=Pr. carbonarius Sem. and Moll. non de Kon.] a nysti de Kon. sp. indet. . Palzolima simplex (Phill. va Aviculopecten clathratus (M‘Coy) . ms planoclathratus ye! 5 sp. indet. Edmondia maccoyt Hind. Conocardium rostratum (Mart.) alxforme (J. de C. Pon) Dentaliid Mourlonia levis (M‘Coy) Murchisonia archiaciana de Kon. Bellerophon hiulcus (Mart. ) tenwifascia J. de C. Sow. Straparollus dionyst de Montfort % ptleopsideus ( Phill.) Loxonema levigatum (Eich.) a sp. a i = anglicum d’Orb. ,9 ge 1 sp Turbonitella oe: ialis (Phill.) Portlockia lacordaireana (de Kon.) se parallela (Phill.) . Orthonychia, sp. - Phillipsia eichwaldi (Fischer) Trilobites, indet. (a) . (0) . : Leperditia okeni (Miinster) . Bairdia eurta (M‘Coy) Cytherella influta (Miinster) . margaritaceus Phill. [Tornquist] . S++ teettetet ++! > —_ t+tte st tet ¢t+B4+4+4+4+8484+4+4+4 -+4+4+ 544) ie) ar) TRANS. ROY. SOC. EDIN., VOL. XLVII. PART I. (NO. 7) +++ +++ + +8+++4++84+ +++ + t+4+4+&4+ +++ ++++44 + +4+4+5 +++ 27 ef. ef. 184 DR. G W. LEE ON EXPLANATION OF PLATES. Puate I. Fig. 1. Lophophyllum cherneyense sp. nov. Transverse section of a mature example. LEpitheca entire except on the right hand side. x 3/2. (The orientation is incorrect, the cardinal fossula being on the right- hand side of the figure, instead of the upper side.) Fig. 2. Median vertical section of another specimen, Showing the vesicular tabule and dissepiments and the thickened columellar septum. x 3/2. Figs. 2a-2d. Serial transverse sections from the same specimen as fig. 2. Epitheca only complete in fig, 2a. The Lophophyllum stage, with the septa extending to the columella, is seen in figs. 2c and 2d. x 3/2. Fig. 3. Campophyllum carinatum sp. nov. Median vertical section of a mature example. The carine, running upwards and inwards at a steep angle, are visible over part of the dissepimental area. Epitheca worn away. x2. Fig. 3a. Transverse section cut immediately below above. The major septa show a maximum extension towards the centre of the corallum. The carinz are best seen in the right-hand bottom corner, Epitheca worn away. x 2. Fig. 4. Transverse section of another example, showing a minimum extension of the major septa towards the centre, another section from this specimen agreeing with fig. 3a. Epitheca almost entire. x 2. Fig. 5. Median vertical section of another example, showing the highly arched tabule. Carine visible at the right-hand top corner. LEpitheca practically entire. x 2. Fig. 6. A young example, showing the general external characters. x 1/1. Fig. 7. Lophophyllum sp. Transverse section, In the upper half the epitheca is entire. x 1/l. Fig. 7a. —— Median vertical section from the same specimen as above. x 1/1. Fig. 8. Carcinophyllum sp. Transverse section just below floor of the calyx. Epitheca preserved in right-hand upper corner. x 1/1. Fig. 8a. Median vertical section from same specimen. Epitheca just worn away, x 1/1. Fig. 8b, ——- —— Transverse section from younger part of same specimen, Epitheca almost entire. eabyih, Fig. 9. Hexaphyllia m‘coyi (Duncan), Median vertical section showing the thick wall (on right-hand side of figure) and the closely set tabule. x 1/1. Fig. 10. Dielasma lenticulare de Koninck. Lateral view. x 1/1. Fig. 10a. Ventral view of same specimen, x 1/1. Fig. 106, ——- —— Dorsal view of same specimen. x 2. Fig. 10c. ? Ventral view of a large specimen. x 1/1. Fig. 11. Spirtferina insculpta (Phillips), var. Dorsal view. The beak is broken. x 1/1. Fig. lla. Frontal view of same specimen. x 1/1. Fig. 116. ——- ——— Lateral view of a fragmentary pedicle-valve, showing beak. x 1/1. Fig. 12. Martiniopsis? (or ?Spirtfer) sp. Ventral view. x 1/1. Fig. 12a. Punctate surface of same, much enlarged. Figs, 13-13¢. Martinia brucei sp. nov, x 1/1. Fig. 13d. -— Surface of same, showing transverse striz, much enlarged. Fig. 13e. A larger specimen provisionally referred to Martinia brucei. x 1/1. Fig. 14. Martinia sp. a (ef. Spirifera planata Phillips]. Ventral view. x 1/1. Fig. 15. Martinia sp. b |aff. rhomboidalis M‘Coy]. Ventral view. x 1/L. Fig. 15a. Surface of same, showing longitudinal striee, much enlarged. Fig. 16 Rhipidomella michelini (L’Eveillé). Ventral view. x 1/1. Fig. 17. Schizophoria resupinata (Martin) var. x 1/1. Fig. 17a. Surface of same, much enlarged. Fig. 18. Squamularia sp. a. Ventral view of a mature specimen. x 1/1. ‘Trans, Roy Soc. Edin™ Vol xn VAbe Piate I—Lre: A Carponirerous Fauna FROM NowaJa SEMLJA. A CARBONIFEROUS FAUNA FROM NOWAJA SEMLJA. 185 Fig. 18a. Squamularia sp. a. Lateral view of same. x 1/1. Fig. 180. —— Dorsal view of same. x 1/l. Fig. 19. Squamularia sp. b. Lateral view of a mature (?) specimen. x 1/1. Fig. 19a. Ventral view of same. x 1/1. Fig. 190. — Dorsal view of same. x 2. Fig. 20. Athyrid. Dorsal view of circular form. x 5/2. Fig. 21. Dorsal view of elongate form. x 5/2. Fig. 22. Cumarospira? sp, Restored outline of largest specimen, slightly enlarged. Fig. 22a. Fragmentary pedicle-valve showing the median septum and spondylium. x 3/1. Fig. 220. Transverse section of a brachial valve, about 2 mm, from umbo, Showing the median septum and the absence of a spondylium. x 2/1. Fig. 22c. Transverse section of a pedicle valve, about 3mm. from umbo. x 6/5. Fig. 22d. ——- —— Transverse section of another pedicle valve, near frontal margin. x 11/7. Prats II. Fig, 23. Orthotetid, indet. Pedicle-valve. x1/l. Fig. 23a. A brachial valve referred to the same species, x 1/1. Fig. 24. Schizophoria sp. Slightly oblique view of a decorticated brachial valve showing the general appearance of the muscle-scar, x 1/1. Fig. 24a, —— Isolated pedicle-valve. x1/1. Fig. 240. Outline of a complete specimen, x 1/1. Fig. 25. Chonetes papilionacea (Phillips). Brachial valve. 1/1. Fig. 26. Chonetes sp. Coarse-ribbed form. Pedicle-valve. x 1/1. Fig. 26a, —— The same, enlarged. x 2/1. Fig. 27. Chonetes sp. Smooth-ribbed form, External view of a pedicle-valve. x 1/1. Fig. 27a. Internal view of another pedicle-valve. x 1/1. Fig. 28. Chonetes sp. Dorsal view of a specimen, showing the ribbing characteristic of Chonetes cf. erassistria [Vaughan]. x 1/1. Fig. 28a. The same, enlarged. x 2/1. Fig. 29. Productus cf. margaritaceus (Phillips). x 1/1. Fig. 30. Productus spinulosus J. Sowerby. Fragmentary pedicle-valve. x 1/1. Fig. 31. Productus longispinus J. Sowerby. Slightly lobate form. x 1/1. Fig. 32. var.? Lateral view of a pedicle-valve belonging to the globose and fine-ribbed form, x1/1. Fig. 33. Productus concinnus J. Sowerby. Brachial valve (partly as a mould). x1/l. Fig. 34. Productus (Proboscidella) nysti de Koninck. Ventral view. x 1/1. Fig. 34a. Outline of same specimen, showing part of the brachial valve in apposition. Fig. 35, Productus coatsi sp. nov. The specimen is slightly distorted. x 1/1. Fig. 35a, Outline of another specimen. x 1/I1. Fig. 36. Productus keyserlingianus de Koninck, var. (The ornamentation has been completed from another specimen.) x 1/1. Fig. 36a. Outline of the same. x1/1. Fig. 37. Productus cherneyensis sp. nov. x1/l1. Fig. 37a. Outline of the same. x 1/1. Fig. 375. ——- —— A small specimen probably representing the juvenile stage of Productus cherneyensis. x 12/5. Fig. 38. Productus elegans M‘Coy. x1/1. Fig. 38a. A young example. x 1/1. Fig. 380. —— The same, enlarged. x 2/I1. Fig. 39. Orthonychia sp. Dorsal view. x 1/1. A CARBONIFEROUS FAUNA FROM NOWAJA SEMLJA. Fig. 39a. Orthonychia sp. Lateral view of the same specimen, enlarged. x 2/1, Fig. 40. Conocardium rostratum (Martin), x 2/1. Fig. 41. Conocardium alzforme (J. de C. Sowerby). x 1/1. Fig. 41a. Posterior view of the same. x 1/1. . Fig. 42. Murchisonia aff. archiaciana de Koninck. x 7/4. Fig. 43. Loxonema sp. b. x 5/2. Figs. 44-446. Mourlonia levis (M‘Coy). x 8/5. Fig. 45. Paleolima att. simplex (Phillips), x 2/1. Fig. 46. Loxonema levigatum (Eichwald). x 12/7. Fig. 47. Turbonitella biserialis (Phillips). Mature example. x 8/5. Fig. 47a. —— A young example, showing the “‘semdsuleatus” stage. x 5/3. Fig. 48. Loxonema sp. a. Outline. x 3/2. Fig. 48a. —— Mature whorl. x 3/1. Fig. 49. Lowonema? sp. [Macrocheilina?]. x 6/1. Fig. 50, Pygidium of undetermined trilobite. x 8/3. Fig. 50a. —— A segment of the same. x 3/1. 1s. Roy. Soc. Edin®™ Vol. XLVII. Puate IJ.—Lee: A CaARBONIFEROUS FAUNA FROM NowaJaA SEMLJA. 24 24b 23 32 36 36a 34a 35 35a 38) 3875 38 37a 4la 40 39a - = © @ 44 44a 446 49 46 47 47a 48a 48 50a je ners) VIII.—Scottish National Antarctic Expedition: Osteology of Antarctic Seals. By Robert B. Thomson, M.B., Ch.B., University of Edinburgh. (With One Plate.) (MS. received April 26, 1909. Read July 4, 1909. Issued separately October 12, 1909.) Through the courtesy of Dr W. S. Brucn, the leader of the Scottish National Antarctic Expedition, and of the late Professor D. J. Cunnincuam, the skeletons of the seals collected in the Antarctic during the voyage of the Scotva have been placed in my hands for study, and I wish to express my gratitude to these gentlemen for their great kindness in placing all the necessary requirements at my disposal. I also wish to state that Dr Bruce has given to the Museum of the University of Edinburgh a choice of the various skeletons, which, together with the specimens already obtained, will form a comprehensive and almost unique collection of the Antarctic seals. The seals obtained during the stay of the Scotia in the Antarctic seas numbered forty-four, and embrace all the Phocide found there, with the exception of the elephant seal. Of the forty-four, thirty-four are Weddell seals (Leptonychotes Weddelli); four are saw-toothed seals (Ogmorrhinus or Lobodon carcinophaga) ; three are leopard seals (Stenorhyncus leptonyx); one is a sea-lion (Otaria Jubata) ; while two are Ross seals (Ommatophoca Rossi). A most careful account of date of capture, sex, probable age, full measurements, and weight was recorded in a special log-book. In some cases the skeletons were cleaned by immersing the carcases beneath the ice, when amphipods and cushion-starfish did the work ; in other cases the carcases were conveyed in special tanks and cleaned after their arrival home. The osteology of the more common species of the Phocide represented in this collec- tion, such as the Weddell seal, saw-tooth seal, sea-lion, and leopard seal, appears to have been pretty fully worked up, and especially so in a minute and careful description by Principal Sir Witt1am Turner; for the present, therefore, the endeavour of this work is to present a detailed description of the Ross seal, as I have not been able to find such of the skeleton of this species, with the exception of the skull. In the second place, I purpose making a summary of the large collection of other seals with regard to such points as dentition and vertebral formula, since the collection is large enough to obtain a satisfactory average. Ross Seat (Ommatophoca Ross). The first specimen of Ommatophoca was obtained by Sir James Ross in 1840, in the pack-ice in the Ross Sea, and is now preserved in the British Museum as the type-specimen. I am indebted to the authorities of the British Museum and to Dr O. Tuomas for the privilege of examining this specimen for purposes of com- TRANS, ROY, SOC. EDIN., VOL. XLVII. PART I. (NO. 8). 28 188 MR ROBERT B. THOMSON: SCOTTISH NATIONAL ANTARCTIC EXPEDITION : parison. Unfortunately, it is not a complete skeleton: both fore-limbs with the exception of one scapula, both hind-feet, one dorsal vertebra, and two pieces of sternum are awanting. A description of the skull was made by Dr J. E. Grey in 1844. For over fifty years it remained the sole representative of its race in natural history collections, as not till 1899 was the Ross seal again captured by the Belgian Expedi- tion (1898-9). Since then numerous specimens have been obtained and the skins and crania brought home by the Southern Cross (1898-1900), Discovery (1901-4), Morning (1902-4) ; while the Scotca captured two specimens—a male and a female, of which the male skeleton is complete with the exception of two segments of the sternum and one patella. The female skull has been unfortunately damaged, the occipital region being wanting ; but a perfect set of teeth, of so much importance in Ommatophoca, is preserved. Further, the whole respiratory apparatus of the male Ross seal has been brought home, and is of great interest—as the voice of this animal, which has been described by Dr Racovirza, with the peculiar inflation of the pharynx, has attracted the notice of, and been recorded by, most Antarctic explorers. The method of describing the skeletons is that adopted by Sir Witt1am TurRNER in Challenger Reports, and I wish to express my appreciation of its arrangement and carefulness. Surface Measurements (from Log-Book). No. 2. No. 43. Date of Capture—6th February 1903. 28th February 1904. Sex—Female. Male. Age—Adult. Adult. Length— Nose to tip of tail, 90”. 89°5”. Girth— Anterior or neck, 41”. 38:5”, Axillary, 52”. 55°5”. Greatest (9 ins. behind axillary), 54”. Tail, 5", Fore flippers— (12 ins. behind axillary), 56:5”. Outer edge, 16”, 2% Base, 83”. Hind flipper— Outer edge, 16”. 18-4”, Inner edge, 143”. RS ire Stretch, 25”. 24", Base, 10°7”. 1S he Hye, 14”x 1". 14x 8". Interorbital space, 6”. 6:5”. Total weight, 450-500 Ibs. 400 lbs. Remarks. Intestines of No. 43 measured from pyloric end of stomach to anus 268”. Cuttlefish beaks, pieces of cuttlefish, and what appeared to be scales of fish were found in the stomach. A great quantity of tape-worms were present in the gut. OSTEOLOGY OF ANTARCTIC SEALS. 89 No. 43 or CoLuection (Mate). The skull is characteristically short and broad. The greatest length was from premaxilla to the occipital condyles, whilst the greatest breadth was between the two zygomatic arches, just at the articulation between its two component parts—the zygo- matic process of the temporal and the malar bones. A comparison of this skull was made with those of an adult Weddell and sea-leopard in regard to length and breadth—an index being framed. Thus, the length-breadth index of the Ross skull was 72°4; of the Weddell skull, 62°18 ; of the leopard seal, 55:07. The nasals measured 79 mm. long, and were completely ankylosed. They articulated with the frontal and superior maxillary bones. The part between the two frontals amounted to about three-quarters of the entire length of the bones (57 mm.), and was triangular in form, while the remaining quarter between the two superior maxillary bones was quadrilateral. The distance between anterior edge of the outer border of the nasals and the tip of premaxilla was 18 mm., the superior maxilla thus forming to this extent part of the outer boundaries of the anterior nares. ‘The length of the section of the boundary formed by one of these bones varies in the different specimens from 9 to 17 millimetres” (Barrett Hamitton, Pésultats du Voyage du S.Y. “ Belgica,” 1897-1899, p. 5). The anterior nares sloped downwards and forwards at an angle of 59° with the hard palate. In Weddell seal this angle was 48°, in leopard seal 35°. The anterior nares were bounded from above downwards by the anterior border of the nasal bones, part of nasal borders of superior maxilla, and by the premaxillary bones. Looking into the anterior nares, one was struck with two points—the thickness of the meso-ethmoid, and the extremely convoluted arrangement of the turbinate bones. The anterior edge of the vomer was received between the meso-ethmoid and the premaxillary bones. The premaxillary bones supported the two incisor teeth ; their palatal parts were triangular in form, and, as before mentioned, their nasal parts did not extend so high as to meet the nasal bones. The ante-orbital process of the superior maxilla was well marked, and lay in the same transverse plane as the infra-orbital foramen, below which is a definite depression from which a groove leads to the orbital floor. The widest part of the hard palate was situated well behind the last molar, and was 7 mm. in front of outer end of articulation of palatal processes of the superior maxilla and palate bone. It was almost flat, showing only a trace of a concavity. The posterior border of the hard palate was concave, and possessed a faint post-nasal spine. On the norma verticalis of the skull, at the junction of the frontal bones, z.c. at the seat of the anterior fontanelle in the young, is situated an opening between the bones. Its margins are irregular and spiculated (see fig. 1). As there is no history of an injury, the animal not being clubbed, but shot in the thorax, it appears to me to be a persistent anterior fontanelle. In the female Ross seal this condition is only faintly represented, but in the plate of the Ross seal of the Belgian Expedition a similar well-marked deficiency is to be 190 MR ROBERT B. THOMSON: SCOTTISH NATIONAL ANTARCTIC EXPEDITION : observed. The occipital bone presented occipital crests which were well marked, as also were the par-occipital processes. Two well-marked supra-occipital venous foramina were to be noticed in the middle line of the occipital squama. ‘The condylar articular surfaces were highly convex and approached close to each other on the inferior aspect (7 mm.) ; above, they were distant 41 mm. The basi-occipital was quadrilateral in form, flat and thick. The tympanic bulla was smooth and rounded. The carotid canal was separated from the jugular foramen by a thick bar of bone. In the interior of the skull the tentorium cerebelli and falx cerebri were partly ossified. In the female skull the nasal bones were not ankylosed, and measured 55 mm. long. The distance between the anterior edge of the nasals and the premaxilla was 20 mm. on right side, 18 mm. on left. The Lower Jaw. The body was long and somewhat slender, due to small size of teeth, with lower border slightly incurved. The ramus formed with the body a moderate angle and possessed a well-marked sub-condyloid process. The condyle, which was convex and elongated transversely, was separated from the coronoid process by a shallow sigmoid notch. The coronoid process was short and pointed. Dentition. The dentition of the Ross seal is remarkably feeble, and varies more especially with regard to the number of post-canines. Dr E. A. Wixson, in vol. il. of Natural History of National Antarctic Expedition, 1907, states: “The food of the Ross seal consists mainly of soft-bodied cephalopods, and to this end has developed the incisors and canines into needle-pointed recurved hooks of great delicacy and has allowed its post-canines to degenerate. The gums presumably can manipulate such food as well as could molar teeth, and so we find in some cases the post-canines are small and insignificant, whereas in others they are loose and useless, and occasionally absent altogether.” He also gives a complete table of the dentition of all the Ross crania which have been brought home, and which shows the great variations one meets in the dental formula. ‘To this list I would add the formulz of the two specimens of the Scotia :-— 2.1.5 No. 2. Femal : ) “emale, 21-5 2s B . 43, No. 43. Male, ST 1-51 No. 2 skull presents for examination a perfect set of teeth, both upper and lower. The upper incisors are more powerful than the lower, but both present, like the canines, the well-marked recurved, needle-pointed character. The post-canines are recurved OSTEOLOGY OF ANTARCTIC SEALS. 191 backwards and three-cusped, the central one being longest. In all but the first the post-canines were double-rooted. In No. 43 the teeth are not so well developed, and in the upper jaw the left outer incisor and canine are broken. The dentition is not quite easily made out. There is no fifth post-canine, but small sockets are to be observed; hence we may presume that they were present, having been partly fixed in the natural state by the fleshy gum, a condition pointed out by WEBER, Bennet, and Wixson (National Antarctic Kaxpedition : Natural History, vol. ii. p. 46). Again, in the lower jaw no central incisor is present, but the two halves of the jaw have been disarticulated. On placing the two halves together, it would seem that no room at all is left for central incisors. A comparison of the measurements of the two lower and upper jaws between outer side of canines shows :— No. 2. Upper diameter, 44 mm. Lower _ 30 ,, No. 43. Upper diameter, 48 _,, Lower - 20055 In No. 2, which has not been disarticulated, there is a fair amount of fibrous tissue, so it is possible to explain the difference in diameters by the absence of this. Hence, with the fibrous tissue in place in No. 43, room would be made for central incisors. Against this may be stated the fact that the upper central incisors are very feeble, and, as the whole of the upper teeth are stronger than the lower, it is posetnte that the lower central incisors may not have developed. Measurements of Skulls. Ross Seal, | Ross Seal, Weddell Leopard No. 43. No. 2. Seal. Seal. mm. mm. mm. mm, Extreme condylo-premaxillary length . ; ; 250 x 272 394 | re inter-zygomatic width . c 181 aR 169 217 se width behind external meatus : : 167 a: 177 198 Greatest width of palate. 76 Ti 58 90 Width between outer side of base of upper canines 48 44 52 75 - lower canines 26 30 30 50 Length of palate i a line of suture to central incisor gl 82 105 157 Height of skull—basion to mid-occipital crest : 83 ade 84 110 Smallest interfrontal diameter . : : : 45 49 30 45 Length of nasals . ‘ : ; : : ; 79 55 61 103 Greatest width, anterior nares. ; 43 34 36 44 Length of mandible. : : : 168 170 cial 301 Width between outer end of condyle ; : : 182 168 57 205 The vertebral formula was :— C, Dy; L; Sy Cadi: 192 MR ROBERT B. THOMSON : SCOTTISH NATIONAL ANTARCTIC EXPEDITION : Cervical Vertebre. The bodies of the posterior six cervical vertebrae were keeled on their ventral aspect, the keel being more pronounced in the anterior than in the posterior series. On con- trasting the bodies of the Ross seal with those of the Weddell and leopard seal, a great difference was noticed with regard to their length and breadth. If the 4th cervical be taken as a typical vertebra, we find the length index of the body to be :— In the Ross seal—Length 33 48x 100 = Index 145°4. Breadth 48 33 In leopard seal—Length 51 47x 100_Figex 99-16, Breadth 47 51 In Weddell seal—Length 38 index 100. Breadth 38 If these indices be compared with the length-breadth index of the skull, one is justified in saying that the short and broad character of the Ross skull can also be applied to the bodies of the vertebrae. The neural ring was triangular. A depth- breadth index shows much the same difference as the bodies in the three species contrasted. In the Ross seal, the index was 35:1; in Weddell seal, 54°5; and in the leopard seal, 50. The transverse processes in all except the 7th were perforated at their roots by a vertebraterial foramen, and possessed at their extremities well-marked tubercles in all except the 2nd and 7th. The spinous processes exhibited a marked difference in the various vertebre. ‘Thus the axis possessed a massive spine with four well-marked tubercles—two anteriorly and two posteriorly. ‘lhe 7th had a well-marked spine, not bifid; the 3rd and 4th had only a trace of a spine, while the 1st possessed a well-marked tubercle representing a spine. The 5th and 6th presented fairly prominent spines, bifid —a character not shown by either the Weddell or leopard seals. The type-specimen in the British Museum also exhibits this character. The anterior articular processes looked upwards and inwards, the posterior downwards and slightly forwards. The atlas has powerful lateral masses supporting anteriorly deeply concave articular surfaces for occipital condyles, while the posterior articular facets for the axis were flat. The transverse processes were plate-like, and projected outwards and downwards. The lamina was perforated on each side by a foramen for the vertebral artery, which foramen was situated internal to the upper ends of the articular facets for the occipital condyles. The axis did not present so well-marked an odontoid process as either the Weddell or leopard seal. On the inferior aspect of the odontoid process was a triangular facet for articulation with the atlas, whilst superiorly it had a small oval facet for articulation with the transverse ligament. The transverse processes were feeble and pointed, whilst their superior roots were mere spicules of bone. The ventral aspect of the body was markedly keeled. OSTEOLOGY OF ANTARCTIC SEALS. 193 Dorsal Vertebre. The bodies of the anterior four were keeled, the keel becoming less pronounced as we proceeded from 1 to 4. At the posterior end of the dorsal region, 14 and 15 pre- sented a slight keel, more accentuated in 15 than in 14. ‘The side of the 1st body had one and a half facets; 2 to 10 inclusive (two half-facets, 11 to 14 inclusive) a whole facet for the corresponding ribs. The 15th dorsal vertebra showed a peculiar arrangement; thus, on the right side of its body a well-marked facet was present for articulation with the 15th rib of that side, while on the left a well-marked plate-like costal process, representing the last rib, projected from the junction of the body and pedicle outwards, forwards, and slightly downwards. It measures 42 mm. in length and 25 mm. in breadth. (See tig) 5.) The spines of the dorsal vertebrze were relatively feeble compared with those of the Weddell and leopard seals. They were low, and projected upwards and backwards as far as No. 11; while 12, 13, 14, 15 projected upwards. The transverse processes of 1 to 10 were prominent and presented rounded facets for articulation with the tubercles of the ribs. In 11, 12, and 13 the transverse processes were feeble but possessed facets. In 14 and right side of 15 they were rudimentary and non-articular, while the character of the transverse process of left side of 15 has already been noticed. From 9 to 15 the mammillary processes were well defined, while accessory processes were faintly marked in 9 to 14. Lumbar Vertebre and Sacrum. Each body of the lumbar vertebrae was elongated antero-posteriorly and was keeled on its ventral aspect. The spines were broad and more pronounced than in either the Weddell or leopard seals. The mammillary processes were rounded and directed forwards and outwards. The transverse processes were thin and flat, and curved out- wards, forwards, and downwards. The sacrum was composed of three segments fused together. The ventral surface in its anterior part was broad and flat, with a faint keel, while its posterior part, corresponding to the 2nd and 3rd segments, was protuberant in the centre, concave laterally, where it presented the openings of the two anterior sacral foramina. The posterior surface showed mesially three spimous processes, while the laminz of the three segments were only fused laterally. The fused articular processes were prominent at the junction of 8, and S,, less so at S, and 8,. The direction of the 1st posterior sacral foramen was horizontally outwards, in the 2nd and 3rd upwards and only slightly outwards. Laterally the sacrum presented a definite ‘U-shaped articular surface for ilium, while the interval between the two limbs of the U and the area posterior to it were hollow for ligamentous attachment. The articular surface for the ilium was confined entirely to the first sacral segment. 194. MR ROBERT B. THOMSON: SCOTTISH NATIONAL ANTARCTIC EXPEDITION : Sacral Indea. Ross seal : 4 : : ; 108 Weddell seal . é 3 : 5 102°9 Leopard seal . : : : : 114°8 The caudal vertebrae were ten in number. The posterior six were represented by elongated bodies alone—the last being cartilaginous. The 1st possessed, in addition to the body, a definite neural arch terminating above in a rudimentary spinous process. The lamine in the 2nd did not unite superiorly, so that the neural groove was not con- verted into an arch. ‘The third possessed on each side of its body rudimentary lamine. The bodies of the first four caudal vertebrae were keeled on their ventral aspect. The sternum was incomplete, two segments being awanting. ‘Their articulations could not be made out definitely, so I refrain from describing them. Ribs. The ribs were fifteen in number on the right side and fourteen on the left, this anomaly being explained in connection with the thoracic vertebre. The Ist, 12th, 13th, 14th, and 15th ribs possessed a single facet each on their heads for articulation with the corresponding vertebree. ‘lhe 2nd to 11th inclusive possessed two facets separated by a distinct ridge. The Ist to 12th ribs inclusive possessed well-defined necks, while the 1st to 13th showed an oval concavo-convex facet for articulation with the corresponding transverse processes. In the anterior series of ribs was a slight depression, in some a roughness, on each side of the articular surface of the head, for attachment of ligamentum conjugale costarum described by Sir Witi1am Turner (Challenger Reports, vol. xxvi., Report on Seals, p. 14). The shafts of the ribs were slightly curved, the most pro- nounced curvatures being observed in the 2nd, 3rd, and 4th, while the greatest in length were the 8th and 9th. The last rib on the right side possessed a head with an articular facet, no neck, and a feeble shaft. Its length was 127 mm. The Fore-Limb. The scapula presented the well-marked sickle shape so characteristic in the Phocidee. The anterior and superior or vertebral borders were convex and thin, the inferior or axil- lary was concave and rounded. On the axillary border at junction of lower and middle thirds in the Weddell and leopard seals was a prominent ridge for muscular attachment. No such ridge was present in the Ross scapule. ‘The dorsum scapule was divided into a pre-spinous and post-spinous area by a feeble spine. This spine was relatively longer than in the Weddell and leopard seals, but not so well developed. The pre-spinous area was concavo-convex, the post-spinous was flat. There was no acromion process, but in the Weddell and leopard seals this was present although feeble. ‘The coracoid process was rudimentary and projected forwards, measuring from the bottom of the shallow OSTEOLOGY OF ANTARCTIC SEALS. 195 notch at its base 18 mm. The ventral surface showed a slight subscapular angle cor- responding in position to the attachment of the spinous process on the dorsal surface. Two faint muscular ridges were present, directed towards the glenoid fossa. The glenoid fossa was pear-shaped, deeply concave, with a prominent margin. Index. Ross seal : . ‘ ; F 85 Weddell seal . : 3 3 F 82°14 In the leopard seal the epiphysis of the posterior angle was wanting, hence the index could not be ascertained. The humerus measured 115 mm. long. The head was convex, and its articular surface was continued forwards on to the base of the lesser tuberosity, which was equally as prominent as the greater one. The shaft was slightly twisted, and pre- sented a strongly marked deltoid ridge. The bicipital groove was in the form of a shallow groove. The lower end presented a well-marked trochlea for the ulna, a slight coronoid fossa, but no radial or olecranon fossee. The capitellum was circular and convex. The internal condyle was more pronounced that the external, the external supra-condyloid ridge more so than the internal, which did not possess a supra- condyloid foramen or even a process. Bones of Forearm. In neither were the distal epiphyses fused with the shaft. The ulna measured 152 mm. and was expanded above from before backwards so as to form a long but narrow olecranon process. A coronoid process was not present. The articular surface for humerus was concavo-convex, while the facet for radius above was flat and con- tinuous with that for the humerus. The shaft was flat, with a rounded anterior border and a sharp posterior border. The lower end was conical and articulated with radius slightly, and with cuneiform and pisiform. The radius was 135 mm. long. The head possessed a deep cup-shaped cavity for articulation with the capitellum of humerus, while the margin of the head was non-articular except on its inner side, where there was a well-marked facet for articulation with the ulna. On the neck a faint tubercle was noticed, while the shaft was paddle-shaped, with a definite roughness on its outer aspect for pronator radii teres. The lower end was grooved on its posterior aspect, and articulated with the ulna and scapholunar. Humero-Radial Index. Length of radius x 100 Length of humerus Ross Seal. Sea Leopard. Weddell Seal. 117-4 107°1 98-1 TRANS. ROY. SOC. EDIN., VOL. XLVII. PART I. (NO. 8). 29 196 MR ROBERT B. THOMSON: SCOTTISH NATIONAL ANTARCTIC EXPEDITION : Carpal Bones. These were seven in number, the scaphoid and semilunar being fused together to form a single bone. The carpal bones were devoid of ridges or processes. The articu- lations were dificult to make out, but I have referred these as follows :— The scapholunar articulated with radius, trapezium, trapezoid, os magnum, and unciform. The cuneiform articulated with the ulna, pisiform, and unciform. The pisiform was a mere nodule, and articulated with the cuneiform and ulna. The trapezium articulated with scapholunar, trapezoid, and 1st and 2nd metacarpals. The trapezoid articulated with scapholunar, trapezium, os magnum, and 2nd meta- carpals, The os magnum was small, and articulated with the 2nd, 3rd, and 4th metacarpals, and with the trapezoid, scapholunar, and unciform. The unciform articulated with the scapholunar, cuneiform, and os magnum, and with the 4th and 5th metacarpals. Metacarpal Bones. The 1st metacarpal was the longest, and possessed both a proximal and a distal epiphysis not united with the shaft. The 2nd was longer than the 3rd, 4th, or 5th, which were of about equal length, and the epiphysis for the head of each was separate from the shaft. The Hind-Inmb. The innominate bone, measuring in length 215 mm., consisted of three parts—ilium, ischium, and pubis. The ilium was short and stunted. The ischium and pubis were long, somewhat narrow, and enclosed between them a very large obturator foramen— larger indeed than the ilium itself. The acetabulum was circular in form, entirely articular except an elongated, hollow area at the bottom with a definite margin. The brim of the acetabulum was deficient in its lower aspect, thus forming a cotyloid notch. The ilium measured from centre of acetabulum to the highest part of the iliac crest 74 mm. The length of its crest was 102 mm. Its dorsal and ventral aspects were flattened and did not possess any definite muscular ridges. At the posterior part of the ventral surface was a U-shaped articular surface for that part of sacrum corresponding to its first segment, while there was a roughened area for the attachment of the posterior sacro-iliac ligament. The os pubis projected backwards and slightly downwards from the acetabulum, and its junction with the ilium was marked by a prominent pectineal eminence. From this tubercle to the symphysis, which was small, the pubis measured 130 mm. The ischium passed backwards and slightly upwards from the acetabulum, and possessed a pointed spine. The femur measured 114 mm. long. The head was entirely articular except in its postero-superior quadrant, where there was a non-articular area continuous with the OSTEOLOGY OF ANTARCTIC SEALS. iL non-articular neck for attachment of the ligamentum teres. Dr Kerrru, in Human Embryology and Morphology, p. 385, states: “The ligamentum teres is isolated from the capsule by the development of the head of the femur, which expands as a wing on each side of the ligamentum teres, and by fusion of the wings isolates it from the capsule.” In the Ross seal femur the wings have not yet met; hence, according to KerrH’s view, the ligamentum teres would still be in connection with the capsule. (See fig. 6.) The trochanter major was a broad, rough prominence projecting from the outer side of the junction of the neck of the bone with the shaft. There was no trace of a trochanter minor, as is to be found in Otaria Jubata; neither was there a third trochanter. The shaft, flat and expanded, possessed an anterior and a posterior surface, both being very slightly rounded and devoid of muscular ridges. The outer border was short and concave, the inner convex and very rough for muscular attachment. At the posterior end of the bone, the trochlear surface for the patella was flattened and ascended as high on its outer as on the inner side. It was quite distinct from the condylar surface, but closely approximated to the external one, being separated by a very narrow groove. The internal condylar surface was flat and circular, the outer one oval and slightly convex. The intercondyloid fossa was shallow. The patella was 34 mm. long and 24°5 mm. broad. It was oval in form, and its articular surface was slightly concave, and did not exhibit any subdivision into special articular areas. Its anterior end was much thicker than the posterior. The tibia and fibula were fused together at their upper ends. The tibia measured 247°5 mm. Its upper end was elongated transversely and possessed two articular areas for the condyles of the femur, separated by a rough intermediate area. The shaft was slightly curved, flat and smooth on its ventral aspect, deeply concave on its dorsal aspect for origin of the tibialis posticus muscle. The nutrient foramen was a large one. The inner border was broad and rough, the outer one thin and rounded. The lower end was not united with the shaft, and from it projected a slight malleolus, which was non- articular. The ventral and dorsal aspects of the lower end were deeply grooved, while the articular area for astragalus was reniform and concave. The fibula was 233 mm. in length. The upper end was fused with the tibia, whilst the lower end was grooved on the outer side by the peroneal tendons, and had on its inner aspect an oval facet for outer surface of astragalus. The shaft was straight, and its surface broad for muscular origins. The lower epiphysis was not united to the shaft. Tilia- Femoral Index. Length of tibia x 100 Length of femur Ross Seal. Weddell Seal. Leopard Seal. 217-1 203°9 213°5 198 MR ROBERT B. THOMSON: SCOTTISH NATIONAL ANTARCTIC EXPEDITION : Tarsal Bones. The astragalus was the largest of the tarsal bones. It consisted of a body and two large processes. ‘The posterior process projected backwards beyond the hinder end of os calcis, and presented a well-marked groove on its under aspect. Its anterior process or head passed as far forward as did the os calcis, and articulated with the scaphoid, cuboid, and fore-part of os calcis. The superior surface of the body presented a quadri- lateral convex surface for articulation with the lower end of tibia, continuous externally with a triangular facet for fibula. Its internal surface did not articulate with the internal malleolus. The inferior surface articulated with os calcis and presented a deep groove for an interosseous ligament. The os calcis possessed an attenuated posterior or peroneal process, and a feeble internal or sustentacular one. It articulated with the astragalus and fibula above, with the cuboid in front, and was grooved on its outer aspect by the peroneal tendons. The cuboid presented on its inferior aspect a well-marked plantar tubercle and a deep peroneal groove. It articulated with the os calcis, astragalus, scaphoid, external cuneiform, and 4th and 5th metatarsal bones. The scaphoid articulated with the astragalus, cuboid, and the three cuneiforms. Its posterior surface was not entirely concave, being convex in its outer part. The internal cuneiform articulated with the scaphoid, middle cuneiform, and 1st and 2nd metatarsals. The middle cuneiform was the smallest of the three, and was completely shut out of the plantar aspect of the foot. It articulated with the cuneiform on either side of it, with scaphoid behind and 2nd metatarsal in front. The external cuneiform articulated with the scaphoid, cuboid, 2nd and 8rd metatarsals, Metatarsal Bones. The 1st was the longest, the 5th next in order, then 4th, 2nd, and 3rd, which was the shortest. None of the epiphyses were fused with the shafts. The base of the 2nd showed the characteristic hook-shaped base. The articulations of their bases were as follows :— lst with internal cuneiform and 2nd metatarsal, 2nd with the three cuneiforms 1st and 3rd metatarsal, 3rd with external cuneiform 2nd and 4th + 4th with cuboid 3rd and 5th bs 5th with cuboid 4th 9 while its base projected outwards as an elongated process. The terminal phalanges of both fore and hind limbs were retained in connection with the skins. OSTEOLOGY OF ANTARCTIC SEALS. 199 Summary. The skulls correspond in their distinguishing characters from other Antarctic seals to those tabulated by Sir Wituiam Turner in the Challenger Reports (Voyage of “Challenger,” Reports on Seals, p. 66). With regard to the rest of the skeleton, one specimen alone does not provide altogether sufficient evidence ; still, I note the following points which may be of interest on further skeletons being ware home and examined :— 1. Much greater length-breadth index of vertebree. 2. Bifid character of 5th and 6th cervical spines, also present in type-specimen in British Museum. . Sacral index. . Scapular index. . Humero-radial index. . Non-articular area for ligamentum teres on head of femur continuous with non- a oP CO articular neck. 7. Tibio-femoral index. NuMBER OF VERTEBRA IN PHOCID. In the Résultats du Voyage du S.Y. “ Belgica”: Zoologie, p. 7, Mr Barrer HamitTon states that in most seals the numbers of the thoracic and lumbar vertebrae appear to be usually 15 and 3 (57), more rarely 14 and 6. In the Antarctic seals without exception I have found the vertebral formula in the adult to be C, D,, L, Ss Cdj OF 43- With a view to obtaining as large a number of specimens as possible, in order to form an average, I have made a careful examination of the seal skeletons in the University Museum here, the Museum of Science and Art of Edinburgh, British Museum of London, and give a table of the vertebral formula in 47 specimens :— Weddell seal: 27 with formula C, D,, L,; 8, Cd, or 4,. Crab-eating seal: 2 with formula C, D,, L, 8, Cd, Sea-leopard : 2 with formula C, D,, L, 8, Cd4o. ie ee i Ora lg Cane Ross’ seal: 1 with formula C, D,, L, S, Cd, . Sea-lion : 2 with formula C, D,, L; 8, Cd, Fur seal: 1 at C, D,, Ly S, Cd,,. Crested seal: 1 at C, D,; L, 8, Cdjo. Common seal: 2 at C, D,, L; 8, Cd,,. Elephant seal: 1 at C, Dy, L; 8, Cdjp. New Zealand fur seal: 1 at C, D,, L, 8, Cd,p. West Indian seal: 1 at C, D,,; L; 8, Cd,,. 200 MR ROBERT B. THOMSON: SCOTTISH NATIONAL ANTARCTIC EXPEDITION : Grey seal: 1 at C, D,, L, 8, Cdjp. Bearded seal: 1 at C, D,x L; 8, Cd,,. Northern fur seal: 1 at C, D,, L; 8, Cd,o. Altogether 46 specimens with formula of C, D,, L, 8, Cd,, to 45. One specimen with formula of C, D,, L, 8, Cd,,. The number of caudal vertebrae seems to vary from 10 to 12, while in some cases one could not be positive, as frequently the terminal vertebre are preserved in the skins. SuprA-ConpyLoip FoRAMEN. With a view to the formation of a percentage of such foramina found in the humeri of the Phocide, I have examined altogether 100 humeri, made up as follows :— 60 Weddell seals. 4 crab-eating seals. 6 leopard seals. 4 sea-lions. 2 Ross seals. 4 elephant seals. South American fur seals. New Zealand fur seals. West Indian seals. common seals. North American fur seals. Nownwvye we Total, 92 In these 92 no foramen was present, but in 4 Weddell humeri there were small supra-condyloid processes. In the following 8 humeri, supra-condyloid foramina were present :— 2 grey seals. 2 bearded seals. 2 crested seals. 2 common seals. Total, 8 thus giving a percentage of 8. SumMMaRY oF DENTITION. As the dentition of the Phocide is of some importance, especially with regard to variation, | have formed a table of the dentition of the seals under my observation. I 4 OSTEOLOGY OF ANTARCTIC SEALS. 201 have also examined the crania of the seals in the University Museum of Science and Arts, Edinburgh, and include such in the following list :— Weddell seal: 21 specimens with formula . - : : 1 specimen : 2.1.6 aarp wl The fourth post-canine in the upper jaw was obviously split into two, thus account- ing for the variation. Sea-leopard: 5 specimens with dental formula ; : ; : i i : 25 Crab-eating seal: 7 specimens with formula airs Sea-lion (Otaria Jubata): 2 specimens with formula aes - : 1 specimen _,, a 3. 1. 6 3. 1.5 LITERATURE. The Zoology of the Voyage of the “ Erebus” and “ Terror,” pp. 7-8, plates vii. and viii. J. E. Gray, Ommatophoca Rossi, 1844. Challenger Reports, vol. xxvi., Sir Wm. Turner, 1888. Résultats du Voyage du S.Y. “ Belgica,” G. E. H. Barrett Hamiuton, 1901. National Antarctic Expedition: Natural History, vol. ii., E. A. Wiuson. DESCRIPTION OF PLATE. Fic. 1.—Ross skull, No. 43. Superior aspect. Fig. 5.—Ross seal, No. 43. 15th dorsal vertebra. Showing costal process on right side, and Fic. 2.—Ross skull, No. 43. Lateral aspect. facet for rib on left side. Fig. 3.—Ross skull, No. 43. Anterior aspect. Fic. 6.—Ross seal, No. 43. Femur showing non-articular area on head. Fic. 4.—Ross skull, No. 43. Inferior aspect. Y Sain PRESEN TED Rages 18 FEB, 1910 Trans. Roy. Soc. Edin. Von. XLVII. Rosert B. Taomson—“ Scottish National Antarctic Expedition: Osteology of Antarctic Seals.” Fic. 1. Fie. 2. Fic. 3. Fie, 4. Fic. 5. 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Wop -ViLEL, ys ; ls. 6d. ls. 1dd. 2 4 MAY. 1910 TRANSACTIONS OF THE ROYAL SOCIETY OF EDINBURGH. VOLUME XLVII. PART II.—SESSION 1909-10. CONTENTS. No. IX. Lower Palxozoic Hyolithide from Girvan. By F. R. Cowrrr Ruzp, M.A., F.G.S. (With Three Plates), . : . : : (Issued October 13, 1909.) a X. Report upon the Anatomy and Embryology of the Penguins collected by the Scottish National Antarctic Expedition, comprising: (1) Some Features in the Anatomy of the Penguin ; (2) The Embryology of the Penguin: a Study in Embryonic Regression and Progression. By D. Warersron, M.A., M.D., F.R.C.S.E., and A. Camppett Geppss, M.D. (With Three Plates), (Issued October 21, 1909.) XI. The Pharmacological Action of Harmaline. By James A. Gunny, M.A., M.D., D.Sc., (Issued December 31, 1909.) XII. The Pharmacological Action of Protocatechyl-Tropeine. By Professor C, R. MarsHALL, (Issued January 7, 1910.) XII. The Pharmacological Action of Tutu, the Toot Plant of New Zealand. By Professor C. R. MarsHatt, (Issued January 29, 1910.) XIV. On the Life-history of Hydrobius fuscipes, L, By Frank Batrour-Browne, M.A. (Oxon.), ° FE.Z.S. (With Three Plates), (Issued February 1, 1910.) XV. Strophanthus sarmentosus : its Pharmacological Action and its Use as an Arrow-Potson. By Sir THomas R. Fraser, M.D., F.R.SS. L. & E., Professor of Materia Medica in the University of Edinburgh, and Auistzr IT. Mackenziz, M.A., M.B., Ch.B., Carnegie Research Scholar. (With Eleven Plates), ; , (Issued February 8, 1910.) EDINBURGH: PUBLISHED BY ROBERT GRANT & SON, 107 PRINCES STREET, AND WILLIAMS & NORGATE, 14 HENRIETTA STREET, COVENT GARDEN, LONDON. MDCCCCX. Price Twenty-three Shillings. 223 245 273 287 317 341 ( 203 ) IX.—Lower Paleozoic Hyolithide from Girvan. By F. R. Cowper Reed, M.A., F.G.8. Communicated by Dr Horne, F.R.S. (With Three Plates.) (MS. received March 5, 1909. Read June 7, 1909. Issued separately October 13, 1909.) INTRODUCTION. The extensive series of specimens of members of the Hyolithide which Mrs HiizaBeTH Gray has collected from the Ordovician and Silurian beds of the Girvan district, and submitted to me for investigation, forms the basis of the following memoir. Through the kindness of Dr Joun Horne and Professor J. W. Gregory, special facilities have been afforded me of examining the other Scotch examples in the Geological Survey Collection, Edinburgh, and the Museum of Glasgow University. My thanks are also due to Dr F. L. Kircnin for the opportunity of studying the specimens in the Jermyn Street Museum. The collections in the Sedgwick Museum, Cambridge, from various localities and many Lower Paleozoic horizons in Great Britain and Ireland, have enabled me to compare the Girvan species with previously described and typical forms. ‘he specimens in the same museum from Bohemia, named by Barranve himself, have proved of much value. Examples in the British Museum have also been consulted. The classification of the Hyolithide drawn up by Hotm is here followed, the genus Hyolithes being divided into two principal subgenera, Hyolithes sens. str. and Orthotheca, but Novax’s Ceratotheca is retained as a third independent group of generic rank for two doubtful species. The genus Pterotheca is also included in the same family, but the reasons for doubting the accuracy of this customary position are mentioned and briefly discussed. Genus Hyolithes, Hichwald, 1840. Subgenus Hyolithes, Kichwald-Holm, 1893. HYoLiITHES ARDMILLANENSIS, sp. nov. (Pl. I. figs. 1-6.) Shell straight, short, broad, rapidly tapering at about 1 in 24, dorso-ventrally compressed. Ventral face moderately convex, rounded or slightly subangular Dorsal face flat. Mouth oblique, with strongly arched dorsal lip. Lateral angles sharp, acute. Ventral face ornamented with 18-20 strong, rounded, straight longitudinal raised lines, of equal or subequal size, and separated by wider interspaces. Dorsal face, with fine concentric arched striz and traces of longitudinal folds, and occasionally with shallow submarginal grooves visible in the cast. TRANS, ROY. SOC, EDIN., VOL, XLVII, PART II. (NO. 9), 30 204 MR F. R. COWPER REED ON Dimensions - Te Jue Length : : : : : : 26:0 c. 23°0 mm. Width at mouth . : ; : ? 10°5 ce. 9:0 ” Horizon and Locality.—Balelatchie Group, Ardmillan. Remarks.—This is a well-characterised form. Generally only the dorsal face is preserved or the internal cast of the shell. The strong, regular longitudinal lineation of the ventral face and the short, broad shape of the shell are distinguishing features. Houm’s H. cymbium,* from the Lituites Limestone, appears to be closely allied in both these features, but has finer longitudinal lines and a slightly curved shell. The species described by Remetit as H. inequistriatus, from the boulders of Orthoceras Limestone in the North German Drift, closely agrees with our species in shape and rate of tapering, and the ornamentation is of the same type, though the longitudinal lines vary in strength. Some unnamed specimens from the Bala beds of Horderley, in the Jermyn Street Museum, bear considerable resemblance to H. ardmillanensis in cross-section, rate of tapering, and ornamentation, but are too poorly preserved to warrant identification with it. An operculum (PI. I. fig. 6) occurs associated with the above shells at Ardmillan which is of the requisite proportions and may probably be referred to this species. It is subelliptical in shape, folded sharply along a line near the ventral margin, parallel to the longer diameter, and is conical and moderately elevated, with the apex situated on the fold close to the ventral margin. The ventral side is short, narrow, very steeply inclined, and almost at right angles to the rest of the surface. The general surface of the cone, though rising to the apex, is somewhat concave, and is marked with three strong radiating grooves on each side, diverging from the apex, and not reaching the margin. As we are dealing with an internal cast, these grooves really represent internal ridges in the operculum, as BARRANDE pointed out. Fine radiating lines occur on each side of the sharp, angular junction of the ventral side with the general surface, and are traceable (though less distinctly) near the margin on other parts of the surface, but over the greater part only very delicate concentric strize can be detected. The transverse diameter of one such operculum is 11°5 mm. BarranvE{ has figured from Stage Ddl a somewhat similar isolated operculum with the internal cast showing three radiating grooves on each side; these he interprets as muscular impressions. It should be noted that none of the Girvan opercula have been found in position, but their relations to the shell are inferred from the Bohemian species described by * Hou, “Sver. Kamb.-silur. Hyol. 0. Conul.” (Sver. Geol. Undersékn., Ser. C, No. 112, 1893), p. 82, t. iii. figs. 6-11. + RemE Lh, Zeitschr. deut. geol. Gesell., Bd. xl., 1888, p. 670, t. xxviii. figs. 4-6. | Barranpe, Syst, Silwr, Bohéme, vol. iii., 1867, p. 98, pl. ix. figs. 16, 17. LOWER PALAOZOIC HYOLITHIDA FROM GIRVAN. 205 BaRRANDE with the operculum in place. In these the apex of the cone is always nearer the ventral side of the shell; but SaurER regarded it as nearer the dorsal side, thus reversing its relations. BaRRANDE’s orientation is here followed. Hyolithes asteroideus, sp. nov. (Pl. I. figs. 79a.) Shell straight, gently tapering at about 1 in 5. Ventral face moderately convex, weakly angulated in centre and obscurely on each side near lateral edge, so that it shows two pairs of flattened surfaces, of which the median pair is the wider. Dorsal face weakly convex. Lateral angles subacute. Section plano-convex. Mouth oblique, with strongly arched, high dorsal lip. Ventral face with one faintly impressed longitudinal line along median keel, and traces of very delicate longitudinal strize over rest of surface, which is also crossed by more or less developed growth-ridges and strie. Dorsal face with strongly arched transverse growth-strie, most marked near mouth ; (longitudinal strize absent 2). Horizon and Locality.—Starfish Bed, Drummuck Group, Thraive Glen. Remarks.—Only imperfect shells of this species are known, but the formation of the ventral face is characteristic. One fragment with an apertural diameter of 9 mm. has a leneth of 20 mm., and the shell, when complete, must have measured about 45 mm. Hicuwaup’s H. acutus,* from the Orthoceras Limestone, has a somewhat similar shape and cross-section, but the ornamentation is different.j H. elegans, Barr.,{ from Stage D, Bohemia, is certainly allied to our species. Hyolithes candidus, sp. nov. (PI. I. figs. 10-13.) Shell straight, short, broad, tapering at about 1 in 2. Dorsal face divided longi- tudinally into three areas, the lateral ones, which are each about one-quarter the width of the whole face, being flattened, gently inclined, and ornamented with 4-6 strong raised longitudinal lines; the median area is of twice the width, slightly concave or flattened, with very fine transverse arched striz and very delicate longitudinal stri, producing a minute cancellation of the surface. Dorsal lip of the mouth strongly arched. Ventral face rounded, gently convex, with rather distant fine longitudinal lines, about 20-25 in number, crossed by much more numerous transverse growth-lines. Lateral angles obtuse, rounded. Dimensions :-— Length . : ; é é : : : 11:5 mm. Width of mouth . . 3 5 5 F 4:5 Horizon and Locality.—Whitehouse Group, Whitehouse Bay. Remarks.—This species is rather imperfectly known, being based on the one * HICHWALD, Leth. Ross., p. 1045, t. xl. figs. 13, 14. + REMELE, Zeztschr. deut. geol. Gesell., Bd. xli., p. 762, t. xxx. } BarRanpbgE, op, cit., p. 81, pl. xi. figs. 14-25, pl. xv. figs, 34-36. 206 MR F. R. COWPER REED ON specimen showing the dorsal face and on several fragments. The dorsal face and shape of the shell resemble the Cambrian species H. sxgnatulus, Novak, * from Stage C, but the longitudinal lineation is different. The ornamentation is more like that found in many species of Orthotheca in Bohemia, figured by Novak, but H. norvegicus, Kjerulf,t from Norway, may be compared with it. The minute cancellation is met with more generally in species of Orthotheca. The shape of the shell resembles the figure of a small unnamed species of Hyolithes from Tyrone, described by Porriock { as the “internal bone of a Cephalopod.” Hypolithes columbanus, sp. nov. (Pl. I. figs. 14-14d.) Shell large, gently curved ventralwards, tapering at about 1 in 4-5, with rather an obtuse blunt tip. Ventral face moderately convex, rounded, not angulated. Lateral angles acute, but not sharp. Dorsal face slightly convex. Mouth with dorsal lip gently arched, not high; ventral lip straight. Cross-section unequally biconvex. Shell thick, with strong transverse growth-strize and ridges, especially on dorsal face, which also shows near lip about 20 impressed, widely distant, regular longitudinal strize, and a very delicate close longitudinal lineation over the whole surface; on the ventral face the longitudinal strize are indistinct. Dimensions :-— Length of fragment : : 55 mm. (estimated length of shell 80-90 mm.). Width of mouth : : ; Sie Horizon and Locality.—Drummuck Group, Thraive Glen. Remarks.—This species is not abundant. Its shape, curvature, and cross-section distinguish it from H. asteroideus, and its ornamentation from H. multipunctatus (q.v.). Eicuwa.p’s H. latus§ is allied in curvature, cross-section, and apparently in ornamenta- tion, but tapers much more rapidly to the apex. Hyolithes gurvanensis, sp. nov. (PI. I. figs. 15-17.) Shell slender, straight, slowly tapering to sharp point at about 1 in 5 or 6. Ventral face angulated in middle, the two sides meeting at about 110°-120°, with narrow, flattened border at sides. Dorsal face flat, or nearly so. Cross-section triangular. Lateral angles acute, sharp. Ventral face with very fine longitudinal striation (rarely visible), and stronger transverse concentric growth-strize. Dorsal face with very slightly arched lip and strong transverse concentric growth-strie. Dimensions :-— Length of shell . : 3 : : ‘ p 17:0 mm. Width of shell . . ‘ : : F i 3°25 ” * Novak, “Rey. Palaeoz. Hyol. Bohm.,” Abh. kin. bohm. Gesell. Wiss., vii., Bd. iv. (Math.-Nat. K1.), No. (1891), p. 23, t. iv. figs. 37-39. + Hou, op. cit., p. 87, t. iii. figs, 1-5. ft Porttock, (Geol. Rep. Londond., p. 470, pl. xxix.a. fig. 6. § Etcuwatp, Leth. Ross., p. 1045, t. xl. fig. 16, EEE |. EEE eel LOWER PALAOZOIC HYOLITHIDAZ FROM GIRVAN. 207 Horizon and Locality.—Balclatchie Group, Balclatchie. Remarks.—This species is allied to H. ommemor (q.v.), from the same horizon, but differs by its more slowly tapering form, the non-chambered apex (so far as known), the more strongly angulated ventral face, and the sharp lateral angles. The longi- tudinal ornamentation of the ventral face is only very faintly seen in one specimen, and its existence is, indeed, open to doubt; it is at any rate different to that of H. wmmemor. H. simplex, Salter,* is allied, but has not a sharply angulated ventral face, and tapers more rapidly; the operculum referred to this species by Saursr bears a con- siderable resemblance to that here mentioned under H. wmmemor, and the species occurs in the Lower Llandeilo of Shelve. H. vaginula, Salter,t is a shorter, broader shell, but has an angulated ventral face. H. triangularis (Portl.),t from Tyrone, is a larger shell, one of the type-specimens measuring just 40 mm. in length, but its rate of tapering and shape of cross-section seem identical ; the dorsal (flat) side is, however, stated to be longitudinally striated, while the ventral face has only transverse strive; but all the specimens are very badly preserved, and the lineation, which is very unsatisfactorily seen, is at any rate coarser. Hyolithes rmmemor, sp. nov. (Pl. ities, 18-2 PE fie. 1.) Shell straight, tapering at about 1 in 33, with the apical portion (z.e. the posterior fourth part of the shell) tapering more gradually and divided up by transverse partitions into 12-15 chambers. Dorsal face gently convex, with traces of narrow border. Dorsal lip gently arched. Lateral angles acute, but not sharp. Ventral face angulated, with strong submarginal raised longitudinal rib on each side, and two median weaker ribs along the central keel, but rest of surface smooth (imperfectly known). Surface of dorsal face marked by fine transverse growth-strie, with traces of 1-2 marginal longitudinal lines occasionally visible. Dimensions :— Length . ; : ° : ; ; ‘ 21:0 mm. Width of mouth . : : , : d ; 5:0 ” Horizon and Locality.—Balclatchie Group, Ardmillan. Remarks.—This small shell has very constant proportions, but in no case is the ventral face well seen; the marginal groove and border on the dorsal face are only visible when the shell is preserved, and then generally only in the apical half. The chambering of the apex and the slender, more tapering shape of this portion are remarkable, and have not been observed in other species from Girvan. The shell tapers less rapidly than H. ardmuillanensis, but is probably closely allied to it, and imperfect casts are difficult to distinguish. Its relations to H. girvanensis have been * SatTEeR, Mem. Geol. Surv., vol. iii., 1866, p. 352, pl. xi.B. figs. 22-26. + Ibid., p. 352, pl. x. fig. 14. { PortuocK, Geol. Rep. Londond., p. 375, pl. xxviii. figs. 3 a-c. 208 MR F. R. COWPER REED ON above mentioned. It is probable that Satrer* refers to H. wmmemor in recording H. reversa (Salter) from the South of Scotland. In the latter species, originally described from North Wales,t there is no mention of a tapering chambered apex, but it seems closely allied in other respects. EicHwaLp (op. cit., p. 1045) believed that the Russian species, H. acutus, HKichw., which Hotm{ has fully described, from the Orthoceras Limestone, was a close ally of H. reversa. Amongst other Scandinavian species, it may be remarked that the ribbing of the ventral face of H. scalprum, Holm,§ is somewhat similar. Probably certain small opercula (Pl. II. fig. 1) occurring in the same beds at Ardmillan are referable to this species. This type of operculum is subcircular in shape, conical, with an elevated excentric apex situated close to the ventral margin; the ventral face is triangular, short, flattened, more or less inclined to the general surface, and has a small median prominent fold below the apex, traversing it and extending to the margin. The general or dorsal surface of the operculum slopes down regularly from the apex and has one distinct regular concentric groove near the margin, and fine concentric strize above and below. In casts, two short, shallow, broad radial grooves, diverging from the apex at about 80° and widening outwards, traverse the dorsal portion of the shell, but do not extend more than two-thirds of the distance to the margin. ‘hese indicate muscular attachments, and correspond to those represented by BaRRaNDE | in opercula from Stage D in Bohemia. Hyolithes multipunctatus, sp. nov. (PI. II. figs. 2-20.) Shell straight, gently tapering at about 1 in 6-7. Dorsal and ventral faces gently convex, the ventral side rather more convex than the dorsal. Lateral edges sharp, acute. Mouth oblique ; dorsal lip slightly arched and swollen at margin. Surface of shell crossed by weak growth-ridges at irregular distances and by finer growth-lines and striz, and covered with small, round, shallow pits, situated at about two to three times their diameter apart. Dimensions :-—_ Length (of broken fragment of shell). : ; : 45:0 mm. Width of mouth . C é ; : , : 16:0 _,, Horizon and Locality.—Balclatchie Group, Ardmillan. Remarks.—Only one broken specimen with the impression of its exterior is known, but this, luckily, has the mouth and ornamentation well preserved. ‘The latter feature is peculiar, and does not appear to be of a secondary origin. Iteasily marks it off from other allied species, such as H. asteroideus. * Satter, Mem. Geol. Surv., vol. iii., 1866, p. 353, and woodcut 14, fig. 6, on p. 347. + SaurerR, in Murchison’s Siluria, 2nd edit., p. 550, foss. 10, fig. 21. {| Hou, op. cit., p. 99, t. ii. figs. 29-36, t. vi. fig. 23. § HoLm, op. cit., p. 96, t. ii. figs. 46-50. || BARRANDE, op. cit., pp. 95-97, pl. xiii. figs. A-B. LOWER PALAOZOIC HYOLITHIDA FROM GIRVAN. 209 Hyolithes penkillensis, sp. nov. (Pl. II. figs. 3-5.) Shell straight, short, broad, rapidly tapering at about 1 in 2%. Ventral face low, angulated (imperfectly known). Dorsal face flattened or very weakly convex, with narrow border distinctly marked off by groove, and with 1-3 strong, submarginal, longitudinal lines. Mouth with dorsal lip strongly arched. Lateral edges acute, sharp. Dimensions :—- Ie ae Length . : : : : : 15:0 mm. 11:0 mm. Width at mouth . : ; : 5 Oi Dios Horizon and Locality.—Penkill Group, Penkill. Remarks.—This broad, short form has much the shape of H. tardus, Barr.,* from Stage G, but the margins of the dorsal fave have the characters of H. norwegicus, Kjerulf.t . obvius, Barr.,t from Stage H, tapers less rapidly to the point, and does not show the narrow borders on the dorsal face, as does the Girvan form. There is one specimen in the Hunterian Museum referable to this species from the same locality, but it tapers rather more slowly, measuring 17 mm. in length and 6°5 mm. in width at the mouth. Hyolithes subcrispatus, sp. nov. (PI. II. figs. 6-8.) Shell short, broad, straight, tapering rapidly at 1 in 23. Dorsal face flattened, very weakly convex. Lateral angles sharp, acute. Ventral face unknown. Mouth with dorsal lip strongly arched. Surface of dorsal face crossed by strong, rounded, fairly regular and equal concentric growth-ridges or strie, and marked by 20-25 regular, straight, equal and equidistant raised thread-like lines, separated by broad interspaces. Occasionally two of the longitudinal lines seem more strongly developed, and divide the dorsal face into three equal longitudinal areas. : Dimensions :-— I II. Length . . ‘ : ; : 24 mm. 21 mm. Width at aperture : : 5 : 10 _~,, (8) gp Horizon and Locality.—Drummuck Group, Thraive Glen. Remarks.—This shell has more the shape and ornamentation of H. decipiens, Barr.§, but not such marked cancellation of the surface, nor a bent tip. H. sulcatulus, Nov..,| has a more slender shape and fewer longitudinal ribs. * BARRANDE, op. cit., p. 93, pl. vi. figs. 17-18 ; Novak, op. cit., p. 27, pl. iv. figs. 33-36. + Houm, op. ctt., p. 87, t. iii. figs. 1-5. { BaRRANDE, op. cit., p. 86, pl. xii. figs. 1-6, § BaRRANDE, op. cit., p. 80, pl. xii. figs. 33-37 ; Novak, op. cit., p. 16, t. iii. figs. 25-28. || Novak, op. cit., p. 26, t. iii. fig. 22. 210 MR F. R. COWPER REED ON Hyolithes sylvestris, sp. nov. (Pl. Il. figs. 9, 9a.) Shell short, straight, broad, tapering at about 1 in 23. Ventral face unknown. Dorsal face weakly convex, with narrow raised border, separated by broad shallow groove; surface ornamented with 30-40 fine, straight, thread-like longitudinal lines, of equal size and at equal distances apart, crossed by very delicate, closely placed, arched transverse strize. Dorsal lip strongly arched. Dimensions :— . Length . : : ; : : ‘ : 12°5 mm. Width at mouth . ‘ : ; : : 5 Or Horizon and Locality—Saugh Hill Group, Woodland Point. Remarks.—In shape this species, of which there is only one specimen, is indis- tinguishable from H. penkillensis, but the ornamentation of the dorsal face is sufficient to separate it. ‘he shell is shorter than H. crispatus (Boll.),* but the ornamentation somewhat recalls it, though the Swedish form shows no raised border. BaRRANDE'S H. columnaris,t from Stages EH and F, has a concave dorsal face, but a similar ornamentation. Perhaps Saurer’s [Hyolithes anceps, from the Silurian of Hastnor Castle, is allied ; the shape and ornamentation appear to be similar, but the species is imperfectly known, and the type-specimen is indifferently preserved. Hyolithes, sp. ind. (a). An imperfectly known species of Hyolithes, somewhat resembling in shape H. immemor, is represented by four specimens from Penkill, in the Hunterian Museum. It is too poorly preserved to warrant giving it a specific name, but it is probably distinct and new. The shell tapers at about 1 in 4; the dorsal face is slightly convex and has a very slightly arched lip; its surface is crossed by transverse strize; there seems to be no border nor marginal longitudinal lines, as occasionally seen in H. immemor. The lateral angles appear to be sharp and acute. The ventral face is angulated or subangu- lated, and marked with a few longitudinal lines, apparently about 12 or rather more in number. One specimen measures 15 mm. in length, with an apertural width of about 4 mm., and another about 12°0 mm. in length, with an apertural width of about 3°25 mm. Hyolithes, sp. ind. (6). Some imperfectly preserved examples of an indeterminable species of [Hyolithes occur in the Mulloch Hill Group of Mulloch Hill. They are rather short, and taper * Hou, op. cit., p. 89, t. ili. figs. 36-52. + BARRANDE, op. cit., p. 78, pl. ix. fig. 13, pl. xii. figs. 13-23. | Saurer, Mem, Geol, Surv,, vol, ii. pt. i., 1848, p. 355, pl. xiv. fig. 1, LOWER PALAOZOIC HYOLITHIDA FROM GIRVAN. 211 at about 1 in 3-4. The dorsal face is flat, with a strongly arched mouth. No shell or ornamentation is preserved satisfactorily. One specimen measures 15 mm. in length and 4 mm. in width at the mouth. Hyolithes, sp. ind. (7). A small species of Hyolithes occurs in the Craighead Limestone, but it is too poorly preserved for specific determination. Itis a broad, short form, tapering at about 1 in 3 ; the dorsal face seems to be flat, with a very slightly arched mouth, but no further features can be made out satisfactorily. The one specimen recognised measures about 7 mm. in length and 2°5 mm. in width at the mouth. Subgenus Orthotheca, Novak, 1887. Hyolithes (Orthotheca) subexcavatus, sp. nov. (Pl. II. figs. 10-12.) Shell straight, slender, slowly tapering at about 1 in 8, with elongated, pointed apical portion showing chambers. Cross-section subtriangular. Ventral face high, sharply angulated, the sides meeting at nearly a right angle. Dorsal face concave, with swollen margins. Lateral angles acute, but rounded. Mouth straight, truncate. Ventral face ornamented with about 30 fine, straight, longitudinal thread-like lines, of equal size and equidistant, crossed by very numerous finer, closely placed stria, giving the surface a cancellated appearance. Dorsal face with similar longitudinal lines and transverse strie. Dimensions :— Length . - : : 3 : : : 18°5 mm. Width at mouth : : ; : : ; HAA) Horizon and Locality.—Balclatchie Group, Ardmillan. Remarks.—In shape this shell much resembles H. girvanensis, but differs by tapering more slowly and by the absence of a dorsal lip and by the character of its ornamentation. In none of the specimens is the surface of the dorsal face well preserved, so that there is a little doubt as to the presence of the longitudinal lineation upon it. The chambering of the apical portion recalls H. ommemor. In cross-section our shell resembles H. (O.) cor., Holm,* and H. (O.) excavatus, Holm,t of the Cambrian, but the ornamentation is somewhat like H. (O.) fluctuosus, Holm,t{ from the Trinucleus Beds, and HH. (O.) trinucleorum, Holm.§ Our H. (O.) thraivensis tapers more rapidly, but otherwise is closely allied. There is a fairly good example of the upper portion of a shell from Desertcreat, Co. Tyrone, in the Jermyn Street Museum, which may be identified with this Girvan species. The shape and ornamentation are identical, and the specimen, when perfect, must have measured about 45 mm. long; its width at the mouth is about 6 mm. * Hou, op. cit., p. 58, t. i. figs. 60-62. + Hom, op. cit., p. 61, t. v. figs. 67-71. { Hou, op. cit., p. 68, t. iii. figs. 58-57, t. v. figs. 88-91. § Houm, op. cit., p. 67, t. v. figs. 79-87. TRANS. ROY. SOC. EDIN., VOL. XLVII. PART II. (NO. 9). 31 212 MR F. R. COWPER REED ON Hyolithes (Orthotheca) subornatus, sp. nov. (Pl. II. figs. 13-15.) Shell slender, straight, slowly tapering at about 1 in 8, triangular in cross-section. Ventral face strongly angulated at nearly a right angle, high, with strong median longitudinal rib and 6-8 fine, straight, thread-like longitudinal lines on each side (with 2-3 very delicate ones between adjoining ones), and a strong marginal or supra-marginal longitudinal rib near the lateral edge. Dorsal face apparently flat or slightly concave and crossed by very fine transverse striz. Mouth straight. Horizon and Locality.—Whitehouse Group, Whitehouse Bay (and Shalloch Mill ?). Remarks.—The peculiar ornamentation in this species distinguishes it from all others in the Bala Beds. One specimen from Whitehouse Bay measures about 30 mm. in length, but the tip is missing, so that its original length must have been about 35 mm., and its diameter at the mouth is about 4 mm. Some poorly preserved specimens from Shalloch Mill probably belong to this species, but seem to taper rather more rapidly. Hyolithes (Orthotheca) thrawwensis, sp. nov. (PI. IL figs. 16-23a.) Shell very slightly and regularly curved dorsalwards, or straight, gently tapering at about | in 6 or 7, markedly triangular in cross-section, nearly as high as wide. Ventral face strongly convex, high, more or less angulated. Dorsal face flat or slightly concave. Lateral angles rounded, obtuse. Mouth straight, not oblique; dorsal lip straight, not arched. Surface of ventral face marked with 30-36 longitudinal equal or subequal raised lines, separated by broader interspaces, and crossed by subequal, more closely placed and more numerous but weaker raised lines, forming granules at the intersections, and giving a cancellated appearance to the surface; the 1 or 2 longitudinal lines on the rounded lateral angles are rather stronger than the rest. Dorsal face crossed by straight transverse lines similar to and continuous with those on the ventral face, but without longitudinal lines. Dimensions -— I. II. Length about . ; : : : 33:0 mm. 37:0 mm. Width at mouth about . ; é j BO) = 0) 5 Horizon and Locality.—Drummuck Group (? including Starfish Bed), Thraive Glen. Remarks.—This beautifully ornamented species seems abundant in the Drummuck 3eds of Thraive Glen, and is usually well preserved and has the shell still attached. H. (Orthotheca) trinucleorum, Holm,* may be compared with this species, and it comes from a homotaxial horizon. The ornamentation in H. (O.) thraivensis is coarser than in H. (O.) subexcavatus, and the shell tapers more rapidly, and is usually somewhat curved. * Hou, op. cit., p. 67, t. v. figs. 79-87. LOWER PALAOZOIC HYOLITHIDA FROM GIRVAN. 213 Hyolithes (Orthotheca) thrawensis, var. granuliferus. (Pl. II. figs. 24, 25.) Shell straight, triangular, slowly tapering at about 1 in 6. Ventral face high, angulated, its sides meeting at about a right angle. Dorsal face slightly concave. Lateral angles obtuse, rounded. Mouth straight. Ventral face bearing one strong median raised longitudinal line along ridge and 6-8 finer raised lines on each side, each line bearing a single row of small equidistant tubercles. The longitudinal lines are not all of equal strength or at equal distances; some are finer than others and are not continued to the point, only extending about half or two-thirds the length of the shell ; the interspaces between the lines are wide, and very delicate transverse lines cross them. The rounded lateral angles bear several similar longitudinal lines, but the dorsal face seems devoid of them, and only shows very faint transverse striz. Horizon and Locality.—Drummuck Group, Thraive Glen. Remarks.—The one internal cast and external impression on which this variety is based differ from the typical form by the granules on the longitudinal lines being fewer and so much larger as to be worthy of the designation of tubercles. The longi- tudinal lines also are fewer and less regular, and the concentric strize much finer and fewer. The fragment measures about 11°75 mm. in length, with an apertural diameter of about 4°25 mm. Hyolithes (Orthotheca *), sp. ind. An elongated, narrow, slowly tapering species from Penkill is represented by one poor and imperfect specimen in the Hunterian Museum, Glasgow. The fragment measures 32 mm. in length, but the tip is broken off; at its broader end it measures 3°25 mm. in width; its true rate of tapering is about 1 in 11. The impression of the one face preserved shows that this face (? the dorsal) must have been gently convex, and there are traces of faint transverse, scarcely arched striz running across it. No other features can be detected. Genus Ceratotheca, Novak, 1891. Ceratotheca ? subuncata, sp. nov. (Pl. II. figs. 26-28.) Shell cornuate, strongly curved back at about half its length, and thence rather rapidly tapering to point, the curvature being such that the posterior part of the shell is bent back in a regular, nearly semicircular curve, so that the apical extremity lies at about two-thirds to three-fourths the length of the shell. Section of shell subcircular or elliptical. Mouth at right angles to shell, not oblique; margin of mouth slightly sinuous. Shell thin; surface ornamented with very fine concentric growth-strie and with very delicate closely placed longitudinal strize (observable near mouth), which on crossing the transverse striz are raised into minute granules, 214 MR F. R. COWPER REED ON Dimensions :— I. Il. TI. Length of shell (to bend) . : : 15:0 22-0 21:0 mm. Diameter of mouth : f : 85 10°5 11:0 mm. Horizon and Locality.—Penkill Group, Penkill. Remarks.—The specimens are mostly rather crushed and imperfect, and the surface ornamentation is rarely visible. The species which most resembles it is Ceratotheca unguiformes, Novak,* from Stage He 1 in Bohemia, but the rate of tapering in the latter is less rapid and the curvature of the shell begins nearer the apex ; the ornamentation also is different. The inclusion of C. subuncata and the following species, C. balclatchiensis, in the genus Ceratotheca, appears to me rather doubtful. However, it is impossible to establish a new genus on such indifferent material, and it seems best to leave them associated in it for the present. The resemblance of C. swhuncata in general shape to some species of Hecyliomphalus may be noted, and even Linpstr6m’s Huomphalus gotlandicus + has a certain degree of similarity. Ceratotheca ? balclatchiensis, sp. nov. (PI. III. figs. 1-3.) Shell conical, slightly curved ventralwards, rather rapidly tapering at about 1 in 3. Cross-section subcireular to subelliptical. Ventral face strongly convex, rounded. Lateral angles rounded, obsolete and indefinite. Dorsal face rather less convex than ventral. Mouth straight or with margins slightly sinuous, at right angles to shell; no arched dorsal lip. Shell lustrous, corneous, thin, smooth, or with irregular faint concentric growth-ridges and strize. Dimensions :-— I. II. III. Length . : ; : 21:5 25°0 18:0 mm. Diameter at mouth 5 ; , 8:0 9-0 7:0 mm. Horizon and Locality.—Balclatchie Group, Balclatchie. Remarks.—None of the specimens are very well preserved or complete, but it is an abundant form at Balclatchie, and the numerous examples enable one to give a fairly complete diagnosis. The affinities of this species are doubtful. It is much less com- pressed dorso-ventrally than Hyolithes aduncus, Barr., from Stage EH; the lateral angles are less evident and the shell tapers less rapidly. The thin, black, corneous nature of the shell makes it doubtful if this organism is rightly referred to the Pteropoda at all, but it hardly seems possible to regard it as annelidan in nature, and it may provision- ally be referred to Ceratotheca. From its shape it looks as if it might be more correctly placed with the Scaphopoda, but the nature of the shell is against this reference. * NOVAK, op. cit., p. 33, t. i. figs. 28, 24. + Linpstrom, Silur, Gastrop. Pterop. Gotland (1884), p. 139, pl. xiii. figs. 19-31. LOWER PALAOZOIC HYOLITHIDA FROM GIRVAN. 215 Genus Pterotheca, Salter, 1852. The genus Pterotheca, which was established by Saurer®* to contain an Irish fossil described by Porrtock as Atrypa transversa, is represented not only by British species, but also by others from America and Bohemia.t Its true position and affinities have been a matter of uncertainty: ZirreL, as recently as 1903, retained it in the Hyolithide, but it is believed by Utricu and ScorreLp { to belong to their family Carinaropside, which is defined by them as follows :—‘‘ Symmetrical, almost patelliform shells, the aperture being greatly expanded: apex small, involute, overhanging the posterior margin, consisting of no more than two volutions. Within the aperture a broad concave septum. Anterior lip with a central emargination.” The type of the genus Carinaropsis § is C. carinata, Hall, of the Trenton Group, and the close affinities of Carinaropsis with Bellerophon and Pterotheca are clearly pointed out by ULRicH and SCOFIELD. Pterotheca agrees with Camnaropsis in a striking manner, and possesses the characteristic internal septum, the broadly expanded mouth, and the carinated back, but differs by the shell being simply arcuate with an incurved beak, instead of being coiled into two volutions, and also by possessing a pair of small vertical septa in the apical extremity, dividing the cavity into three portions. This peculiar septate apical region is considered by the same authors to be of generic importance only, but sufficient to make it doubtful whether Pterotheca is a Pteropod, and should not rather be classed with Carinaropsis as a family of the Docoglossa. Novax|| considered that his genus Pterygotheca somewhat recalled Saurer’s Pterotheca, but it is difficult to see any structural affinity, or even to recognise much superficial resemblance. . Satrer’s definition of the genus Pterotheca is short :—‘‘ Shell transversely oval, bilobed, with wavy sides and a strong median keel. Ventral plate short, narrow, flat.” Only a few species have been described, and the type is Pé. transversa (Portl.).{ The genus is stated to occur in Bohemia and North America, and it ranges through the Ordovician and Silurian. The best-known British species is Pt. corrugata, Salter, 1852, which is recorded from the “Caradoc” rocks of Dolbenmaen, near Criccieth.** Pt. undulata, Salter,tt from Carnarvonshire, has apparently never been figured or described. Pt. avirostris, Salter (MS.), is from the Woolhope and Upper. Llandovery Beds. * SALTER, Brit. Assoc. Reports, 1852, Trans. Sect., p. 61. + BARRANDE, op. cit., p. 99, pl. xv. figs. 1-6. { Unricu and Scortexp, “ Lower Silur. Gastr. Minnesota” (Geol. Nat. Hist. Surv. Minnesota, vol. iii. pt. ii., Final Rep., “ Palezont,” 1897, chap. x.), pp. 857, 926. § Hatt, Paleont. New York, vol. i., 1847, p. 183. || Novak, “Rev. Palseoz. Hyolith. Bohmens,” Abhandl. kén. bohm. Gesell. Wissen., vii., Bd. iv. (Math.-nat. K1.), No. 6, 1891, p. 45, t. iii. figs. 36-47. ‘| Porriock, Geol. Rep. Londond., p. 455 (Atrypa transversa) ; SaureR, Brit. Assoc. Rept., 1852, Trans. Sect., p. 61; Morcuison’s Stluria, 5th ed., 1872, foss. 41, fig. 4, p. 199. ** Saurer, Mem. Geol Surv., vol. iii., 1866, p. 353, fig. 18. tt SALTER, op. cit., 1866, p. 274. 216 MR F. R. COWPER REED ON Pterotheca drummuckensis, sp. nov. (Pl. III. figs. 4, 5.) Shell subcircular to transversely subcordiform, rather broader than long. Apertural margin emarginate, with broad, rounded lateral lobes, and broad, angular median notch between them. Median fold narrow, sharp, acutely angular, not projecting on apertural margin, but corresponding with angular notch. Lateral folds very low, wide and inde- finite, passing into flattened lateral portions of shell. Posterior margins of shell meeting at apex at about 100°-110°. Pair of narrow shallow grooves diverging forwards from apex at about 45°, halfway between median ridge and lateral margins of shell. Surface marked with regular strong corrugations, concentric to apertural margin, and bending back sharply in median notch so as to form re-entrant angle of about 50° on ridge of keel. Dimensions :— I Le. Length : . : : 5 20:0 15:0 mm. Width : : : A ; : 20:0 180: 5, Horizon and Locality.—Drummuck Group, Thraive Glen. Remarks.—This shell, though only known by imperfect specimens, differs from the next described species, Pt. gurvanensis, by its shape, its non-projecting median keel, the angle of divergence of the grooves on the surface, the acutely emarginate apertural margin, and the regular corrugation. Pterotheca girvanensis, sp. nov. (Pl. IIL figs. 6-13.) Shell transversely subelliptical, widest towards the front. Apertural margin sinuous, projecting in middle in short subangular tongue, excavated slightly on each side, and with rounded, more or less projecting lateral lobes, forming the angles and merging into the lateral margins, which are rounded. Apex small, sharp, incurved. Surface of shell gently convex, elevated into sharp median longitudinal fold, proceeding from apex with increasing height, width, and acuteness to the apertural margin, where it stands up prominently and projects as the median tongue; a pair of broader but low, rounded, indefinite lateral folds run forwards from the apex divergently, almost dying out on the apertural margin, where they form the lateral lobes. Two shallow definite grooves, slightly curving outwards, diverge from the apex at about 60°--70°, and run forwards towards the outer lateral angles of the apertural margin, nearly or quite reaching it, and they lie on the outer slopes of the lateral folds. These grooves correspond internally with low ridges or septa. Internal concave plate present, extending about two-thirds the length of the shell. Surface of shell minutely punctate, and with more or less de- veloped concentric corrugations and strize following the outline of the apertura] margin. Dimensions :-— i ie IE Length : : : ; ; 15:0 9:0 13:0 mm Width ‘ : : : : 20:0 12°5 20°0 ,, LOWER PALAOZOIC HYOLITHIDA FROM GIRVAN. 217 Horizon and Localities.—Balclatchie Group, Balclatchie and Ardmillan. Remarks.—This species is undoubtedly closely allied to Pt. corrugata, Salter. The median slit which the latter species is represented as possessing near the anterior margin is apparently due merely to the breaking off of the acute carina, as precisely the same appearance is met with in damaged specimens of this Girvan form. There are three lateral folds represented by SaurER on each side of the central one, and the con- centric striz are said to meet the keel abruptly at a right angle, whereas in ours they pass over it continuously, following the outline of the apertural margin. Pterotheca mullochensis, sp. nov. (Pl. III. figs. 18-20.) Shell transversly elliptical to subcircular. Apertural margin arched forwards, not emarginate, with small projecting triangular tongue in middle. Surface gently convex. Apex small, curved. Median fold angulated, acute, narrow and weak near beak, but increasing in height and width to front margin, where it projects slightly as tongue. Lateral lobes on apertural margin and lateral folds on surface obsolete, but halfway between median fold and lateral margin of shell the surface is angulated, being bent down rather suddenly on each side, along lines diverging from the apex at about 60° ; the angulations die out towards the anterior margin, curving gently outwards, or running nearly straight forwards; they correspond near their origin to the course of the thin internal divergent septa which start from the apex and extend about one-third to one-half the length of the shell. Internal plate gently concave, extending forwards . rather more than half the length of shell. Surface marked with numerous moderately strong concentric corrugations and strie. Dimensions :— I, II. Ill. Length . : ; ‘ : 24:0 22°5 23:0 mm. Width =. ; ; : ; 26:0 27-0 c. 25:0 ,, Horizon and Locality.—Mulloch Hill Group, Mulloch Hill; and (?) Saugh Hill Group, Newlands. Remarks.—The shape of the anterior margin, the obsolescence of the lateral lobes and folds, the angulation of the lateral surfaces and general shape of the shell, dis- tinguish it from Pt. gurvanensis and Pt. drummuckensis. Porriock’s Pt. transversa from Desertcreat, the type of which is in the Jermyn Street Museum, much resembles Pi. mullochensis in surface characters, but is broader and more transversely elliptical in shape, its length to its width being as 2-3; and it possesses, near the beak, a small additional pair of very short septa between the two long divergent ones, and the latter bend away at rather a wider angle, though this may be due to crushing. The con- centric rugz are less developed, the concentric markings being rather of the nature of fine wrinkles and strize. 218 MR F. R. COWPER REED ON Pterotheca multidecorata, sp. nov. (Pl. III. figs. 14-16.) Shell semi-elliptical, with long, strongly arched apertural margin and short postero- lateral margins meeting at apex at wide angle (about 130°-150°); apertural margin narrowly and not deeply, but acutely, emarginate in centre. Surface of shell gently convex; median fold high, angulated, acute, with narrow raised keel, increasing eradually in width and somewhat in height to anterior margin, where it ends in the median notch; lateral folds low, rounded, curving slightly outwards and projecting as rounded lobes on margin, well marked off from median ridge and from lateral portions of shell, which form broader and lower outer rounded folds; the grooves separating these outer from the inner lateral folds diverge at rather less than 45°. Surface of shell marked (1) by a few concentric corrugations and strize bending back acutely at a re-entrant angle of 45°-50° on median fold, and (2) by fine radial striz, slightly wavy, and increasing in number towards the margin, containing along them a row of closely set minute puncte. Internal septa indistinct, thin, gently curved, less than the length of shell, diverging from apex at about 30°—45°, and running in outer lateral grooves. Dimensions :-— I. IL. Length : : : : : ; 9°5 15:5 mm. Width 5 é : ; : é 11:5 21:0 mm. Horizon and Locality.—Penkill Group, Penkill. Remarks.—The ornamentation of this shell easily marks it off from all the other species, somewhat resembling that in the brachiopod Porambonites. The two pairs of lateral folds and shape of the shell are additional distinguishing features. Pt. avirostris, Salter (MS.), from the Upper Llandovery of Bog Mine, Shelve, and Damery Bridge, Tortworth, is only known as internal casts. It is almost circular in outline, and does not show clearly two pairs of lateral folds, but the high angulated narrow keel and the small angle of divergence between the internal septa constitute points of resemblance between it and Pt. multedecorata which are worthy of notice. Pterotheca simplex, sp. nov. (PI. III. figs. 17, 17a.) Shell small, transversely semi-elliptical, with strongly arched apertural margin. Surface gently convex, with one broad, slightly elevated subangular median fold, increasing rapidly in width from the beak to the apertural margin, where it projects slightly and has a width equal to nearly half the shell; it is bounded laterally by weakly impressed shallow grooves, curving gently outwards to margin, and diverging from apex at about 45°. Lateral folds obsolete. Concentric corrugations and strie few and faint, strongest near apex. Dimensions :— Length . . ADDED 4 _ 5-25 mm. Width : ; ‘ 70 ” LOWER PALAOZOIC HYOLITHIDA FROM GIRVAN. 219 Horizon and Locality.—Balclatchie Group, Balclatchie. Remarks.—The simplicity of the lobation of this small shell distinguishes it from Pt. girvanensis, and it does not seem possible to regard it as merely the young of that species. It more resembles the Bohemian species, Pt. bohemica, Barr.* The American species, Pt. expansa (Emmons),t may also be compared. List oF SPECIES. Hyolithes ardmillanensis, sp. nov. 53 asteroideus, sp. nov. 3 candidus, sp. nov. - columbanus, sp. nov. = girvanensis, sp. nov. sn immemor, Sp. NOV. As multipunctatus, sp. nov. Fs penkillensis, sp. nov. oa subcrispatus, sp. Nov. ‘ sylvestris, sp. nov. a sp. ind. (a). = sp. ind. (f). ee Sp. indin(y), Hyolithes (Orthotheca) subexcavatus, sp. nov. subornatus, sp. Nov. thraivensis, sp. nov. var. granuliferus. ” by) ” LP) ” 4 i tsp. ind. Ceratotheca ?subuncata, sp. nov. e 1 balelatchiensis, sp. nov. Pterotheca drummuckensis, sp. nov. 5 gurvanensis, sp. nov. - mullochensis, sp. nov. re multidecorata, sp. nov. a simplex, sp. nov. STRATIGRAPHICAL DISTRIBUTION OF THE SPECIES. Craighead Limestone— Hyolithes, sp. ind. (y). Balclatchie Group— Hyolithes ardmillanensis. 3 girvanensis. FA immemor. - multipunctatus. 35 (Orthotheca) subexcavatus. Ceratotheca *? balclatchiensis. Pterotheca girvanensis, > simples. Whitehouse Group— Hyolithes candidus. » (Orthotheca) subornatus. Drummuck Group— Hyolithes asteroideus. 34 columbanus. * BaRRANDE, op. cit., p. 104, pl. xv. figs. 1-6. . Drummuck Growp—continued. Hyolithes subcrispatus. 5 (Orthotheca) thratvensis. 55 P rr var. granuliferus. Pterotheca drummuckensis. Mulloch Hill Group— Hyolithes, sp. ind. (f). Pterotheca mullochensis. Saugh Hill Group— Hyolithes sylvestris. 5 sp. ind. (a). t Pterotheca mullochensis. Penkill Group— Hyolithes penkillensis. ‘ (Orthotheca 2) sp. ind. Ceratotheca ? subuncata. Pterotheca multidecorata. + Emons, Geol. Rep. 2nd Distr. New York, 1842, p. 397, figs. 109-112; WELLER, Geol. Surv. New Jersey, Paleont. iii. (1902), p. 179, pl. xii. fig. 35. TRANS. ROY. SOC. EDIN., VOL. XLVII. PART II. (NO. 9). 32 220 MR F. R. COWPER REED ON REMARKS ON THE STRATIGRAPHICAL DISTRIBUTION. It is noticeable that the greatest variety of species is met with in the Balclatchie Group. In the underlying Stinchar Group only one species, represented by a single individual, is so far known. The Whitehouse Group has yielded no more than two species, while the Drummuck Group is particularly rich in members of the family. In the Silurian Beds it is remarkable that the Penkill Group contains a greater abundance than the earlier Silurian horizons. This irregular development of specific types does not appear to be directly related to the lithological characters of the strata, for though the argillaceous Balclatchie rocks have the maximum of species, the sandy Drummuck Beds have many more types than the fine-grained shaly Whitehouse Group. On the other hand, the arenaceous Mulloch Hill and Saugh Hill Groups are very poor in representatives of the family, while the Penkill mudstones contain a compara- tive abundance of forms. Other factors than the coarseness of the sediments must have been at work in determining the relative abundance of species. It cannot be alleged that their absence or rarity is due to their non-preservation in a coarse matrix, for the evidence of the Drummuck Beds proves that sandy deposits lend themselves just as well as clays to their conservation. Their independence of the rock-composition is as noteworthy in Bohemia as in the Girvan district. In addition to the relative number of species on the various horizons, we must have regard also to the relative number of individuals of each species, as evidenced by the specimens in the collections. In this case, also, the maximum is found in the Balelatchie and Drummuck Groups. On the former horizon, H. ardmillanensis, AA. girvanensis, and H, immemor are represented by so many specimens that they easily outnumber all the other forms. In the Drummuck Beds the commonest species are H, (Ortho.) thrawensis and H. asteroideus, but the former is by far the most abundant. With respect to the Silurian species, there are very few examples of any of them, and less than a dozen have come under my notice from the Mulloch Hill and Saugh Hill Groups. In the Penkill Group individuals are less rare, but pteropods do not seem to be at all abundant members of the fauna. We are generally precluded from making any estimate, even with approximate accuracy, of the actual relative abundance of species and individuals of any zoological group of fossils in the beds deposited at a given period, the obstacles being the paucity of material and the unsystematic method of collecting, or accidental manner by which most museums have obtained their fossils. But in this case Mrs Gray’s regular visits and careful collecting from the sgme district and horizons for many years past justify us in considering the above conclusions as to the proportional distribution of the Hyolithide in the beds of the Girvan area to be a closer approximation to the truth than usual. LOWER PALA OZOIC HYOLITHIDA FROM GIRVAN. 221 RELATIONS OF THE GIRVAN HYOLITHIDH TO THOSE OF OTHER AREAS. It is a remarkable fact that all the Girvan species, except one or possibly two which occur in Co. Tyrone, appear to be peculiar and new. After a careful comparison with all the well-known English and Welsh species, it does not seem possible to regard any as identical, and the relationship of any of them is not very close. Many, however, of the Girvan forms are allied to Scandinavian ones, of which we have such a full and detailed knowledge owing to Hotm’s memoir. A few show affinities with Bohemian species. It may be noticed that amongst the Girvan Conularie, Miss SLaTER * recognised one species identical with a Scandinavian form, and more than one possessing considerable affinities with those from that region. CoNCLUSION. By the foregoing study of the Hyolithide the Girvan area is seen to contain an unusually rich and varied assemblage of members of the group. The interest of them is heightened by the local restriction and peculiarity of the species, and by the frequently good state of preservation of the material, which renders it possible to define their characters with greater precision than is usual in dealing with members of the same group from other British localities. EXPLANATION OF PLATES. Puate I. Figs. 1-6. Hyolithes ardmillanensis. 1, ventral face; 2, dorsal face; 2a, cross-section; 3, dorsal face ; 4, ventral face ; 5, dorsal face ; 6, operculum. (All x 2.) » (-9a. Hyolithes asteroideus. 17, ventral face; 7a, cross-section; 8, dorsal face; 9, ventral face ; 9a, cross-section, (All x 2.) » 10-13. Hyolithes candidus. 10, dorsal face; 11, 12, ventral face (all x 24); 13, ornamentation of ventral face (x 5). », 14-14d. Hyoltthes columbanus. 14, dorsal face; 14a, side view ; 140, ventral face ; 14c, cross-section (all x 2); 14d, portion of dorsal face near lip (x 3). », 15-l7a. Hyolithes girvanensis. 15, ventral face; 16, dorsal face; 17, 17a, dorsal and ventral faces of same shell. (All x 24.) », 18-21. Hyolithes immemor. 18, 18a, dorsal face, impression and cast; 19, dorsal face; 20, ventral face (three-quarter view, right side) ; 21, dorsal face. (All x 24.) Prate II. Fig. 1. Hyolithes immemor. Operculum (x 24). Figs. 2-26. Hyolithes multipunctatus, 2, ventral face; 2a, cross-section (both x 2); 2b, ornamentation (x 3), », 3-5. Hyolithes penkillensis. Dorsal faces (x 24). * Starter, Mon. Brit. Conularie (Paleont. Soc.), 1907, pp. 16, 21, 31. »” ” ” 9 LOWER PALAOZOIC HYOLITHIDA FROM GIRVAN. Hyolithes subcrispatus. Dorsal faces ( x 2). Hyolithes sylvestris. 9, dorsal face (x 24); 9a, portion of surface (x 5). Hyolithes (Orthotheca) subexcavatus. 10, dorsal tace; 10a, cross-section; 11, side view of ventral face (all x 24); 12, portion of surface of ventral face (x 5). Hyolithes (Orthotheca) subornatus. 13, ventral face ( x 24); 13a, portion of surface of ditto (x 5); 14, interior from dorsal side (x 24); 15, ditto (x 2). Hypolithes (Orthotheca) thraivensis. 16, side view; 17, dorsal face; 18, ditto; 19, ventral face; 20, side view of ventral face; 21, cross-section; 22, ventral face; 23, ditto; 23a, cross-section. (All x 2.) Hyolithes (Orthotheca) thraivensis, var. granuliferus. 24, impression of ventral face (x 23); 24a, portion of surface of ditto (x 5); 25, cast of same specimen (x 25). Ceratotheca ‘subuncata, Side views (x 2). Puate III. Ceratotheca % balelatchiensis. 1, side view; 2, ventral view; 3, ditto, showing apex. (All x 2.) Pterotheca drummuckensis. 4, cast; 5, impression showing internal plate. (Both x 2.) Pterotheca girvanensis. 6, showing internal plate ; 10, showing marginal projection of carina ; 11, internal cast ; 12, impression of part of exterior; 13, showing internal plate (all x 2). 6-10 from Balelatchie ; 11-13 from Ardmillan. Pterotheca multidecorata. 14, 15, 16 (all x 2); 15a, portion of surface ( x 5). . Pterotheca simplex. 17a, cross-section; 17, 17a (all x 23). Pierotheca mullochensis. (All x 2.) Vol. XLVIL. ” “ee vans .Roy. Soc.Edin. We PLATE MR F.R.COWPER REED ON LOWER PALAOZOIC HYOLITHIDZ FROM GiRVAN. J.Green del. lith.et imp. Vol. XLVII. ” “ MR F.R.COWPER REED ON LOWER PALA OZOIC HYOLITHIDA FROM GIRVAN cans. Roy. Soc .Edin. PLATE Ill. — a BR tg ac TR a eer) LET REXIET EM hous, UASeTee TAR Neer is J.Green del. lith.et imp. ae e Roy. Soc.Edin. Vol. XLVIL. | M® F.R.COWPER REED ON“LOWER PALAZOZOIC HYOLITHIDE FROM GIRVAN” PLATE III. J.Green del. hth.et imp. (ia22371.) X.—Report upon the Anatomy and Embryology of the Penguins collected by the Scottish National Antarctic Expedition, comprising: (1) Some Features in the Anatomy of the Penguin; (2) The Embryology of the Penguin: A Study in Embryonic Regression and Progression. By D. Waterston, M.A., M.D., F.R.C.S.E., and A. Campbell Geddes, M.D. (From the Laboratory of the Anatomical Department, ‘he University, Edinburgh.) (With Three Plates.) (MS. received March 11, 1909. Read February 3, 1908. Issued separately October 21, 1909.) Among the specimens collected by the Scottish National Antarctic Expedition and brought back to this country was a series of the embryos of penguins of various species, collected at different stages of development by Mr R. N. Rupmosr Brown and Dr J. H. Harvey Pirin. These embryos were, for the greater part, of the species Gentoo (Pygoscelis papua, Forst.), the remainder of the species Adelia (Pygoscelis adelia, Hombr. and Jacq.). Two Emperor penguins, which had been preserved for examination by the injection of the blood-vessels, were also brought home. Through the kind- ness of Dr W. 8. Bruce these specimens were entrusted to one of us (D. W.) for examination and report, but as the work involved in this proved to be very extensive, Dr A. CamMppELL GEDDES was asked to undertake a share in it, and this he agreed to do. This report contains the results of the combined investigations. PART f. ON THE ANATOMY OF THE ADULT PENGUIN. INTRODUCTORY. Upon the return of the Challenger Expedition to this country, the late Professor Morrison Watson had handed over to him the valuable collection of adult penguins which had been made, Upon these he based his classical memoir, ‘“‘A Report on the Anatomy of the Spheniscide collected during the Voyage of H.M.S. Challenger” (Challenger Reports, vol. vii. p.1). To that report little, if anything, can be added. There are, however, two points which a detailed dissection of the adult penguin has brought to light, which seem to us to be worthy of being placed on record. These are, first, the peculiarities of the cervical portion of the vertebral column ; second, some points in connection with the arrangement of the fascial layers. in relation to the pectoral muscles. In all the other points illustrated by our dissections we can merely corrobo- rate the description given by Professor Watson. TRANS. ROY. SOC. EDIN., VOL, XLVII. PART II. (NO. 10). 33 224 DR D. WATERSTON AND DR A. CAMPBELL GEDDES ON EMBRYO PENGUINS Section I.—-Some PECULIARITIES OF THE CERVICAL PORTION OF THE VERTEBRAL CoLUMN. The cervical vertebree are thirteen in number: the portion of the vertebral column which they make up is remarkable for the development of its antero-posterior curves. In the lower half there is an extraordinary antero-posterior curve the convexity of which is directed forward ; in the upper half there is a second antero-posterior curve the convexity of which is directed backward. These curves are present in all birds ; in none, however, do they approach in intensity the curvatures found in the penguin. The lower curvature is so pronounced that the bodies of the vertebrae come in contact with and fill up the angle formed by the limbs of the clavicle, and, in the living bird, actually cause a projection on the front of the neck. Asa result the trachea and cesophagus, instead of passing into the thorax in front of the vertebral column, are pushed away to the right side and actually le on a plane posterior to the vertebree of the convexity. The knowledge of this extraordinary condition is not new; it is fully described by Professor Watson, but the point seems to have been entirely overlooked, for in all the specimens in museums that we have had the opportunity of seeing the cervical vertebree are incorrectly mounted. Professor Morrison Watson was of opinion that these curvatures were associated with the maintenance by these birds when on land of the erect attitude, and that they served to bring the centre of gravity of the head and neck over the base of support formed by the feet. To us it appears more probable that the curvature is useful to the bird when in the water, giving it a greatly increased displacement forward and reducing to a minimum the fatigue of carrying the head and heavy beak in the long ocean voyages which it undertakes. These may apparently be very prolonged, for the birds have been seen no less than 600 miles from land. Section [I].—Somer PECULIARITIES IN THE ARRANGEMENT OF THE FascraL LAYERS IN RELATION To THE PECTORAL MUSCLES. The most striking feature with regard to the pectoral muscles is their enormous size and power. There are four muscles belonging to this group: first, the pectoralis major; second, the dermo-humeralis (muscle des parures) ; third, the pectoralis medius ; fourth, the pectoralis minor. ‘The origins and insertions of these muscles are given in detail and with great accuracy by Professor Morrison Warson. Their action is not without interest. The pectoralis major is divided into two distinct parts: an anterior which arises from the clavicle and from the outer surface of a strong aponeurosis which separates it from the medius and is inserted through a special tendon into the whole length of the anterior or radial margin of the bones of the wing, and by an aponeurotic extension from the tendon which covers both surfaces of the wing and conceals the blood-vessels and nerves, and a posterior which terminates in a V-shaped OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION. 225 tendon inserted into the humerus. These two portions have different actions; the anterior carries the wing forward to a position at right angles with the trunk, while the posterior draws the limb back in the effective part of the swimming stroke, and by its insertion into the anterior margin of the humerus it “ brings about that rotation of the wing round its long axis which, combined with the backward stroke, gives rise to the screw- like motion of the organ observable when the bird is progressing through the water.” * The nerve supply is provided for by a special branch from the large cord of the brachial plexus. The dermo-humeralis arises from a strong fascia which covers the external oblique muscle ; at the posterior margin of the sternum it is reinforced by a number of fibres arising from the subcutaneous tissue covering the knee joint. It passes forward parallel to the outer border of the pectoralis major to be inserted along with the posterior fibres of that muscle into the anterior margin of the humerus. Its action apparently is to co-operate with the external part of the pectoralis major in depressing the wing. It is supplied by a twig from the nerve to the pectoralis major. The pectoralis medius passes over a pulley formed by the bones of the shoulder and acts as the great elevator of the wing; by some authors it has been called levator humeri. It also is supplied by a twig from the nerve to the pectoralis major. The pectoralis minor is inserted into the outer margin of the tricipital fossa ; its chief action appears to be to rotate the humerus in a direction opposite to that of the rotation produced by the action of pectoralis major. In addition, however, it must help in producing the effective, propulsive stroke of the wing. Its nerve of supply is a twig from the nerve to pectoralis major. The arrangement of the fascia in regard to these muscles is striking and, peculiar. Tracing the deep fascia of the neck downwards, it passes deep to the clavicle, to which it is lightly attached; and, passing to the thoracic region, it lies superficial to the pectoralis medius and minor and becomes continuous with the strong fascia which covers the external oblique. Superficial to this fascial layer lie the pectoralis major and dermo-humeralis, and from it they both obtain a portion of their origin. The super- ficial aspects of pectoralis major and dermo-humeralis are in close contact with skin—so close, indeed, that they are marked by regular rows of pits which accommodate the rounded ends of the feather quills. From this arrangement it appears to us possible that the dermo-humeralis, pectoralis major, and clavicle are in reality cutaneous structures. That the dermo-humeralis is in series with the dorsal cutaneous muscle and panniculus carnosus is certain; that it is closely associated with the pectoralis major in position, in action, and nerve supply is also certain; but whether the pectoralis major can justly be associated with the same group is, on the present evidence, not certain, although it appears to us probable. Un- fortunately the evidence from nerve supply is so weakened by the association of pectoralis medius and minor with the same nerve as to be valueless. * Quoted from Professor Morrison WaArSON’s memoir. 226 DR D. WATERSTON AND DR A. CAMPBELL GEDDES ON EMBRYO PENGUINS The suggestion that the pectoralis major is in reality a cutaneous muscle seems to us to be not without interest in view of the occasional occurrence in man of the abnormal muscle, sternalis. Sir Wituiam TurNeER has suggested that it is to be regarded as a vestige of the panniculus carnosus. Professor D. J. CunnincHAM has pointed out that there is considerable evidence to show that it is formed by a devia- tion or dislocation of a portion of the pectoralis major; Mr F. G. Parsons has shown that in rodents the panniculus carnosus possesses two strata, and that there is evidence to show that in man the deeper stratum of the panniculus forms the fascial sheath over the external oblique and that possibly a portion of the sterno-mastoid is derived from the same stratum. THe has also shown that there are good reasons for regarding the pectoralis major as derived from the panniculus. To these observations we now add the facts of the anatomy of the penguin, which appear to us to suggest that the apparently antagonistic views of Sir Writ1aM TuRNER and Professor CunNINGHAM are in reality not opposed, but complementary. PART II. ON THE EMBRYOLOGY OF THE PENGUIN. INTRODUCTORY. As has been shown, the anatomy of the adult penguin has been previously very completely described, and little remains to be added to complete our knowledge of it. Of the embryology, on the other hand, the existing knowledge is incomplete, owing presumably to the great difficulty in bringing back the necessary material in a con- dition which allows of a detailed examination being made. A small number of embryos was collected by the Challenger Expedition, but the condition in which they were received by Professor Morrison Watson rendered them useless for description. The material which was put into our hands consisted of a number of specimens each one of which had been removed from its egg upon a different day of incubation. It would, therefore, appear that we had a specimen for each 24 hours of development. In one sense this was so, in another not. The method of collection which was of necessity adopted was as follows :—The nest was watched, and the eggs were marked with the date of their first appearance and were subsequently collected upon the desired day. The difficulties of so doing and the fallacies necessarily attendant upon it are obvious. First, it was quite impossible for the observer to know in which of the 24 hours preced- ing the marking the egg had been laid; secondly, it is known to be no uncommon thing for the males and females to fight for possession of their egg and for the privilege of incubating it. In the course of these struggles the eggs are apt to be dropped and to lie for some time directly upon the ice. The result of this must be to retard for a time the processes of development, and therefore, although an egg may have been laid for 3, OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION. 227 6, 9, or 12 days, it does not follow that the development has been proceeding for the same number. Be this as it may, we found it necessary, especially in dealing with the early specimens, to revise the sequence of specimens so as to avoid the absurdity of embryos of an obviously earlier stage of development being classified as older than specimens of a later stage. The number of days of incubation is, according to the series of specimens, thirty. This is surprisingly short, and as there are some obvious gaps in the series, we are not prepared to express an opinion as to whether it is correct or not. The embryos were preserved in 24 per cent. formalin, and we desire to place on record our sense of real obligation to the collectors for the care and trouble which they took, and for their great skill in packing the specimens, which alone is responsible for the excellent state of preservation in which many of these exquisitely delicate specimens were when put into our hands. As a result of the long time which elapsed between the date of collection and examination and of the prolonged immersion of the specimens in formalin, it has not been possible to carry out, satisfactorily, any very fine histological examination nor to determine with absolute accuracy the exact date of the first appearance of the primary ossific centres, the formalin, apparently, having exercised a slight decalcifying action. Our investigations have, therefore, of necessity been somewhat restricted, and many important problems in connection with the development of the penguin remain unsolved. In dealing with the embryology of the penguin the following points seemed to us important :— 1. The enormous length of time through which the penguin has persisted without undergoing important modification, In the Quart. Journ. Geol. Soc., vol. xv. p. 670, Professor Huxtey described a penguin of large size whose remains were obtained from the pliocene strata of New Zealand. For the reception of this bird he established the genus Palzeudyptes. It is exceedingly doubtful whether the establishment of a new genus was necessary, and Professor Morrison Watson in his report on the anatomy of the Spheniscide (Challenger Report, vol. vii. p. 46), expresses. the opinion that this pliocene bird might very well be regarded as belonging to a species of the existing genus Eudyptes. From the outset, therefore, we were prepared to meet with some peculiarities of development of a more primitive avian type than that obtaining in modern birds. 2. The enormously high specialisation undergone by the forelimb of the penguin in the course of which it has been transformed from an organ of erial to an organ of sub- aqueous flight. 3. The modification undergone by the cervical spine to secure sufficient water dis- placement anteriorly to carry the head and heavy beak without fatigue during the long ocean voyages which the penguins undertake. 4. The skeletal arrangements necessitated by the quasi-erect attitude. 228 DR D. WATERSTON AND DR A. CAMPBELL GEDDES ON EMBRYO PENGUINS Metuops or EXAMINATION. The earlier specimens were prepared for examination by being stained in borax carmine or alum carmine and mounted entire. Duplicate specimens for examination by section were not available. The later specimens were cleared so far as possible, but owing to the long immersion in formalin the clearing did not yield very satisfactory results. A figure illustrating the development of the limb bones is included among the illustrations (fig. 26). The decalcifying action of the formalin has, unfortunately, made any definite statements as to date of appearance and order of appearance of the ossific centres impossible, and we have had to content ourselves with the negative observation that no unusual or abnormal processes or sequences were to be observed, although such were looked for as carefully as the material available would permit. In order to carry out a comparison regarding the dates of.acquisition of the external adult features and the progressive development of the limbs and trunk during the later stages of development, we found it necessary to provide a control by carrying out observations on the same lines upon the embryo of another genus of birds. Data for this purpose were not available in the literature, and we therefore prepared and examined, by methods identical with those used for the penguins, a series of duck embryos at every 24 hours of incubation. The period of incubation in the duck (28— 30 days) approximates sufficiently closely to that of the penguin (30-32 days %) to enable comparisons to be drawn with accuracy between embryos of corresponding age taken at each 24 hours. The data which we obtained appear to be new, and we have, therefore, included an account of the external form and the measurements of the embryo-duck with those of the penguins. Section I].—Haruy Sraces oF DEVELOPMENT. EMBRYOS SHOWING EARLY STAGES, COMPRISING SPECIMENS UP TO THE TWELFTH DAY. Specimen I.—In the first of these, which was the youngest specimen examined, there was an oval area pellucida measuring 3 mm. in length on the surface of the blastoderm. This specimen illustrates the earliest stage after the close of segmentation before the formation of the primitive streak, and it closely resembles in size and shape the corresponding area in the ovum of the chick and the duck. No area opaca could be made out, nor was there any sign of the embryonic shield. The shape of the area pellucida is comparable to that seen in the blastoderm of the chick at about 18 hours, but in the absence of primitive streak and embryonic shield it resembles the chick blastoderm at the commencement of segmentation. Specimen IT. (fig. 1).—The second specimen showed a more advanced stage. The germinal area was somewhat pear-shaped. At its wider end was a narrow, semi- OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION. 229 circular, deeply marked line running round the periphery of the clear area, This line appeared to be the commencement of the amniotic fold. Behind this line was a narrow area in which there were a few dark spots, which marked apparently an early stage of the formation of a vascular area. ‘his area, in turn, merged into a crescentic clear area which was limited on the opposite side by a short crescentic dark line parallel to the first line. This line occupied the central part of the wider end of the clear area, and lay across it, and it appeared to be the rudiment of a commencing head fold. From the concavity of this fold a darker streak passed for a short distance backwards towards the narrow part of the clear area, and this structure appeared to us to be an indefinite stage in the formation of the primitive streak. Specomen ITT. (fig. 2).—This specimen resembled that of the chick of 22 hours. The head fold was well formed, and behind it lay the medullary folds. These diverged from one another at their posterior extremity, and behind them lay the remains of the primitive streak stretching to the posterior end of the clear germinal area. The primitive streak had the form of two narrow dark lines, enclosing between them a narrow Clear streak. Specomen IV. (fig. 3).—Labelled as 3 days.—The embryo was now 3 mm. in length and presented well-formed medullary folds, a notochord, seven pairs of somites, and also the remains of the primitive streak which was separated by a narrow clear area from the posterior end of the notochord and extended for a distance of 1 mm. behind it. The appearances were very similar to those of the chick at 25 hours. The medullary folds were as yet ununited, but showed at the cephalic end evidence of distinct thickening, while at the posterior ends they diverged from one another in a V-shaped manner. The notochord could be traced as far forward as the cephalic enlargements (but this part of the embryo was rather damaged), while posteriorly it terminated in a club-shaped enlargement. Specumen V. (fig. 4).—The succeeding specimen, labelled as being 5 days, showed a slightly more advanced stage of the same condition, closely similar to the chick at 26 hours. The embryo measured 3°5 mm. in length, and nine pairs of somites were present. The posterior end of the notochord was enlarged and shaped like an Indian club, and extended beyond the termination of the medullary folds, which diverged outwards on either side of it. Behind this extremity was a clear area, and behind it again lay two short parallel dark lines, the representatives of the primitive streak. The cephalic ends of the medullary folds were considerably enlarged, to form the cerebral vesicles. These vesicles were not distinctly marked off from one another, but the anterior part was bent forwards, to form the earliest cephalic flexure. In front of this again was a narrow clear crescentic area—the pro-amniotic region. Specomen VI. (fig. 5).—Labelled 6 days.—The next specimen was somewhat larger, measuring 6 mm. in length. Twelve pairs of somites were present, and the neural folds were larger, and had apparently united with one another at the fourth 230 DR D. WATERSTON AND DR A. CAMPBELL GEDDES ON EMBRYO PENGUINS somite. The folds approached one another closely at the posterior end, and in that region they enclosed a clear area, shaped like an arrow head, in which lay the expanded posterior end of the notochord. The end of the notochord projected backwards beyond the medullary folds in the form of a bulbous extremity. Behind it there was no distinct evidence of a primitive streak. The whole of the posterior portion was enclosed in a clear oval area. The somites were well formed, and their internal structure showed them to consist of a clear central portion and a periphery or cortex arranged in columns radiating outwards. At the outer margin of the somites was a clear area in which lay a narrow darker line, extending along the side from the third anterior somite backwards to beyond the hinder somites. In position this structure corresponds to the primitive cardinal vein. The neural folds showed evidence of segmentation, being constricted opposite the centre of the somites, and they were united about the middle. The specimen described was one said to be of the sixth day, and the subsequent specimens, which showed stages of gradual advance, were in harmony with this statement. If these dates be accepted, it is obvious that the early changes in the penguin embryo are very much slower than the corresponding changes in the chick, for the same degree of development is shown in the chick of 40—44 hours. Specimens VII., VIII, IX., and X. (fig. 6).—Labelled 7-11 days.—These specimens showed that the next stages of development are very similar to corresponding stages in the development of the chick, and do not require detailed description, except in so far as they show differences from it. The head and trunk show gradual increase in length and in thickness. At the hinder end of the notochord a dark area shaped like a spear-head persists for some time, but eventually becomes merged in the hinder end of the trunk. Figs. 7, 8, and 9, which illustrate the development of the duck at 5, 6, and 7 days, are introduced for comparison with fig. 6, which is a photograph of a penguin embryo labelled as 8 days. Section I].—Rate oF GRowTH OF THE TRUNK AND LIMBS FROM THE TWELFTH Day ONWARDS. Our observations were directed first to an examination and measurement of the developing trunk and limbs. The observations begin with the twelfth day of incubation, which is the date of appearance of the limb buds upon the surface of the body. The details of the measurements are recorded in the appended table (Table No. L.), OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION, 231 NovTes upon AND CoMPARISON OF THE MEASUREMENTS. The measurements were analysed along several different lines. 1. General Growth of the Trunk. A, Penguin.—The vertex to coccyx length of the specimen of 12 days (fig. 10), taken as it lay in the egg, 7.e. in a flexed position, was 15°5 mm., while that of the duck of corresponding age was practically twice as great, namely, 30°6 mm. In the penguin the rate of growth from this date onwards was fairly uniform and progressive, and the largest embryo, that of 30 days, measured 61°2 mm. in the same position. The specimen labelled as being 24 days was larger than those of 25 and 26 days, and among the other specimens similar anomalies were present. Taking the maximum measurements of specimens which seemed to be typical in their rate of growth, we found that the length attained at 12 days was doubled at 19 days and doubled again at 27 days, the progression being thus almost in geometric ratio. The same ratio is found to exist when measurements of the breadth of the specimen are taken. The maximum breadth attained at 12 days was rather more than doubled at 19 days, and rather less than doubled again at 27 days. B. Duck.—The maximum length of the embryos of this species was uniformly greater than that of the penguin of same age. Even the largest fully developed penguin embryo was smaller than the largest duck embryo. At 12 days (fig. 20) the duck embryo was almost exactly twice as long as that of the penguin of corresponding age. The rate of growth from this date onwards, as ascertained by the same criteria as in the case of the penguin, was, however, much slower than in that creature. | At 19 days the length had increased only by rather less than one-half, 7.c., was 4°36 cms. as against 3°06 ems., and not until the 23rd day of incubation was the length doubled, and from this time until the end of incubation the increase again was rather more than half, from 43°6 mm. to 76°38 mm. The breadth figures showed a rate of growth very similar to that found in the penguin. The breadth at 12 days was 10 mm., and this figure was practically doubled at the 19th day and almost doubled again at the end of incubation. 2. Rate of Growth of Different Segments of the Trunk. Pengwin.—Analysing the rate of growth of the different segments which make up the vertex to tail measurement, the head segment is found to increase slowly at first, but grows rapidly in the last few days, while the neck increases rapidly through all stages. The growth of the trunk and tail follows the general rate of the whole body, the length attained at 12 days being doubled at 19 days, and doubled again at 27-28 days. The figures for the head are modified by the fact that measurements were taken TRANS. ROY. SOC, EDIN., VOL, XLVII. PART II. (NO. 10). 34 » 232 DR D. WATERSTON AND DR A. CAMPBELL GEDDES ON EMBRYO PENGUINS TABLE Adelia. Days 12 13 14 15 16 17 18 18 P| By eaten ie DP) a ape: aD | P D: | a D:; | Pe | Vertex to coceyx length, flexed . | 1-55 | 3-06 | 1-94 | 3:36 | 1°03 | 3-32 | 1-94 | 4-07 | 2°42 | 3-95 | 2°58 | 4 3°35 | ... | 2°86 | 445 Maximum breadth . . -| 64 | £28 PAS | gor | 89.) 76 | *91 | tH | 4-09 | 7S | 1-53 | a's | 1-46) |) ieee Length of head. : : . -| 76 | 2°47.) <84 | 2:24") 1:01 | 2:3 94 | 7-34 | 1°05 | 2°28 | 1-49 | 2-6 | 1-25 | |) eon een » mek . . « .- +) 48 | 24% | 156 | 65 | “68 | t5 | ea | £24.) 67 | 2-82 | “82 | 7-85 | 1-06) (an) Pose » trunk, including tail. -| 1:32 | 2-22 | 1:44 | 2:65 | 1:41 | 8:09 | 1:35 | 3:55 | 2:01 | 3°74 | 2°18 | 3:82 | 2-71 | ... | 2°42 | 4-45 Cloaca to furcalangle . . =. = «| 78 | 1:8 | 1°08 | 1:82 | 114 | 1:92 | -98 | 2-22 |} 1:36 | 25 | 1:44 | ays |192 | ... | 1:65 | seg (Tip of beak to occiput . .| +78 | 2-05 | 1-21 | 2-88 | 1:29 | 26 | -98 | 2°63 | 1-68 | 2°86 | 1°93 | 3-96 | 2-28 | ... | 2-06 | 3:28 | » sy toear - 6) +] 52 | 299 | °7 | W82 | 82 eog | “7 | Bt || 1:8 | 28s | 4B | 2-8) Ire » »» topostangleofeye .) -63 | 1-44 | ‘86 | 1-64 | 84 | 2-7 | +74 | 187 | -97 | 1°97 | 1-33 | 2 1:68). so. | 1°3 Soe Head{ ,, ,, to mouth Sy 2 | a) 2) | ee ed "El £2 | Ue78i | Dae Ae 208 781 78 »» 93 to nostril f Git. 24a bee ‘6 x oy x 64 x TR Nt 8 x x “87 ome between eyes. «| 64 | 1:23 11:5 | 1:84 | 89 | 2:85 | +91 | 185 | 1-09 | 1°85 | 1°33 | 1°45 | 1-46 1°35 | 164 go COTS) ET RP EB | RZ) BEBE Aero | BRAD SE gr ei07 | MGRy tel Mom 1-04 | 15 ae ee a a MB eg) BE | res. | GeB ren) “eB, eB tare es een) Zee ier 1:28 | 1°74 Width ee =) SUPE SB eee) ae Pepe age: ee Bat ESP tay ars cra ere Me 53 Upper arm, anterior border a Cita 44 | 19 “4h | “28 “46 18 5 BY 56 "85 oo) "b4 ate 35 | “64 iz yg, | posterior y,° “my, fe TT) re 4 ei py aa") eZ eon! 7 og ey ee 28 | 76 Forearm, anterior border . af 2G Te eae St Sega s edpi ll eaaryr ad 6 |t79 | 6 | +91 | «79 Heat |... feto4 | °78 | » posterior ,, we Hand ‘67 | 58 | +6 | +52 | 62 | $48 | “82 | 84 | “78 | 14 | 8 | 1-95) | eee ro | aa | limb | Blbow, width... 13.) 81 | -16.| ye) 17 | 88 |, 16 | 68 | 24) 87 | 26°) oe |) 28 | een | » thickness - - + |, 08. | “29'| 09 | <25 | -07 | -22 | 08 |) 26 | -o9 | “22| 09°) ‘27 |.0°9 | oom ae of hand es Gee en ee ee le ee aero mins | Width ap cn, Coyle Pel ete | cen |) B00) | otal ae lade Ol eee) lune Wh eet or \ Length of pollex Sr eee 8 | ore eg eres | Pen eel Mee er ii] gee Leg, anteriorborder . 8. =. 2. | $°54 | 1:04 | 3-72 | -98 | $-78 | 2-05 | $61 | 2-08 | “57 | 228) +55 | 1-36 | 77 | eee »» posterior ,, SD eee BBL eee YR) vee | GE wae! [SPH | V1) “82 18.) 86 | 28 | ere cae to ond of third) ... | 226 | 16 | 2-46 |. | 062 | 4 | £65 | 754 | 2-97,) -69 | 2:08 | -91 | .. ene | Plantar, length of phalanges «| .., | 65). | 65 | .. | "78 | «. | 76) 28 | 96 | 28) 225 | -45 | 5 Foot J Plates ee between outer and] .., | ... - Cally aa yal... 47 | 18 | 3 wi o7 19 17.| +96 gee p web between inner and] .,,, ee sa ane 2 "4D ike ‘4 12) 5644) STA 65a) ate "12 || sada Dorsal, aukle to tip of nail of). ...)| 204 || tacdeaw | cbs. deemed) Army tea)! 0d) a8" || (eed) ero eae 78 | 2:25 \ Dorsal, length of nail P pd iene ida ies ies a <3 wes a e- LG Al aces 28) 1 09 | ‘28 ” Over eyes, t Elbow to tip. + Knee to tip. al OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION. 238 20 21 22 23 24 25 26 27 28 2 ng | aS eS ee ee eee D P D P D P. Ds-|, .P. De D P D P D P D 6:25 | 3°62 | 5:93 | 3°49 | 5:05 | 4:09 | 6:39 | 4°64 | 7:04 | 4:13 | 6-06 | 3:99 | 7-68 | 6:07 5-71 2-73 |1°75 | 2:95 | 168 | 2:8 | 1:89 | 3-22 | 2°24 | 3:35 | 2°21 | 38 | 218 | 4-46 | 2°35 3°05 LE | 1°85 | 2-48 | 1-24 | 1-55 | 1°48 | 1-97 | 1-47 | 2-48 | 154 | 2:85 | 1°55 | 1°95 | 1°91 2°55 1:94 | 1:28 |-1-:95 | 1-4 | 2°15 | 1°66 | 2-34 | 1°63 | 2°62 | 1°58 | 2:93 | 1:39 | 2-88 | 1:92 1°85 5:25 | 8:47 | 5-01 | 3°38 | 5:4 | 4°06 | 6:02 | 4°16 | 6 3°55 | 5:88 | 3:91 | 6:32 | 4°91 551 4:16 | 3°66 | 3:46 | 2°52 | 3°23 | 3°21 | 4:56 | 3°41 | 447 | 2°81 | FOL | 3°38 | 44 | 4°15 3:45 | 277 | 663 | 2°69 | 364 | 21 | 3-9 | 82 | 4:02 | 3:12 | £01 | 319 | 44 | 3°48 3°98 2-78 | 218 | 2:73 | 2:08 | 2:75 | 2°26 | 2:9 | 2°57 | 2:95 | 2-48 | 3 2°48 | 3:27 | 2°72 3°07 23 | 1°79 | 2:43 | 1:71 | 225 | 1°85 | 26 | 1:98 | 2-47 | 1:97 | 2°56 | 1:95 | 2:87 | 2°07 2°39 1-48 | 1:23 | 1:42 | 128 | 143 | 129 | 7:58 | 15 | 26 | 1-44 | 166 | 1°38 | 1-82 | 1°66 1:91 in| 43) || =978)|| 52) <9 By) 940) 58") 2 Bee pecOs e825 | 359 7 remcegul arse Wi i72)) 15S (1-7 9) t6 i 1°88 |) 7°77 |) 8. | 2:68 | 1-82 | Leg | 1:87 2°02 Pepe dalbn| 2-45) 1-13 | £48 | 1-27 | foe) 1-54 | 766 | 1-42, | 154 | 1°32 | 1:78 | 1-69 1°83 1-94 | 1:28 | 1:95 | 1:4 | 2-25 | 1°66 | 2:34 | 1°63 | 2:62 | 1:58 | 2:93 | 1:39 | 2:88 | 1°93 1°85 1 | a8) |) a2 63 | 1 8 | 1:08 | 1:09 | 1:05 | -88 | Z 91 02eh 31-93 1:25 it 6£ 6S). 465 |) <5: We) bee Seeley WASBGE | 258i) 564 4) 7 Uy iN 9 77 Sule E46. | es6e: Ae | SS 0) az | 7d yh No; BB te7 We P72) 23 4, 58 81 ‘OY 152 | 98 1-45 | 64 165 | Lg 187 | 9 | +78 | 1:08 |t1-73 | 118 ft2-17 |... 68 1°23 | 159 | 1:07 |t1°51 | “96 |t1:72 | 1:07 |+2:07 | 2-2 |+1°84 | 1:23 HH1°93 | 2-32 [42°18 | ... | 1°01 6 ‘4 Coole Ata erate) aR oer ce ES. VbRe | Se 1 be eho 64 || vas ‘76 2g acon eso 12nlr 25a Gn 26s lel4 |b Be) 27 1-29 i a5 89) 99 || 23 LS ou Wee | eee e2 ano || LMS | pos |) PON coe | A Gos RIA oes soo | LS 4a 4 37 “43 52 34 52 43 5 21 ‘56 73 64 64 Seal Oiles | 2015 1-07 CSam ee eo | On 2S. Vida) 12-9601 157. | S08 | 1-61. | .... | 2°18 154 | 37 | 14 82 | 1:68 || <48 | 1:55 | °46 | 2:22 | -46 | 7-88 | -78 | 2:74 | 66 | ... ‘73 3°56 | 1°31 | 3:24 | 115 | 3:42 | 1°25 | 4-2 | 1°73 | 3:93 | 1:5 | 4:28 | 1°56 | 462 | 1°85 | ... | 2°32 19 ei menOuemoom|) 561 Wetor | “80 | 175 1 e=66 e203) 62 | B08 f -034. ot) 1°00 ee SAI etePS ana 25) | “84 | 1°56 | "55 | 1:64 || 47 | g62)| ba | 27, 86514 Tsu ‘81 Lita oer tems 125 || 23) 146) Bl | 76% | 28.) 252) 42 |2:72) 41. | 53 29 | 1°15 | 3-07 | 1:08 | 3-23 | 1:34 | 3:62 | 1°58 | 3-64 | 1°47 | 3-62 | 1°32 | 42 | 1°79 | ... | 2°02 “Soa eelbn | Sel 19a) 85 25 | “88 | 29 | we) «26 | gs | <29 | 7 | 39) ... “46 * Over eyes. + Elbow to tip. t Knee to tip. 234 DR D. WATERSTON AND DR A. CAMPBELL GEDDES ON EMBRYO PENGUINS from the tip of the beak, and consequently the increase in total length is found to be greatest in the last few days, when the beak elongates rapidly, while the figures obtained for the growth of the head in width show a more gradual and regular increase, comparable to the rate of growth of the trunk. 3. Growth of the Inmbs. A, Penguin.—Fore Limbs.—The first appearance of the fore limb had the form of a bud-like projection from the lateral aspect of the trunk (fig. 10). This process at first was somewhat spatulate, and then rather bulbous at its free end (figs. 11 and 18). This shape was in turn soon lost, and the limb rudiment assumed the characteristic sickle-shaped form found in the adult bird (figs. 15 and 17, 19 and 23). The two seoments of the arm were soon clearly to be recognised and were separately measured. The measurements taken were those of the anterior border of the upper segment, together with the anterior border of the forearm from the elbow to the tip. At 12 days the total length was 4 mm., made up of 1'1 mm. of upper segment and 2°9 mm. of the lower. At 19 days the length was 13°9 mm. and at 27 days 29°7 mm. ‘The rate of growth, therefore, had been such that the length of the limb was more than trebled in the first period, and was more than doubled in the second. Both segments shared the growth, and did so in fairly equal proportions throughout. Lower Iimb.—The portion of the lower limb which was free from the coverings of the trunk was measured in one or in two pieces, as the size permitted. At 12 days it measured 5°4 mm., at 19 days 18°1 mm., and at 27 days 36°2 mm. In the first period the length was more than trebled, and in the second it was again doubled, showing a rate of growth almost exactly similar to that found in the upper limb. B. Duck.—In the duck corresponding measurements showed that at 12 days (tig. 20) the fore limb measured 15°3 mm. and was nearly four times as long as in the penguin. At 19 days the length was almost doubled (28°5 mm.), and at the end of incubation it had only added one-third to its length (37°5 mm.). The lower limb, measured in a similar way at 12 days, was 22 mm. in length, again four times as long as in the penguin, and in the specimen of 20 days it was more than doubled, and at the end of incubation it was almost doubled again. If the proportion which the length of the limbs bears to the total length of the body be examined, it is found that in the penguin the fore limb at 12 days is almost one-fourth of the body length, that it increases rapidly for a few days until it measures nearly one-half of the length of the body, and maintains approximately this ratio tiil near the end of the incubation period, when it progresses rapidly and assumes a ratio of rather more than two-thirds of the body length. In the duck the ratio at 22 days was one-half, and this proportion became rather larger, but eventually again became nearly one-half. OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION. 235 Tasie I].—Vertex to Coccyx Lenetru, THE LenctH or Limps AND THE RATIO BETWEEN THEM, THE FORMER BEING TAKEN As l. Length of Lower eee Tener eae yadon: Sant Index, Days. Penguin. | Duck. | Penguin, | Duck. | Penguin, Duck, | Penguin. | Duck. Penguin. Duck. 12 1:55 3°06 ‘40 1538 25, 50 “54 2°20 “34 ‘71 13 1-94 3°36 60 L54 “30 “45 ‘72 243 ‘on, 72 14 1:93 33k 68 £79 "35 53 ‘73 257 37 an 15 1:94 407 D9 2 30 “49 “61 2-68 “31 65 16 2°42 3°95 1:16 2°37 “47 60 1-11 3°20 “45 80 17 2°58 4 1:26 2:58 “48 64 1:26 343 “48 85 18 2°85 445 1-75 262 “61 D8 171 412 ‘60 92 19 | 2-99 436 1-39 2°95 “46 “67 1°81 363 60 83 20 3°37 6°25 1°85 S00 BA Roe 1°84 5°36 54 85 21 3°62 5:93 2°16 301 D9 D0 2°42 5°39 66 ‘90 22 3°49 5°05 2-10 224 60 44 = |* 2°38 572 68 113 23 4:09 6:39 2°42 2°38 “59 52 2°62 6°73 63 1:05 24 4°64 7 OL 2°98 3-21 64 “45 3°52 V73 ‘76 1:09 25 4°13 6:06 te 347 aoe ‘DT sor 719 sia 1:18 26 SFE) 768 2°45 B75 “61 “48 3°21 779 80 101 27 6:07 nh 2°97 wil “48 530 3°62 ee 59 ae 28 571 aoe 3°53 61 bt 4:71 500 "82 30 6°12 4-24 69 4:79 ‘78 Synopsis oF RESULTS FROM COMPARISON OF THE MEASUREMENTS OF THE PENGUIN AND Dock. 1. Length of Trunk. At early stages the length of the trunk of the penguin is much less than that of the duck of corresponding age—the proportion being that the penguin at the 12th day is almost exactly one-half of the length of the duck at the same age. As development proceeds the penguin grows more rapidly, until at the end of incubation the penguin is only fractionally shorter than the duck. 2. Length of Fore Inmb. At corresponding dates after the limbs first appear the fore limb of the penguin is only about one-fourth of the length of that of the duck. Its growth subsequently is much more rapid, and at the three weeks it is almost the same size, and at the end of incubation it exceeds in length the duck’s fore limb, in some specimens by as much as one-quarter. 3. Lhe Ratio of Length of Upper Limb to Trunk. At the 12th day of incubation the fore limb of the penguin measures approxi- mately in length one-fourth (index, *25) of the length of the trunk, while in the duck 236 DR D. WATERSTON AND DR A. CAMPBELL GEDDES ON EMBRYO PENGUINS the limb is equal to one-half of the length of the trunk (index, °50). The ratio of growth of the fore limb in the penguin is continuous and progressive in relation to the ratio of growth of the trunk. Towards the end of the third week it measures one-half of the length of the trunk (index, 21st day, *50), and at the end of incubation it measures two-thirds (index, °69) of the length of the trunk. In the duck, on the other hand, the length at first increases relatively to the trunk, attaining in one specimen, at the end of the third week, a maximum ratio of over two- thirds (index, 19th day, °67); but thereafter it slowly loses ground, and at the close of incubation the fore limb is less than one-half the length of the trunk. 4, Hind Inmb. A. Absolute Length.—In the penguin at the 12th day of incubation the hind limb is also one-fourth of the length of the hind limb of the duck, the ratio being very similar to that between the upper limbs of the two animals. In the penguin’s limb, however, growth is more rapid, with the result that at the end of the third week the hind limb has reached a length equal to nearly one-half of the length of the hind limb of the duck of corresponding age, and at the end of incubation it is rather more than half as long as the corresponding part in the duck. 5. Ratio of Length of Lower Inmb to that of the Trunk. At the 12th day in the penguin the length of the lower limb is approximately one-third of the length of the trunk (index, °34), but the rate of growth is more rapid, so that at the end of incubation it is nearly as long as the trunk (index, 28th day, °82). In the duck the limb at 12 days is rather more than two-thirds of the length of the trunk (index, ‘71), at the 18th day it is nearly as long as the trunk (index, °92), and thereafter it continues to increase at a rate relatively rather greater than the trunk, so that at the end of the third week it becomes actually longer (index, 22nd day, 1°13), and retains this superiority to the end of incubation. In this respect, @.e. in the rate of growth from the 18th day, it is the exact reverse of the fore limb—it continues to increase in length relatively to the trunk, whereas the fore limb actually shows a relative decrease. ANALYSIS OF RESULTS. Among the four limbs examined there are, therefore, two types of development. First, there is a progressive type, and to this the development of the fore and hind limbs of the penguin and of the hind limb of the duck belongs. Second, there is a partially regressive type, and to this the development of the fore limb of the duck belongs. OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION. 237 Now, if it be true that ontogeny is an abbreviated recapitulation of phylogeny we are forced to conclude that the fore limb of the penguin is a progressive structure, the fore limb of the duck, partially at least, a regressive. The reason for this, doubtless, is that the mesoblastic portion of the duck’s wing has, with the advancing specialisation and adaptation of the epiblastic structures, become less valuable. To all flying birds feathers, not bone and muscle, are of prime importance in securing wing spread for aerial flight: for the penguin, on the other hand, a rigid bony paddle has been evolved, adapted to resist the pressure of the water to which it is subjected when the bird’s great pectoral muscles are in strenuous action. And we thus are led to the interesting conclusions :— 1. That the mesoblastic structures in the penguin’s epiblastic regressive. 2. That the mesoblastic structures in the duck’s wing are regressive, the epiblastic progressive : for in the development of the duck we find that the developmental energy is suddenly on the 17th day switched off, as it were, from the mesoblastic structures on to the epiblastic ; for from that day onward the down rapidly develops and the meso- blastic framework loses ground. _ With regard to these developmental facts the question arises :— Is the duck’s or the penguin’s wing the more direct descendant of the common ancestor ; or have they both diverged from the common stock approximately equally, but in opposite directions ? Embryology alone cannot answer this question, but the evidence is clear in this, that the fore limb of the penguin in its development goes through a progressive and continuous series of stages along one unbroken line. The mesoblastic portion of the fore limb elongates, but its characters do not alter. It elongates, however, with a relatively greater rapidity towards the end of development, whereas the duck’s fore limb, after being relatively longer than the penguin’s ever is, regresses rapidly. So that the answer to our question, so far as the embryological evidence is concerned, must be that the wings of both these birds are different from the ancestral wing, and that the differentiation has been in opposite directions and that the common ancestor was a flying bird of a somewhat primitive type depending in large measure for the spread of its wing upon bone and muscle. ‘¢ Rorsra}, : wing’ are progressive, the Section II[].—ExTernaL ConFIGURATION OF THE EMBRYOS FROM THE TwetrrH Day ONWARDS. 1. Gentoo Penguin Embryo, 12 Days. (Fig. 10.) The embryo was not fully formed, the brain was covered by the thinnest of mem- branes, the beak was short and soft, the eyes were very prominent, with dense white opaque pupils. The fore limb well developed ; the elbow flexure completely marked, 238 DR D. WATERSTON AND DR A. CAMPBELL GEDDES ON EMBRYO PENGUINS It ended in a somewhat spatula-shaped extremity. The hind limb formed a spatula- shaped protrusion. Duck Embryo, 12 Days. (Figs. 16 and 20.) The embryo was fully developed, the integument was strongly marked, and there were down papillze on two short lines at either side of the tail. The eye prominences were very marked; the head and beak were soft and pliable except at the point of the beak, where there was a small white nodule of keratin. The fore limb showed distinct division into upper arm, forearm and hand. The pollex was prominent. 2. Gentoo Penguin Embryo, 13 Days. (Fig. 11.) The embryo was not fully formed. The head was soft and the brain showed through it. The beak was short, the eyes were prominent, the fore limbs were short, but showed plainly the elbow joint, and had a flattened, spade-like tip: to the naked eye there was no sign of digitation. The hind limbs were short, the ankle was unformed, the end was spatular and showed signs of three digits, the tail was relatively long and was much curved ventrally. Duck Embryo, 18 Days. (Fig. 18.) The embryo was fully formed. Down was appearing in lines on the back and sides of the embryo, and laterally on the neck ; otherwise as for 12 days. 3. Gentoo Penguin Embryo, 14 Days. The embryo was not fully formed, the head was soft, the brain showing through it. The beak was short, the eyes prominent; the fore limbs short, but well formed; the hind limbs were short, spatular protrusions. The heart showed distinctly, shiming through the thin anterior wall of the thorax. The tail was relatively long and much curved ventrally. Duck Embryo, 14 Days. (Fig. 21.) Down papillee were seen all over the embryo, especially along the dorsal tracts and on the tail. The eyes were widely open. 4. Gentoo Pengun Embryo, 15 Days. (Fig. 13.) The embryo was small, the beak was beginning to grow forward. The fore limbs were small and soft, but fully formed. The hind limb showed no division into the toes, but ended in flat, spatular extremities. There was no sign of the fourth toe independent of the flattened extremity. The tail was relatively long and curved ventrally. Duck Embryo, 15 Days. (Fig. 22.) s Down was appearing upon the head along a well-defined tract which looped round the eye from behind, | OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION. 239 5. Gentoo Penguin Embryo, 16 Days. (Fig. 14.) The embryo was small, the nails were not developed, so far as could be seen on naked-eye examination. There was a well-marked ridge down the centre of the abdomen and thorax. The toes did not appear beyond the web, but looked more as if they were embedded in it. The fourth toe appeared as if the web were continued round it. Duck Embryo, 16 Days. As for 15 days. Down more marked. 6. Gentoo Penguin Embryo, 17 Days. (Fig. 15.) The embryo was in good condition and the fully formed integument was complete. The skin was marked by papillee regularly arranged. On either side of the tail there were several down fibres varying from 1°7 mm. in length to a length just visible to the naked eye. Duck Embryo, 17 Days. The down was well developed. 7. Gentoo Penguwn Hmbryo, 18 Days. (Fig. 19.) This specimen was somewhat macerated. Duck Embryo, 18 Days. Embryo completely down-covered. 8. Adela Penguin Embryo, 19 Days. The embryo was fully formed, the integument was complete. There was a well- marked median groove over the abdomen and thorax, and from it on either side extended twelve transverse grooves. The body was completely naked except for eight fibres on the tail (four on either side of the median line). 9. Gentoo Penguin Embryo, 19 Days. The embryo was fully formed, and the integumentary covering was complete. There were no transverse grooves on the abdomen. 10. Gentoo Penguin Embryo, 20 Days. The embryo was fully flexed and completely devoid of down. The integument over the abdomen was not quite complete. A bare area in length ‘28 em. and in breadth at base ‘07 em. extended forward in front of the umbilicus. There were no transverse grooves*on the abdomen. TRANS. ROY. SOC. EDIN., VOL. XLVII. PART IT. (NO. 10). 35 240 DR D. WATERSTON AND DR A. CAMPBELL GEDDES ON EMBRYO PENGUINS 11. Gentoo Penguin Embryo, 21 Days. The embryo was completely flexed; the limbs were completely formed. The integument over the abdomen was incomplete; a triangular area 1'23 cms. in length, ‘07 em. in breadth, extended forward in front of the umbilicus and was continued over the front of the thorax as a well-marked groove from which on either side extended twelve transverse grooves. 12. Gentoo Pengun Embryo, 22 Days. (Fig. 17.) The limbs of the embryo were complete; the integument over the abdominal regions was incomplete. An area 1:04 cms. in length and ‘09 cm. in breadth at the umbilicus, gradually tapering to a point, was undeveloped. The head and body were entirely naked. The eye prominences were very marked; the skull and beak were soft and pliable. There were five transverse grooves on the abdomen. Flexion was incomplete. 13. Gentoo Pengun Embryo, 23 Days. The embryo was not quite fully developed, the integument being incomplete. On the anterior abdominal region—anterior to the umbilicus—a very acute isosceles triangle persisted uncovered, the sides of which measured 1°19 cms., the base *12 em. The eye prominences were very marked, the skull and beak were soft and pliable. A few short pieces of dark down had developed on the head; the rest of the body was absolutely naked. There were no transverse grooves on the abdomen; flexion was incomplete. 14. Gentoo Penguin Embryo, 24 Days. (Fig. 23.) The embryo was fully developed. The prominence of the eye-balls was well marked and the eye-slit was open, 18 ems. long; its width at its broadest part when stretched was ‘08cm. The head was covered with dark down, and on the back and body there was a sparse covering of light-coloured down considerably more than on the 25-days’ embryo. The middle line of the front of the abdomen showed a deep groove from which extended across the abdomen, on either side, six well-marked transverse grooves. The feet and tail had now assumed the adult relations, and formed the peculiar flat base which is characteristic of the adult bird and permits of its adopting the quasi-erect attitude habitual to it. 15. Gentoo Pengun Embryo, 25 Days. (Fig. 24.) The embryo was fully formed. The prominence of the eye-balls was well marked. The eye-slit was small, the beak was soft, and the head was sparsely covered with short black down on the back. The body was fully flexed ; the spine formed a semicircle from tail to head. There was a well-marked median groove on the front of the abdomen and thorax, from which extended transversely on either side eight well-marked grooves. OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION. 241 16. Gentoo Penguin Embryo, 26 Days. The embryo was fully formed ; the eyes looked large and prominent ; the beak was soft. The head was covered with short black down ; the rest of the body was practically naked. Along the middle line of the abdomen, from the umbilicus to the root of the neck, was a well-marked groove. 17. Gentoo Penguin Embryo, 27 Days. (Fig. 25.) The limbs of the embryo appeared fully formed. The feet were flexed over the front of the abdomen, plantar surface outwards. The upper limbs lay close to the side and over the abdomen under cover of the feet. The neck was flexed so that the beak lay along the right side of the abdomen immediately internal to the right superior extremity. Down was plentifully present on the head and back, shading from almost black over the head to a light slate grey over the sacral region. The down on the abdomen and anterior aspect of the thorax was sparse and light in colour. The middle line of the body in front was marked by a very evident ridge in the lower part, flanked on either side by a groove. These grooves coalesced in the upper part of the abdomen and were continued over the thorax as a median depression. ‘This could be traced as high as the root of the neck, where it was lost. 18. Gentoo Penguwn Embryo, 28 Days. The embryo appeared fully formed and was in a fully flexed position, so that the under surface of the mandible was pressed firmly against the thorax and abdomen ; the beak lay in a groove formed by the right fore limb and left hind limb. The head was covered with a dark grey, almost black, down ; the back with slate coloured, the abdomen with white. The back of the head had been flattened by pressure. Duck Embryos, 19-28 Days. The adyance was purely one of body bulk. The amount and directions of growth are shown by the measurements of the specimens. (See Table I.) 19. Gentoo Pengun Embryo, 30 Days. (The specimen in fig. 25 closely resembled this older specimen, but was rather smaller.) The embryo appeared fully formed and was in a fully flexed position, so that the under surface of the left side of the head was pressed against the thorax, the beak lying to the right side of the abdomen between the right fore limb and the foot of the left hind limb anteriorly and the right hind limb posteriorly. The head and back were covered with slate-coloured down, the belly with white down. In the flexed position the greatest length was 5°71 cms., the greatest breadth 3:05 cms. In the extended position the length from the crown of the head to the tip of the tail was 8°41 ems. When measured separately the head was 2°55 cms. in length, the neck 1°85 cms. ; the body and tail 242 DR D. WATERSTON AND DR A. CAMPBELL GEDDES ON EMBRYO PENGUINS together 5°51 cms.—giving a total length of 9°91 cms. The 1°5 cms. of difference (9°91 ems. — 8°41 ems.) is accounted for by the marked spinal curves which immediately asserted themselyes on extension, a lumbo-thoracic curve ventrally concave, a cervical curve dorsally concave. SUMMARY AND CONCLUSION. Part I,—Anatomy of the Adult Pengwn. 1. The cervical portion of the vertebral column is possessed of a curve of extra- ordinary convexity which causes the vertebral bodies tu form a visible bulging on the front of the neck. This convexity we regard as useful to the birds by increasing water displacement anteriorly and permitting of the head and beak being carried without undue fatigue on the long ocean voyages which the birds undertake. The furcal angle being occupied by the vertebral bodies, the trachea and cesophagus enter the thorax on a plane posterior to the cervical spines. This curvature of the column and arrangement of the viscera is permanent and characteristic of the adult bird. 2. The arrangement of the fascial layers in relation to the pectoral muscles suggests that the pectoralis major and clavicle are in reality cutaneous structures. It was found impossible from the embryological material at our disposal to prove or to disprove this suggestion. Part II.—Embryology of the Pengwn. 1. The number of days of incubation is, according to the series of specimens, thirty. This is surprisingly short, and as there are some obvious gaps in the series, we are not prepared to express an opinion as to whether it is correct or not. 2. The early stages of development are exactly comparable to the corresponding stages of the chick or duck, though, if the dates of our specimens be correct, they take place more slowly. 3. The later stages of development are exactly comparable to the corresponding stages of the duck except with regard to the down, the fore limb, and the foot. 4. The foot of the penguin is, throughout its development, exceedingly clumsy and primitive, as is the foot of the adult; the foot of the duck, from its earliest appearance, assumes a lighter and neater appearance, and is adapted as a paddle, which the foot of the penguin is not. 5. The fore limb of the penguin is, in its mesoblastic structures, definitely progressive, whereas the mesoblastic portion of the duck’s wing is relatively regressive. Both, however, have varied from the ancestral type, but their variation is in opposite direc- tions: the wing of the penguin has required rigidity to subserve its function as a paddle, and it has acquired it, throughout its length. The wing of the duck, on the other hand, has required lightness, and it has acquired it—the reduced spread of bone being compensated for by an increased development of feathers. OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION. 243 6. The evidence from embryology is in favour of the early divergence of the Aptenodytidee from the main stem of the flyimg birds, and of the common ancestor of both having been a flying bird, though of a clumsy type. 7. We were unable to verify the facts recorded by other observers in the ossification © of the limbs, as we were handicapped by the decalcifying action of formalin even in weak solutions, so that our observations in this direction could not be considered altogether satisfactory. ILLUSTRATIONS. Prats I. Fig. 1. Gentoo penguin, embryonic area, early stage. Fig. 2. Gentoo penguin, within first 24 hours, showing head fold, medullary folds, and primitive streak. This stage resembles that of the chick at 22 hours. Fig. 3. Gentoo penguin, said to be 3 days. This stage of development resembles the chick at 24 hours. Fig. 4. Gentoo penguin, said to be at 5 days. The stage of development resembles the chick at 26 hours. The photograph shows specially the cephalic flexure, the hinder ends of the medullary folds enclosing the bulbous extremity of the notochord, and the remains of the primitive streak. Fig. 5. Gentoo penguin, said to be 6 days. Comparable to the chick at 40 hours. The photograph shows the bulbous expanded posterior end of the notochord, and evidence of a segmentation of the neural folds. Fig. 6. Gentoo penguin, about 7 days. Puate II. Fig. 7. Duck embryo of 5 days. Compare fig. 4 of penguin of same age. Fig. 8. Duck embryo of 6 days. Compare fig. 5 of penguin of same age. Fig. 9. Duck embryo of 7 days. Compare fig. 6 of penguin of same age. Fig. 10. Gentoo penguin embryo of 12 days. Both upper and lower limbs have appeared on the surface of the trunk. Fig. 11. Gentoo penguin embryo of 13 days. Fig. 12. Duck embryo of 8 days. Compare character of limbs with those of penguin of 13 days (fig. 11). Figs. 13, 14, and 15. Successive stages of Gentoo penguin embryos at 15, 16 and 17 days, to illustrate stages in the development of the sickle-shaped fore limb of the adult. Fig. 16. Duck embryo of 12 days, im situ, to show how the embryonic attitude is maintained, and is not dependent upon external pressure. Fig. 17. Gentoo penguin embryo of 22 days, to show advancing change in the position of the feet. Compare figs. 15 and 23, which show other stages in the development of the peculiar flat base of the adult. Puats ITT. Fig. 18. Duck embryo of 13 days. Figs. 19 and 20. Gentoo penguin embryo of 18 and duck embryo of 12 days, photographed to the same scale, to show altering characters of the limbs, when the trunks are of the same size. Fig. 21. Duck embryo of 14 days. Fig. 22. Duck embryo of 15 days. Compare with fig. 15 of gentoo penguin embryo of the same age, TRANS. ROY. SOC. EDIN., VOL. XLVII. PART IL. (NO. 10). 36 244 EMBRYO PENGUINS OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION. Fig. 23. Gentoo penguin embryo of 24 days. Notice the character of the fore limb, and the attitude of the hind limb and tail. Fig. 24.-Gentoo penguin embryo of 25 days. Fig. 25. Gentoo penguin embryo of 27 days, to show the stage of development reached before leaving the shell. Fig. 26. Photograph of fore limbs of gentoo penguin embryo of 22 days, to show ossification. The clear portions of the shafts are composed of bone, and the dark extremities are of cartilage. Both extremities of the radius and ulna are cartilaginous. Beyond them is a darkish area, in which are the’ cartilaginous rudiments of the carpus. Beyond that region are the metacarpal bones and phalanges. Vol XEY i. Prare, I. ‘rans. Roy. Soc. Edin* Scottish NATIONAL ANTARCTIC EXPEDITION. EMBRYO PENGUINS: TON AND GEDDEs TERS Drs Wa ies A ING, Bt z| ks| <2] oy | 2 wi Fa | | yy E = | | trans. Roy. Soc. Edin* Prusre, 1. Vol. XLVIL. | Drs WATERSTON AND GEDDES.—EmMBRYO PENGUINS: ScoTTISH NATIONAL ANTARCTIC EXPEDITION. Fic. 7 Fic. 8. Fic. 9. Fic. 10. Bie. 11, Fic. 12. Fic. 13. Fic. 14. Fig. 17. Fic. 16. M'Farlane & Erskine, Edin. | (rans. Roy. Soc. Edin" Puate III. Vol XLVIL | Drs WATERSTON AND GEppDES.—EmBryo Pencurins: Scorrish NATIONAL ANTARCTIC EXPEDITION. Fic. 19. Fic. 20. Fic. 21. Fic. 24. Fic. 25. Fic. 26. M‘Farlane & Erskine, Edin. Sc ee ( 245 ) XI.—The Pharmacological Action of Harmaline. By James A. Gunn, M.A., M — 90) 7120 after injection No. ee a 54 BA BT «158 «EBC 199" One ome per minute When injected into the dorsal lymph sac of winter frogs, 0°04 mg. per gramme body-weight caused slight slowing of respiration, but no other obvious symptoms ; 0°08 mg. per gramme body-weight induced marked slowing of the respiration, which also took on a periodic character, groups of about six respirations being followed by an interval up to half-minute duration of no breathing; there was also some muscular depression; 0°16 mg. per gramme body-weight caused almost complete cessation of respiration and muscular paralysis; and 0°24 mg. per gramme body-weight eventually produced complete paralysis of the respiration and death. As an example, the effect after the injection of 0°16 mg. per gramme body-weight may be taken. In this experiment the frequency of the respirations fell to one-half in ten minutes, to one- eighth in twenty minutes, and to one-twentieth in seventy-two minutes after the injection ; the frog was then pithed. ACTION ON VaGAL ENDINGS. This was determined by observing on the blood-pressure the effect of electrical stimulation of a divided vagus before and after injection of the drug, or the effect of the drug after the injection of pilocarpine. When electrical stimulation was employed the procedure was as follows: the vagus was stimulated at intervals until a practically constant effect was obtained, the drug was then injected into the external jugular or facial vein, and the vagus again stimulated at intervals until an effect as near as possible identical with that previously obtained was reached ; or the injection was made during continued stimulation of the vagus (fig. 2). “UU 0% PeSlel OUI] ese ‘| “Sy UI Sv sxoqyoT ‘qno sndeA JUST “TOYA “yeOQ ‘snsea jo uoryepNUIs Sutimp ATsnouearyut opiso;yoorpAy outedo1y4-[Ayooyeo0jo1d “3 [0.0 Jo Jooyy—'s “D1 275 ‘Ieout] € x “qooyja yeordAy poos ve peonpoid uooolut oy 19qje “SUIT ZT PUE aLoJod SNSVA Jo UOTR[NUIG ‘spuooes UI oul = 7 ‘(WU ZE paste) OUT, eseq = Q { cInsserd-poo[q pryoavo = 9 { uoTyeaIdsut syueserder exorysdn ‘ (pioder ydersousiyd) uoyeatdser = « ! uoroelut Jo aULI} SeABOIpU 7 BUTT UO UOISsaIdaq + “aplo,yoorpAy eutedosy-[Ayoozeo0jo1d “3 [9.0 Jo UorjoolUT SnOUAARIZUT Jo JOaYA “ynotsea yJog “IYIA “3000S 42O—'T “SIA WAIT TIT ITT SLL of/ >of -—— Ao bony Tt ; THE PHARMACOLOGICAL ACTION OF PROTOGATECHYL-TROPEINE. | 276 PROFESSOR C. R. MARSHALL ON By these methods 0°01 g. protocatechyl-tropeine hydrochloride was found to paralyse the terminations of the vagus in the heart for five minutes. At the end of this period stimulation of the vagus generally produced a slight effect, and the effect gradually increased with succeeding stimulations until the former normal condition was reached, usually about twelve minutes after the injection. Occasionally the same strength of stimulus failed to induce any action, and consequently it had to be slightly increased. ‘Thus, after the injection shown in fig. 2, stimulation with the secondary coil at 20 em. produced no effect up to eight minutes after the injection, whereas stimulation with the coil at 16 em. produced at this time a very good effect. I have obtained the same result with other tropeines. When pilocarpine is used as a stimulus the paralysis of the vagal terminations does not last so long. Thus the injection of 0°01 g. protocatechyl-tropeie hydrochloride, after a previous injection of 0°01 g. pilocarpine, totally abolished the pilocarpine effect for one and a half minutes only. ‘The pilocarpine action then reappeared and in three minutes had reached its previous form. The shorter period of paralysis in these experiments as compared with that obtained when the vagus is stimulated electrically is due to the fact that pilocarpine and protocatechyl-tropeime are to some extent mutually antagonistic. EFrect oN BLoop-PRESSURE. The commonest effect on the blood-pressure is a gradual fall, usually commencing after the effect on the respiration has begun, and continuing when the respiration spontaneously recovers, after this has become practically normal (cf figs. 1 and 4). The fall of blood-pressure continues for 14—3 minutes, rarely more, and then the pressure gradually returns to its previous height. It is not usually severe in extent in rabbits; after a dose of 0°01 g. intravenously it generally amounts to about one- third the normal blood-pressure. In cats the fall is often greater, and not infrequently is preceded by a slight rise. ‘The extent of the fall in the last-named animals is shown in the following table, which expresses the result of two successive injections :— Minutes after injection . : : 5 : 0 3 9 0004 g. Bp.in Mm. Hg. . ; ‘ . 135 (f) 130 0002 gz. Bp.inMm. Hg. . ; : . 136 Bal 130 Occasionally in cats a rise of blood-pressure has been the most prominent feature. This effect is seen in fig. 3, which illustrates the greatest rise of blood-pressure I have obtained with this substance. There was no fall below the normal subsequently. The vagal terminations in the heart were found to be paralysed between A and B, and to be again irritable between B and C. Two injections of 0°01 g. each were made in the same animal later, and in both instances a rise of blood-pressure to nearly the height of that shown occurred. The rise was more prolonged in each case, and the vagal endings were paralysed for a longer time than after the injection illustrated. The respiration was not graphically recorded in this experiment. i THE PHARMACOLOGICAL ACTION OF PROTOCATECHYL-TROPEINE. “Weal, | X “4[NSed sULeS at[} YFIM UPATS Udeq peY suttedo.14-[AYoe7Rv00}01d Jo asop oULes oY} BL'LL7V ‘seinsy snoraead UL SB SI9}J9] 18Y3Q “LaJeuouRW-Tepey Aq Sutoky = f (‘SuLOeIT} Jo yuaTMedUEMUIOD aIOJaq SpUOdaS GE ‘MMOYS OU YsIY YIIM [eoIyuepr AT[eooeid uUoMe[NUIIYs &e Jo JoaTo [BULOLI9} sMOYsS Sutowy jo 41ed qsitq) ‘sew $Z x JeAe_ ‘quit, uo yooyo = wm “uO ZI 4 [loo Avepuooss YIIM SAIOU Jo UOYR[NUIIYS Jo euIT} o1BOIPUT SOUT, oITYM AavoyT “pasodxe eALou [RANI 4JoT “TOYA “3 OOPG “MAGeY ‘sSurpue oAreu ojou uo ATsnousarxyur optiopyoorpAy outedo.4-[Ayoozeo0j0ad “3 [0.09 Jo ENA —"F “O1q b62-// Z, ae a ol UL “UBeUl[ — X “WW gg pasivl sul] seg ‘salnoy snotacrid UI se sieq{jeT + “OH pur g UsdeMyoq a[qezIIUI A[QYSI[S ‘gq pue Y Uoemyoq e[qeyIIIIUN sndeA ‘qo snsea qysiy “euIg “3 gaze 4% einssoid-poojq uo Ajsnousaeut epiiopyoorpAy outedo.r-[Ayooqeooqoid “8 ¢Q0.0 Jo Jooye [eucIseA0Q—"e “‘DIq 278 PROFESSOR C. R. MARSHALL ON When the respiration is paralysed and does not recover spontaneously the blood- pressure after a brief interval falls rapidly to a few millimetres. That this fall is mainly asphyxial is proved by artificial respiration, which quickly raises the pressure to the level it would probably have reached if respiration had recommenced. If the substance is injected after the administration of pilocarpine or during stimulation of the vagus a rise of blood-pressure always occurs. The fall in blood-pressure is probably cardiac in origin, since it is accompanied by a diminution in the frequency of the heart-beats; and as the vagus nerve endings are paralysed, the action of the drug would seem to be on the muscular tissue of the heart itself. The rise of blood-pressure seen occasionally in experiments with cats is accompanied by an increase in the number of the heart-beats due to paralysis of the vagal terminations. The cardiac muscle in these cases is not depressed sufficiently to counteract the effect of paralysing the vagus, but that some depression is produced is shown by the rise of blood-pressure being less than that obtained with a correspond- ing physiological dose of atropine. Moreover, increase in dose leads to a somewhat smaller rise of pressure. Why this prolonged rise should occur in isolated cases is difficult to explain. Probably a certain concentration of the drug in the heart acting for a brief interval only is necessary, but the condition of the heart itself may also be a factor. Action oN VoLunTARY MuscLeE anp Moror NERVES. The activity both of motor nerves, or more probably their terminations, and of voluntary muscle, is depressed by this drug. In rabbits the nerve endings appear to be more susceptible than the muscle; in frogs the muscle is at least as powerfully affected as the nerves. After an injection of 0°24 mg. protocatechyl-tropeine hydrochloride per gramme body-weight into the dorsal lymph sac of a frog, muscular weakness appeared within five minutes—the head began to droop and the animal became unable to turn over when placed on the back. A few minutes later the frog was quite flaccid. This condition and the paralysis of the respiration referred to previously was maintained during the period of observation (the frog died during the night). Although apparently completely paralysed, the frog occasionally made a slight voluntary movement, and a slight reflex was several times obtained on pinching a toe moderately strongly for some seconds. After 0°16 mg. per gramme body-weight was injected similar symptoms were observed. The head commenced to droop in twenty minutes and the animal gradually sank on the table. The reflexes, however, remained distinct, and when placed on the back an hour after the injection the frog recovered once, but not twice, from this position. The frog was pithed seventy-two minutes after the injection and the irritability of the sciatic nerves and gastrocnemii muscles tested. The sciatics reacted with the secondary coil at 35 cm., the muscles with the coil at 28 cm. ‘The injection of 0°08 mg. per gramme body-weight produced only slight muscular depression. THE PHARMACOLOGICAL ACTION OF PROTOCATECHYL-TROPEINE. 279 On immersing a nerve-muscle preparation (gastrocnemius) in a 1—1000 solution of protocatechyl-tropeine hydrochloride in 0°6 per cent. sodium chloride and stimulating (one Daniell’s cell) the nerve and muscle alternately every three minutes, the contractions were found to fall with the same regularity in each case, and both muscle and nerve became inexcitable forty-five minutes after immersion. In a rabbit no muscular paresis was observed after 0°062 g. p. kg. intraperitoneally. Depression but not always paralysis of the nerve endings is obtained after intravenous administration to an anesthetised animal. ‘This is shown in fig. 4. The rabbit was prepared in the manner described, the right crural nerve was laid bare, and a hook was put into the right foot and connected by means of a thread working over pulleys to a weighted lever. After a practically constant effect had been obtained from stimulation of the nerve (one accumulator cell; secondary coil 12 cm.), 0°01 g. protocatechyl- tropeine hydrochloride was injected into the right facial vein. The contraction of the muscles produced by stimulation of the nerve gradually diminished until it disappeared two minutes after the commencement of the injection. Two and a half minutes later the limb muscles again responded to stimulation of the nerve, but the contractions did not assume their previous (normal) form until ten minutes after the injection of the drug. The tracing reproduced records a second injection of the drug; the same dose was administered at 11.19 and produced a similar effect. In a second experiment, in which both the right crural and sciatic nerves were exposed, it was found that the muscles, when stimulated directly, contracted in the neighbourhood of the electrodes at a time when the nerves were inexcitable. It would seem, therefore, that in the rabbit the nerve endings are more susceptible to this compound than the muscular substance. An interesting point is the want of correlation between the rep eTitOn and the neuro- muscular effects; at a time when the latter are most manifest the respiration is practically normal. The explanation is probably to be found in the stimulus of the respiratory centre being physiologically more powerful than the electrical stimulus used, and thus able to overcome any tendency towards paresis of the respiratory muscles. As bearing upon this point, evidence was obtained showing that the paralysis of the muscles and nerve endings was only relative, z.e. only for minimal or moderate stimuli. The effects on the blood-pressure and the neuro-muscular system, on the other hand, are correlative, the fall of blood-pressure being mainly, if not solely, due to an action on the cardiac muscle. REMARKS ON THE ACTION OF THE DRUG ON THE RESPIRATION. Rapid cessation of respiration in the expiratory phase has been described by ‘'apPEINER”® after the intravenous injection of methyl-phenyl-isoxazol-methochloride, diphenyl] - methyl - pyrazol- methochloride, dimethyl] - phenyl - pyrazol - methochloride, * Arch. f. exp. Path, u. Pharm., xxxvii., p. 325 [1896]. 280 PROFESSOR C. R. MARSHALL ON and tetra-methyl-ammonium chloride; and by Pox * after the intravenous injection of papaverin-methochloride, papaveraldin-methochloride, and papaverinol-methochloride. None of these substances produced the effect when administered hypodermically. As the result of his experiments, TAPPEINER came to the conclusion that methyl- phenyl-isoxazol-methochloride stimulated the terminations of the trigeminal nerve, and that the effect produced was, in reality, a Kratschmer-Hering reflex. Thus he found that cocainisation of the nasal mucous membrane prevented the respiratory paralysis ; and that while cocaine administered intravenously also antagonised the action of the isoxazol, the dose required was larger than when the cocaine was applied directly to the nose. ‘The action of tetramethyl-ammonium chloride and the other substances investi- gated by him was explained in the same way. IopLBavER,t who investigated tetramethyl- ammonium chloride more minutely under TAPPEINER’s direction, came to the same conclusion. PoHL, however, still obtained the characteristic stoppage of the respiration Fic. 5.—Effect of 0°01 g. protocatechyl-tropeine hydrochloride intravenously on decerebrate rabbit with both fifth cranial nerves cut. Tracing of respiration taken by connecting one limb of tracheal cannula to a tambour ; other limb left open. Time in seconds. after cocainisation of the nose and after division of the ophthalmic branch of the fitth nerves, both in the case of papaveraldin-methochloride and of tetramethyl-ammonium chloride, and he therefore maintains that the action is a central one. The action of protocatechyl-tropeine is also upon the centre. The typical effect on the respiration is still obtained when the brain above the pons is excised and both the fifth cranial nerves are cut within the cranium (fig. 5). Moreover, the same action is obtained in cats (fig. 1), animals in which the typical Kratschmer-Hering reflex is not present. And since the effect on the neuro-muscular system is not coincident in time with the effect on the respiration, the cause of the latter would seem not to be peripheral. It has, however, been shown that the motor nerve endings in the diaphragm may be affected differently, either as regards time or degree, from those of many other voluntary muscles, and it therefore seemed desirable to determine whether the diaphragm played any part in producing the respiratory paralysis. Accordingly, in an anesthetised * Arch. Internat, de Pharmacol., xiii., p. 479 [1904]. + Ibid., vii., p. 183 [1900]. THE PHARMACOLOGICAL ACTION OF PROTOCATECHYL-TROPEINE. 281 animal, both phrenic nerves were cut and the respiration registered by a connection with the tracheotomy tube. On injecting 0°01 g. protocatechyl-tropeine hydrochloride into the external jugular vein the respiration ceased thirteen seconds after the commencement of the injection (fig. 6). In connection with the production of temporary cessation of the respiration, the question of dose is of paramount importance. The concentration of the drug producing this effect in the blood supplying the medulla can apparently vary only within narrow limits. Hence not only the amount of drug injected but also the time occupied by the injection is of moment. ‘his explains why the effect does not result from hypodermic or intraperitoneal administration. It is probably owing to this fact that the effect has not been observed more frequently. It can certainly be produced by a number of UU UU Fic. 6.—Effect of 0°01 g. protocatechyl-tropeine hydrochloride on respiration of rabbit with both phrenic nerves divided, Tracing taken as fig. 5, but with air-exits closed. Time in seconds. drugs, but the doses of those which induce it fall within very narrow limits. In many of these cases the effect is probably merely that of a protoplasmic poison, and the respiration is most obviously affected because the respiratory centre is the most sensitive of the medullary centres to this class of substances. This hypothesis, however, does not seem sufficient to explain the actions of the substances already mentioned. In a preliminary report on the physiological action of pukateine, an alkaloid isolated by Aston * from pukatea bark, Maucoum says: ‘“ Intravenous injection of 10 mg. into an anesthetised rabbit caused sudden stoppage of respiration without convulsions: the heart continued to beat vigorously for a long time afterwards.” As Mr Aston had given me some years ago a specimen of the mixed alkaloids isolated by him from pukatea bark, I was able to investigate this point. The alkaloids were first converted into the hydrochlorides. On injecting 0°01 g. intravenously (time occupied by injection, seven seconds) into a rabbit (wt. 2100 g.) the respiration was not * Annual Rep., New Zealand Dept. of Agriculture, 1908, Chemistry Division, p, 226. TRANS. ROY. SOC. EDIN., VOL. XLYII. PART II. (NO. 12). 42 282 PROFESSOR C. R. MARSHALL ON paralysed; it began to get shallower ten seconds after the commencement of the injection, and reached its lowest point five seconds later, from which time it very gradually returned to the normal. The blood-pressure commenced to fall before the respiration was affected, and fell from 87 mm. before the injection to 58 mm. twelve seconds after the commencement of the injection. From this point it gradually rose in three minutes to 107 mm. The effect of a subsequent injection of 0°02 g. caused almost complete cessation of respiration for half a minute, and then a very gradual return to normal breathing. The fall in the respiratory curve, however, was less abrupt, the blood-pressure was more quickly and more powerfully influenced, and the return of normal respiration was later than occurs after protocatechy]-tropeine. The other synthetic tropeines I have investigated do not distinctly influence the respiration in doses which depress the vagal endings, but doses of both atropine and homatropine may be found which will cause temporary cessation of the respiration. The dose of atropine necessary for rabbits is about 0°0001 g. per kg. body-weight intravenously. After this dose, however, the return of normal respiration is less rapid than after the administration of protocatechyl-tropeine. Homatropine, in doses of 0'005 g. per kg. body-weight intravenously, produces a more transient effect on the respiration and a greater effect on the blood-pressure than atropine in the dose mentioned. I have also succeeded in producing temporary stoppage of the respiration with the allied alkaloid cocaine, and with §-eucaine and other local anesthetics. I failed, however, to produce the effect with ammonium hydroxide. A certain degree of recovery may, with appropriate doses, be obtained after intravenous administration of this substance, but this recovery is not maintained, and the respiration fails again unless artificial respiration is performed. The best effect I have obtained was from the injection of 0°025 g. ammonium hydroxide per kilogramme body-weight. Larger amounts of ammonium hydroxide may be followed by a few spontaneous respirations, but they paralyse the respiration permanently unless it is restored by artificial means. Doses less than 0°01 g. per kilogramme generally increase the respiratory movements ; above this dose the movements are diminished. COMPARISON OF EFFECTS ON BLOOD-PRESSURE. All the drugs previously mentioned, in doses which temporarily abolish the respira- tion, diminish in rabbits the frequency of the heart-beats. The effect is greatest in the case of methyl-pheny]-isoxazol-methochloride and tetramethyl-ammonium chloride, and least in that of the tropeines. In most cases there is also a fall in blood-pressure. Methy1]-pheny]-isoxazol-methochloride, however, notwithstanding the marked slowing of the heart produced by it, causes, after a preliminary slight fall, a rise of blood-pressure, which is due, according to TappEINER,* to stimulation of the medullary centres. The slowing of the heart and fall of blood-pressure produced by tetramethyl-ammonium * Loc. cit. THE PHARMACOLOGICAL ACTION OF PROTOCATECHYL-TROPEINE. 283 chloride, and probably also by the alkaloid pukateine, are due mainly to stimulation of the vagal terminations. According to lopLBauzER,* the effect of tetramethyl-ammonium chloride is less after section of both vagi, and consequently he concludes that its action on the circulation is partly, if not mainly, central. He also obtained a marked rise of blood-pressure when this drug was injected after atropine, a result he attributes to stimulation of the vasomotor centre. I have obtained a rise of blood-pressure on cutting both vagi a few minutes after the injection of tetramethyl-ammonium chloride, but have not observed any difference on the frequency of the heart-beats after injection of this substance whether the vagi were divided or not. After atropine had been given, the usual slowing of the heart did not occur, nor was there any rise of blood-pressure. The slowing of the heart’s frequency and the fall of blood-pressure produced by the papaverin derivatives investigated by Pout are attributed by him to the stoppage of the respiration. He does not, however, refer to any investigations bearing on this point in his paper, and he gives no tracings from which the action on the circulation might be inferred. The action of protocatechyl-tropeine on the circulation differs from that of most substances producing temporary paralysis of the respiration, in that it paralyses the vagal endings within the heart. Hence the diminution in the frequency of the heart- beats and the fall of blood-pressure are less than are produced by most members of this class of respiratory paralysants. As previously stated, the fall of blood-pressure caused by these tropeines is probably due to a depressant action on the cardiac muscle, an action which in all probability is present in the case of the other substances also. From these statements it is evident that no uniformity, such as exists in the case of the respiration, characterises the action of these different substances on the circulation, and this further supports the statement made previously that their action on the two systems is not correlated. REMARKS ON THE RELATION BETWEEN CHEMICAL CONSTITUTION AND PHARMACOLOGICAL ACTION. Protocatechyl-tropeine, as previously stated, was one of a series of tropeines in- vestigated for the purpose of determining the difference between a lactone and the corresponding hydroxy-acid. As it is not a lactone it was used as one of the control substances, and it was found that, unlike pilocarpine and the lactone-tropeines, it does not lose its action when a molecular quantity of caustic alkali is added to its aqueous solutions. Even the addition of distinct excess of caustic alkali and prolonged standing do not influence its pharmacological action to any appreciable degree. It is evident, therefore, that the replacement of the hydrogen of the phenolic hydroxyls by an alkali metal is of no pharmacological importance. A point of interest on which the action of protocatechyl-tropeine eos a little * Loc. ctt. 284 PROFESSOR C. R. MARSHALL ON light is the part which the constituent groupings play in the pharmacological action of tropeines. BucHueim,* LapensurG,t and others have found that various aromatic acidic groups could be attached to the base tropine, and an atropine-like action be obtained ; whereas Crum Brown and FRasmr,{ and more recent observers, have shown that alterations in the tropyl radical generally produce profound changes in the pharmacological action of a tropeine. This suggests that the fundamental action of these tropeines depends on the tropyl radical, and that the interaction with the so- called nerve endings occurs through the mediation of the acidic grouping. Whether the acidic group has any other action than a connecting or haptophoric one with the tissues is difficult to decide. Many of the tropeines, in so far as they have been investigated, differ little except quantitatively in pharmacological action from atropine, and it is open to assume that in these compounds the acidic groupings are purely haptophoric in character, since their different quantitative effects might be explained by differences in the power with which the acid group is able to attach the tropyl group to the nerve endings. GorTLiEB,§ however, showed that certain tropeines act very differently from atropine—acetyl-tropeine causes convulsions and succinyl-tropeine paralysis, and neither affects the vagus endings or the pupil; and in view of the comparatively-simple nature of the acid groups of these compounds, it seems questionable whether any distinctive action can be attributed to the constituent groupings of tropeines. In the case of protocatechyl-tropeine we have a compound which differs materially in action from other tropeines, yet most of its actions can be demonstrated with atropine. But while it acts much less powerfully than atropine upon the vagal endings in the heart, it acts more powerfully on voluntary muscle, and has an action on the respiratory centre which atropine shares to a relatively slight degree. It may be that these differences might be explained by the presence of ditferent receptive substances in the different tissues, but this view would scarcely explain why, in small doses, atropine exerts a stimulating effect on the respiratory centre, and protocatechyl-tropeine no such stimulant action. Un- fortunately, it is not possible to determine the action of the two constituent groups of these compounds by investigating the products of their hydrolysis. It was early shown by Fraser? that tropine and tropic acid produce no atropine-like action, and it was consequently not surprising to find that simple protocatechyl compounds possess no action like that of protocatechyl-tropeine. The substances I investigated were proto- catechyl-aldehyde,|| methyl-protocatechyl-aldehyde (vanillin), and di-methyl-proto- catechyl-aldehyde (piperonal). Intravenous injections of these compounds produced no effect on the respiration resembling that produced by protocatechyl-tropeine. There is thus no evidence leading us to assume that the actions of protocatechyl-tropeine can be attributed with any certainty to the two cunstituent groupings individually. * Arch. f. exp. Path. u. Pharm., v., p. 463 [1876]. + Annalen d. Chemie, Bd. 217, p. 82 [1883]. t Trans. Roy. Soc. Edin., xxv., p. 693 [1869]. § Arch. f. exp. Path. u. Pharm., xxxvii., p. 218 [1896]. “| Proc. Roy. Soc. Edin., 1869, p. 558. || Prepared by acting on piperonal with phosphorus pentachloride. THE PHARMACOLOGICAL ACTION OF PROTOCATECHYL-TROPEINE. 285 All the substances previously investigated which have been shown to paralyse temporarily the respiration are quaternary bases. The tertiary bases corresponding to these did not, where tried, affect the respiration. Protocatechyl-tropeine, however, is a tertiary base, and chemically has nothing in common with the other substances pro- ducing the same respiratory effect. But it possesses another physiological action common to many of them, namely, the power of paralysing the motor nerve endings to voluntary muscle. It is questionable, however, if this connection is more than a co- incidence, since trimethyl-ammonium chloride, triethyl-ammonium chloride, and tetraethyl-ammonium chloride, which paralyse motor nerve endings, have no paralysing action on the respiratory centre;* and the papaverin bases which do produce re- spiratory paralysis do not paralyse motor nerve endings. SUMMARY. Protocatechyl-tropeine paralyses the vagal endings in the heart, but is much less powerful than atropine, or even homatropine. It depresses muscular activity, especially in frogs, and interferes with the con- ductivity of motor nerves, or more probably the myo-neural junctions, in rabbits. In certain doses, it paralyses temporarily the respiration, and causes a gradual fall of blood-pressure ; but the two effects are not apparently correlated. In cats, a rise of blood-pressure sometimes occurs, owing to the paralysis of the vagal endings. The temporary stoppage of the respiration is due to paralysis of the respiratory centre, and not to a peripheral action. No evidence indicating separate pharmacological actions of the two component groupings of the compound was obtained. ; * TAPPEINER, loc. cit. TRANS. ROY. SOC. EDIN., VOL. XLVII. PART ITI. (NO. 12). 43 ( 287 ) XIII].—The Pharmacological Action of Tutu, the Toot Plant of New Zealand. By Professor C. R. Marshall. (MS. received November 20, 1909. Read June 21, 1909. Issued separately January 29, 1910.) CONTENTS. : PAGE | PAGE Introduction : , : F 3 ; . 287 | The Convulsions : : ; : : . 300 Active Principle . é , ( : : . 288 | Influence of Anzesthetics on the Convulsions . 302 Previous Pharmacological Investigations . . 289 Muscles affected by the Convulsions . 5 . 3804 General Effect of Tutin : : : . 290 Seat of Origin of the Convulsions : : . 3807 on Rabbits . 6 0 , 3 : . 290 | Action on the Higher Centres. : : . 309 on Guinea-pigs . : ; . é . 294 Influence on Temperature . j : > . 310 on Frogs. : , 3 6 5 . 294 | Effect of Decomposition Products : : . 312 on Trout and Tadpoles : ; : . 295 Comparison with other Convulsants . : . 812 Action on Medullary Centres. : : . 295 | Summary . 4 : : : ‘ E . 314 INTRODUCTION. Under the name Tutu, three species of Corraria—C. ruscifolia (L.), C. thymifolia (Humb. & Bonp.), and C. angustissima (Hook.) *—are usually included. Other species have been described, but they appear to be varieties only. All the forms are poisonous, and in times past this plant has proved a serious hindrance to the rearing of stock in New Zealand. Linpsay,t in the course of a tour through the country during the latter part of 1861 and the earlier months of 1862, “‘ was everywhere struck by the abundant evidences of the devastation produced among flocks and herds from their feeding on the toot plant”; and he further states that ‘‘ he seemed a fortunate farmer or runholder who had not lost more than 25 per cent. of his stock from toot-poisoning; while in some instances the losses were so high as 75 per cent.” In order to gauge the extent of its ravages at the present time, a circular letter, dated 25th July 1900, was sent by Professor KasteRFIELD and Mr Astron to every medical man in New Zealand, and to such laymen as they thought might take an interest in the matter. Very few replies were received (4 from medical men, 10 from laymen),{ and none of them indicate the extent of the malady. On the whole, the correspondents corroborate the observations of Linpsay (J.c.). The following account summarises the statements made in the letters and in the extracts from the lay press submitted to me. * C. ruscifolva is commonly known as “tree-toot.” It grows to a variable height, sometimes to 6-8 metres, and has glossy, succulent leaves and shoots. C. thyméfolia or “ground-toot” usually grows to 30-50 cm. in height, and has small leaves and flowers. C. angustissima is herbaceous, and grows about 50 cm. high. Its slender branches terminate in very fine branchlets which carry numerous leaves, thus giving the plant a fern-like appearance. It is compara- tively rare. + “On the Toot Plant and Poison of New Zealand,” Brit. and For. Med.-Chir. Rev., July 1865, p. 153. { These were kindly sent to me by Mr Aston. TRANS. ROY. SOC. EDIN., VOL. XLVII, PART II, (NO. 13), 44 288 PROFESSOR C. R. MARSHALL ON The plant seems to be most dangerous in spring. At this season it puts forth thin, succulent shoots before other vegetation appears, and it is consequently liable to be eaten in large quantities as soon as the animals are turned out to graze. Later in the year, when fodder is abundant, poisoning is less common and the symptoms less severe. The symptoms observed after tutu is eaten are very similar in both cattle and sheep. Unsteadiness, stupor, and convulsions are the chief, and they usually occur in this order. Any one of these may be predominant, but the convulsions are the most characteristic. After bemg “tooted” the animals may show no symptoms until they are aroused, when unsteadiness and staggering expose the condition. The animals, it is said, frequently run about as if wild, often against obstacles, and if any water is near at hand they may rush into it and be drowned. If such an accident does not occur, convulsions of a clonic type develop, and the animals die in these or during the period of exhaustion which follows. Other animals are affected in a similar manner. Fowls have been poisoned by eating the seeds, and an elephant by eating the young shoots. Pigs and rabbits + are said to be immune, but the statement is untrue. This was proved for the pig by Mr J. A. GitrutH (v. p. 290), and for the rabbit by myself. Men, and especially children, have frequently been poisoned by eating the fruits (so-called “‘ berries”). The juice of these is innocuous, and was made into a beverage by the Maoris; the seeds, however, are poisonous. The main symptoms noted have been vomiting, giddiness, stupor or excitement with delirium, convulsions, and coma; but cases appear to have varied considerably in the clinical picture they have presented. A notable effect is loss of memory: in most of the cases of recovery described, there has been no remem- brance of eating tutu, or of the convulsions and other symptoms which followed. ACTIVE PRINCIPLE. The earliest recorded chemical investigation of the toot plant was made by Mr W. Sxey,{ Analyst to the Geological Survey of New Zealand, and was published in 1869.§ By exhausting an alcoholic extract of the seeds of C. ruscifolia with ether, he obtained a pale green viscid oil, five minims of which, administered to a cat, caused almost immediate vomiting, followed in half an hour by uneasiness and convulsive twitches of the ears and eyes accompanied by forward jerking of the head. These symptoms culminated in a convulsive fit about one hour after the substance was administered. The animal eventually recovered. He concluded that the active principle existed in the oil, if it was not the oil itself; and he conceived the idea that * Trans. N.Z. Instit., vol. 1. p. 399 [1869]. + Ibid., vol. iii. p. 242 [1870]. { Ibid., vol. ii. p. 153 [1869]. § Dr Lauper Linpsay (/.c.) mentions that Dr Murray THomson of Edinburgh undertook a chemical investigation of the specimens brought home by him, but owing to Dr Thomson’s removal to India the research was not completed. Sxey also states that “a great many experiments have from time to time been made upon the toot plant with the object of extracting the formidable poison known by sad experience to exist therein”; but as no other mention is made of these, they had probably ended in failure, THE PHARMACOLOGICAL ACTION OF TUTU. 289 the oil was a fixed oil, and consisted of esters of peculiar fatty acids, to one or more of which acids he ascribed the poisonous effects obtained. In the following year Hucuss * endeavoured to show that the active principle was an. alkaloid; but the crystalline principle he isolated was not properly examined, and his experiments are otherwise unconvincing. Nothing further was done until 1900, when HasrerFieLp and Aston, t by extracting a purified decoction of the plant with ether, obtained a crystalline glucoside from each of the three kinds of tutu. This substance, which they termed tutin, and to which they ascribed the formula C,,H,.0,, has proved to be the active principle of the plant. It was sent to me for pharmacological investigation, and | desire at this point to express my indebtedness to them for the material with which they have supplied me. Other substances were isolated from the three species of Coriama investigated : quercetin from C. thymifolia ; acetic, gallic, succinic, and a few other well-known acids from C. ruscifolia; and a volatile acid, C,H,0,, which has not been identified, from C. angustissima ; but these are obviously not the cause of the physiological activity of the toot plant. Previous PHARMACOLOGICAL INVESTIGATIONS. Apart from the published descriptions of accidental cases of poisoning in man and animals, and the solitary experiment of Skry already mentioned, our knowledge of the action of tutu is limited to a few experiments made by Mr Acurson{ and to an investigation of Dr W. LepincHam CuristIE.§ AcHESON administered an aqueous extract prepared by Hucuxs to dogs and cats, and noticed, after giving 15 grains or more, increased frequency of respiration, with slight twitchings of the extremities followed by severe attacks of convulsions. He found that the extract rapidly decomposed, and after a few days gave practically no pharmacological action, and that lime decomposed the poison and seemed to act as an antidote. CHRISTIE'S investigation included some observations on the effect produced by different doses of a crude extract, and some experiments to determine the mode of action of the drug. After the administration of small doses of an aqueous extract by the mouth or hypodermically to dogs and cats, he observed salivation and apparent nausea, and after somewhat larger doses vomiting also. Still larger amounts (corre- sponding to 40-50 grains of the leaf) caused in addition, and commencing within half an hour, “ twitching of the smallest muscles, as those of the eyes, nose, and ears. Then the muscles in front of the neck began to jerk, the fore paws began to follow suit, and soon the hind limbs and tail were affected.” LHarly and late dyspnea, erection of hairs, and increased frequency of micturition and defecation were noted. The vomiting he believed to be partly gastric but mainly medullary in origin; the early dyspncea * Trans. N.Z. Instit., vol. iii, p. 237 [1870]. t Trans. Chem. Soc., vol. 1xxix. p. 120 [1900]. t Trans. N.Z. Instit., vol. iii. p. 241 [1870]. § N.Z. Med. Journ., 1890, July and October, 290 PROFESSOR C. R. MARSHALL ON he attributed to the nausea, and the late dyspnoea to the convulsions. Rabbits he found insusceptible ; but the method of administration adopted—injection of an ethereal solution, at first hypodermically, then into the pleural cavity, and later into the peritoneal cavity—in the two experiments mentioned was unfortunate. And _ his experiments on birds are similarly vitiated. He endeavoured to localise the action of tutu by the following experiments. After obtaining convulsions from an intrapleural injection of two minims of oil of tutu, he sliced off the grey matter of the deft motor area, and noted that when the effect of the aneesthetic had passed away the movements were limited to the rzght fore and hind paws. From this experiment he concluded that ‘‘ when the cortex is absent there are no convulsions in the corresponding muscular area.” In a second experiment he injected an ethereal solution of oil of tutu subdurally in the right parietal region, and obtained symptoms which were undoubtedly due to the ether injected and the increased intracranial pressure produced; but he obtained also, as late symptoms, twitchings of the left eyelid and later of the left fore and left hind limbs. Eventually, the movements became more symmetrical. In a third experiment the ethereal solution was applied to a (decorticated ?) portion of the left cortex, and was also “inserted deep towards the base posteriorly.” ‘‘ Blindness and impaired intellectual perceptions ” were marked, and a convulsion subsequently supervened. Convulsions are said not to occur below a section of the spinal cord, but no details of the experiment are given. From these experiments he concludes that “the cortex of the motor area is the chief part affected ; but the sensory and cerebellar as well as the optic lobes, etc., or corpora quadrigemina, seem to be irritated, for movements of the eyes occurred in some cases.” Harlier in the paper he doubts if the basal ganglia take any part in producing the convulsions. GENERAL Errect oF TuTIN. The effect of tutin was tried, soon after its isolation, on a pig by Mr A. J. GinrutH, M.R.C.V.S., and on two cats by Mr A. R. Younc, M.R.C.V.S. The pig (35 lbs.) received two grains dissolved in water with its food. In half an hour spasmodic closure of the mouth was noted, and somewhat later accelerated respiration and vomiting, which increased in severity until convulsions set in. The animal died four and a half hours after the administration. The cats which were given tutin in solution suffered mainly from convulsions followed by coma. Death occurred in forty minutes. My own observations have been made on rabbits, guinea-pigs, frogs, tadpoles, and young trout. Im all, the effects produced were very similar, so that, taking into consideration the accidental poisoning observed in many of the higher animals, it is evident that the toot plant is a poison to all classes of vertebrates. On Rabbits.—In rabbits, after a lethal dose there is a variable period of quietude, accompanied by a diminution in the number of the heart-beats and an increase in the frequency of the respirations, which is followed by tonic and clonic convulsions. The THE PHARMACOLOGICAL ACTION OF TUTU. 291 earlier convulsions usually commence in a tonic spasm. The animal presents a frightened appearance, the muscles of the limbs increase in tone and become rigid, and opisthotonos develops. ‘This tonic spasm terminates in a clonic one, which in turn ends in exhaustion. During the convulsion, respiration is absent, and it may not return until the period of exhaustion has lasted for two minutes or more. A few minutes later the animal sits up and appears normal. This alternation of convulsions and exhaustion with a normal appearance may occur many times; but after a few convulsive attacks, clonic movements of the limbs, usually accompanied by retraction of the head, generally become constant. Salivation, dilatation of the pupils and of the vessels of the ears, are frequent symptoms. Death may occur in the first or a subsequent convulsion, or may result from exhaustion. After non-lethal doses the convulsions are generally less severe but more character- istic. There is, at first, usually an appearance of clinging to the ground, often accompanied by muscular tremors, and the gait becomes hesitating and spasmodic ; but the convulsions themselves commence with twitches about the eyes or mouth, or ‘ with erection or retraction of the ears, and extend to the whole head and fore part of the body. Frequently these movements rapidly terminate, especially in the earlier attacks, in a tonic convulsion; but later the convulsions are almost solely clonic. At this period, most frequently in animals which have been given a relatively small dose, a curious series of movements often occurs. The animal commences to push backwards with its forefeet, the hind parts being kept fixed, until it may, when the whole move- ment is carried out, perform a somersault over its tail. The movement may be abandoned at any point, and the animal may remain in the constrained position or assume a normal attitude. The movement may also be preceded, or accompanied in its early stages, by clonic spasm of the face or head, and it may terminate after the somersault in a convulsion. CHRISTIE mentions somersaulting round a transverse axis in the protocol of one of his experiments on cats. Tooted rabbits frequently lay, often for long periods, with the hind limbs more or less outstretched and with toes outspread as in the final stages of a spring. This position was so common that I termed it “ characteristic position.” Doses which do not produce convulsions have an effect on the medullary centres which will be described later. The lethal dose* for rabbits is about 0°0025 grm. tutin per kg. body-weight administered by hypodermic injection. After such a dose, death usually occurs from exhaustion. The lethal dose cannot be determined with exactitude, as it varies some- what with external influences. Thus a dose which is usually non-lethal may be * In a “Report on the Physiological Action of Tutin” by Dr Fircaerr and Professor Maucoum, published in the Annual Report of the New Zealand Department of Agriculture for 1908, which has reached me since the above was written, the minimal iethal dose of tutin by hypodermic injection (unless otherwise stated) is given in mg. per kg. body-weight as follows :—cats, 0°75 ; guinea-pigs, 2; rabbits, 2°5 ; lizards, 3-4 ; birds, 10°5 (by oral adminis- tration) ; frogs, 10°5 ; fishes, 50 (when immersed in the solution), An extended report by FrrcnHerr and Matcoum has appeared as this paper is passing through the press (see note at end). 292 PROFESSOR C. R. MARSHALL ON converted into a lethal one by turning the animal on its back a few times, thereby inducing convulsions. The minimal dose causing convulsions in rabbits is about 0°001 grm. per kg. body- weight. Larger doses may be given with impunity if the animal remains quiet, and Exprriment I.—0-002 gim. Tutin per kg. Rabbit. Weight, 585 grm. Time. Observations. 12.19 0°58 c.c 1 in 500 tutin solution injected under skin of back. 12.29 Lay down and stretched hind limbs out. Respiration very rapid. 12.38 Moved backwards somewhat spasmodically ; on resting, fore limbs commenced to slide for- wards. Respiration rapid. 12.44 Occasional slight noddings of head. 12.49 Head-nodding more marked. Tremors affecting whole fore-half of body. 12.59 Convulsion, affecting almost solely right side of face and head—retraction of head with rotation to right, closure of right eye. Quickly recovered. 1.4 Mild tremors over whole body. Moved about very carefully for a short time, and after- wards assumed a crouching attitude as if about to spring. A few spasmodic movements forwards were made later, but the same crouching position was maintained for fifteen minutes. Occasionally tremors over the whole body occurred. Pupils dilated. 1.27 Clonic convulsion, mainly left-sided, and limited to fore part of body—the animal stood up on its fore limbs, the left ear was drawn back, the right remaining erect ; both sides of head, except the eyes, were next affected with coarse clonic movements. Convulsion lasted one and a half minutes. Slight salivation present. 1.33 Another mild clonic convulsion ; limited at first to the mouth, then affecting the whole head (30” after commencement), and finally involving the hind feet (90” after commencement), which, however, only moved slightly. Rotation of head to left occurred (120” after commencement). Respiration, which had been markedly quickened, became slower. 1.39 Laid on venter; fore feet pushed out in front ; ears retracted. Almost constant tremor. 1.41} | A third severer convulsion lasting 135”. At one time slightly right-sided. Salivation continues. 1.46 An attempt to lick a part on the right side produced clonic movements which ended in a convulsion. Marked winking occurred in this. 1.57 Clonic convulsion. In this convulsion the animal fell over on its side, and then showed very marked clonic movements—the legs moving very rapidly, as in running. 2.1 Similar clonic convulsion. 2.8 Convulsion affecting mainly the fore part of the body, especially the mouth and head. The animal repeatedly fell over on its side, but recovered its position each time. The con- vulsion lasted three minutes. 2.144 | Convuision similar to last. 2.254 | Quiet since last note. Suddenly made several high springs, and entered into a tonic convul- sion (slight opisthotonos): a transient rest was followed by clonic movements (running, biting, ete., movements). Head retracted in the main. Salivation. 2.55 Clonic movements of whole body have continued without cessation since last uote. There has been an occasional spasmodic cry. 3.0 Very transient tonic contractions. | 3,2 Tonic contractions of longer duration ; several cries, 3.10 More powerful clonic movements ; more frequent cries, with rapid respirations. 3.30 Improving ; sitting up better, 4.0 , Convulsion ; fell on side, and exhibited running movements. Afterwards gradually improved. bigs } Practically well. morning. f | < probably smaller doses will induce convulsions if the animal is repeatedly stimulated,* but below this point the medullary effect of tutin is generally alone apparent. * It is well known in New Zealand that tooted sheep may not show any abnormal symptoms if left alone, but if driven they become excited and enter into convulsions. THE PHARMACOLOGICAL ACTION OF TUTU. 293 The course of the intoxication in rabbits after different doses of tutin is shown in Experiment I. gives the effect of a nearly lethal dose, and the following protocols. Experiment II.—0-001 grm. Tutin per kg. Rabbit. Weight, 950 grm. Time. 12.0 12.5 12.16 12.23 12.28 12.35 12.54 12.58 Hm CODD Rt ocownhbwon (Stoo) Heart- | Respira- Tempera- @isereaniane beats. tion. ture. 276 72 391° | Temperature of room 16° C. oe Bac 0°48 c.c. 1 in 500 tutin solution injected under skin of back. 192 132 Slight dyspneea. Lar vessels somewhat dilated. Animal has been resting since injection. 180 144 192 168 ae Lay down and stretched hind legs back. 174 156 37°6° | When lifted up the hind feet clung to the fingers. 198 132 ace Slight trembling occurred at intervals, and terminated in a convulsion two minutes later, which was mainly tonic, and seemed to prove almost fatal. Tremors of mouth, and later of fore limbs, recommenced and gradually increased in intensity. The convulsion lasted two minutes. Afterwards the animal lay on venter with hind legs stretched backwards as before. 240 84 371° | Fine tremors present apparently over whole body. 236 144 ace Tremors more marked ; otherwise same. 216 108 37°6° 276 108 38°1° | Somewhat better. 276 108 387° | Much better; has not moved since last note. 258 84 39°2° | Apparently well. Experiment IJ. the effect of a minimal convulsant dose. Experiment III. shows the result of administering a non-convulsive dose. P refers to the number of heart-beats, and R to the number of respirations per minute; T refers to rectal temperature. ExpERIMENT ITI.—0-0005 orm. Tutin per kg. Rabbit. Weight, 1100 grm. Heart- Respira- Tempera- eames beats. tion. ture. OPsenvations: 12.0 252 66 39°5° | Temperature of room 16° C. 12.4 Ye ne 0:28 e.c. 1 in 500 tutin solution inje-ted under skin of back. 12.10 252 S02 Animal moving about. 12.15 216 156 Appears slightly depressed. ar vessels dilated somewhat. 12.21 192 180 Dyspneea. Heart-beats feel more forcible. 12.28 186 228 bist Seems distinctly depressed, and shows no inclination to move. 12.33 156 228 37°6° | When animal was lifted up the movements of the hind legs were | found to be weaker and inco-ordinate. | 12.44 168 144 Trembling slightly. Ear vessels more contracted. 12.53 156 156 1.3 168 108 38°0° 1.18 204 144 1.33 228 96 38°7° | Trembling still present. | 2.0 264 84 39°6° | Apparently well. | 3.3 252 72 294 PROFESSOR C. R. MARSHALL ON To a rabbit weighing 700 grm. a dose of tutin corresponding to 0:0002 grm. per kg. body-weight was given without any effect beyond a slight one on the pulse, respiration, and temperature. The pulse fell from 228, the rate before the injection, to 192 thirty minutes after the injection; the respiration rose from 54 to 60; the temperature fell in the hour succeeding the administration from 390° C. to 382° C. Three days afterwards a dose corresponding to 0°0003 grm. per kg. body-weight was given to the same rabbit. The heart-beats fell from 276 to 180 forty minutes after the injection, and the temperature from 39°3° to 37°6° one hour after the administra- tion. The frequency of the respirations rose from 60 to 120 in thirty minutes. On Guinea-pigs.—Similar symptoms to those described as occurring in rabbits occurred in guinea-pigs. These animals appear, however, to be more susceptible than rabbits to tutin, as the following experiment shows :— Experiment 1[V.—0-001 grm. Tutin per kg. Guinea-pig. Weight, 458 grm. Time. Observations. 12.35 0°46 c.c. 1 in 1000 tutin solution injected under skin of abdomen. 1.14 | Commenced to jerk head. After this ceased, animal began to move about carefully, the feet appearing to cling to bench. 1.28 | Coarse tremors of head, which extended to whole body. 1.324 | Convulsion commencing in fore part of body. The animal then rolled over several times, and finally stood up on toes and remained almost in a tonic spasm. The convulsion lasted 90”. Afterwards the animal moved about carefully and somewhat stiffly. _Convulsion similar to previous one ; lasted one minute. Clonic movements; these caused the animal to run about the bench. They were followed by tonic contraction of the head muscles—eyes closed, ears drawn back, ete.—and by further clonic movements of limbs. On taking up the animal it entered into a tonic convulsion which lasted about a minute, and then the animal died—there was sudden and complete relaxation of the muscles, but the heart beat slowly for nearly a minute. Frogs.—In frogs also the most prominent symptoms are tonic and clonic convulsions. After a short period, during which the animal is apparently normal, it appears to be depressed and weaker, and gradually sinks down on the bench. The respiratory movements are usually deeper. If a large dose has been given, repeated opisthotonic convulsions, sometimes accompanied by a cry apparently resulting from spasm of the laryngeal muscles, follow. The convulsions may come on suddenly or may commence in retraction of the head, or, more rarely, in spasm of the hind legs. In the intervals, if these are short, the animal lies exhausted, but if of long duration it may move inco-ordinately and assume bizarre positions. Usually exhaustion supervenes and the animal appears dead. After smaller doses the convulsions tend towards a clonic type, although with freshly caught frogs, transient opisthotonos, more rarely emprosthotonos, is common even with small doses. In frogs kept in confinement for some time, the convulsions develop in a more definite manner—the eyes are closed, the head retracted, then the fore limbs and lastly the whole body (opisthotonos) are affected. THE PHARMACOLOGICAL ACTION OF TUTU. 295 EXPERIMENT V.—O0-066 mg. Tutin per grm. Frog (&. temporaria—fresh caught). Weight, 15 grm. Time. Observations, 11.43 1 c.c. 1 in 1000 tutin solution injected into dorsal lymph-sac. 11.55 Sunk on to venter ; creeping about. 11.56 Assumed bizarre position, hind legs being stretched out at right angles to body. 11.57 Very transient opisthotonos. 12.0 Blowing on the animal produced transient opisthotonos. Afterwards the frog tried to walk, but was unable to raise itself from the ground, hence it merely slid along on its venter. Hind-leg movements very stiff. 12.5 Still sliding along ground for short distances. 12.10 Made a few short jumps (clonic convulsion) ending in a slight opisthotonic attack. 12.11 Another similar attack. 12.12 Severe clonic convulsion (back arched) with cries. Belly appears blown out. 12.13 Repeated clonic convulsions without cries. The frog remained on its back after the attacks ceased. 12.15 Slight movements—mainly attempts to turn over—with mild cries. 12.16 Placed on venter. At once assumed a peculiar position closely resembling emprosthotonos. 12.18 Made few attempts to spring, croaking each time. Whole body presents a blown-out appearance. 12.20 Moving about, but movements inco-ordinate. Croaking. Eyes opened ; corneal reflex sluggish. 12.25 | Placed on back. Remained motionless at first, then made unsuccessful attempts to turn over. 12.35 Has not succeeded in turning over. Placed on venter. 12.40 Moving about a little, with nose on ground. Movements inco-ordinate and apparently ex- hausting. Lymph-sacs blown out. 1.0 Has been quiet since last note; now made repeated futile attempts to move. Presents a less blown-out appearance. Placed on back. 2.0 Quiet. Has made a few attempts to turn over. Placed on venter. 3.0 No distinct improvement. Has moved inco-ordinately at times. Pithed. All stiffness im- mediately disappeared. Heart beating well. Trout and Tadpoles.—Trout, a few weeks old, when put into 1 in 1000 tutin solution, showed tremors of the tail in eleven minutes, marked twitches in thirty minutes, and tetanic convulsions in less than an hour. Tadpoles were affected similarly. AcTIoN oN MEDULLARY CENTRES. The protocols given on page 293 show that in rabbits tutin diminishes the rate of the heart-beat and increases the frequency of the respiration, and that the two effects are in each case nearly reciprocally proportional. This relative effect is seen better when the experiments are plotted as graphs (figs. 1 and 2). Fig. 1 shows the action of a minimal effective dose of tutin, and demonstrates the slightly irregular reciprocity in the effect on the pulse and respiration which is invariably seen after such small doses. After moderate doses of tutin the curves of frequency of respiration and pulse become more uniformly reciprocal—in the graph shown (fig. 2, Experiment II.) each curve is almost a mirror curve of the other. This greater uniformity in the case of moderate doses is probably due to the greater stimulus assuming more effective control of the medullary centres, thus rendering them less liable to accidental variation from without. That this influence on the heart-beat and respiration is due to an action on the medullary centres TRANS. ROY. SOC. EDIN., VOL, XLVII. PART II, (NO. 13), 45 296 PROFESSOR C. R. MARSHALL ON is suggested by the curves, and is supported by the fact that division of the vagi annuls the action of tutin on the circulation but not on the respiration. Systematic observations of the respiration and pulse have not been made after the administration of large doses mainly owing to the fear of exciting convulsions by taking the pulse ; but from the occasional notes made, it would seem that the increase in the frequency of the respiration and the diminution in the rate of the heart-beats are not proportionately, and may not be actually, as great after large as after relatively small doses. Heart Resp. EE soot | | I Ben aAe Ro Sale oes SERRE DEERE APPA esa pL eee i eT ig a peed aca ae tT Medea Tae T 1 ede er Tl op Hf ER ee ae EH AMEE REP SORA Sees sews calla —| 100 WEE SEE LEE nEnEneeanennneeee Hl) a eae ae DAE Reese ees SSE Seas sl cE Sg US ee 1 EZ See Us Pee Reese See eeSsee eke ee SEE EERE ee TST AND Sea PVE SS TST LS pa Et et fo ccf 12 IN Tages cal N72 a el cal ni a IRD BERR ET EERARPERE TP AnD ADRES oes Be TTS T AN AUCH ee 2a TS 7 a A Sa: Sannns cep desen cee 0h a rnp anaes Enea aaeeeae 70 oP SENS a Boose =D Cee BARBS eames we AIMS! See siete sea esse eeeeoso BREE RE EEE EEE EE HE ob PPS ZED TPCT SN PSS ST i H+ + +++ SENDERS TASES eo Minutes. 40 60 80 100 120 140 160 Fic. 1.—Diagram bt ale effect of 0°3 mg. tutin per kg. body-weight on the frequency of the respiration and the heart in a rabbit (weight, 750 grm.). Continued line = number of heart-beats per minute. Broken line = number of respirations per minute, During deep anzsthesia the reciprocal relationship between the frequency of the respirations and the heart-beats after tutin is lost. In this condition the administration of tutin usually diminishes the number of respirations per minute, while the circulation remains unaffected or is affected to a relatively slight degree. With chloroform as the anesthetic, slowing of the respiration after the injection of tutin was invariable. When ether was employed, increased rapidity of the respiration followed if the anesthesia was light, and slowing if the anesthesia was deep. The following protocol shows the effect of tutin on the blood pressure, pulse rate, and respiration during chloroform anesthesia :— THE PHARMACOLOGICAL ACTION OF TUTU. 297 Expertment VI.—Rabbit. Chloroform. Blood Pressure from Right Carotid Artery. Respiration recorded by Head’s Method. Ti Blood Pressure | Pulse Rate | Respirations ime. i ” ” Remarks. in mm. per 10”. per 10”. 3.34 61 27 14:3 3.35 ae se a 0:00025 grm. tutin injected into left ext. jugular vein. 3.40 61 24°5 13:7 3.45 61 24 138 3.50 62 23 13 3.53 65 2255 11-5 3.55 66 23 8 3.57 65 23°2 10°8 4.5 66 22 16 4.7 a he a. 0:00025 erm. tutin injected. 4.12 65 22 13°5 4.17 65 21 13 4.22 65 21 12°3 4.25 63 21 12°3 4.27 57 20 10-7 4.30 52 20 13 Subsequently, a small dose of coriamyrtin (0°0001 grm.), and afterwards two doses of tutin (0°0005 grm. and 0:001 grm.), were injected. In each case the blood pressure and pulse rate fell slightly, and the fre- quency of respiration very distinctly. : Besar +H iy } aCe eee xs | | |} 50 3 50 es a EME BEE Z Ld i Pisis a [SIs SELES cae SREB Pas HEE ba eitel fi i Minutes 0 10 20 Fic, 2.—Similar diagram showing effect of 1 mg. tutin per kg. body-weight up to occurrence of convulsion. Rabbit, 950 grm. 298 PROFESSOR C. R. MARSHALL ON The slowing of the respiration produced by tutin during anesthesia is not altered by division of the vagi. In a rabbit anzsthetised with chloroform the injection of 0°0005 grm. tutin intravenously before division of the vagi reduced the number of respirations from 115 to 60 per minute; after section of the vagi the same dose caused the number of respirations to fall from 85 to 50 per minute. If the brain is excised and the administration of the anesthetic afterwards stopped, the injection of tutin produces an effect similar to that observed in an uninjured animal. The action of a large dose up to the onset of convulsions is seen in the follow- ing experiment :— Expermment VII.—Rabbit. Ether. Section through Posterior Part of Mid-brain. Brain above Level of Section removed. Anesthetic stopped. Blood Pressure from Right Carotid Artery. Respirations counted. Time. Blood Pressure Pulse Rate per 10”. Respirations in mm. per 10°. 3.30 65 44 9 3.35 0:01 grm. tutin injected | into right facial vein. 3.36 al 44 3.37 83 38 11 3.38 88 34 12°5 3.40 93 36 15:5 3.42 96 36 17 3.434 Clonic convulsions. The effect, particularly on the pulse rate, is much less than in a normal animal, but this is probably due to the medullary centres being still influenced by the previous administration of the anzesthetic. In a normal animal the dyspncea produced by tutin is largely inspiratory in character, and in the slow respiration occurring during anesthesia it is the inspiration which is mainly affected. It is not only increased in depth, but, when completed, is also maintained for a relatively long period. The effect is shown in fig. 3, which also illustrates the action of an allied substance, coriamyrtin, injected subsequently. The latter tracing further illustrates the irregularity of the respiration which not infrequently occurs after the administration of these substances. ‘The respiration in this experiment and in most others, owing to the late appearance of the effects, was recorded by the diaphragm method described by Hkap; but the same result was obtained when a record of the respired air was taken by connecting the tracheal cannula with a tambour. The slow breathing produced by moderate doses of tutin or coriamyrtin can usually be abolished by pushing the anesthetic, but if a large dose has been ad- ministered deeper anesthesia causes further slowing, which is quickly succeeded by cessation of the respiration. If constant anesthesia is maintained, the respiration after a variable period generally assumes its previous frequency ; but a certain degree of 299 THE PHARMACOLOGICAL ACTION OF TUTU. *soUIORI} OATZOOdSeL OT} Jo Sutuurseq oyy 4e WMOYS oUt oy} 03 dn ssnip ey4 Jo uorjoalur ayy Aq pezoayeun o10M o1nsserd-poo[q pue uoreitdser oy, ‘spwooes UI eu} = 7 ‘“SUTORI} IOMOT Ul “WM FT ‘Surory seddn ur ‘wut GZ pester oul, eseq {(prjoreo) einssord poojq = g ‘(poyjeur s,peez{) uorerdsur syueserder ayoujs-dn { uoierdsea = yw ‘Ajsnousaerzur untdteri0o “sur 1.0 ‘9g'p ‘Ajsnoueaegur un, ‘sur ¢z.o ‘cee “[OHO ‘Maqey ‘uorestdser oy} uo ulyiAueIIon pue utyNy Jo yooBy—"s “SIA Sy ee | ROE HO BSH wAyabartey mantle Walaa d a SANNA NANA Onan 300 PROFESSOR C. R. MARSHALL ON slowing may reappear for short intervals independently of any obvious convulsive action (fig. 3). Later, if the dose has been sufficient, the respiration becomes irregular in extent, although it usually retains a fairly regular rhythm. ‘This condition may continue for a long period, but finally it leads to deficient aeration and to cessation of the respiration. Variations in the respiratory tracings have occurred. In two instances a regular alternation of shallow with slower deep respirations developed ; in other cases the respirations during the absence of convulsions were characterised chiefly by prolonged inspirations. The influence of the convulsions on the respiration during anesthesia was relatively slight. Sometimes a diminution in the frequency of the respirations accompanied the convulsion, but generally no distinct change in the respiratory rhythm occurred. The convulsions in these experiments were purely clonic in character. During anesthesia the circulation also is very little influenced by the administration of moderate doses of tutin, or even by the convulsions produced. When chloroform was employed as the anesthetic a slight rise of blood pressure and slowing of the heart’s frequency followed the injection of the drug and coincided with the effect on the respiration, but not infrequently neither blood pressure nor heart rate was changed. With ether as anesthetic a rise of blood pressure and a diminution in the frequency of the heart were more common, and were greater in extent than with chloroform. The difference in all probability is mainly due to the lighter degree of anesthesia produced in rabbits and cats by ether. Of the three chief medullary centres, the vasomotor appears to be least susceptible and the respiratory the most susceptible to tutin injected during anezesthesia. THE CONVULSIONS. The convulsions produced by tutin are typically epileptiform in character. After a non-lethal but convulsant dose the earliest manifestations of the convulsant action of the drug are rapid head-noddings, or tremors of other parts of the body, and an increase in tone of the muscles, and especially of the limb muscles. ‘These tremors are followed by clonic movements terminating in a tonic spasm. ‘The clonic spasms commence, in the early convulsions generally, and in the later convulsions always, in twitches about the face and head—in movements of the mouth, or winking of the eyes, or twitching of the ears—which may at first be unilateral, There may be also rotation of the head to one or other side. Then follow clonic movements of the fore limbs and later of the hind limbs, and the termination of the clonic spasms in a tonic convulsion, which may or may not be followed by clonic movements. In some fits clonic movements only occur. After large doses a tonic fit may start the series of convulsions, but if the animal recovers from the early fits clonic movements always form part of the convulsion. Still later the animal loses consciousness, and the periodic fits become replaced by continuous clonic movements of the limbs, which only THE PHARMACOLOGICAL ACTION OF TUTU. 301 terminate with the death of the animal. This may occur from exhaustion, but as the movements may apparently continue for long periods (an hour or more) the animal in these cases was generally killed by chloroform. In the clonic movements occurring in rabbits, both fore limbs and both hind limbs act in concert. The two hind limbs also act symmetrically, flexion and extension movements occurring in the two limbs at the same time. The fore limbs may act symmetrically or alternately. Clonic movements of one limb in a normal animal I do not remember to have seen. In anesthetised animals, however, clonic move- ments of a single limb, and even independent movement of all the limbs, are not uncommon. The development of the convulsions in a decerebrate animal after a moderate dose of tutin is seen in figs. 4-6. The animal (rabbit, weight 1400 grm.) was anzsthetised Pee EEE EE ee 4.53 Fic. 4,—Initial contractions of left fore foot in decerebrate rabbit after tutin. Rabbit, 1400 grm. Ether. 3.25, section just in front of pons; brain above removed; cther stopped. 3.55, 2 mg. tutin intraperitoneally. 4.3, twitches commenced in all limbs. Time in seconds, x # linear. with ether. At 3.25 the brain was cut through at the junction of the mid-brain and pons, and the part above the level of the section excised. The administration of ether was then discontinued. Hooks were put into each fore foot and the right hind foot, and were connected by cords working over pulleys to weighted writing levers in such a manner that the levers trebled the extent of the limb movements. It was found impossible to maintain the two fore limbs in the same relative position to cords and pulleys, and this is the explanation of the complete want of uniformity in the tracing shown in fig. 6. At 3.55, 0°002 grm. tutin (1 c.c. 1 in 500 solution) was injected intraperitoneally. Hight minutes later (4.3), convulsive twitches appeared in all the limbs at once. At first the twitches were single (fig. 4), but later they showed a tendency to doubling, and from this state developed into short tonic contractions lasting one and a half to two seconds with clonic contractions at their height. These occurred every minute for the next few minutes, and were followed by an interval of 302 PROFESSOR C. R. MARSHALL ON one and a half minutes terminated by a few irregular clonic contractions and a longer interval of rest (nine minutes). At 4.21, convulsions similar to those shown in fig. 5 commenced, and continued with great regularity for three and a quarter minutes. (A second dose of tutin, 0°001 grm., was injected at 4.234.) The convulsions then gradually became longer owing to the development of an increasing terminal clonus until, at 4.28, aA Lidl dd ddd dala dbdbelda 4,294 Fic, 5.—Later contractions of right fore limb in decerebrate rabbit aftertutin. Sameanimal. x # linear. they became similar to those shown at the beginning of fig. 6. They were now continuous, a tonic stage with superposed slight clonic contractions of about three seconds’ duration alternating with a clonic stage lasting about fifteen seconds. As this state seemed likely to continue, the effect of ether was tried, and it will be convenient to describe it here. INFLUENCE OF ANASTHETICS ON THE CONVULSIONS. The effect of inhaling ether on the convulsions of the decerebrate animal is shown in fig. 6. At the point marked by the arrow the animal was made to breathe ether vapour of the strength used for inducing anesthesia for operations, and this was continued for eighty seconds. The convulsions ceased almost immediately after the commencement of the inhalation, and very slight contractions recommenced two minutes after its discontinuance. From this point the contractions, which at first were regular in rhythm, increased in intensity, and finally assumed the alternating tonic-clonic THE PHARMACOLOGICAL ACTION OF TUTU. 303 character seen at the commencement of the tracing twenty-two minutes after stopping the ether inhalation. Subsequently the effect of a very small concentration of ether vapour was tried. The anesthetising tube was merely placed on one of the mouths of the Woulfi’s bottle, and a second limb of the tracheal cannula was left open. The effect, although less marked than in the preceding case, was strikingly sudden. Only one tonic-clonic convulsion occurred after the commencement of the inhalation. Within 4,42 Fic, 6.—Effect of inhalation of ether on tutin convulsions in decerebrate rabbit. Same animal as preceding figures. Anzsthetising concentration of ether vapour given at point marked by arrow. & = right, Z = left forelimb. Timein seconds. x # linear. twenty-three seconds true convulsive movements had ceased and become replaced by single contractions, occurring at first in small groups, and later alone. These con- tractions ceased eighty seconds after the commencement of the inhalation, which was continued for two and a quarter minutes. Regular small contractions commenced ninety seconds later, and the typical tonic-clonic convulsions quickly developed and became distinct in one minute. A second similar inhalation of ether produced practically the same result. It is evident, therefore, that the convulsions occurring in decerebrate animals are TRANS. ROY. SOC, EDIN., VOL. XLVII. PART II. (NO. 13). 46 304 PROFESSOR C. R. MARSHALL ON very susceptible to the action of ether. When the cerebrum is intact, ether appears to be less active in arresting convulsions. Even with fairly deep anesthesia, clonic convulsions are produced by relatively small doses of tutin; and after large doses of tutin the convulsions do not cease when the concentration of ether inhaled is increased, until shortly before the respiration is paralysed. As will be shown later, when one cerebral hemisphere is excised the injection of moderate doses of tutin causes convulsions only on the same side; and since the mid-brain and pons are still intact in this experiment, it would seem as if the ether had prevented the development of the convulsions from this part of the brain. The effect of diminishing the quantity of ether administered has been observed in experiments in which the contractions of individual muscles were being recorded. To take an example. With a certain concentration of ether vapour the contractions of the Tibialis Anticus were fairly uniform, and registered 3 mm. in height. After reducing the concentration of ether vapour about 40 per cent., the contractions assumed a more convulsive type in twenty-seven seconds, and many at thirty seconds registered 27 mm., and at fifty seconds 35-40 mm. in height. The weaker ether inhala- tion was continued for sixty-six seconds, when the concentration was raised to its previous level. The contractions commenced to diminish in eighteen seconds, and in forty seconds they had almost reached their previous form. This occurred twelve seconds later. The effect of chloroform on the convulsions was not graphically recorded, except indirectly in tracings of the respiration by Head’s method; but from these and other experiments it appeared to be more powerful, for apparently the same degree of anesthesia, than ether in arresting the convulsions. Muscies AFFECTED BY THE CONVULSIONS. A few experiments were made to determine the way in which various individual muscles are affected by the administration of tutin. They were undertaken primarily with the object of determining the cause of the bizarre positions and curious movements which sometimes occur after tutin. In the experiments the contractions of the following muscles were registered :—Digastric, Mylohyoid, muscles of pinna of ear, Biceps, Triceps, Flexor Carpi Radialis, Flexor Sublimis Digitorum, Extensor Communis Digitorum, Rectus Femoris, Gracilis, Tibialis Anticus, Extensor Longus Digitorum, Flexor Longus Hallucis, Plantaris, and Diaphragm. The muscles of the shoulder and hip were not isolated, but their movements were registered in several experiments by more distally situated muscles which had been isolated. The movements of the muscles of the abdominal wall and the prevertebral muscles were observed but not registered. So far as could be determined, all the voluntary muscles of the body are influenced by tutin, although not simultaneously. The muscles were isolated from the surrounding tissues as far as possible without disturbing their blood and nerve THE PHARMACOLOGICAL ACTION OF TUTU. 305 supply, and their distal tendons were connected by hook or ligature to cords working over pulleys and ending in weighted levers. All these experiments, with the exception of those on the diaphragm, which were made on rabbits, were made on cats under the influence of ether. The records of the different convulsions are far from uniform: no convulsion either in the same or in different animals was exactly similar to another. After small doses, and generally after large doses, the early convulsions commenced, as previously described, in the muscles of the head and neck, and affected the fore limb muscles before those of the hind limbs; but in later convulsions great irregularities appeared, and the posterior limb muscles were not infrequently affected first. Even muscles of the same limb serving similar functions appeared at times to be variously affected, and the form of the convulsions in different muscles was not infrequently diverse. Some of these points are shown in figs. 7 and 8. Fig. 7 illustrates in a marked degree the usual condition of a convulsion commencing in the muscle of the anterior limb before that of the posterior limb, and it also shows, what is frequently the case, the greater tonus in the muscle of the hind limb. The cat received an injection into the peritoneal cavity at 3.25. witches occurred in the Flexor Carpi Radialis at 3.49 and 3.51, and short tonic contractions in the Tibialis Anticus with occasional contractions in the first-named muscle at 3.55. At 4.5, short convulsions, clonic and tonic-clonic in character, and lasting six to eighteen seconds with intervals of three to five seconds, commenced in both muscles and continued until 4.16. A longer interval of rest then occurred, followed by a convulsion similar to that shown. This was succeeded by a rest of fifty-five seconds, which in turn was followed by the convulsion illustrated. Fig. 8 shows a convulsion commencing in the Plantaris earlier than in the Extensor Longus Digitorum of the same limb or the Flexor Sublimis Digitorum of the same side, and it also serves to illustrate the variations in form of the convulsions in different muscles. The experiments seem to indicate a marked loss of co-ordination in the motor areas, and they explain in some measure the irregular movements and want of control seen in the earlier stages of severe tutin poisoning, as well as the irregular clonus of the late stages. In an experiment in which the contractions of antagonistic muscles—viz. Biceps, Triceps, Gracilis, and Rectus Femoris—were registered, inhibition of the extensor muscles occurred during the early twitches of the flexors ; but later, when more prolonged tonic contractions and convulsions developed, no distinct inhibition was present. The muscles of the trunk, shoulder, and hip appeared from direct observation to be more easily affected by tutin than the more distal muscles of the limbs, but the diaphragm seemed less easily influenced. The irregularity of the contractions of the diaphragm seen in many of my tracings was undoubtedly caused mainly by contractions of the muscles of the abdominal walls and the prevertebral muscles, but in some instances MARSHALL ON PROFESSOR C. R. 306 ‘ et ‘Ayjeouojytederqur uryny “su g ‘zg OCT ‘OUI “WIS OGHs “VeVO *spUodas UT OWL, ‘opis eules jo (7) SnoIyUY sIpeIqry, pue (27) stjeipey tdieD 1oxe[q Jo Suotjoe4U0) snosuey{NUIG—" THE PHARMACOLOGICAL ACTION OF TUTU. 307 the diaphragm also appeared to take part in the clonic fit. No prolonged tonic contraction of the diaphragm was obtained or observed. SEAT OF ORIGIN OF THE CoNVULSIONS. The spinal cord is slightly, if at all, affected by tutin. In frogs pithed through the occipito-atlantal membrane, the injection of tutin produced no convulsions; and if the tutin was injected and the con- vulsions allowed to develop before pithing, this operation caused the convulsions to cease, and they did not recur. In rabbits anzesthetised with ether or chloroform, or with the brain above the quadrate bodies excised, section of the spinal cord previous to the injection of tutin prevented the occurrence of convulsions or of increased reflex movements below the point of section ; and section of the spinal cord after the development of convulsions caused complete cessation of these below the point of section, and they did not recur.* * GorrLieB (Arch. f. exp. Path. u. Pharm. vol. xxx. p. 21) has shown that in dogs kept for several days to allow the shock of the operation to pass away, con- vulsive movements can be produced in the hind limbs by picrotoxin after section of 7) ) AY the spinal cord, and that in certain fishes and amphibians and very young mammals convulsive movements can be induced by this drug behind a spinal transection soon after the operation. He failed, however, to get any evidence of spinal stimulation in pithed frogs. See also note at the end of this paper. 4.49 Fic. 8.—Simultaneous tracings of Flexor Sublimis Digitorum (S$), Extensor Longus Digitorum (Z), and Plantaris (P) of same side, Cat, 8250 grm, Ether. Ame x 2 linear. 3.57, 10 mg. tutin intravenously. 308 PROFESSOR C. R. MARSHALL ON In frogs pithed anteriorly to the optic lobes, convulsions similar to those occurring in a normal frog were produced by tutin, but after destruction of these lobes muscular stiffness only resulted. In rabbits and cats which had the brain above the pons removed convulsions were still produced by tutin (figs. 4-6); and when the brain was divided just behind the pons, muscular stiffness and increased reflex excitability were produced by its administration, but less readily than convulsions with the pons intact. With the quadrate bodies in place, convulsive movements seemed to be somewhat more easily induced than when the section was made immediately in front of the pons. One cannot, however, be dogmatic on this point. In an experiment (rabbit, 1575 erm.) in which the brain was cut through in front of the corpora quadrigemina, and the anzesthetic subsequently stopped, the convulsions induced by 10 mg. tutin intraperi- toneally were diminished but not abolished by section between the anterior and posterior bodies (the hind limbs still twitched), but all limb movements ceased after section behind the posterior bodies. There can be little doubt that shock played an important part in producing this result ; but nevertheless it gives some support to the view expressed that the presence of the mid-brain increases to some extent the susceptibility to tutin. Notwithstanding the susceptibility of the lower centres of the brain to this substance, the cortical centres are probably mainly affected under ordinary circumstances. This is suggested by the following experiment. A cat (1750 grm.) was aneesthetised with chloroform and the anesthesia maintained with ether, and at 3.30 the left cerebral hemisphere was removed. The fore limbs were connected by cords working over pulleys to weighted levers which registered their movements. At 4.0, 0001 grm. tutin, and at 4.16, 0°0005 grm. tutin, were injected intravenously. Slight movements of the left fore foot occurred at 4.19; and at 4.20 short tonic contractions with superposed clonus developed. These continued at irregular brief intervals for seventy seconds, when a well-marked and prolonged tonic-clonic convulsion occurred, which gave place to the convulsions shown in fig. 9. These continued until 4.29; afterwards, intervals of rest, at first of fifteen seconds and later of longer duration, separated the convulsions, which also became more severe in type. The animal was finally killed with the anzes- thetic. The convulsions on the left side were identical in character with those observed in other experiments with the cerebrum intact. No convulsions and no movements occurred on the right side. Some slight movements were recorded from this side in the earlier convulsions (see fig. 9), but these were found to be transmitted from the left side. When the shoulders were steadied, no movement was recorded on the right side and no contractions were felt. In view of the fact that convulsions may readily be produced by tutin after removal of both cerebral hemispheres, the absence of convulsions on the left side in this experiment can only be explained by assuming a greater susceptibility on the part of the lower centres affected by tutin to anesthetics. This susceptibility has been shown to exist (p. 302). THE PHARMACOLOGICAL ACTION OF TUTU. 309 The chief seats of origin of tutin convulsions in rabbits and cats would therefore appear to be the cortical centres and the pontine nuclei. The share which each of these centres takes in producing the convulsions in unanesthetised and unmutilated animals is difficult to prove; but from the nature of the anxiety of the preconvulsive stage 4,27 Fic. 9.—Effect of tutin on fore limbs after excision of left cerebral hemisphere. Cat, 1750 grm. Ether. 3.80, left cerebral hemisphere removed. 4.0, 1 mg. tutin intra- venously. 4.16, 0°5 mg. tutin intravenously. The small contractions of right fore limb (2) were transmitted from left shoulder muscles. Timein seconds. x # linear. and the character of the early convulsions I am inclined to believe that the earliest action is upon the cortical centres. The continued clonus of the late stages, which is associated with complete unconsciousness, is in all probability due to the activity of the lower centres. ACTION ON THE HIGHER CENTRES. Owing to the ease with which reflex phenomena are obtained after the administra- tion of tutin, it is impossible to say definitely, from the observation of rabbits, that tutin in small convulsant doses has an action on the sensory and other higher centres. Decided depression and even sleepiness are seen in these animals soon after the injection of such doses; but, although [ am of opinion that this condition is the result, in part if not wholly, of an action on the higher centres, I was unable to convince myself that it is not secondary to the changes in the respiratory and circulatory systems, or due to a desire to keep the muscles at rest as long as possible. It is noteworthy, however, that the same degree of depression was not noted after coriamyrtin, which acts more rapidly but otherwise produces the same kind of convulsions as tutin; and it is not seen after strychnine when given in sufficient doses to cause convulsions. After the administra- 310 PROFESSOR C. R. MARSHALL ON tion of a lethal or nearly lethal dose of tutin to rabbits, a comatose condition usually develops in the later stages of the intoxication, and is in all probability a direct effect of the poison on the higher centres. The most convincing evidence of the depressant action of toot poison comes from cases of accidental poisoning in men. In all the cases of recovery described there has been no remembrance of eating tutu berries or of the convulsions which followed, and in some patients normal memory is said not to have returned for many months. Dr GILEs, in a letter to Professor EASTERFIELD and Mr Aston, describes the case of a soldier who suffered from violent convulsions, which he successfully treated by chloroform inhalations. ‘The next morning the man was sore from the violent strain the muscles had undergone, but was otherwise quite recovered, with the single exception that his memory was totally obliterated. The past was a perfect blank to him: he did not know his own name, who he was, or where he came from. This condition gradually passed away, and in a day or two he was as well as ever.” Another example cited by the same medical man is that of two men who, having lost their way in the bush, had eaten tutu berries. One died; the other was found gesticulating and behaving like a maniac. When rescued, ‘‘ his memory was said by the witnesses to be a blank, and it only returned gradually.” He also states that the late Dr Cari FiscHer, who had commenced to investigate the poison, “ had been obliged to discontinue his experiments because his memory became so impaired that he found himself constantly forgetting his appointments.” Somewhat similar symptoms are described by Curistiz* as the result of an experiment on himself. After taking 84 grains of an extract (representing about 800 grains of leaf) in two doses, vomiting commenced about four hours after the first dose, and slight twitches were felt in the arms and legs two hours later. A convulsion which was noted two and a half hours later was not remembered by him, but half a grain of morphine had been administered to stop the vomiting one and a half hours before. After recovery, “sensation and discrimination were dulled,” and a month later pareesthesia and numbness were felt in the fingers and toes. These last-named symptoms continued for a month. There appears to be little doubt, therefore, that tutin, even in relatively small doses, exerts a decidedly depressant effect upon the higher functions of the brain. INFLUENCE ON TEMPERATURE. In my experiments with rabbits, tutin invariably caused a fall of the body temperature, and after non-convulsant or small convulsant doses the fall was in most cases roughly proportional to the dose administered. This is seen in the following graph, more especially in the curves plotted for 0°5 mg., 1 mg., and 1°5 mg. per kg. body-weight respectively. The remaining curve of 0°3 mg. per kg. body-weight shows a greater proportionate effect than would be expected of this dose, but this is due to the * Loc. cit. THE PHARMACOLOGICAL ACTION OF TUTU. 311 fact that a younger animal was used for this experiment than for the others represented in the figure. The temperature appears to be more easily affected by tutin in young than in adult or old animals. After lethal doses of tutin the temperature in the few cases I have investigated fell from the first, and showed no tendency to recover, notwith- standing the continued convulsions. A fall of temperature has been shown by Harnack * and his pupils to occur after the administration of most convulsants. Harnack+ was inclined at first to the view that they acted by stimulating the inhibitory mechanism of the heat-regulating centre in the brain, but later, as the result of further investigations with sodium santoninate and santonate,t he came to the conclusion that the antipyretic action of these Temp. me | | 39 37 Mins. 30 60 90 120 150 180 Fic. 10.——Influence of tutin on temperature of rabbits. I = 0°3 mg. per kg. body-weight ; II = 0°5 mg. per kg. body-weight ; III = 1 mg. per kg. body-weight ; IV = 1°5 mg. per kg. body-weight, administered hypodermically. convulsants is due to loss of heat owing to dilatation of the cutaneous vessels. This dilatation is brought about in all probability by stimulation of the vaso-dilator mechanism in the medulla. He does not think it follows that all convulsants which cause a fall of the body temperature act in this way, but he withdraws his former explanation of their mode of action. More recently Hayasut§ has shown that convulsants temporarily reduce the fever caused by heat-puncture, and in consequence he concludes that the action of these substances is a depressant one on the heat- regulating centre. Part of the evidence brought forward by Harnack in support of dilatation of the * Archiv f. exper. Path. u. Pharmak., xxxviii. p. 397 [1897]; xl. p. 151 [1897]. + Ibtd., xxxviii. p. 421; xl. p. 152. t Ibid., xlv. p. 272 [1901]. § Ibid., 1. p. 247 [1903]. TRANS. ROY. SOC. EDIN., VOL. XLVII. PART II. (NO. 13). 47 312 PROFESSOR C. R. MARSHALL ON vessels being the cause of the fall of temperature is the appearance of the vessels of the ear. ‘These were seen to be dilated when the temperature was falling, and contracted when the temperature was rising. My observations after administering tutin largely confirm those of HaRNack with sodium santoninate on this point. Unfortunately my experiments were made before the publication of his paper, and | did not pay special attention to this connection, but, where noted, dilatation of the vessels of the ear occurred during the fall and contraction during the rise of temperature. In a few instances the ear vessels were noted to be more contracted before the rise of tempera- ture commenced. Errect oF DrEcomposITION PRODUCTS. The products of hydrolysis of tutin possess no peculiar pharmacological action. Insuttcient hydrolysis yields a product with a variable tutin action dependent upon the undecomposed tutin. After heating with excess of 0°2 per cent. hydrochloric acid for fifteen minutes on a water-bath, neutralising with sodium bicarbonate, evaporating, extracting with alcohol, and re-extracting the inspissated alcoholic extract with ether, a colourless syrup was obtained which gave the physiological action of tutin, and from which crystals of tutin separated later. After heating the hydrochloric acid solution on the water-bath for an hour and extracting in a similar manner, a colourless syrup was obtained which did not deposit crystals of tutin, but which caused tutin convulsions after administration. On heating a solution of tutin in 1 per cent. hydrochloric acid on the water-bath for three hours, neutralising and extracting as before, a reddish oil insoluble in water and a brownish substance soluble in water were obtained, but neither produced any physiological effect. The sugar produced by hydrolysis has not been identified. I can corroborate HASTERFIELD and Aston’s* statement that it does not form a crystalline osazone. In itself, however, the sugar is of little pharmacological importance. Dilute solutions of alkalies appear to decompose tutin much more quickly than dilute acids. A solution of tutin in 0°1 per cent. sodium hydroxide, and one in 0:05 per cent. calcium hydroxide, after raising to the boiling-point, cooling, and neutralising, produced no pharmacological action in doses which should have given, if any active principle had been present, a marked result. COMPARISON WITH OTHER CONVULSANTS. The best-known of the substances comparable with tutin are picrotoxin, coriamyrtin, and essence of absinthe. Many others cause convulsions by acting on the brain, but most of them produce other actions not given by tutin. In so far as their convulsant action has been investigated it would seem not to differ materially from that of the substances mentioned. * Loc. cit. THE PHARMACOLOGICAL ACTION OF TUTU. 315 The form and course of the convulsions produced by the first-named substances seem to be the same. Even somersaulting is a symptom common to all. The main differences are in the time of appearance of the convulsions and the duration of the intoxication. Absinthe causes an earlier appearance of convulsions, but after non-lethal doses a shorter intoxication than tutin; and the same is true of coriamyrtin and, to a less extent, of picrotoxin. The two last-mentioned substances are more powerful than tutin, in that they produce convulsions with smaller doses. This is shown in the following comparative experiments on frogs. The frogs were of the same weight (15 grm.), and the substance, dissolved in 0°6 per cent. sodium chloride, was injected into the dorsal lymph-sac in each case. The figures given indicate the number of minutes between the injection and the onset of the first distinct convulsion. Dose. Coriamyrtin. | Picrotoxin. Tutin. 1 me. 1 7 15 0-5 mg. 3 06 38 0:08 mg. 10 19 No convulsion. 0:04 mg. 40 No convulsion. No convulsion. The substance I have compared most frequently with tutin is coriamyrtin. This principle was isolated by Ripan* from a species of Comaria (C. myrtifolia) which is common in the South of France and other coasts bordering the Mediterranean. The relative effects of the two substances will be seen in the following brief protocols of experiments on the same animals. One set gives the results of hypodermic and the other of intravenous administration. A period of fourteen ‘days in the first case and five days in the second occurred between the two experiments. Rabbit, 2000 grm. Time Time after | Remarks. after Remarks, Injection. Injection. 0 _ 0°75 mg. coriamyrtin hypodermically. 0 2 mg. tutin hypodermically. 4’ _Dyspneea. Head tremors. 14’ Dyspnea. 5 | Clinging to table. 60’ Had no convulsive symptoms beyond o Tonie and clonic convulsion lasting head tremors. one minute. 64’ 2 mg. tutin hypodermically. No further convulsions occurred. 653’ Clonic convulsion. | The animal rapidly recovered from 90’ Slight clonic convulsion. | the intoxication. 107’ Clonic-tonic convulsion. 108’ Convulsive movements of head, 110’ Opisthotonos. 125’ Tonic clonic convulsions. 135’ Chloroformed. * Recherches ecpérimentales sur le principe toxique du redoul, Paris, 1863. 314 PROFESSOR C. R. MARSHALL ON Rabbit, 950 grm. R=number of respirations per minute. T= rectal temperature. Time Time after Remarks. after Remarks. Injection. Injection. 0 R60. = 297e 0 R=48. T3975: 144 mg. tutin into marginal vein of 0:24 mg, coriamyrtin into marginal vein ear. of ear. 4’ Breathing more rapidly. i Very rapid breathing. 5’ R= 108. 1’ R= 204. 6 R=168. MHead-noddings apparently 5’ R=204. Head-noddings. synchronous with respiration. Slip- i Powerful clonic terminating in tonic ping down on venter, convulsion. Lasted one minute. 8’ R=204. Laid on venter with hind Quickly recovered. Then R=84. limbs partially out. 12’ R=204. Some twitching of ears. 25’ Head-nodding less marked, but slight 15’ Slightclonic convulsion of head, followed general tremors, Afterwards assumed by early stage of somersaulting move- characteristic (exaggerated springing) ment. posture. This strained position was 17! Clonie convulsion of head. maintained for one hour. 119° Quiet. R=96. 90’ R= 160. 25’ R= 108. 104’ Moved very carefully for a few steps. 45’ R=60. Not moved since last note. Hind-limb movements somewhat Apparently normal. inco-ordinate. 60’ R60.) l—as.0e 120’ R=108. T=36°8°. Recovering. These experiments show that in similar physiological doses tutin takes longer to act and exerts a more prolonged effect than coriamyrtin, but apart from these differences the two substances produce almost identical symptoms. The difference in time of action is important in relation to the question of tolerance, and it explains in some measure why, on the one hand, a second injection of tutin, administered after a long interval, should so rapidly induce the convulsions which failed to appear as a result of the first injection ; and, on the other hand, why relatively large quantities of coriamyrtin, if administered in divided doses at short intervals, can be given without inducing convulsions. I hope to publish experiments on this point later. SuMMARY. 1. Tutin, the active principle of the toot plant, causes epileptoid convulsions in various classes of animals, After large doses to rabbits the earliest convulsions commence with a general tonic spasm which is followed by clonic movements. After small convulsant doses the convulsions begin with clonic spasms of the muscles of the head. These extend to the fore and then to the hind part of the body, and may terminate in a tonic spasm. The later convulsions after larger doses also assume this form. After the initial convulsions following small doses, somersaulting movements are not uncommon. 2. The convulsions are mainly of cortical and pontine origin. Convulsive move- THE PHARMACOLOGICAL ACTION OF TUTU. 315 ments can be obtained when the brain is divided below the pons, but they are less characteristic and are more difficult to induce than when the pons is intact. In frogs the optic lobes seem to be the most important centres affected. _Unequivocal convulsive movements were not obtained below a section of the spinal cord (see, however, note below). 8. The convulsions arising in the pons are very susceptible to anesthetics. This probably explains why convulsions are limited to one side after excision of one cerebral hemisphere during the continuance of the anesthesia. 4. In unanesthetised rabbits the two fore limbs act together, and the two hind limbs act in concert and synchronously during tutin convulsions ; in the aneesthetised animal the limb movements are often asynchronous. Individual muscles of the limbs often act vicariously, and this probably explains the irregular clonus which the move- ments of the limbs show. 5. Preceding the onset of the convulsions the medullary centres are stimulated. This effect is also produced by non-convulsant doses. 6. All doses which produce an obvious action cause in rabbits a fall of body temperature. 7. When heated with dilute caustic alkali solutions, tutin rapidly decomposes, the product being pharmacologically inactive. Prolonged heating with dilute acids leads to the same result. 8. The substance most closely allied to tutin, namely coriamyrtin, is a more powerful convulsant, and, for similar physiological doses, is more rapid and more transient in action than tutin. Note.—Since the preceding paper was accepted for publication a research on the Physiological Action of Tutin by Fitcuerr and Matcoum has appeared in the Quarterly Journal of Experimental Physiology (vol. ii, p. 335; Oct. 1909); an extended description of the research, with protocols, being given by Fircuett in the Transactions of the New Zealand Institute (vol. xli. p. 286). The effect of tutin on various classes of animals is described, and, in so far as our experiments are common, our results are in general agreement. On a few physiological points we are not in accord. Fircuerr and Matcoitm have apparently not obtained any evidence of stimulation of the cardio-inhibitory centre, but they appear to have investigated the matter only in animals under anesthetics. As I have shown, no decided stimulation of this centre occurs during surgical anesthesia. They also attribute to the motor area a smaller and to the spinal cord a larger share in the production of the convulsions than I have done. They state that after inducing convulsions in a cat under chloroform by a large dose of tutin injected hypodermically and removing the right cerebral hemisphere, ‘the con- vulsions continued just as before, affecting both sides equally,” and after scooping out the left cerebral hemisphere ‘‘the convulsions seemed to be quite unaffected.” ‘The spinal cord was then divided in the mid-dorsal region. The movements below the level of section were absent or abolished at first, but four minutes after the section clonic and tonic spasms of the hind limbs were observed, and these continued intermittently until death ensued, a quarter of an hour later.” The continuance of the convulsions after removal of the cerebral hemispheres is explained by the fact that, according to the protocol of the experiment (No. 152 in Fircnurr’s paper, p. 356), the anesthetic was stopped when the tutin was administered—twenty- two minutes before the appearance of any symptoms. The only difference in our results is the effect obtained behind a transection of the spinal cord. All my transection experiments were made on rabbits, TRANS. ROY. SOC. EDIN., VOL. XLVII. PART II. (NO. 18). 48 316 THE PHARMACOLOGICAL ACTION OF TUTU. but since the appearance of Matcoum and Fitcuert’s paper I have repeated two of the experiments on cats. In both cases the animals were anesthetised with chloroform and the anzesthesia maintained with ether. In one animal 0°02 grm, tutin was injected into the external jugular vein, and five minutes after the hind limbs were seen and felt to twitch the spinal cord was completely divided at the level of the 11th dorsal vertebra. The convulsions in the parts innervated beyond the section immediately ceased, and they did not recur, Apparent convulsive movements in the hind parts appeared, but, as in experiments on rabbits, they were synchronous with the clonic convulsions occurring in the fore part of the body, and by careful palpation were proved to arise in the muscles innervated anteriorly to the spinal section. The muscles of the hind limbs were palpated for forty minutes after the transection, but no twitch was felt in them. In the second experiment the mid-brain was cut across just above the pons, and the brain above the section removed. The administration of ether was then stopped. Twelve minutes later (3.28) 0°02 grm. tutin was injected intravenously. ‘witches of the eyelids and ears occurred at 3.37, and slight convulsive movements of the limbs at 3.42. These quickly became more severe, and at 3.51 the cord was completely divided at the level of the last dorsal vertebra. Slight independent movements of the hind limbs occurred at 3.54, and became more marked and assumed a more convulsive type later. Pressure on one hind foot excited convulsions in both hind limbs. The convulsions, however, were not so characteristic as those in the fore part of the body. The animal died at 4.15. This experiment, therefore, confirms the observation of Mancotm and Fircuerr that tutin acts on the spinal cord in cats; and the first experiment seems to show that this effect, like that upon the lower cerebral centres, is very susceptible to anzesthetics. (207) XIV.—On the Life-history of Hydrobius fuscipes, L. By Frank Balfour-Browne, M.A. (Oxon)., F.Z.S. (With Three Plates.) (MS. received July 6, 1909. Read July 12, 1909. Issued separately February 1, 1910.) In following out the life-histories of some of the water-beetles I have been struck by the remarkable differences in habits and in structure between the members of a family, for instance, between Hydrophilus and Hydrobius, two fairly closely related genera of the Hydrophilide. The description of the life-history of Hydrophilus as given by Micrr, Lyonnet, and others will, only to a small extent, apply to Hydrobius, although the suggestions which I am able to make with regard to certain facts in the life-history of the latter may throw light upon somewhat similar facts in the life-history of the former. Hydrobius fuscipes, L., isa very common water-beetle found throughout Great Britain and Ireland in almost every stagnant piece of water where there is plenty of aquatic vegetation, the favourite habitat appearing to be ditches or ponds where there is abundance of grass growing in or floating out on the water. The beetle is small, measuring about 6 or 7 mm. in length, slightly elongate, of a shining blue-black or brown-black, although often in bright light showing a beautiful velvety-green lustre. The legs in the typical form are reddish brown, the tibize being tipped with darker brown or black. é There is a variety of this species which is either not very common in Britain or of which collectors have not recorded their captures. This is H. eneus of Souter (Ann. Soe. Ent. France, p. 314, 1834), and according to Rye is H. chalconotus of Leacu (1814) and of SrepHens (Mand., ii. 128, 1829) (vide Ent. Mo. Mag., vii. p. 36, 1870-71). Rye records having taken this variety at Barnes (Surrey), and Mortgy, C. (Coleoptera of Suffolk, 1899) records it from “chalk pits at Little Blakenham (Suffolk, H.).” There is a specimen in the collection of the late Mr Arraur Currry which he took in 1887 at Cuckfield Park (Sussex, W.), and recently Mr DonistHorPE took a specimen at Tottenham (Middlesex) (Hnt. Rec., xx., July 1908, p. 184), and he mentions that it has been taken also at Sheppey (Kent, E.) and Woking (Surrey). It is recorded from Upper Teesdale (Yorks, N.W.) (Vict. County History, 1907). Mr T. Srarnrortu tells me it is common on the coast near Hull (Yorks, 8.H.). I took a single specimen at Cambridge in 1904, and it seems to be not uncommon at “The Kinnegar,” Holywood, Co. Down. This variety apparently differs only in colour from typical H. fuscipes, and is of a metallic blue or green on its upper surface, but the brightness of the colouring varies ereatly in different specimens and fades considerably after death. THomson (Ann. de la Soc. Ent. de France, 1883, p. cxxxi.) separated as a distinct TRANS, ROY. SOC, EDIN., VOL. XLVII, PART II. (NO. 14), 49 318 MR FRANK BALFOUR-BROWNE ON species, which he named H. pzczerus, a form which is less elongate and more rotund than the typical H. fuscipes and has thicker and more pitchy tibie. The pubescence of the hind femora also is not quite so extensive, and the punctuation of which it is the accompaniment is not quite so dense and fine (SHarp, D., Ent. Mo. Mag., xxi., 84, 5, 1884-85). The elytral strize are more strongly punctured and therefore appear to be rather deeper than in H. fuscipes, and the punctuation of the upper surface is, as a rule, more coarse. The penis is also certainly broader than in the typical form. Although some Coleopterists have followed THomson in regarding HZ. picicrus as specifically distinct, others, and probably the majority, have regarded it merely as a variety of H. fuscipes. After carefully comparing a number of specimens of H. fuscipes and picicrus, I am inclined to think the latter is not entitled to specific rank, chiefly because there are all grades of intermediate forms between the two extremes. H. picicrus seems to prefer a different environment from H. fuscipes, although there are perhaps few places where either form exists to the complete exclusion of the other. H. picicrus is more common on peat mosses than is H. fuscipes, while it is certainly much scarcer on salt-marsh areas than is the latter. I think also H. fusczpes is less common in the north and prcicrus less common in the south, but of this I am not altogether certain, and even if it is so, the distribution may be owing to the distribution of peat mosses. CHARACTERS OF THE IMAGO. Without entering into a detailed description of the imago, I wish to refer to certain characters which are of special interest. The head bears the two antennze which are of the type found in all the Hydrophilide (fig. 34). They consist of nine segments, the basal one long and slightly curved, the next four short and simple. The sixth from the base is somewhat saucer-shaped, the concave surface being outwards and the attach- ment being central. Above the saucer-shaped segment are two more or less gobular seoments each flattened above, and upon them is the apical gobular segment. The three terminal segments are dark brown in colour and are thickly covered with short stout hairs, while the upper face of the saucer-like segment bears a number of short tubercles from the apex of each of which stands up a long fine hair. The colour of this segment, and of all those below it, is pale testaceous. The antennee are attached to the head immediately in front of each eye, and when at rest lie backwards along the ventral margin of each eye and just below the sides of the head. THe MoutH Parts. The labrum is long and narrow, being slightly indented on the free border in the middle. It can be seen moving up and down when the beetle is feeding. The right and left jaws (figs. 1 and 2; 32 and 33) differ somewhat from one another. They are THE LIFE-HISTORY OF HYDROBIUS FUSCIPHS, L. 319 of the type characteristic of the group, serving the double purpose of tearing and grinding. Seen from above, both jaws show a two-pointed apex, the outer point of the left jaw being slightly thickened and having also a double point slightly marked. On the inner edge of the left jaw just below the apex isa thick tuft of long bristles, and immediately below this tuft is a second series of bristles of which those nearer the apex are longer than those nearer the base. ‘The uppermost bristle of this second series has a thickened base. The inner edge of the right jaw shows only a very small tuft of a few bristles in place of the upper tuft on the left jaw, but the second series is similar in both jaws. Seen from above, the base of each jaw ends in a triangular projection of which the surface is smooth, as is also the margin of that projection in the right jaw, but in the left the margin is “feathered” by a series of short hairs. Seen from below, the jaws do not show much more than when seen from above. They are, however, three-sided, possess- ing not only a dorsal and ventral but also an inner face, broad at the base and narrow- ing to a point at the apex; and the inner face of the one jaw is, of course, opposed to the inner face of the other. The basal part of this inner face of the jaw is the grinding region, the food being brought here by the action of the maxille and the points of the jaws and then ground down between the bases of the two jaws before being finally swallowed. And this grinding apparatus is very efficient, as the two halves of it differ from one another in accordance with different functions. The grinding region of the left jaw is somewhat concave, and has a series of strong ridges running down it parallel to the length of the mandible. ‘hese ridges do not cover the whole of the concavity, but taper off towards the dorsal side of it and give place to what appear to be a number of minute tubercles. The grinding region is fringed all round with short hairs; and it is these which are partly visible in the dorsal view of this mandible. On the right jaw the corresponding region is somewhat convex, obviously to fit the concavity on the left jaw. There are no fringing hairs or deep grooves or ridges, but the whole surface is sculptured like a fine file. In a general way this description would probably apply to the grinding apparatus of all the Hydrophilidze, but there are probably slight variations in the different genera. For instance, in Hydrophilus piceus, L., most of the concavity of the left mandible is file-like in sculpture, the strong ridges being confined to a small portion of the upper border of the concavity. The maxille (fig. 3) are brush-like organs. The galea of each is crowned by five or six rows of stiff hairs incurved towards the mouth. These rows are placed one behind the other and no doubt make a very effective brush. The lacinia also bears a strong tuft of hairs, and there are also stiff hairs on the free edge of the stipes along the upper third of its length. The rest of the free edge of this part gives off a very fine trans- parent membrane as a narrow strip along it. The palpi are long and composed of four simple segments, the basal one being very short. The labium (fig. 4) has the galeze and. lacineze feathered along their free edges with 320 : MR FRANK BALFOUR-BROWNE ON fairly long hairs formimg a more or less continuous row right across the front. The palpi are three-segmented, the middle segment of each bearing a thin row of long hairs up its inner side, while the apical segment bears one or two very long hairs on its outer side about one-third of its length from the apex. When the beetle is out of the water the maxillary palpi are withdrawn and lie folded against the maxille under the sides of the head, the antenne being turned forward and constantly waved about. In the water, on the other hand, the antennz are withdrawn and the maxillary palpi extended, obviously being the subaquatic organs of touch and possibly also of smell and hearing. THe THORAX. Only two or three points about the thorax call for remark. The mesosternum bears in the median line a small pointed projection such as is found in certain other members of the Hydrophilide, e.g. Anacena limbata, ¥. ; but whereas it is in the genus Anaceena useful for distinguishing A. limbata, F., and ovata, Reiche, from A. globulus, Payk., where it is represented only by a small tubercle, it does not help in Hydrobius to distinguish H. picicrus from H. fuscipes. The three pairs of legs have the femora covered with a dense mass of short hairs except in the apical region, which is smooth and shining. On the mesothorax and attached both to the tergite and to the elytra is a pair of small alule. It has been suggested that these organs, which occur in the Dytiscide and Hydrophilide, may be responsible for the sounds which some of the species produce, but this suggestion is discounted by the fact that, although both sexes of Hydrobius JFuscipes possess them, only the male produces a noise. With regard to the abdomen, the only part of special interest for this paper is the sexual armature, especially that of the female. THe Mate SexuaL ARMATURE. The penis (figs. 10 and 11) is of the general form common to all the Hydrophilide, possessing two side pieces or valves which are said to be at least partly for the purpose of retaining the penis within the vagina at the time of copulation (Hmennecugy, Les Insectes, 1904, p. 185). Iam inclined to think that they are rather for the purpose of spreading the walls of the vagina so as to allow the seminal fluid to escape rapidly from the duct, and no doubt the valves also function as a penis sheath when in a state of rest. I have used the term “ penis” in connection with this organ, but it is perhaps open to question whether it is correctly so used. Some authors speak of it as “‘ the edeagus.” There seems to be some difference of opinion as to what abdominal segments are represented in the male of Hydrophilus, and since Hydrobius is very similar to Hydro- philus as to its male sexual armature, the remarks apply equally to both genera. THE LIFE-HISTORY OF HYDROBIUS FUSCIPES, L. 321 Packarp (1898, p. 181), although not describing his views, gives the figures of KoLBE (vide figs. 203 and 204) according to which the sclerites of segment 8 have disappeared, the segment being only represented by a length of connective tissue between the sclerites of seoments 7 and 9. PryrourEAU (1895), on the other hand, only recognises as seoments those regions bearing one or more sclerites, and he considers segment 8 to be represented by both tergite and sternite, while segment 9, the terminal one, is represented only by asternite. Packarp treats PeyrourHAU’s segment 8 as being really the tergite of segment 9 (which he considers has no sternite) and the sternite of segment 10, which has no tergite, and he calls Peyrourgav’s 9th sternite the external lobes of the penis. Hennecvuey (1904, p. 185) has followed Pryrournau, as he states that the repro- ductive organs of the males of Coleoptera are always found upon the posterior border of the 9th sternite, and he gives one of PEyrourEau’s figures. It is difficult to find any justification for the view of Koupe and Packarp, since the connective tissue region which they treat as segment 8 is no better marked than a similar region between their seoment 10 and the sclerites of the penis. There is, however, this to be said for their view, the sclerite they describe as the external lobes of the penis, which is PEyrouREAU’s 9th sternite, certainly functions as part of the penis. In copulation the posterior seoments of the abdomen and also the penis—which in a state of rest are telescoped within the other abdominal seements—are protruded, and the penis, together with the external lobes (Kops), z.e. the 9th sternite (PEYTOUREAU) is bent downwards and to the right or left according to the position of the male on the back of the female, and a distinct kink is visible at the bend. ‘The region beyond the kink, 2.e. penis and external lobes, bends right round until it comes to lie beneath the abdomen; that is, it turns through an angle of 180° and the dorsal side remains dorsal during copulation. Mr DonistHorpz (nt. Rec., xii. 291, 1900) observed the same fact in Hydrophilus prceus, L. THe Femae SexuaL ARMatuRE. (Figs. 12-14.) PryToUREAU (/.c., plate xx.) gives a drawing of the female armature of Hydrophilus in what he describes as a state of activity, and he says (l.c., p. 154) that the 8th sternite “porte a son bord postérieur mamelonné deux apophyses simples, courtes, dirigées en arriere, armées a leurs extrémités de poils sensitifs, cachées par le huititme urite et deux autres apophyses latérales plus longues également terminées par des poils sensitifs” ; and he further says (/.c., p. 155), ‘C'est apres le septiéme sternite que s ouvre Yoviducte simplement membraneux. Larticulation tergo-sternale du huitiéme urite a lieu au point # (plate xx. fig. 8); elle est assez lache, grace & la présence d’une membrane d’union.” If this is a correct statement for Hydrophilus, then Hydrobius appears to differ very decidedly, as, from my observations, it is difficult not to associate the armature with the 8th rather than with the 7th segment.—a condition which, according to PEYTOUREAU, exists in Dytiscus. 322 MR FRANK BALFOUR-BROWNE ON Segment 8 is represented by a partially chitinised tergite, immediately underneath which opens the anus. On either side of this tergite (8’), extending round the sides of the segment, is a narrow chitinous band (y). The two bands extend to the ventral face of the segment and then turn through about a right angle and pass backward, lying more or less parallel to one another. ‘hese chitinised bands do not seem to represent any recular sclerite of the segment, but are rather a specialised carrier for the sexual armature which is pivoted on the apices of the two parallel ventral chitinous rods (a2). The bands are, however, undoubtedly part of the 8th segment, and the part which, in Hydrophilus, PryrourEau regards as the 8th tergite (c), is situated posteriorly with regard to them. The sexual armature consists of two swollen masses (aa) from the upper side of each of which a long tapering filament (bb) about 1 mm. in length projects backwards. These two filaments, which correspond with Pryrourzav’s “deux autres apophyses latérales plus longues . . .”, are the spinnerets, and consist each of three portions, a long basal segment and a short apical one bearing a long fine and tapering hair. On the ventral side of the armature, between the swollen bases of the spinnerets, projects a thin chitinised membrane (c)—(which, from Peyrourgav’s drawing (plate xx. fig. 3, viii.), I gather he regards as the sternite of the segment)—which, in a ventral view, has a deep groove running up its median line on either side of which the membrane fits round the base of a spinneret, the middle region of the posterior free margin projecting into two horn-like processes between the spinnerets and one on either side of the median groove. ‘The posterior free edge of this membrane is fringed with hairs and the two horn-like processes are PryTouREAU’s “deux apophyses, simples, courtes dirigées en arriere,” etc. ; The vagina opens between the spinnerets and just dorsal to this ventral chitinous plate, so that if PeyrourEav is correct in describing this ventral plate as the 8th sternite, the genital aperture is, on his own showing, not between the 7th and 8th segments, but posterior to the 8th. I have said that the armature is pivoted on the two chitinous rods (ax) previously described. When the genital armature is in a state of rest, the swollen bases of the spinnerets are drawn back by muscles attached to them dorsally, by reason of which they turn on the two pivots already referred to, and under these circumstances the spinnerets are concealed for about half their length by the grooved ventral plate which wraps round them, since the ventral plate is fixed and does not move on the pivots (vide fig. 14). When the armature is in a position of activity the bases of the spinnerets are thrown forward on the pivots and the spinnerets thus project farther beyond the grooved ventral plate. The pivots upon which the spinnerets swing are apparently what Pryrourgav describes as “Jarticulation tergo-sternale du huitieme urite. . . .” Prytournav’s figure (plate xx. fig. 1) represents what appears to me to be a dis- section of the parts i a@ state of rest. THE LIFE-HISTORY OF HYDROBIUS FUSCIPES, L. 323 SECONDARY SEXUAL CHARACTERS. There seems to be no character by which the sexes can be distinguished at first glance, in fact the only differences I have been able to detect between the sexes are that the antennz and maxillary palpi of the male, when compared with those of the female, are distinctly stouter. In Hydrophilus the sexes are easily distinguished by the anterior tarsi, which are simple in the female and have a dilated apical segment in the male. In Hydrocharis the male has the claws of the anterior tarsi “sharply bent like a grappling hook,” * but there is no such distinction in Hydrobius fuscipes. Hapits. The food of the imago Hydrobius consists chiefly of plant tissues, although it is also very ready to eat up dead insect larvee, snails, ete. Its favourite food appears to be the filamentous algze and also algze growing upon stems of other water-plants, ete. It also readily devours dead plant tissues, for instance, decaying blades of grass, leaving the fibrovascular bundles, or at any rate stripping them first of the softer tissues. Apparently only when driven to it for want of other food will it attack living higher plants, and then it seems to prefer the epidermis. It was quite extraordinary the amount of filamentous algze which it caused to disappear when kept in an aquarium. During the colder months of the year Hydrobius disappears more or less completely from its usual habitats but, as with regard to other water-beetles, there seems to be no definite knowledge as to where it disappears to. The beetle passes the winter as an imago. I have never found the larve in the autumn, although Mian (1895, p. 93) says that eggs and larvee are to be found in August. I have, however, occasionally found the eggs of Helochares, a related genus, in August, but I fancy that autumn eggs are exceptional and that the larve do not survive the winter. I think that the imago burrows in the mud at the bottom of the pond and passes the winter in the torpid state, but this is only theory. The only evidence I have is that in the aquaria in cold weather very few beetles are visible, and as they cannot get out of the water they must be amongst the stones at the bottom. Also some specimens of Bidessus minutissimus, which I have kept through the winter in a tumbler, have spent the whole time amongst the fine gravel, and I have not seen them once for many weeks. With regard to breeding habits the usual egg-laying period seems to be from March to July, although Mratu (/.c., p. 93) mentions eggs in August. This year I found several egg-cocoons on 21st February, but I think that was exceptionally early owing to the long spell of mild weather in this district (near Belfast). Seeing that the larval period lasts from twelve to fourteen weeks, I doubt if eggs laid later than the end of July would have much chance of getting through before the cold weather, and the absence of larvze during the winter seems further evidence of the limits of the egg-laying period. * Fow.er, W. W., British Coleoptera, i., p. 220. 324 MR FRANK BALFOUR-BROWNE ON I am not sure as to when the males reach maturity, but they are sexually excited early in February. Possibly the females are “fertilised” in the autumn, as, so far as I have observed, they will not take a male at the beginning of the season. The only time I have seen them accept a male is immediately after having completed an egg-cocoon. DonisTHorRPE (/.c., p. 291) refers to the refusal of the female Hydrophilus to accept a male as a case of sexual selection. So far as my experience goes, it is not a question of the choice of a male by the female, but of the male courting the female at the wrong time, and if a female has her spermotheca full of spermatozoa, she will not unnaturally reject any male. Immediately after completing an egg-cocoon a Hydrobius will accept any male which offers, and I expect the same is the case with Hydrophilus. The courtship is a somewhat strange phenomenon, and it is the only period at which I have heard the male stridulate. I think that the males are only excited during the daytime, at least in the earlier months of the year, and require suitable light and warmth to arouse them. For instance, in February or early March in my aquaria, kept in a warm room, there are few beetles visible at 9 a.M., most of them being amongst the stones at the bottom. About 11 or 11.30 the sun reaches the aquaria, and then all the beetles come to the surface and move about rapidly and feed and the males are then excited. If at 9 a.m. the temperature of the water is raised from 50° F. to about 60°, the same phenomena are observed as occur later in the morning under the influence of the sunshine. The male sits on the back of the female rather in the position of a jockey on a race- horse, that is, well forward, his metasternum being about over her prothorax. His head is bent right over hers, and his courtship consists in drumming upon her labium with his maxillary and labial palpi. His maxillary palpi pass round outside the bases of hers. She bends her head downward as if trying to avoid his attentions, and her maxillary palpi wave about. Her jaws can be seen working, and she occasionally succeeds in biting the palpi of the male. The front tarsal claws of the male hold on either in front of the eyes or behind them or even to the eyes themselves; the mid- tarsal claws catch the posterior margin of the female prothorax, while the posterior tarsal claws hold the borders of the elytra. If the female is very annoyed by the attentions of the male she signifies it by swaying rapidly from side to side, or by running quickly over the submerged vegetation, and, as a rule, after a short struggle the male desists. Otherwise she may feed quietly, quite regardless of his wooing. He makes occasional attempts at copulation by rapidly moving backwards and extending his abdominal segments and the penis, but the female usually sweeps the latter away with her posterior tarsi. After making several ineffectual attempts the male usually desists and goes in search of another mate. During courtship the male keeps up a faint but distinct stridulation, reminding one of a corncrake at a great distance. At the moment the noise is produced the ventral side of the abdomen collapses like bellows, as if expressing air, and at each collapse of the abdomen the apex bends downwards. I ean find no stridulating file. The noise is not THE LIFE-HISTORY OF HYDROBIUS FUSCIPES, L. 325 produced by the rubbing of the tips of the elytra against the abdominal tergites, as these regions are quite smooth. The other British species of the genus, Hydrobius oblongus, Herbst, stridulates much more distinctly, and I believe in that case the noise is made by both sexes, and it certainly is not specially confined to the periods of courting. THE SPINNING OF THE Ecace-Cocoon AND OVIPOSITION. The cocoon of Hydrobius differs somewhat in form from that of Hydrophilus, and Lyonnet’s description of the method by which the female of the latter spins her cocoon would not apply to the female Hydrobius. Whereas Hydrophilus builds a more or less spherical cocoon with a short “mast” projecting unsupported from the water, Hydrobius builds a cocoon attached along one side to its support, and with a flat, ribbon-like process extending to the water surface in place of the ‘“‘ mast.” According to Lyonnet (1829-32), Hydrophilus builds her cocoon in two parts ; that is, having built so much, she deliberately turns round and constructs a replica of the first part and then weaves the two parts together. Under normal circumstances Hydrobius builds her cocoon straight through without turning round. The favourite situation for the cocoon is on the under side of a growing blade of grass floating upon the water. The female first takes up a large supply of air so that her subelytral space is full, and she has also a large bubble attached to her under side. She lies lengthwise along the under side of the blade of grass, and, projecting her spinnerets fully, she brings them and the air-bubble ito contact with the grass and the silk begins to flow as a fine thread from the apex of each of the spinnerets. The whole cocoon is formed on the film of the bubble which is on the under side of the insect. The spinnerets work the whole time in the air, pressing against the film and weaving the cocoon on its curved surface. The silk will not form threads in water, but comes away from or adheres to the apices of the spinnerets in the form of whitish lumps. It seems to be unusual for the beetle to build the cocoon far below the water. I tried the experiment of submerging the grass blade while the cocoon was being built. After a time the beetle turned to the side of the blade to reach the surface, but failing, she continued to work for a minute or so. She then made more determined attempts, and as she was in danger of asphyxiat- ing, she very reluctantly broke the air-bubble which united her with the cocoon and reached the surface. Having renewed her supply she returned to the cocoon, and by moving her abdomen over it endeavoured once more to take up her work, but after two or three failures she gave it up and later started a new cocoon nearer the surface. The silk threads are first spun backwards and forwards across the grass blade and the insect then begins to spin on the film immediately behind her and so raises a wall which soon takes the shape of a blunt cone. She gradually moves forward, spinning all round the film, and thus extending the cone. She never turns over, but continues facing the same direction all the time, occasionally turning to the side of the grass blade and TRANS, ROY. SOC, EDIN., VOL, XLVII. PART II. (NO. 14). 50 26 MR FRANK BALFOUR-BROWNE ON oo elevating her head to the surface and renewing her air-supply. She also occasionally slightly raises her elytra and separates them and then closes them down again, thus enlarging her ventral bubble from her subelytral air-supply. The building of the body of the cocoon occupies an hour or more, but oviposition always commences before the body is complete and often quite early in the building process. I removed one female after she had been spinning for twenty minutes, by which time she had only formed a small cone of silk and she had already deposited three eggs. An ege passes rapidly from the oviduct, scarcely interrupting the spinning. It passes out between the spinnerets and rubs against the fringing hairs of the grooved ventral plate. The spinnerets deliberately place it in the cocoon, and then rapidly spin threads backwards and forwards over it, fastening it down. The spinnerets are remarkably sensitive. On several occasions I ripped gashes in partially formed cocoons and the spinnerets quickly turned in the direction of the rent and closed it up. The number of eggs laid in a cocoon is from thirteen to nineteen, and possibly may vary beyond these figures. Once they are laid the female spins a layer of silk over the top and then continues the sides of the cocoon, leaving a certain amount of space unoccupied by eggs but filled with air. From time to time she squeezes the cocoon with her hind tibiz, shaping it ; and, as a rule, this squeezing is specially noticeable where the cocoon ends off with a more or less flattened surface rather like a lid in appearance. Once this flat surface is complete the female continues to spin a narrow band on the air-bubble, every now and again pressing it against the grass blade and thus attaching it. She carries this ribbon to the edge of the blade and then spreads it out on the water and on the upper surface of the grass. Its form and length depend upon the position of the cocoon with regard to the surface of the water, and sometimes, instead of being spread out on the surface, it is carried up the support into the air, but under normal circumstances the ribbon always reaches the surface. As to the process of spinning, the silk comes off from the apex of each spinneret as a fine thread, and the two spinnerets work side by side backwards and forwards, attach- ing the threads at the end of each stroke. But although the silk thread comes from the apex of the spinneret there is not, so far as I can discover, any canal running up within that organ. The long apical hair, although not solid, has no aperture from which the silk could flow, and the apical and basal segments are, I believe, filled up with tissues. So far as I can make out, the silk flows along the outer surface of the spinnerets, which appear wet all the time that spinning is going on. During spinning the spinnerets are fully extended and the grooved ventral plate fits around their bases. The fringe of hairs on the free edge of this plate appears, like the spinnerets, to be wet, and I think the silk fluid flows off the two horn-like processes of the plate which are pressed between the spinnerets and runs out along these. I found that once the beetle had fairly started making the cocoon it was possible to lift the grass blade completely out of the water and even to place it under the micro- THE LIFE-HISTORY OF HYDROBIUS FUSCIPES, L. 327 scope without disturbing her. ‘The cocoon is formed just as easily under these circum- stances as under water, provided that the temperature is not too high or as long as there is sufficient moisture in the surrounding air. I found by placing a spinning female in the direct sunshine that, after a short time, the spinnerets, although continuing to work backwards and forwards, produced no silk. At first the silk spun became thicker than usual and then it became lumpy and stuck to the spinnerets, and I think this was due to the heat drying up the silk fluid on the spinnerets. It must be admitted that if this explanation is correct it is difficult to explain how, when the beetle is upside down and working on the under side of the grass, the silk runs out the length of the spinnerets against gravity ! About half way up the basal segment of each spinneret and on its inner face is a transparent region of more or less oval shape. This region is beyond the “horns” of the grooved ventral plate when the spinnerets are extended, but is completely covered by them when these organs are withdrawn. Under high magnification (x 400) this region shows a structure different from that of the rest of the spinneret, and I at first thought that the ducts of the silk glands might open here, but I cannot detect any pores. I think therefore that the silk glands must open to the exterior somewhere below the fringe of the ventral plate, that the silk flows out on the hairs of this and passes on to the spinnerets from them. The posterior end of the abdomen of the female is extremely flexible during the spinning of the cocoon, segment 7 moving very freely upon segment 6, and the spinnerets also waving about with independent movement. As arule, shortly before a female has completed her cocoon, a male takes up his position on her back and commences stroking her labium. Within two minutes of completing her cocoon the female accepts the male and copulation lasts about a minute. The male ceases to stridulate the moment he begins to protrude the penis. THE DEVELOPMENT OF THE Hac. (Figs. 15-31.) The development of the egg of Hydrophilus has been described by several authorities such as HeIpER, GRABER, KowaLevsky, etc., and that of Hydrobius appears to be very similar, but Hydrophilus has been studied chiefly from the histological point of view, and I propose to describe the development of the embyro of Hydrobius as seen from day to day through the shell of the living egg. There are certain points which are not referred to by the above authors which are of-some interest. The egg of Hydrobius is oval in shape, from 1°1 to 14 mm. in length, and bears at one end a small pedicel, this being the end of the egg at which the head of the embryo develops. The newly laid living egg shows no structure under the microscope beyond a shell membrane and granular contents which completely fill the shell. After twenty-four hours the contents have shrunk so that there is now a broad, perivitelline space, and on the contents there are at this time certain irregularities which suggest that cell-division 328 MR FRANK BALFOUR-BROWNE ON has commenced. By careful focussing and manipulation of the light a median groove, the “ primitive streak,” is to be seen, which extends for about half the length of the ego from the posterior end. On the third day the groove is more visible, and at the posterior end of the egg the commencement of the amnion and the amniotic cavity is apparent. There are also indistinct traces of segmentation of the yolk, and twenty-four hours later these have developed into complete segmentation. By this time—the fourth day—the amniotic fold has extended forwards over the median groove, but has not yet completely enclosed it, so that the blastopore is to be made out over certain irregular outgrowths which constitute the beginnings of the embryonic head. The groove has extended round the posterior end of the yolk and along the dorsal side so that the “tail” end of the developing embryo lies almost over the head. Indistinct traces of somites are visible in the embryo which, by transmitted light, is more transparent and _ less granular than the yolk. On the fifth day the form of the embryo is quite distinct, the yolk having consider- ably diminished. The embryo lies round the yolk, its ventral side towards the shell. The appendages of the head, thorax, and abdomen are recognisable as buds. It is to be noticed that the tail end of the embryo does not now extend quite so far round the yolk as it did in the previous stage. On the sixth day the mouth parts, the maxille, and the labium (which at present consists of two separate parts) have folded on themselves and the maxillary palpi are recognisable as buds. The embryo has still further contracted in length. The legs, although recognisable, are at present only partly defined, but on the seventh day they are more distinct. The segmentation of the abdomen, which on the sixth day has only just commenced, is also more distinct on the seventh day, and the embryo is still shorter in length. In the fifth-, sixth-, and seventh-day stages the embryo is seen to be surrounded by a membrane—the amnion—within the shell. On the eighth day this has disappeared—it having burst along the ventral side and shrunk back to lie as an irregular double ridge on the dorsum. It goes by the name of “the dorsal organ” or “dorsal tube.” The embryo has now contracted so much that it lies straight out within the shell. At this stage the eyes first become visible as six small pigment spots on each side of the head. The labium is now a lilobed structure lying backwards on the ventral side of the head, and the maxillee lie in the same direction on either side of it. During the next twenty-four hours the abdomen begins to lengthen out, but this time it curves ventrally instead of dorsally. The legs, which up to now have lain close together in the median line, have separated, leaving a space between those of the two sides of the body. The “dorsal organ” has by the ninth day completely disappeared, having sunk into the yolk and been absorbed. As soon as the dorsal organ disappears the dorsum of the embryo closes completely over the yolk, which up to that time has remained exposed. The trophi have lengthened and show traces of segmentation. THE LIFE-HISTORY OF HYDROBIUS FUSCIPES, L. 329 Within the posterior abdominal segments are traces of the lateral tracheal trunks which are just forming. By this time all trace of the abdominal appendages has disappeared. On the tenth day the trophi and legs show segmentation and have lengthened out considerably, while the segmentation of the body somites is also well advanced, proceed- ing forwards from the posterior end of the abdomen. The lateral tracheal trunks are now quite distinct in the posterior part of the abdomen as colourless tubes. The eleventh day seems to be the muscle-forming period, as at this stage muscle fibres are visible in various parts of the body. ‘The lateral tracheze which have not yet developed through the whole length of the embryo show, posteriorly, rudimentary branches, and in the median dorsal line faint traces of the heart are visible in the posterior segments. The most marked change on the twelth day is the extension forward and the pigmentation of the lateral tracheal trunks. These now reach into the head and show indications of the main branches and of the future “ setose gills.” . On the thirteenth day the tracheal system is almost completed, the dorsal network having been formed, and the branches of the head can be seen tapering off to fine threads, but, so far as I can make out, the ‘“‘setose gills” * themselves do not appear until the fourteenth day. ‘There are, however, only small differences noticeable in the living egg of the thirteenth and the fourteenth day. ; The fourteenth day is the last stage, the embryo now being complete within the shell. One point to notice is that, almost from the commencement of development the embryo has never completely occupied the shell, there having all through been a fair amount of free space. On the fourteenth day, however, the embryo commences to swell up two or three hours previous to its escape, so that at the time of the bursting of the shell there is no visible unoccupied space within the shell. While watching this process of swelling up of the embryo I found in the head, situated in the median line and immediately below the dorsal aorta, a pulsating organ. It pulsates much more slowly than the heart, only commences to pulsate a few hours before the embryo escapes, and does not, so far as I have observed, pulsate con- tinuously from the time it first starts. It begins with a few beats, after which there is a pause of, may be, many minutes, but as the time of emergence approaches the pauses are fewer and the pulsations rather more rapid, being about 40 per minute, while the the heart is beating at quite 100 per minute. This pulsating organ is driving fluid backwards, but it was not possible to see where the fluid came from or was driven to. I discovered a ligament suspending the organ, and as it passes on one side of the aorta, I expect that it is one of a pair. There are no blood corpuscles visible in the fluid pumped through this pulsating organ. In conjunction with its pulsations certain movements of the gut are visible, and these become more noticeable as the beats of the pulsating organ become more frequent. * Vide infra. 330 MR FRANK BALFOUR-BROWNE ON The position of this organ in the head (immediately below the aorta and just at the point where the aorta divides into two vessels passing down the frontal region), and the apparent relationship between its pulsations and the peristaltic contractions of the gut, lead me to think that there is no special embryonic organ, but that the pulsations are due to the working of a pharyngeal pump. The larva, as we shall see, feeds by suction, and there is no reason why the pharyngeal pump should not be of service before the embryo escapes from the egg. I imagine that the fluid which surrounds the embryo within the shell is taken in by the mouth and pumped by the pharynx into the gut. Thus the embryo swells up and space is afforded it within the shell by the removal of the surrounding fluid.* For a short time before the shell bursts the embryo is so tightly packed that it is difficult to make out the limits of head and tail (vede fig. 31). The head lies well into the anterior end of the egg, while the first, second, and third abdominal segments occupy the posterior end. The shape of the egg is often somewhat altered by the packing of the embryo which, although usually as described, is sometimes turned in other positions. The shell ultimately bursts somewhere in the region of the maxillary palpi and the . larva immediately straightens out, the shell falling off its back as it does so. All the trophi and legs at once assume their normal positions, the labium folding forwards and coming to lie immediately behind the mouth, the maxillee likewise folding forward and lying on either side of the labium. I have described the maturation of the egg as taking fourteen days from the time it is laid until the escape of the larva. ‘This seems to be the usual time in April or May, but the rate of development varies with the temperature. Thus eggs laid in February—which is exceptionally early —took twenty days to mature, while in one or two cases during the summer thirteen days were sutticient. KowaLevsky (1871) and Heimer (1889) found that the eges of Hydrophilus developed in twelve days, while MicER (1809) gives twelve to fourteen days as the usual period. In my experience with this species twenty days and more was the usual time from the building of the ege-cocoon to the appearance of the larva, but as my specimens were kept in the north of Ireland—far beyond the normal range of the species—this may account for the slower rate of development. I have not followed the development of the embryo of Hydrophilus, but the statements of Kowa.nvsky, etc., indicate that the stages are practically identical with those of [ydrobius. None of these authors, however, describe the swelling of the embryo prior to the bursting of the shell nor do they refer to the cephalic pulsating organ. Tue Larva. (Figs. 5 to 9.) The larva which emerges from the egg is a small white worm-like creature with a pale testaceous head and darker eyes, the eyes being the most visible part, and is about * On this point see “ The Life-history of the Agrionid Dragonfly,” Proc. Zool. Soc. London, Aug. 1909. THE LIFE-HISTORY OF HYDROBIUS FUSCIPES, L. 331 one and a half times the length of the egg from which it comes. But such larve are never seen under normal circumstances. As it comes from the cocoon it is distinctly larger and darker and the head, with its extended jaws, seems also larger in proportion than in the newly hatched individual. Lyonner states that the larvee of Hydrophilus remain a day in the cocoon before breaking out. Those of Hydrobius also remain in the cocoon for some time, but I do not know how long. ‘They ultimately make their way out by breaking through the cocoon, usually along the edge of the flat, lid-like surface which is at the base of the ribbon and just about on the water-level. I hatched a number of larvee from eggs which had been taken from the cocoon and allowed to sink in a tumbler of water. The larve crawled about aimlessly for two or three days, but a number of them died. I took some out of the tumbler and placed them in a drop of water upon a slide to examine them under the microscope, and they immediately began to crawl about and at last succeeded in getting their heads out of the water. Once this was accomplished the head and anterior end of the body was raised up, and the larvee remained in this attitude for thirty seconds or more. During this short time they visibly expanded, and I found, when I replaced them in the water, that they now floated at the surface instead of smking. Examination under the microscope revealed that the gut contained through most of its length a series of air-bubbles. These larve had, in fact, taken a “drink” of air at the first opportunity and had expanded in consequence, and I found by an examination of larve of all stages that the gut almost always contained these air-bubbles. If a larva lying quietly out of the water or at the surface is suddenly submerged, it quite probably sinks, but if it is first gently disturbed it at once raises its head and takes an “air-drink,” after which it is secure against drowning if it is placed in the water. From the fact that larve can be taken unawares and sunk, it is obvious that the air in the gut disappears and has to be renewed from time to time, but whether such air is of any use in respiration I do not know. It is possible, then, to account for several things by this peculiarity of the larva, First, the air-space within the cocoon is probably for the purpose of enabling the larvee to get an “‘air-drink.” A similar space occurs in the cocoon of Hydrophilus, and it has been suggested that it is to enable the cocoon to float. It no doubt makes the cocoon buoyant, but neither in the case of Hydrophilus nor in that of Hydrobius is the cocoon ever built free in the water; it is always attached to some floating object. There seems, therefore, no good reason for regarding the air as specially for the purpose of buoying up the cocoon. It has also been suggested that the air is for the eggs, but the eggs of Hydrobiws—and probably also those of Hydrophilus—hatch perfectly well if taken from the cocoon and submerged. Lyonner observed that the larvee of Hydrophilus swell up three or four times the size of the egg from which they emerge before they have taken any food. This also is probably to be accounted for by assuming that they, like the larvee of Hydrobius, take an “‘ air-drink” in the cocoon. 332 MR FRANK BALFOUR-BROWNE ON We may, perhaps, also account for the ‘‘ mast” on the cocoon of Hydrophilus and the ribbon on that of Hydrobwus as connected with this peculiar habit of drinking air, as so far no satisfactory explanation has been given of these structures. They have been described as the result of excessive zeal on the part of the female—a sort of using up of silk, and the mast of the Hydrophilus cocoon has also been described as an air-funnel to aerate the eggs. The mast always projects above the water, while the ribbon is either spread out upon the surface or upon some object above the surface. The larvee of one batch of eggs do not all necessarily hatch at the same time, some of the eggs—probably the upper ones—hatching sooner than the others. Now the earliest hatched larvee presumably take their drink of air, and as they remain some time in the cocoon, they no doubt tend to use up the limited supply within the cocoon. When they escape they open the cocoon below or on the water-line, and there is there- fore a risk of the remaining air escaping. I think the mast and the ribbon are provided against all these possibilities, and to make certain of other larve getting to the surface to obtain their supply of air. The head of the larva has two eyes made up of about six more or less oval patches of pigment. The antennz are short, three-segmented organs, the apical segment being much finer than the others. The other two segments are, in the youngest stage, about equal in length and twice that of the apical one, but in the growth of the antennz the basal seement grows more rapidly than the middle one, so that, in the final stage, the basal segment is about twice the length of the middle one. On the middle segment and just external to the insertion of the apical one isa small tubercle. The apex of the terminal segment bears two long hairs. The mandibles are large, and end in a sharp incurved point and bear three sharp teeth on their inner margin, the basal one being small. The two mandibles are quite similar to one another. The maxille are long and are presumably to be described as consisting each of a cardo and stipes bearing a minute inner lobe—the undifferentiated galea and lacinia—and a large three-seemented palpus. The middle segment of the latter bears at its apex, just external to the insertion of the apical segment, a long fine hair. The labium consists of a broad submentum and mentum, the latter bearing a pair of two-segmented palpi, and between them a small process presumably corresponding to a ligula. The submentum on its upper surface bears series of closely set minute outwardly projecting points. The labrum is peculiar. Its free edge is strongly serrated in the middle, but the organ is not bilaterally symmetrical. The central part of the anterior margin bears about five teeth. Some slight variations occur in the arrangement of the teeth, and they also change slightly at the moults, but the general arrangement is that there are five teeth of which the four on the right are more or less similar and evenly spaced, while the one on the left is widely separated from the rest of the series. This want of bilateral symmetry of the labium is not unique in Hydrobius, as it occurs also in ~ —— THE LIFE-HISTORY OF HYDROBIUS FUSCIPES, L, 333 Helochares, where, however, it takes a slightly different form, and is perhaps to be found elsewhere. There seems to be no reason for such irregularity, which is quite constant, and it is difficult to see how it can be of any advantage to the larva. On the posterior region of the head, on that part which telescopes under the tergite of the first thoracic segment, are two small sclerites. The prothoracic segment is entirely chitinised, and has no characters calling for remark, ‘The mesothoracic segment is much shorter, and has a large chitinised tergite. On either side of it and at its anterior border a small projection exists into which runs a tracheal tube. Packarp (/.c., p. 471) mentions that the American species of Hydro- bius possess seven pairs of short “setose gills.” The larva of H. fuscipes has, on either side of the mesothorax, this setose structure, and on either side of each of the first seven abdominal segments is a similar structure also containing a tracheal tube communicat- ing with the main lateral trunk. Hydrophilus also possesses the abdominal “ sete,” but 1 amcnot sure as to whether the mesothoracic pair are present. The larve of both Hydrophilus and Hydrobius drown comparatively easily ; they are never long away from the surface of the water, and are constantly to be seen renewing their air-supply by the posterior spiracles, and I do not see why these structures, which may be aborted or closed stigmata, should be described as “ gills.” The dorsum of the metathoracic segment is less strongly chitinised than that of the mesothorax, and on either side of the median line are two minute papilla each giving off a long fine stiff hair. Both meso--and meta-thorax tend to be bilobed at the sides, the one lobe lying above the other. The first‘abdominal segment is as short as the meso- or meta-thorax, but is distin- guished from these—apart from the absence of legs—by possessing two well-marked, though small, dorsal sclerites. The tracheal papillee or setose structures already referred to project on either side of the dorsum and near the anterior margin, and below this the sides of the segment are distinctly bilobed, the lobes being one above the other and somewhat resembling parapodia in appearance. Immediately behind the two sclerites and nearer the posterior margin of the segment are two papille each with a long hair, and a similar pair lie, one on either side, just behind the tracheal papille. The other seven abdominal segments are all very much longer from in front backwards, and are apparently composed of at least two annuli. The first annulus of each segment is plain, possesses no sclerites and no papille. There is a slight constriction at its junction with the posterior part of the segment. The posterior part of each segment from 2 to 7 is exactly similar to the anterior abdominal segment, since it possesses two distinct sclerites, smaller, however, than those of the first abdominal segment, a pair of tracheal papillz, and two pairs of papille bearing long hairs. The anterior annulus of each of these segments is apparently the one which is specially contractile. Hach of the papille is apparently slightly chitinised, and perhaps their bases are correctly described as additional sclerites. The second part of segment 8 is somewhat peculiar. It extends posteriorly into TRANS. ROY. SOC. EDIN., VOL. XLVII. PART,IL:(NO. 14). _ bl 334 MR FRANK BALFOUR-BROWNE ON a three-lobed expansion, below the middle lobe of which the anus opens, and above which is a depression bounded anteriorly by a more or less perpendicular wall of tissue in the form of a slightly lobed ridge. Right and left of this ridge and just posterior to it is a small lobe bearing a flattened chitinised projection on which are three long hairs. The trilobed expansion is fringed with several long hairs and bears on its upper surface, just at the junction of the middle with the lateral lobes, two dark-coloured papillee each with two long hairs at its apex and one long hair projecting from a small papilla on its outer side.* On the posterior face of the anterior ridge and at either side of the median line is a narrow aperture—the spiracle—leading into a small chitinous vestibule into which opens the lateral tracheal trunk. The trunk is constricted where it opens into the vestibule, and there appears to be some apparatus of small chitinous pieces controlling this opening. The lateral lobes and the trilobed part of the segment are capable of spreading outward and of closing in, thus enlarging and diminishing the dorsal depression accord-_ ing as the larva is obtaining air at the surface and holding on to the surface film or moving under water. On the dorsum of this second part of the last abdominal segment is a large semi- circular chitinised tergite, developed perhaps for the protection of the functional spiracles which open just below its posterior margin. The surface of the skin of the larva is covered with scattered hairs arising out of small papillee amongst which are numerous small pores leading into short pits each of which gives off into the skin numerous minute hair-like processes. Amongst these pits are still more numerous minute pointed pits which look very like minute inverted spines. The larvee are metapneustic, the only functional spiracles being those opening into the dorsal depression of the eighth abdominal segment. The expansion of the lobes of this segment enable the larva to hold on to the surface film, the rest of its body being submerged. By moving the body rapidly from side to side when thus attached to the surface, the larva moves fairly rapidly ; otherwise its only method of progress is by creeping on the water plants, as it has no power of free swimming like the young larva of Hydrophilus. The position of the larva in relation to the surface film has nothing to do with the state of buoyancy, since an individual with no air in its gut can and does travel suspended to the surface film as easily as one filled with air; but whereas the former sinks if, from any cause, it loses its hold, the latter floats and can regain its attachment to the film. The method of feeding of the larva is peculiar. Its habit is to lie half out of the * Between these two papille is a small median pit from which bubbles of air seem to come off if, for instance, the larva is placed in alcohol. I have seen the same phenomenon in the case of the larva of Hydrocharis caraboides, L., and it would seem to indicate that there is a median spiracle or pair of spiracles opening at this point, although I cannot see any trachez leading there. But by putting the larva in water under the microscope and flooding with alcohol, air- bubbles seem to come off from numerous places. For instance, in one or two cases I distinctly saw bubbles escaping from the tracheal papille of the first abdominal segment, and in many cases bubbles come from the sides of the other segments, but apparently not from the tracheal papillae, Where these latter bubbles come from I cannot determine, and at present I can merely mention these observations, leaving the matter for a future occasion, THE LIFE-HISTORY OF HYDROBIUS FUSCIPES, L. 339 water, head downwards, on the side of the tumbler, and with the jaws widely expanded. Under natural conditions it may be found in a similar position on projecting water plants, or it is often to be seen enveloped in spirogyra with only its head projecting out. Its food consists largely of Entomostraca, daphnids being the chief item, though it also attacks small insect larvee and is quite ready to devour one of its own kind. The prey sooner or later comes within reach of the jaws and is at once seized. The larva then rapidly wrigeles backwards out of the water, and having once got to a fairly dry spot, it raises its head and thoracic segments off the substratum and, holding the prey with antenne, maxille, and labium, it proceeds to puncture it through and through by opening and closing the jaws, turning the captive round and round by movements of the antennze and maxille. The outwardly projecting points on the upper surface of the submentum and the row of small spines on the cardo of the maxilla assist in holding the prey and prevent the solid parts from being sucked down, as only the juices, with a large quantity of air, are absorbed. When there is nothing left but a crumpled mass of solid substance the larva lays this down beside it and either rests for a while or returns to the water in search of more food. It is apparently absolutely essential that the larva should get at least its head out of the water for feeding. By preventing larve from so doing I found that they died in spite of being in the midst of daphnids. They would seize one and carry it about for quite a long time, always endeavouring to get out of the water. Quite recently I discovered a larva with its head buried in an egg-cocoon devouring a batch of fresh-laid eggs, but whether this is a usual occurrence or was a special discovery on the part of this individual I do not know. The cocoon was floating on the water, but I imagine that the eggs must have been submerged by the time the larva had penetrated the wall of the cocoon. The larvee seem to spend a great deal of their time out of water. I have often found them on the muddy sides of ponds, where they must be largely devoured by small birds and in the tumblers they were constantly to be seen on the sides. I have seen them seizing one of their own kind when out of water, but as a rule they return to the water for their food, and if they wander too far from it they are apt to suffer from drought. The duration of the first larval stage varies considerably. In specimens hatched in the middle of April the usual time until the first moult was about fifteen days, but later on, in warmer weather, the first stage only lasted from six to ten days. A second moult takes place at the end of a period about as long as the first. The two moults have, as far as external characters are concerned, produced no change in the appearance of the larva. In the young larva the head and jaws are much larger in proportion to the body than they are in the older larva, but this change is merely due to a more rapid growth of the body as compared with that of the head and is not connected with the moults. Having moulted twice the larva is in its last stage, and this occupies five to seven or even eight wecks. At the end of that time the full-grown larva crawls out of the 336 MR FRANK BALFOUR-BROWNE ON water and burrows into the ground and after several days becomes a pupa. My larvee, reared in tumblers, indicated their maturity by great activity and by many futile attempts to get out. As soon as they were placed in small glass dishes containing moist earth they burrowed into it. In the earth the larvze spins a silken cocoon. My larvee burrowed to the bottom of the glass dishes in perhaps an inch of soil, and there formed the cocoons in contact with the glass. In some vases these were sufficiently thin to allow the larva to be seen within. I was surprised at the little difficulty I had in getting the larve to burrow into the soil. Lyonner found the larvee of Hydrophilus very particular in the choice of soil for pupating in, and only sueceeded in getting pupa after several attempts. So far I have always failed to get the larvee of Hydrophilus to pupate, although possibly the fact that I have reared them beyond the limits of natural distribution may have affected them. The larvee of Hydrobius accepted garden soil without démur. Once the cocoon is complete the larva rests and does not at once change to a pupa, as I have found larvee unchanged after eight or nine days in the cell. THe Pupa. The pupa is of the typical Coleopterous form, but bears on the dorsal side of its thoracic and abdominal segments series of stiff hairs. In Hydrophilus only a few hooked spines exist in place of these hairs, and Lyonner discovered that they were supports upon which the pupa rests within the cell. The Hydrobius pupa, I found, was very active and turned in all positions in the cells, not by any means always lying passively on its back. The pupal stage lasts but a short time. I cannot give the exact duration, as I omitted to note the times at which the larvee changed to pupe, but, from the time the larva burrowed into the ground, it was three weeks before I saw any imagines in the cells, and as at least some of the larvee remained unchanged for eight or nine days, the pupal condition lasted about eleven or twelve days. There are, no doubt, variations in the length of this stage, just as there are in that of the maturation of the egg and of the larva, according to temperature conditions. THE Lire-Cyc ie. M1auu (l/.c., p. 98) states that “very soon after the emergence of the beetle the cocoon is formed and the eggs are laid,’ and “all the stages may be found together about the end of August.” So far as my experience goes, Hydrobius completes its life- cycle during the one season, the whole cycle occupying about twelve weeks. I have never been able to find eggs or young larvee in August or later, yet, if Mrat’s state- ment is correct, larvee should be found during the winter, as are the larve of the Dytiscide. It is possible that some of the later larvae may pupate in September and - THE LIFE-HISTORY OF HYDROBIUS FUSCIPES, L. 337 that the pupze may lie dormant through the winter, or possibly the imago may emerge from the pupal skin in the autumn and remain within the cell until the spring. Under ordinary circumstances the beetle remains some time within the cell before breaking out, but a newly emerged beetle is always recognisable on account of its comparative soft- ness, and I have no record of having found newly hatched imagines before June. Whereas a large number, perhaps all of the Dytiscidex, pass the winter in the larval as well as in the imago condition, I think that normally the Hydrophilidee are then only imagines, and that they all usually complete their life-cycle within a single season. This certainly is generally the case with Hydrophilus and, I think, with Phalhydrus maritimus, Thoms, and, were it not for Mrauu’s statement, I should also include Hydro- bius fuscipes, L. FECUNDITY. The egg-laying period of a single female apparently continues through the egg- laying period of the species. I have isolated a number of females from the time they built their first cocoon, and up to the end of June some have built as many as seven ege-cocoons. In these cases no male has been with the females since they started their first cocoon, with one or two exceptions, in which they received a f after the completion of the first cocoon. The average number of eggs in these later cocoons is rather smaller than in the earlier ones, there being only about twelve; but each of these females has produced from eighty to a hundred eggs up to the end of June, and there may yet be one or two cocoons built by them before the end of the season. The eggs are fertile, the development of the embryos being quite normal. Hydrobwus fuscipes is certainly a common species, but there must be a tremendous mortality among the larve. SUMMARY. Hydrobius fuscipes, L., is a common water-bettle of which H. picicrus, Thoms, is apparently only a variety, the two forms being extremes of a series and tending to prefer different environments. The food of the imago is chiefly filamentous and confervoid alow, but it also devours dead insects, ete. The jaws, as in all the Hydrophilide, have grinding surfaces at their bases, but the two jaws differ from one another in accordance with slight differences in function. The courtship of Hydrobius is peculiar, the male on the back of the female bending forward over her head and tapping upon her labium with his palpi. There are apparently only certain times at which a female will accept a male. The peculiar twisting of the penis in the process of copulation, which was observed in Hydrophilus by DonisrHorpE, is also seen in Hydrobius. 338 MR FRANK BALFOUR-BROWNE ON The female genital armature is apparently part of the eighth abdominal segment, so that PryrourEAv’s statement that, except in Dytiscus, in the Coleoptera this is always on the posterior margin of the seventh segment, is apparently incorrect. The spinning apparatus, which is somewhat different from that described for HHydrophilus, is described in some detail. The spinnerets, from the apices of which the silk runs, do not appear to have any duct running through them, and it is suggested that the silk flows out on their surfaces. The method of building the egg-cocoon seems to be quite different in Hydrobius from what it is in Hydrophilus, and whereas in the latter case it is stated (LyonNET) that the egos are laid after the body of the cocoon is completed, in Hydrobius, the egg-laying commences very soon after the cocoon is begun. The development of the embryo is followed in the living egg. In the final stage the embryo swells up so as to completely occupy the shell, and a cephalic pulsating organ, which is probably a pharyngeal pump, appears. It is suggested that this absorbs the periembryonal fluid and thus enables the embryo to completely occupy the shell. The larva shows certain peculiar habits, one of which is that it normally has its gut partly filled with air-bubbles, and as it takes its first “ air-drink” shortly after hatching, it is suggested that the ribbon attached to the egg-cocoon, and which normally reaches the surface of the water, is for a pathway by which the larvee may get to the air. In the same way perhaps the mast of the Hydrophilus cocoon may be useful. Another peculiarity of the larva is that it can apparently only feed out of the water. The prey, which consists chiefly of daphnids, is sucked dry, the solid portions being left as useless. BIBLIOGRAPHY. Heuer, K., 1889, Die Embryonalentwicklung von Hydrophilus piceus, L. I. vol., Jena. Hennecuey, L. F., 1904, Les Insectes, Paris. Kowatevsky, A., 1871, “ Embryologische Studien an Wiirmen u. Arthropoden,” Mém. Acad. St, Pétersbourg, (7), be evil. Lakzr, A. G., 1881, “The Cocoons of Hydrophilus piceus and Hydrobius fuscipes,” Entomologist, xiv., pp. 82-84. Lyonnet, P., 1829-32, “Recherches sur l’anatomie et les métamorphoses de différentes espéces d’Insectes,” Mém. du Muséum @ Hist. Nat. Paris, xviii.—xx. Mratt, L. C., 1895, Natural History of Aquatic Insects, Macmillan & Co. Micer, F., 1809, ‘‘Mém. sur les larves des Insectes Coléoptéres aquatiques. I. Mém. sur le grand Hydrophile,” Assoc. du Muséum @ Hist. Nat., xiv. Pacxarp, A. S., 1898, Zext-book of Entomology, Macmillan & Co. PryrourBav, A., 1895, Morph. de ? Armure génitule des Insectes, Paris. Scui6pre, J. C., 1861-3, “De metamophosi Eleutheratorum Observationes. Bidrag til Inseckternes Udviklings-historie,” Naturhistorisk Tidsskrift, Kjobenhavn. THE LIFE-HISTORY OF HYDROBIUS FUSCIPES, L. 339 DESCRIPTION OF PLATES. Puate I. Fig. 1. Right mandible of imago, showing the grinding region (a). Partial view of the lower side and of the “inner” face. Fig. 2. Left mandible, showing concavity (6) into which the grinding portion of the right mandible fits. Partial view of the lower side and of the “inner” face. Fig. 3. First maxilla of left side of imago showing the series of ‘‘brushes” which assist in the feeding process. (a) cardo, (0d) stipes, (c) palpiger, (d) galea, (e) lacinia, (/) the transparent membrane of the stipes. m., muscle attachments. Fig. 4. The labium from below. m., mentum; sm., submentum. Fig. 5. Dorsal view of a larva recently hatched ( x 42); dotted portions are chitinised. Fig. 6 represents the maxille and labium of larva as seen in a moulted skin by removing the frontal selerite with the antenne and mandibles. Note the upper face of the submentum (s.) with its outwardly pointing teeth. m., first maxilla; 7, labium. Maxilla composed of (1) cardo; (2) stipes bearing (3) undifferentiated galea and lacinia, and (4) palpus. Fig. 7. The frontal sclerite of head of larva with antennz and mandibles as removed from fig. 7. Ir., labrum. Figs. 8 and 9 represent dorsal and lateral views of the posterior abdominal segment of the larva. (a) anus ; (0) dorsal sclerite ; tr., lateral tracheal trunks; v., their vestibules (these are seen through the dorsal sclerite and open below its posterior ridge) ; (c) the lateral appendages or so-called “ cerci.” Puate II. Fig. 10. Dorsal view of penis. The abdominal segments are only partially extended. In numbering the abdominal segments I have followed Pryroursav. 7’ and 8’, the tergites of these segments; 7, 8, and 9, the sternites ; a., accessory lobe of penis. (x 60.) Fig. 11. Ventral view of penis, the connective tissue regions between segments 7 and 8 and 8 and 9 being fully extended. Lettering as in fig. 10. (x 60.) F Fig. 12. Ventral view of ? genital armature. 7 and 8, the abdominal segments. (a) Base of one of the spinnerets (6); [for b’ see text]; (v) the ventral grooved plate; (x) one of the two pivots upon which the spinnerets swing ; (y) the chitinised lateral portion of segment 8. (x 60.) Fig. 13. Lateral view of the ? genital armature, both vagina and rectum being prolapsed. Lettering as in fig. 12. (x 60.) Hig. 14. Ventral view of ? genital armature in a state of rest showing the spinnerets retracted. Puate IIT, Figs. 15-31. The development of the egg. Fig. 15, 2-3 days stage, showing primitive streak (p.s.) and commencement of the amniotic cavity (am.c.) at posterior end of egs. per., perivitelline space. Figs. 16 and 17, 3-4 daysstage. a.f., amniotic folds. The amnion (am.) has grown over the posterior part of the primitive streak towards the head, the lateral folds of which (fl.) are to be seen backed by the lateral folds of the amnion. 01., blastopore. Fig. 18. 5 days stage. (There isa stage wanting between this and that of the last figure owing to the three earlier drawings having been taken from an egg at a different time of year showing a different rate of development.) The embryo is seen bent backwards round the yolk, the appendages being visible as buds. Figs. 19 and 20. 6 and 7 days stages showing the gradual shortening of the embryo within the egg and the development of the appendages. Fig. 21. 8 days stage. The embryo showing the ruptured amnion twisted on the back to form the dorsal organ (d.0.). 340 THE LIFE-HISTORY OF HYDROBIUS FUSCIPES, L. Fig. 22. Ventral view of same embryo, Figs. 23 and 24. Lateral and ventral views of the 9th day stage showing the ventral curvature of the embryo. Fig. 25. 10 days stage showing the segmentation of the appendages and the commencement of the formation of the lateral tracheal trunks (é7.). Fig. 26. 11 days stage. The first sign of the heart (.) and of the muscular system. Fig. 27. 12 days stage. The pigmentation of the lateral tracheal trunks. Fig. 28. 13 days stage. The building up of the tracheal system. Fig. 29. The 14th day, some hours before the embryo is to emerge. Note the amount of free space within the shell. Figs. 30 and 31. The same egg shortly before and within a few minutes of the escape of the embryo, p.o., pulsating organ, The tracheal papille (‘‘setose gills”) of the second thoracic and first abdominal segments are seen in fig. 30, and all these are visible in fig. 31. Note the tightly packed embryo in fig. 31. Figs. 32 and 33. The right and left mandibles of the imago seen from above. j Fig. 34, The right antenna of the ¢ of the imago. ans. Roy. Soc. Edin" Vol. XLVII. F. Batrour Browne: Lire History or Hyprosius rusciees, L.—Puare I. Poa a << PIE MsFarlaue & Erskine, Lith. Edin* Tgos. Roy. Soc. Edin* Vol. XLVII. F. Batrour Browne: Lire History or Hyprogius ruscieses, L.—Puare II. x\ EE VEX X N : Na MeFarlane & Erskine Lith Edin® ans. Roy. Soc. Edin*™ Vol. XLVII. F. Batrour Browne: Lire History or Hyprogrus Fuscipes, L.—Puare III, WiFarlane & Erskine. bith. Edin? ( 341 ) XV.—Strophanthus sarmentosus: its Pharmacological Action and its Use as an Arrow-poison. By Sir Thomas R. Fraser, M.D., F.R.SS. L. & E., Professor of Materia Medica in the University of Edinburgh, and Alister T. Mackenzie, M.A., M.B., Ch.B., Carnegie Research Scholar. (With Eleven Plates.) (MS. received July 2, 1909. Read May 3, 1909. Issued separately February 8, 1910.) CONTENTS. PAGE Introductory.—Sources of Materials used, Follicles C. Effects on the Cerebro-Spinal Nervous aoe "364 and Seeds, Extract, Arrow - poison, D. Effects on Skeletal Muscle . ; 374 Poisoned Arrows . : ‘ ; peel E. Effects on the Circulation . : ; . 376 Pharmacological Action— F. Effects on Respiration . : F : ~~ A07 A. Lethality or Toxic Power . ; : . 349 G. Summary . : : : : : . 408 B. General Effects. ; A 3 , . 351 | Explanation of Plates . : : : : - A409 In the course of an endeavour, which was successful only after a number of years, to obtain specimens for the purpose of identifying the species of Strophanthus that produces the smooth seeds which had been chemically and pharmacologically investigated by one of us several years ago, the follicles, flowers’ and other parts of a number of different species of Strophanthus were obtained from Africa. Among them, __S. sarmentosus was represented, and in the course of time a sufficient quantity of seeds ‘of this plant was collected to allow an examination to be undertaken of their chemistry and pharmacology. In this communication, a description will be given chiefly of the pharmacological portion of the investigation, with only such brief reference to botanical and chemical facts as may be useful for the identification of the seeds, and of the preparations from them that were used in the experiments. Further, as there has come into our posses- sion much information regarding the use of the seeds of this plant as an arrow-poison, and an unusually complete collection of illustrative specimens, a few statements will also be made on this subject. It has a special interest inasmuch that arrows poisoned with these seeds are used against British troops in the punitive expeditions that are still found to be necessary in Nigeria and other parts of West Africa. ’ The first specimens of S. sarmentosus that were obtained consisted of a few follicles from Gambia, in West Africa, sent to one of us in 18938, by Sir Roperr LLEWELYN, the then Governor of that Colony. From Gambia, also, specimens of flowers as well as of follicles and seeds were received in 1903, collected by Dr Durrton, of the Liverpool School of Tropical Medicine. In the same year Sir Wituiam Macerscor, K.C.M.G., sent some seeds and flowers from Lagos, of which Colony he was at that time Governor. In 1905, a large number of follicles was received from the Acting-Secretary to the Northern Nigeria Administration, and they were followed, TRANS, ROY. SOC, EDIN., VOL. XLVII, PART IL. (NO. 15), fs 52 342 SIR THOMAS R. FRASER AND MR ALISTER T. MACKENZIE ON in 1906, by several follicles and a small quantity of seeds collected by Dr Jonn M. DauzieL, Colonial Medical Officer of Zungeru, Northern Nigeria. As these were supplemented, in 1908, by a large supply of follicles and seeds, also collected by Dr DauzigeL, ample material had now been obtained for the purposes of investigation. We have to express our obligations to the above colonial officials, and especially to Dr DatziEL, as well as to the Colonial Office in London, by whom the co-operation of several of the above gentlemen and of the Administration of Northern Nigeria was secured. In order to obtain official confirmation of botanical identification, the whole, or specimens of each, of the above consignments were submitted to the authorities at the Royal Botanical Gardens, Kew, and their confirmation, and especially that of Dr Srapr, author of The Flora of Tropical Africa, may be accepted as a guarantee that the seeds used in this investigation were those of S. sarmentosus. In addition to the above-mentioned localities, this species of Strophanthus has also been found in Senegal,* Senegambia and Guineat in West Africa, and a variety of the species, distinguished as S. verrucosus, in Zanzibar and districts adjoining Mombassa and Delagoa Bay in Eastern Africa.{ The plant has also received the names of S. Senegambia, S. pendulus and S. lawrifolius. In the Lagos territory, the native name of “‘ Isa- (or Isha-) kekere” has been given to it, that of “ K wankwanni” or “Kunkunni” in the Zungeru district of Northern Nigeria, and that of ‘‘ Kunna Nombo” (shuttle vine) in the Gambia Colony. Follicles.—The follicles, which are generally fusiform in shape and have a shallow bifid apex, were received in a dry state, and either entirely closed or opened in various degrees at their ventral surfaces. In a considerable number (a) the carpel coverings are retained in their natural state; and in others (b) the epi- and meso- carpels have been scraped off, no doubt to facilitate the drying of the contained seeds. (a) The non-scraped follicles are rough on the surface, with fine, longitudinal, linear projections ; and are of an irregularly mixed brown and green colour, the lower parts being chiefly brown and the upper parts chiefly green. In three or four out of 120 examined, there was a number of small, slightly elevated, pale brown spots, about 4 to 4mm. in length. The follicles vary in dimensions from 180 to 312 mm. (about 7 to 124 inches) in length, and from 20 mm. to 50 mm. (about ~ to 2 inches) in greatest width, the greater number being of or about the larger of these dimensions. The weight varies from 24 to 96 grams (about # to 32 ounces), the usual weight being about 60 or 70 grams (about 2} to 24 ounces), although a considerable number are heavier than the latter weights. (b) The scraped follicles are of a very pale fawn colour, and many of them have darker longitudinal markings, where the mesocarp has not entirely been removed. .* PayRav, Recherches sur les Strophanthus, 1900, p. 86. + Srapr, Flora of Tropical Africa, vol. iv., 1904, p. 180. { Payrav, loc. cit., p. 86, STROPHANTHUS SARMENTOSUS: ITS PHARMACOLOGICAL ACTION. 343 Their dimensions range between 180 mm. and 320 mm. in length (74 and 12} inches), and 25 and 28 mm. in greatest width (1 and 14 inch); and their weight between 11 and 39 grams (2 to 12 ounces), the greater number approximating to the larger dimensions and the higher weight. Seeds.—The seeds, which are provided with the plumose appendages common to the genus, are narrow-ovate or spindle-shaped. They vary in size from 10x 3 to 18x4 mm. (about ‘4x°12 to ‘5x°‘15 inch), the greatest number being about 12x 4 mm. (‘47x ‘15 inch); and they vary in weight from 0°014 to 0°028 gram (22 to ‘44 orain), having an average weight of about 0°024 gram (‘37 or 3 of a grain). They are of a dark fawn colour and have a velvety appearance caused by the short brown hairs with which they are covered,* and their appearance is distinctive when contrasted with that of the seeds of many species of Strophanthus with which we have had an opportunity of comparing them. The seeds have a bitter taste, and, when reduced to powder, they have a feeble woody odour with slight acridity. When the surface of a transverse section is moistened with 80 per cent. sulphuric acid, a faint red tint appears round the cotyledons, in a few minutes the cotyledons assume a dull purple and the albumin a red tint, and, ultimately, the cotyledons become blue, and remain so for several hours.t When 80 per cent. sulphuric acid is applied to the seed divested of its testa, a lavender tinge is produced, which passes into violet and then into pale blue. ~Both of these reactions markedly contrast with the green colour produced in the seed of S. daspidus. Hydrochloric acid applied to a transverse section produces a greenish-blue colour in the interior of the seed, whereas the interior of the seed of S. hispidus is unaffected by this reagent. Extract of Seeds.—¥or the purpose of determining the pharmacological action of S. sarmentosus we used an alcoholic extract of the mature seeds. It was prepared by reducing the seeds to a fine powder, percolating the powder, in the first place, with ether so long as ether removed anything, and then percolating the residual powder with 90 per cent. alcohol in the proportion of five parts of alcohol to one part of the original powdered seeds. The etherial and alcohol percolates were separately distilled to a small bulk and evaporated at a low temperature on a water-bath, and the latter percolate finally dried 7m vacuo over sulphuric acid. By this process there was obtained 34°7 per cent. of ether extract and 23°5 per cent. of alcohol extract. The latter extract, with which chiefly our experiments were made, is a brittle, shining, yellowish-brown and slightly hygroscopic substance, freely soluble in water and moderately so in 90 per cent. alcohol. The watery solution of the extract is strongly bitter in taste and slightly acid in reaction. It contains an active principle * A few of the seeds that had protruded to the outside of opened-up follicles, and many of the seeds that had been removed from the follicles and dried in Africa, are of a very pale fawn or deep cream colour, having hecome blanched, apparently, by exposure to weather. + On submitting seeds to Mr Houmas, of the Pharmaceutical Society of Great Britain, who originated this test and has applied it to the seeds of many species of Strophanthus, he states that he obtained very similar colour-changes, and he expresses the opinion “that there can be no doubt, therefore, that your seeds are those of Strophanthus sarmentosus,” 344 SIR THOMAS R. FRASER AND MR ALISTER T. MACKENZIE ON having the chemical properties of a glucoside. In many of its colour reactions it closely resembles the extract of S. hispidus, as these have been described by one of us in a paper communicated to this Society.* It differs, however, in several colour reactions. For example, in the case of the dry extract, strong sulphuric acid produces a dark brown which slowly becomes violet; and dilute nitric acid a light yellow, which remains unchanged when the temperature is raised to 130° F. In the case of a 2 per cent. solution in water, phosphomolybdic acid causes an abundant yellow precipitate, which is dissolved by heating, and the solution on cooling becomes dark blue in colour with a greyish deposit; and the solution of the extract is made bright yellow by the addition of molybdate of ammonium. Arrow-poisons.—In common with the seeds of several other species of Strophanthus, those of S. sarmentosus are employed in producing an arrow-poison, which is in extensive use in the districts of Africa where the plant occurs. Dr Dauzret has placed at our disposal specimens of this arrow-poison, the preparation of which he has had an opportunity of witnessing at Kantagora and at Zungeru, in Northern Nigeria. Accord- ing to Dr Dauzret, the natives place about a quarter of a pound of coarsely ground S. sarmentosust seeds in an iron pot with about two pints of water and, after heating to the boiling point, add a small quantity of a mixed powder containing five ingredients, chiefly the bark and roots of several other plants. After boiling for a few minutes, the contents of the pot are percolated through a rough filter of twigs, leaves and husks, and the pea-soup-like percolate is again boiled until it has acquired the consistence of a thin syrup. At Kantagora, the powdered head of a small venom- ous serpent and a few drops of cock’s blood are also added to the poison. The arrow- heads are poisoned by being several times dipped into this syrupy liquid, and at Zungeru they are finally dipped into some “animal substance.” It is stated that a poisoned arrow-head may be used several times without renewal of the poison. The poison is a brownish-grey, grumous fluid of thin syrupy consistence, and holding minute, flaky, solid particles in suspension. After being at rest for several weeks, a small quantity of a brown, clear liquid collects at the upper part, and a considerable cohesive, grey sediment at the bottom. This sediment was found to be insoluble in water. When the thoroughly mixed poison was dried, it became a dark brown, brittle and slightly hygroscopic solid, 10 c.c. of the original poison yielding 10°14 grams of solid substance. When this dried poison was tested with the reagents giving fairly distinctive colour-changes with the extract of S. sarmentosus seeds, no conclusive results were obtained, no doubt because of the many ingredients, other than Strophanthus extract, which are present in the arrow-poison. The poison in its original form, the liquid that spontaneously separates from it, and the several times washed sediment were each tested in regard to lethality and effects * Transactions of the Royal Society of Edinburgh, vol. xxxv., part iv., 1890, pp. 1001-3. + Some of these seeds were secured by Dr Dauziut, and on examination were found to be seeds of S. surmentosus, STROPHANTHUS SARMENTOSUS: ITS PHARMACOLOGICAL ACTION. 345 on the heart. While the former two were found to be active, though less so than the extract from the seeds, the third (sediment) appeared to be inert. The experiments are epitomised at pp. 347 and 348. There has also been sent by Sir Freprrick Lucarp, K.C.M.G., High Commissioner of Northern Nigeria, a description, obtained from a “ professional native poison-maker ” by Dr G. W. THomestonz, P.M.O., of the manner in which the poison is prepared. This coincides very closely with the description furnished by Dr DauzieEt, even to the extent of there being used in the preparation, besides Strophanthus seeds, parts of five plants, the blood of a “red fowl” and, occasionally, the head of a venomous serpent. The “animal substance” referred to in Dr Dauzixrt’s description of the manufacture of the arrow-poison is, according to this account, obtained, though only rarely, by sticking the arrows into the abdomen of a dead man and leaving them there for three days. The native name of the arrow-poison is ‘‘Gunguma,” and arrows poisoned with it are in use at Zaria, Kantagora, Bida, Zungeru and Lapai. While, however, at each place, Strophanthus seeds are known to enter into its composition, it was not known if S. sarmentosus is the species, or the only species. ‘The poison appears to be less fluid than that whose preparation was witnessed by Dr Dauzigt, and, further, it is applied by being smeared on the arrow-heads with a feather and not by several times dipping the heads into the poison. It is said to remain active for three years, and to cause death, according to one account, in not less than twelve hours, and, in another account, in not less than thirty minutes ‘after a mere scratch.” Poisoned Arrows from Sir Frederick Lugard (Plate I., figs. 4a and 4b):—Arrows have not been received upon which had been applied the poison whose preparation had been observed by Dr Dauziet, but in 1904, Sir Freperick Lucarp most kindly sent to one of us four leather quivers containing a large number of poisoned arrows obtained for him by Dr Lever, one of the medical officers of Northern Nigeria. These arrows are not feathered. They vary in length from 685 to 787 mm. (27 to 81 inches), a large proportion of them being 762 mm. (30 inches). The shaft consists of bamboo cane, about 8 mm. (?; of an inch) in diameter, which is notched at one extremity for receiving the bow-string, while about 38 to 51 mm. (14 to 2 inches) of the iron stem of the arrow-head is inserted into the other extremity without being tied or fixed with any adhesive substance. In the majority of. the arrows, the head is ovate in shape, and in others it is lanceolate (see fig. 4, Plate I.). The iron arrow-head with its stem is about 178 to 200 mm. (7 or 8 inches) in length, the stem being 128 to 165 mm. (5 to 63 inches) in length, and the head 25 to 50 mm. (1 to 2 inches) in length and 12 to 22 mm. (4 to % inch) in greatest width, a common width being about 19 mm. (? of an inch), and on each surface of the head one wing is concave and the other convex. From the base of each wing, a pointed barb projects downwards at an angle of about 20 degrees with the long axis of the arrow. The barbs are 11 to 15 mm. long (2 to 3 inch), and 2 to 2°5 mm. (5 to +5 inch) thick at their attachment to the arrow- head. In two of the arrows that were received, there are also iron spikes projecting 346 SIR THOMAS R. FRASER AND MR ALISTER T. MACKENZIE ON from each side of the iron stem, in one arrow two pairs and in the other three pairs (see fig. 4b, Plate I.). The long portion of the iron stem of the arrow-head that is not inserted into the cane shaft, and nearly always a considerable part also of the head itself, is thickly and unevenly covered with the poison, the iron stem having previously had a vegetable fibre wound spirally round it, apparently for the purpose of increasing the adhesion of the poison. When the arrows were first received the poison was slightly soft, but, in the course of time, it has become hard and brittle and greyish on the immediate surface, and almost black in the interior. The poison scraped from two arrow-heads, the one small and the other large, weighed 2°5 and 4°5 grams (38°5 and 69°4 grains) respectively. On shaking the powdered poison in a test-tube with a little distilled water, a pale brown, slightly viscous fluid was produced, in which were numerous nearly white, flaky particles, which remained in suspension for several hours. Under similar treat- ment, 70 per cent. alcohol produced a brownish-yellow, clear solution, and 90 per cent. alcohol a very pale yellow solution. In each case, much the greater part of the poison remained undissolved. Neither ether nor chloroform appeared to exert any solvent action upon it. Although by chemical reagents definite proof could not be obtained that extract of S. sarmentosus seeds forms an ingredient of this poison—any more than with Dr Dauzret’s arrow-poison, in which this extract is definitely known to be present—the manner of its preparation and the locality in which it is prepared are in favour of this being the case. This probability is strengthened by the results of experiments that we have made to determine its lethality and its effects on the heart, which are recorded at p. 348. Arrow-heads from the Chibbuh Hill Campargn.—We have also had an opportunity of examining two arrow-heads, obtained by Dr A. C. Parsons, M.O., Hast Bornu, and transmitted to one of us by the Acting High Commissioner of Northern Nigeria, which had been removed from the dead bodies of two members of the Chibbuh Hill Campaign of 1906-7. Both arrows are stated to have caused death in half an hour. They have the same characters as those sent by Sir Freverick Lucarp. The lethality of the poison closely corresponds with that of the arrow-poison obtained by Dr Datziex and of the poison on the arrows received from Sir FRepERIcK Lucarp, and it affects the heart in the same way (pp. 348 and 349). It may, therefore, be assumed as being probable that the seeds of S. sarmentosus form a main ingredient of the poison in these arrows also. With each of these arrow-poisons and with several of their products a number of experiments was made with the objects of determining their lethality and the nature of their action. Before being used, each substance was completely dried mm vacuo over sulphuric acid. The experiments are summarised in the following Tables (I. to IV.). STROPHANTHUS SARMENTOSUS: ITS PHARMACOLOGICAL ACTION. 347 TasLe I.—Dr Datzien’s ARROW-POISON PREPARED FROM S, SARMENTOSUS BY THE Natives oF NortHern NicERta. Minimum-Lethal Dose for Rabbits by Subcutaneous Injection. ane pe OD eee Result. in grams, Brel in grams. 2050 ~ 0:00615 0:003 Recovered. Slight effects. 1860 0:00651 0:0035 Reeovered. Distinct effects. 1600 0:0064 0-004 Death in 5 hours 18 minutes. 1200 0-009 0:0075 Death in 45 minutes. 1950 0:039 0:02 Death in 47 minutes. The minimum-lethal dose per kilogram of this poison by subcutaneous injection in rabbits was found to be 0:004 gram, or 22 times larger than the minimum-lethal dose of the extract of S. sarmentosus. The arrow-poison was not completely soluble in water. The symptoms and post-mortem appearances caused by its administration were identical with those produced by the extract of S. sarmentosus, but they did not occur so soon after the injection. While death occurred within two and a half hours after the injection of the minimum-lethal dose of the extract, it did not occur until five hours after the injection of the minimum-lethal dose of this arrow-poison. TasLe II.—Liguip Part or Dr Dauztew’s ARRrow-Polson. Minimum-Lethal Dose for Rabbits by Subcutaneous Injection. ag A EEE pn tail Result. in grams, in grams, 2600 0:0052 0:002 Recovery. Severe effects. 1290 0:0029 0:00225 Death in 2 hours. 1600 0:004 0:0025 Death in 3 hours. 1500 0:0045 0-003 Death in I hour 10 minutes. The minimum-lethal dose per kilogram of this part of Dr DauziEt’s arrow-poison by subcutaneous injection is, in rabbits, about 0°00225 gram per kilogram, or 1} times the minimum-lethal dose of the extract of S. sarmentosus. Death occurred as rapidly after injection as in the case of the extract of S. sarmentosus, and similar symptoms and post-mortem appearances resulted. As it contains only soluble matter and presumably the Strophanthus products present in the arrow-poison, its greater toxicity is easily accounted for. The difference of lethality between it and the entire arrow-poison indicates that all or several of the other ingredients which enter into the composition of the entire poison diminish its lethality, and are introduced to 348 SIR THOMAS R. FRASER AND MR ALISTER T. MACKENZIE ON render the poison more viscous and adhesive, or under some misapprehension of their potency, or with a superstitious intention. That there are ingredients in the entire arrow-poison which diminish its lethality as contrasted with that of extract of S. sarmentosus was made apparent when the insoluble substance in the arrow-poison was separately examined. It was then found that when rabbits received by subcutaneous injection doses of this portion of Dr DauziE.’s arrow- poison up to 0°02 gram per kilogram (or equivalent to 13 times the minimum-lethal dose of extract of S. sarmentosus), no definite symptoms were produced with even the maximum dose. TabiLe I]],—NortHern Nigeria PoisoN FROM ONE OF SIR FREDERICK LUGARD’s ARROWS. Minimum-Lethal Dose for Rabbits by Subcutaneous Injection. Nasal eae its Fan GA diton Result. in grams. 2 r in grams, 1370 0:00411 0:003 Recovery. Slight effects. 1700 0°00544 0-0032 Recovery. Slight effects. 1320 0:00462 0:0035 Death in 3 hours. 1380 0:00552 0-004 Death in 1 hour 17 minutes. The minimum-lethal dose of this poison for rabbits is the same as that of the poison sent by Dr DauzIxrt. those produced by extract of S. sarmentosus. When a dose of 0°004 gram per kilogram was injected subcutaneously into the The symptoms and post-mortem appearances are identical with flank of a rabbit, distinct symptoms were manifested in 50 minutes, and the animal died in 1 hour 17 minutes. When the same dose per kilogram (0°004 gram) was inserted in a dry state under the skin of the flank of a rabbit, distinct symptoms As the poison removed from one of the smaller and one of the larger arrows weighed 2°5 and 4°5 grams respectively, the one arrow carried sufficient poison to kill ten men and the other eighteen men. appeared in 50 minutes, and the animal died in 1 hour 5 minutes. Taste 1VY.—ARRow-PoIson FROM ONE OF THE CHIBBUH Hitt CampaiGn ARROWS. Minimum-Lethal Dose for Rabbits by Subcutaneous Injection. Weight of | Dose per kilo- Animal Actual Dose gram af Animal Result. in grams, PI aatee in grams, 1700 0:0051 0:003 Recovery. Slight effects. 1300 0:00455 0:0035 Recovery. Distinct effects. 1650 0:0066 0-004 Death in 3 hours. STROPHANTHUS SARMENTOSUS: ITS PHARMACOLOGICAL ACTION. 349 The minimum-lethal dose of this arrow-poison is 23 times as great as that of extract of S. sarmentosus. The symptoms and post-mortem appearances are identical with those caused by extract of S. sarmentosus. In the investigation of the toxic power, the general effects and the effects on the more important functions and structures of the body of S. sarmentosus, we made use of an alcoholic extract of the seeds which had been deprived of all substances soluble in ether, and whose mode of preparation and chief characters have already been described (p. 343). A. Letaatiry or Toxic Power. The experiments made with the extract for the purpose of ascertaining the minimum quantity capable of producing death when administered by subcutancous injection to frogs, rats, rabbits and cats, have been arranged in Tables V., VI., VII. and VIII.* TasLe V.—Minimum-Leraat Doss or Extract ror Frocs (Rana temporaria and R. esculenta). ‘ Dose per kilo- Gcperinent | Anincl‘n groms. | “ingreme:” | etm of Animal Rent 1 30 R, ese. 0:000045 0-:0015 Recovery. Slight effects. 2 24 R. temp. 0:00006 0:0025 Recovery. Slight effects. 3 28 R. temp. 0:000077 0:00275 Recovery. Slight effects. 4 38 R. esc. 0:000114 0:003 Recovery. Distinct effects. 5 42 R. esc. 0:000126 0-003 Recovery. Slight effects. 6 32 R. temp. 0-:0001024 0°0032 Recovery. Slight effects. 7 33 R. ese. 00001089 | 0:0033 Recovery. Severe effects. 8 38 R. esc. | 0:0001254 0:0033 Death after 6 and before 21 hours. 9 41 BR. ese. 0:0001394 0:0034 Recovery. Severe effects. 10 41 R, temp. 0:0001394 0:0034 Recovery. Distinct effects. 1 24 R. temp. 0:000084 0:0035 Death after 9 and before 104 hours. 12 42 R. temp. 0:000147 0:0035 | Death in 34 hours. 13 42 R. ese. 0:0001596 0:0038 Death in 4 hours. 14 30 &. temp. 0:000112 0-004 Death in 2 hours 20 minutes. 15 34 R. esc. 0:00017 0-005 Death in 1 hour 50 minutes. * In the case of warm-blooded animals—rats, rabbits and cats—the weight was taken and the extract was administered always at the end of the same interval of time (18 hours) subsequent to the last reception of food, in order to eliminate, as far as possible, variations in weight due to differences in the amount of the contents of the alimentary canal. TRANS. ROY, SOC. EDIN., VOL. XLVII. PART II. (NO. 15). 53 350 SIR THOMAS R. FRASER AND MR ALISTER T. MACKENZIE ON Tante VI.—Minimum-LerHat Dose or Extract ror Rats. | No. of Weight of Actual Dose ee pees Result , Experiment. | Animal in grams. in grams, in grams. : ra | 18 240 00072 0:03 Recovery. Very slight effects. 19 205 0:041 0-2 Recovery. Slight effects. 20 185 0:0555 0:3 Death in 4 days. 21 205 0:123 0°6 Death in 4 days. 22 130 0:0975 0-75 Death in 3 days. 23 180 0:27 15 Death in 5 hours. TasLeE VII.—Minimum-Leruat Dose or Extract ror Rassits. No. of _ Weight of Actual Dose er Wire Result. Experiment. | Animal in grams. in grams. in grams, 24 2000 0-001 0:0005 Recovery. Very slight effects. 25 2000 00015 0:00075 Recovery. Very slight effects. 26 2210 000177 0:0008 Recovery. Very slight effects. 27 1940 0:001745 0:0009 Recovery. Very slight effects. 28 1870 0:00187 0:001 Recovery. Distinct effects. 29 2500 0:00275 00011 Recovery. Distinct effects. 30 1970 0:002364 0:0012 Recovery. Distinct effects. 31 1750 0°002275 0:0013 Recovery. Distinct effects, 32 1640 0:002296 0:0014 Recovery. Severe effects. 35 2330 0003262 0-0014 Recovery. Severe effects. 34 1460 0:00219 0-0015 Death in 1 hour 25 minutes, 35 2200 0:0033 0:0015 Death in 2 hours 2 minutes. 36 2150 — ~—0°00438 0:002 Death in 1 hour 17 minutes. Taste VIII.—Mrnimum-Leraat Doser or Extract ror Cats. : Dose per kilo- ; ae pees pots piper 8 a af Anil SSE Me 38 2950 0:00295 0:001 Distinct effects, Recovery. 39 2230 0°00446 0:002 Death in 1 hour 15 minutes, The following table (IX.) gives the results following the injection of the same extract into the marginal vein of the left ear in rabbits. STROPHANTHUS SARMENTOSUS: ITS PHARMACOLOGICAL ACTION. 351 TasLte [X,—InrrRavenous Minimum-LetHat Dose or Extract ror RasBits. : | Dose per kilo- Experiment | Aviat ngams. | ‘ingrama” | 64m of Anima Result 40 2180 0:00218 0-001 Recovery. Severe effects. 41 2290 0:00229 =| 0-001 Recovery. Severe effects. 42 1830 0-002 | 0:0011 Recovery. Severe effects. 43 1800 000216 | 0-0012 Death in 1 hour. 44 2950 0:0059 0:002 Death in 4 minutes. From the foregoing Tables, it appears that, when the extract is injected into the subcutaneous tissues, the minimum-lethal dose per kilogram is in frogs about 0°0035 gram; in rats, about 0°3 gram; in rabbits, about 0°0015 gram; and in cats, about 0002 gram.* In rabbits, the intravenous minimum-lethal dose is about 0°0012 gram per kilogram. With reference to the subcutaneous administration of the extract, in general terms rabbits and cats are equally susceptible, frogs are twice as resistant as, and rats 200 times more resistant than, rabbits. A detailed account of several of the experiments in the Tables is given below to illustrate the symptoms produced by large non-lethal and by lethal doses. { B. GENERAL EFFECTS. (a) Experiments on Cold-blooded Animals.—Frogs. Hapervment VII.—Into the dorsal lymph-sac of a male frog (Rana esculenta) weighing 33 grams, 0'0001089 gram of extract, dissolved in Ringer’s solution, was injected (equivalent to 00033 gram per kilogram, or 14 of minimum-lethal dose). Before the injection the throat and flank respirations were regular and at the rate of 20 per 10 seconds ; the nose reflex, conjunctival reflex and sacral reflex were acute. During 45 minutes succeeding the injection, no symptoms appeared. In 1 hour 35 minutes, the throat respirations were 25 per 10 seconds, and irregular in amplitude; the flank respirations were 16 per 10 seconds and irregular in time; the flanks were distended ; the frog jumped well and would now remain on its back; and the cardiac impacts were 6 per 10 seconds. In 3 hours 45 minutes, the throat and flank respirations were 23 per 10 seconds, regular and shallow ; the nose, conjunctival and sacral reflexes were acute ; the frog’s head and throat rested on the floor of the tray; its limbs were flexed ; it could jump well, but landed clumsily, and remained when placed on its back ; the flanks * The extract of S. sarmentosus used in these experiments is thus shown to possess only one-fifth of the lethal power in frogs and one-half in rabbits of the similarly prepared extract of S. hispidus which was used in the experi- ments with the latter substance made by one of us (Trans. R.S.E., vols. xxxv. and xxxvi., 1890 and 1891). + In this and in the other sections of the investigation, several other experiments were made, which gave results concording with those in the experiments that have been recorded. 352 SIR THOMAS R. FRASER AND MR ALISTER T. MACKENZIE ON were distended and the pupils large; and the cardiac impacts were 6 per 10 seconds, regular and shallow. In 5 hours 27 minutes, the throat respirations were 23 per 10 seconds ; the flank respirations were not visible unless the frog was disturbed ; the three reflexes previously referred to were acute; the muzzle rested on the floor of tray ; the limbs were loosely flexed ; the frog could not jump well, the left anterior extremity appeared to be powerless ; the pupils were widely dilated; and the animal remained on its back without any effort to recover the prone position, but when the abdomen was stroked, the prone position was rapidly resumed. Nineteen hours after the injection, the throat respirations were 22 per 10 seconds; there were no visible flank respirations while the frog was undisturbed ; the nose reflex was dull, the conjunctival reflex almost gone, but the sacral reflex was acute; when the point of a blunt needle was lightly drawn across the ocular conjunctiva the frog breathed more deeply and moved feebly, but there was almost no movement of the eyelids; the pupils were dilated and the limbs flexed ; when the posterior extremities were passively extended, there was feeble twitching of the toes, but the limbs were not drawn up, even on pressing the toes ; when the frog was laid on its back, the only movements resulting were slight tremors of the toes, and, when the abdomen was stroked, contraction of the muscles of the abdominal wall occurred ; the cardiac impacts were very distinct, and 8 per 10 seconds; and the flanks were much distended. The frog now weighed 33 grams., Twenty-four hours after injection, the conjunctival reflex was absent, and the condition of the muscular system as last noted. Twenty-seven hours after the mjection, the frog was lying flaccid; the conjunctival reflex was present but sluggish; the nose reflex and the sacral reflex were acute ; if one of the posterior extremities was extended passively it was drawn up again, but only after several efforts; and the voluntary movements of the hind-limbs were feeble, and accompanied by twitching of the toes. Two days after the injection, the throat and flank respirations were 20 per 10 seconds and regular; the conjunctival reflex was acute; the abdomen and flanks were much distended; and the frog could jump, but not well, and at each effort some urine was ejected. Afterwards, the flanks became less distended, and the animal recovered the prone position rapidly when laid on the back. It now weighed 32 grams. On the following day, the only symptoms were slight distension of the flanks and a failure to jump as high as before the injection. The frog weighed 31 grams. Four days after the injection, the frog weighed 33 grams, and seemed in normal health. The laboratory temperature was 57° F. Experiment XI.—0°000084 gram of extract dissolved in Ringer’s solution was injected into the dorsal lymph-sae of a male frog (Rana temporaria) weighing 24 grams ( = 0°0035 gram per kilogram, or the minimum-lethal dose). Before the injection, the throat and flank respirations were 28 per 10 seconds and regular in time ; the nose, conjunctival and sacral reflexes were acute; and the posture was normal and the anima] jumped well. STROPHANTHUS SARMENTOSUS: ITS PHARMACOLOGICAL ACTION. 353 Thirty minutes after the injection, the thorax was raised off the tray and the anterior extremities were fully extended. One hour after the injection, the throat respirations were 18 per 10 seconds and regular, while the flank respirations were 8 per 10 seconds, and irregular in time and feeble; the thorax was raised off the tray, with the fore-limbs partly extended and widely abducted; and the flanks were more distended than previously. In 1 hour 25 minutes, the throat respirations were 19 per 10 seconds, and the flank respirations 10 per 10 seconds ; the general attitude of the animal was as last noted ; and the flanks became more and more distended with each flank respiration, and, when considerably distended in this way, they became flattened completely during a single expiration. Five hours after the injection, the throat respirations were 20 per 10 seconds, feeble and irregular, no flank respirations were visible and the flanks were permanently distended ; the nose and conjunctival reflexes were acute, but the sacral reflex was less acute ; the fore-limbs were flexed and slightly abducted ; the muzzle and throat were raised off the tray and the thorax partly raised off; and the frog jumped well and would not remain on the back. Nine hours after the injection, spontaneous active voluntary movements took place. In 10 hours 30 minutes after the injection, the frog was quite motionless and flaccid, and the pupils were contracted. When the heart was exposed, it was found to be motionless and inexcitable, with the auricles large and dark and the ventricle small and pale ; and a section of the ventricle gave an acid reaction to litmus paper. Twenty-four hours after the injection, the muscles of the upper part of the body, to the level of the anterior extremities, were in rigor ; and stimulation of any part of the surface with the secondary coil of a Du Bois Reymond’s apparatus at zero, and a single bichromate cell in the primary circuit, gave no response. Stimulation of the exposed sciatic or direct stimulation of the muscles themselves with the secondary coil at zero also elicited no response. The temperature of the laboratory during this experiment was 56° F. Expermment X V.—0'00017 gram of extract was injected into the dorsal lymph-sac of a male frog (Rana esculenta) weighing 34 grams (equivalent to 0°005 gram per kilogram, or 7,° of the minimum-lethal dose). Before the injection, the throat and flank respirations were 20 per 10 seconds and regular in time. Fifteen minutes after the injection, the throat and flank respirations were 24 per 10 seconds. In 30 minutes after the injection, the throat and flank respirations were 26 per 10 seconds, and the thorax rested on the tray and all four limbs were flexed. In 50 minutes after the injection, the flank respirations were irregular in amplitude, the nose and sacral reflexes were increased, the thorax was raised off the tray, and the anterior extremities were extended. One hour after the injection, the throat respira- tions were 8 per 10 seconds and regular, and the flank respirations were 8 per 10 seconds and irregular in amplitude. In 1 hour 12 minutes after the injection, the throat 354 SIR THOMAS R. FRASER AND MR ALISTER T. MACKENZIE ON respirations were 11 per 10 seconds, irregular in time and amplitude, and the flank respirations were 3 per 10 seconds and deep ; and the nose reflex was dull, the conjunctival reflex almost gone and the sacral reflex absent. The frog would now remain quietly on its back for 10 seconds ; it jumped feebly and clumsily ; and the pupils, which were dilated before the injection, were now contracted. . In 1 hour 30 minutes after the injection, there were no throat or flank respirations seen in 60 seconds, but occasionally a single throat respiration occurred at longer intervals ; the conjunctival reflex was almost absent ; the thorax was raised off the tray, the fore-limbs being extended and parallel; the frog jumped with difhculty ; and when the posterior extremities were extended passively, they were not drawn up, and when laid on its back the frog made no effort to recover. One hour 40 minutes after the injection, examination of the web under the microscope showed that the circulation had ceased. One hour 50 minutes after the injection, the heart was exposed. It was motionless; the auricles were large and dark; the ventricle was moderately contracted but not very pale, and none of the heart’s chambers responded to mechanical or electrical stimulation. When the skin of the abdomen was pinched, there followed a general con- traction of the muscles of the abdomen and posterior extremities. (b) Expervments on Warm-blooded Animals. (1) Rats. Expervment XIX.—0:041 gram of extract was injected subcutaneously into the left flank of a male rat of the weight of 205 grams (equivalent to 0°2 gram per kilogram, or 3 of minimum-lethal dose). Before the injection, the respirations were 27 per 10 seconds and regular; and the animal was alert. One hour 40 minutes after the injection, the respirations were 27 per 10 seconds, regular, heaving and of greater amplitude ; and the rat was drowsy and lethargic, and when disturbed, its movements were unsteady. In 5 hours after the injection, the respirations were 25 per 10 seconds, regular and heaving ; the cardiac impacts could not be felt, and when the animal moved about the tray its gait was unsteady, and there appeared to be general muscular tremors accompanying voluntary movements. In 22 hours after the injection, the respirations were 27 per 10 seconds, regular and slightly heaving in character, and the rat was sitting quietiy in a corner and showed no signs of uneasiness, but when disturbed, he ran well and seemed mentally alert. Twenty-four hours after the injection, the respirations were 40 per 60 seconds, and irregular in time ; and at regular intervals there occurred sudden respiratory spasms, during which the thorax was fixed and the head was jerked forwards in an abrupt manner —a condition which lasted for about three minutes, after which the rat sat quietly in the tray. STROPHANTHUS SARMENTOSUS: ITS PHARMACOLOGICAL ACTION. 355 Two days after the injection, the rat was quite bright and active. On the third day after the injection, the respirations were 24 per 10 seconds, the rat was active and mentally alert when disturbed, but at other times it slept much. It now weighed 200 grams. No further symptoms were manifested. The temperature of the laboratory was 54°F. during this experiment. Experiment X XII.—0:0975 gram of extract was injected subcutaneously into the left flank of a female rat, of the weight of 130 grams (=0°75 gram per kilogram, or 2°5 times the minimum-lethal dose). Before the injection, the respirations were 18 per 10 seconds and regular, and the rat sat quietly in a corner of the tray. In 5 minutes after the injection, the rat ran round the tray, dragging its hind-limbs along in the extended position; the left hind-limb seemed paralysed ; thereafter the rat sat down, and the posterior part of the body and the hind-limbs trembled violently for a short time. During the next two hours no symptoms were manifested. In 2 hours 10 minutes after the injection, the respirations were 15 per 10 seconds and regular, and the rat sat quietly. In 2 hours 35 minutes after the injection, the respirations were 16 per 10 seconds, slightly irregular in time, and the head moved gently with respiration; the eyes were closed; occasionally the head trembled violently ; and when disturbed, the rat seemed dull, but it was able to run well. In 3 hours 45 minutes after the injection, the respirations were 14 per 10 seconds, slightly irregular in time, and the rat was sleeping. In 5 hours 50 minutes after the injection, the respirations were 16 per 10 seconds, noisy and regular; the rat was still sleeping and showed no signs of uneasiness; when disturbed, it ran round the tray as if feeling rather than seeing its way ; and it was unsteady in its movements, and when running it sometimes fell over on to the left hip. In 22 hours 40 minutes after the injection, the respirations were 16 per 10 seconds, but sometimes the respiratory movement of the thorax was replaced by a rapid forward movement of the head, which would occur three or four times in succession rhythmically with the respirations ; and the rat was sitting drowsily in a corner with the eyes closed, but when disturbed it ran about actively. In 26 hours 10 minutes after the injection, the respirations were regular at the rate of 26 per 10 seconds, and the rat was asleep placidly. In 46 hours 40 minutes after the injection, the respirations were 21 per 10 seconds and slightly irregular in time; and there was no evidence of discomfort. Fifty hours 40 minutes after the injection, the rat became very restless, sprang to the roof of its box and gnawed the wire-netting ; this was repeated several times, and finally the rat sat on the floor of the box breathing heavily. In 51 hours 40 minutes after the injection, the respirations were 6 per 10 seconds, and both inspiration and expiration were accompanied with loud wheezing ; and the rat sat with the eyes closed. In 58 hours 40 minutes after the injection, the respirations were 7 per 10 seconds, and were still accompanied with noisy wheezing ; expiration was much longer than inspiration; the 356 SIR THOMAS R. FRASER AND MR ALISTER T. MACKENZIE ON cardiac impacts were strong, regular, and numbered about 36 per 10 seconds; and voluntary movements were accompanied by well-marked tremors. Seventy-eight hours after the injection, the rat was found dead and general vigor mortis was present. When the thorax was opened, the lungs appeared to be quite healthy, the ventricles of the heart were firmly contracted, and the auricles were dark and distended and contained clotted blood; the liver was healthy; the stomach contained food; the cecum and large intestine were greatly distended with gas, and contained no solid matter. ‘The animal now weighed 120 grams. The laboratory temperature was 50° F. Experiment X XIII.—0'27 gram of extract was injected subcutaneously into the left flank of a male rat weighing 180 grams (=1°5 eu per kilogram, or 5 times the minimum-lethal dose). Before the injection, the respirations were 18 per 10 seconds and regular. One hour 30 minutes after the injection, the respirations were 22 per 10 seconds, regular but heaving in character; the rat sat with the head lowered and the eyes almost closed, and when disturbed it was not active ; and the cardiac impacts were masked by the heaving respirations. ‘T'wo hours after the injection, the respirations were 20 per 10 seconds and there was much movement of the head and shoulders with respiration. ‘Two hours 50 minutes after the injection, the rat became suddenly restless and tried to climb over the edge of the tray. In 2 hours 55 minutes, the respirations were 18 per 10 seconds and very laboured and heaving, and the eyes were completely closed. In 3 hours 35 minutes, the respirations were 18 per 10 seconds and expiration was prolonged and accompanied by a hissing sound apparently produced in the nostrils, and occasionally expiration was suddenly arrested for about five seconds. The rat was now lying with the nose, mouth and thorax resting on the floor. In 3 hours 45 minutes after the injection, the rat began to move about, but the head moved unsteadily and clonic spasms occurred. It soon afterwards fell on the side and then walked with much difficulty, dragging the thorax and abdomen along the floor. In 4 hours 12 minutes | after the injection, the respirations were 18 per 10 seconds and laboured; the muzzle, throat, thorax and abdomen were on the tray, and when the head was raised it trembled violently. In 4 hours 35 minutes after the injection the respirations were 6 per 10 seconds, deep and heaving. In 4 hours 58 minutes after the injection, several general convulsions occurred and the animal died. (2) Rabbits. Expervment XX VITI.—0:00187 gram of extract was injected subcutaneously into the left flank of a doe rabbit whose weight was 1870 grams (=0'001 gram per kilo- gram, or 3 of the minimum-lethal dose). Before the injection, the respirations were 30 and the daltie impacts 35 per 10 seconds and strong, the left pupil measured 5 mm. transversely, the conjunctival reflex was acute, and the rectal temperature 38°5° C. In 30 minutes after the injection, the respirations were 21 per 10 seconds and the cardiac STROPHANTHUS SARMENTOSUS: ITS PHARMACOLOGICAL ACTION. 357 impacts 32 per 10 seconds. In 1 hour after the injection, the respirations were 24 per 10 seconds and irregular in time; the cardiac impacts were 35 per 10 seconds, less distinct and slightly irregular in time; the rectal temperature was 37°5° C.; and fine tremors were occurring in the muscles of the neck. In 1 hour 25 minutes after the injection, severe dyspnoea occurred, lasting for three minutes, during which the rabbit's head was projected forwards, the mouth partly open, the nostrils dilated, and sucking noises were made, the thorax appearing to be fixed; and the cardiac impacts were rapid and indistinct. In 1 hour 45 minutes, the respirations were 21 per 10 seconds and regular; the cardiac impacts were 40 per 10 seconds, slightly irregular and much more distinct ; the pupil was unchanged, the conjunctival reflex was acute; and the rabbit sat in a normal attitude and ate food. In 2 hours after the injection, the respirations were 21 per 10 seconds and somewhat irregular in time; the cardiac impacts were 36 per 10 seconds and markedly irregular in time and force ; the rectal temperature was 38° C. In 2 hours 30 minutes, the respirations were 22 per 10 seconds and irregular in time; the cardiac impacts were 45 per 10 seconds, strong and slightly irregular in time ; and the left pupil measured 6 mm. transversely, the conjunctival reflex was acute, and there was no sien of discomfort. In 3 hours after the injection, the respirations were 29 per 10 seconds and slightly irregular in time; the cardiac impacts were 42 per 10 seconds, regular and tapping in character; the conjunctival reflex was acute ; the rectal temperature 37°5° C. ; feeces had been passed ; and the rabbit was alert, but not rest- less. In 3 hours 10 minutes after the injection, the respirations were 21 per 10 seconds, and regular; the cardiac impacts were 50 per 10 seconds, feeble but regular; the left pupil measured 5 mm. transversely ; and the rabbit was restless, and sat very erect, with the head nodding with the respiratory movements. ‘Three hours 30 minutes after the injection, the respirations were 18 per 10 seconds and the cardiac impacts were 42 per 10 seconds, regular and abrupt, and the animal ate food. Four hours 30 minutes after the injection, the respirations were 21 per 10 seconds, the cardiac impacts were 39 per 10 seconds, reeular and easily felt, and the rabbit sat upright with ears erect. Four hours 53 minutes after the injection, there were tremors of the neck muscles. Five hours after the injection, the respirations were 17 per 10 seconds and irregular in time, the cardiac impacts were 40 per 10 seconds and irregular in force; the left pupil measured 4 mm. transversely and the conjunctival reflex was acute; the rectal temperature was 37°5° C. ; and the rabbit sat upright with ears erect and back well arched. Five hours 40 minutes after the injection, the respirations were 12 per 10 seconds and regular; the cardiac impacts were 38 per 10 seconds and thumping in character ; the transverse measurement of the left pupil was 4 mm., the conjunctival reflex was acute ; tremors were present in the neck muscles and in those of the cheeks; and occasionally a faint sucking noise was made and chewing movements of the lower jaw occurred. Seven hours 30 minutes after the injection, the respirations were 14 per 10 seconds and irregular; the cardiac impacts were 40 per 10 seconds, regular and easily felt; the rectal temperature was | 38°5° C.; and the rabbit was alert. Twenty-four hours 30 minutes after the injection, TRANS, ROY, SOC, EDIN., VOL, XLVII. PART II. (NO. 15). 54 358 SIR THOMAS R. FRASER AND MR ALISTER T. MACKENZIE ON the respirations were 30 per 10 seconds, the cardiac impacts 41 per 10 seconds and easily felt; the left pupil measured 6 mm. transversely ; and the rectal temperature was 38° ©. The rabbit weighed 1880 grams and seemed to have quite recovered. Experiment XX XVI.—0:0043 gram of extract was injected subcutaneously into the left flank of a doe rabbit whose weight was 2150 grams (= 0°002 gram per kilogram, or + of the minimum-lethal dose). Before the injection, the respirations were 32 per 10 seconds and regular; the cardiac impacts were 30 per 10 seconds, regular, but not easily felt; the transverse diameter of the left pupil was 7 mm. and the con- junctival reflex was acute; and the rectal temperature was 38° C. Ten minutes after the injection, the respirations were 30 per 10 seconds and regular; the cardiac impacts were 40 per 10 seconds, regular and more easily felt; and the rabbit was restless and there were occasional fine tremors in the muscles of the body and limbs. In 20 minutes after the injection, the respirations were 35 and the cardiac impacts 34 per 10 seconds; the left pupil was unchanged ; the conjunctival reflex was acute; the rectal temperature was 37°5°C.; and the rabbit was more alert, but its back was less arched. Thirty minutes after the injection, the respirations and cardiac impacts were each 30 per 10 seconds and regular; the left pupil measured 8 mm. transversely ; and the conjunctival reflex was acute, and the skin reflexes over the posterior part of the trunk and the posterior extremities were greatly exaggerated. Thirty-eight minutes after the injection, the rabbit was very nervous and the panniculus carnosus muscle contracted over a wide area when any sudden movement was made near the animal. Forty minutes after the injec- tion, the respirations were 22 and the cardiac impacts were 35 per 10 seconds, and both were irregular in time ; and fine muscular tremors occurred in the body and limbs. Fifty minutes after the injection, the respirations were 26 per 10 seconds and irregular in time, the cardiac impacts were 38 per 10 seconds and irregular in time and in force, and the transverse diameter of the left pupil was 8 mm. One hour after the injection, the respirations were 29 per 10 seconds and regular, the cardiac impacts were 46 per 10 seconds, regular and very distinct ; the left pupil measured 7 mm. transversely ; the rectal temperature was 37° C.; each respiration was accompanied by an abrupt movement of the head ; and the animal did not refuse food. One hour 5 minutes after the injection, the respirations were 26 per 10 seconds, and the cardiac impacts 46 per 10 seconds. For about 15 seconds, the rabbit sat well back on its haunches with the back very con- vex, the ears erect and the head shaking violently ; it pawed the floor with each anterior extremity and thereafter it sank forward with the thorax on the tray and the cervical portion of the spine very concave. One hour 10 minutes after the injection, the respira- tions were 16 per 10 seconds and the cardiac impacts were 46 per 10 seconds, and feeble ; the ears were quite erect ; and alternately the head was jerked upwards and then sank slowly down. One hour 15 minutes after the injection, the rabbit raised itself on its extended fore-legs and arched the spine strongly, a large amount of urine was then freely passed, and thereafter it sank down and its head rested on the floor of the tray. One hour 17 minutes after the injection, the respirations were 4 per 10 seconds, but the cardiac . STROPHANTHUS SARMENTOSUS : ITS PHARMACOLOGICAL ACTION, 359 impacts could not be felt. General convulsions occurred and the pupils contracted greatly and then rapidly dilated to 10 mm. transversely. One minute later, gasping re- spirations occurred and the animal expired, fine tremors occurring at the time of death in the muscles of the anterior extremities. One minute after death, the pupils had con- tracted to 3 mm. transversely and the conjunctiva had become insensitive. Two minutes after death, fine twitches were present in the neck muscles but not elsewhere, and the rectal temperature was found to be 37° C. Ten minutes after death, a more powerful electrical stimulus was required to cause muscular contraction through the exposed sciatic nerve than on direct stimulation of the thigh muscles. Eighteen minutes after death, the heart was exposed ; it was motionless, the left ventricle was small, pale and hard, the other three chambers were dark and dilated, and the right auricle alone re- sponded to local electrical stimulation. Sections of the ventricular and thigh muscles were found to be distinctly acid to litmus paper. Expervment XXX VII.—0°00462 gram of extract dissolved in Ringer’s solution was injected subcutaneously into the left flank of a doe rabbit whose weight was 1540 grams (=0'003 gram per kilogram, and equivalent to twice the minimum-lethal dose). The solution of extract had been kept at a temperature of 100° C. for 30 minutes, and then allowed to cool before being injected. Before the injection, the respirations were 18 per 10 seconds and regular; the cardiac impacts were 35 per 10 seconds, regular but not easily felt; the conjunctival reflex was acute ; and the rabbit’s attitude was normal. One hour 30 minutes after the injection, the respirations were 18 per 10 seconds, regular and panting, the cardiac impacts were 36 per 10 seconds and irregular in time and force; the conjunctival reflex was acute, urine and faeces had been passed ; and the ears were erect, but the head drooped slightly and moved with respiration. ‘T'wo hours and 10 minutes after the injection, the respirations were 25 per 10 seconds and regular: the cardiac impacts were 48 per 10 seconds and feeble; the conjunctival reflex was acute ; and occasionally the anterior extremitiesyielded and glided forwards. Two hours 12 minutes after the injection, violent convulsions occurred, some being opisthotonic in character ; the conjunctival reflex was absent and the pupils were dilated ; and after a brief pause more convulsions succeeded ; and expiration became forced and noisy, and no cardiac impact could be felt. Two hours 14 minutes after the injection, the rabbit was dead and the pupils had contracted to 1 mm. Four minutes after death, the thorax was opened. The thoracic muscles twitched strongly while being divided. When the sternum was removed and placed on the tray, the muscles attached to it continued to twitch violently for a short time. The exposed heat was found to be contracting very feebly and superficially—quite insufficiently to maintain the circulation. Eleven minutes after death, the cardiac contractions were 25 per 10 seconds, very feeble and superficial, the ventricular movement being larger than the auricular, but the organ was not engorged, nor was the left ventricle contracted. Sixteen minutes after death, the heart was twitching feebly, and peristalsis was active in the stomach and intestines. Seventeen minutes after death, the auricles alone continued 360 SIR THOMAS R. FRASER AND MR ALISTER T. MACKENZIE ON to twitch. On section at this time of the ventricles, neither cavity was found to be contracted, both contained dark fluid blood, and there were no clots in them. The abdominal and thoracic organs were healthy. Notwithstanding the prolonged subjection to a high temperature, this extract re- mained active. There were, however, some modifications in the symptoms which appear to indicate that the activity of the extract was lessened, and this is supported by the fact that death supervened at a considerably longer time after the administration than occurs with equal or even considerably smaller lethal doses of the unheated extract. ‘Ihe exposure to a high temperature, which occurs in the preparation of the native arrow-poison, therefore, probably lessens the lethal activity of the products of S. sarmentosus seeds that enter into its composition. (3) Cats. In experiments upon cats, the observations are necessarily incomplete owing to the restlessness and irritability developed in these animals. The general symptoms that were observed are illustrated in the following experiment. Experiment XX XIX.—0'00446 gram of extract was injected subcutaneously into the left flank of a male cat, weighing 2230 grams (=0°'002 gram per kilogram, and equivalent to the minimum-lethal dose). Before the injection, the respirations were 12 per 10 seconds and regular, and the pupils were semi-contracted. Five minutes after the injection, the cat was restless and frequently licked the site of injection. Twelve minutes after the injection, the respirations were 14 per 10 seconds and regular, and the cat was still restless. Fifteen minutes after the injection, feeces were passed. Twenty minutes after the injection, the cat was licking its lips and mewing ; and immediately after this it was very sick and vomited a large amount of dark brown semi-solid matter. Twenty-one minutes after the injection, vomiting again occurred, the vomited matter being dark brown and more fluid. Twenty-two minutes after the injection, the cat retched about a dozen times, and then vomited a small quantity of dark brown fluid matter. Twenty-five minutes after the injection, the respirations appeared to be 8 per 10 seconds, deep and irregular in time; the cat retched 9 times and then vomited a small quantity of dark brown fluid. Twenty-nine minutes after the injection, the cat retched but did not vomit. Thirty-two minutes after the injection, the cat vomited clear frothy fluid after retching for some time, and this was repeated 35 minutes after injection. Retching occurred and clear frothy fluid was vomited 39, 45 and 55 minutes after injection. One hour after the injection, the respirations were 6 per 10 seconds, the cardiac impacts appeared to be 38 per 10 seconds and regular, and the thorax and abdomen rested on the floor. One hour 5 minutes after the injection, the cat was lying on the right side. One hour 10 minutes after the injection, the animal was still on the right side, and retched frequently ; and the respirations varied from 6 to 20 per 10 seconds. One hour 12 minutes after the injection, the pupils dilated widely ; the conjunctiva became inseusitive ; a single gasping respiration occurred, no cardiac STROPHANTHUS SARMENTOSUS: ITS PHARMACOLOGICAL ACTION. 361 impact could be felt; and death occurred, and on its occurrence muscular twitches took place at the shoulders. Three minutes after death, the exposed heart was found to be motionless, the left ventricle being small, the right ventricle large and dark ; the left auricle small and of a bright red colour, and the right auricle large and dark. In response to mechanical stimulation of the ventricles, all the heart’s chambers gave several feeble twitches. A section of the ventricular muscle was found to be acid to litmus paper, as also were sections of the thigh muscles. There were active vermicular movements of the intestines. Intravenous Injection.—Rabbits. The following experiment illustrates the effects produced in rabbits when the extract is injected into a vein. Experiment XLIV.—0:0059 gram of extract was injected into the marginal vein of the left ear of a doe rabbit whose weight was 2950 grams (=0'002 gram per kilogram, equivalent to 3 of the intravenous minimum-lethal dose). Before the injection, the respirations were 23 per 10 seconds, and the cardiac impacts were 28 per 10 seconds, the conjunctival reflex was acute, and the transverse diameter of the left pupil was6 mm. One minute after the injection, coarse chewing movements of the jaws occurred and the rabbit was very restless. ‘Two minutes after the injection, the head was protruded forwards and inspiration consisted of a prolonged sucking movement of the mouth, during which the cheeks were indrawn and the thorax apparently did not expand. Three minutes after the injection, no cardiac impact could be felt. Four minutes after the injection, general convulsions occurred, the pupils became dilated and in- sensitive, the mouth opened widely and air was inspired with a loud sueking noise, and death occurred. Two minutes after death, the eyes became pallid and the pupils contracted rapidly, so that at 4 minutes after death, they were contracted to pin-point size. Five minutes after death, the heart was exposed ; it was motionless and did not respond to mechanical stimulation. All the chambers were of moderate size, but the left ventricle was smaller than the right; the right side of the heart contained dark-coloured fluid blood and the left side bright red fluid blood; and there was no evidence of air embolism. Thirteen minutes after death, the cut surface of the left ventricle was distinctly acid to litmus paper. Summary of General Effects in Frogs. From the foregoing experiments on frogs, it appears that the subcutaneous injection of a moderately large lethal dose of the extract (sufficient to cause death within four hours) produces the following effects both in Rana temporaria and in R. esculenta :— Soon after the injection, the thoracic extremities are abnormally extended and in most cases adducted ; the frog's head is held very high, and reflex movements may be per- formed more actively. The respiratory movements are usually slowed, and they become regular; the flank respirations are more distinctly affected than those of the throat ; 362 SIR THOMAS R. FRASER AND MR ALISTER T. MACKENZIE ON the flanks become alternately distended by several inspirations and then flaccid with a single expiration; and respiratory movements of the throat occur after those of the flanks have ceased, and can be observed when no cardiac impact is visible. Signs of muscular weakness are soon manifest, all the extremities become loosely flexed, the thorax and abdomen rest on the surface of the tray and the muzzle sinks down. When this weakness first appears, jumping can still be performed fairly well, and the frog recovers the prone position rapidly when placed on the back. Subsequently, jumping is effected with difficulty and the frog is unable to raise its body completely off the surface of the table, and it makes no visible spontaneous effort to recover the prone position when lying placed on the back. Reflex movements may still be obtained, but the reflexes are less acute. Careful examination of the precordia does not reveal any cardiac impact, and, if the thorax be opened, the heart is usually found to be motionless, the ventricle is pale and small and does not respond to mechanical stimulation, the auricles are usually very large and dark, and they may be quite motionless or feebly contracting upon their contents, but their movements fail to produce any visible change in the condition of the ventricle. Soon afterwards, the heart is found to be motionless, pale and small, its contractility quickly disappears and the ventricular muscle gives an acid reaction when tested with litmus paper. The conjunctival reflex may be present after the heart has ceased to contract, and the pupils are at that time of small size. The skeletal muscles are contractile for a short time after death, but become acid in reaction soon after the heart muscle, and general rigor very soon sets in and is well marked, especially in the anterior extremities and the anterior parts of the trunk. Summary of General Effects in Rats. As already indicated, rats are peculiarly resistant against even large subcutaneous injections of the extract. When the dose is suchas is sufficient to cause death in four hours, restless movements occur at first, and recur at considerable intervals, and between them drowsiness and inactivity are manifested. The respirations become slower, more deep, and laboured and heaving in character, and expiration is more prolonged than inspira- tion and is often interrupted. In some cases (Experiments XIX. and XXIII.), sudden interruptions of expiration occur in the course of the experiment, and the thorax appears to be fixed during attacks of dyspnoea. These symptoms may be compared with the ballooning of the flanks in frogs. Signs of muscular weakness develop early, and the rat becomes unable to climb up the side of the tray, its muzzle rests on the surface of the tray and, when the animal moves, it drags its hind-limbs ; still later, voluntary move- ments become more impaired and are accompanied with clonic spasms; and, finally, the animal becomes paralysed, general convulsions occurring immediately before death. Summary of General Effects in Rabbits. When a dose sufficient to cause death within four hours is given subcutaneously to a rabbit, the first change usually observed is a temporary flattening of the back, which STROPHANTHUS SARMENTOSUS: ITS PHARMACOLOGICAL ACTION. 363 takes place within a few minutes after the administration and may be repeated several times afterwards. Restless movements also occur early, and the rabbit seems to become more alert. The respiratory movements soon become irregular, and although their rate is not distinctly affected, their type becomes abdominal. Spasmodic interruptions of inspiration occasionally occur. Muscular weakness is displayed ; at first the anterior extremities yield, but the rabbit can yet recover its position; later, the thorax rests continuously on the floor; then the head begins to sink forward, at first for brief periods only, but, afterwards, the muzzle rests continuously on the tray and the back loses its convexity. Still later, the side of the head rests on the tray and the rabbit falls over on to one side, and, finally, the animal lies quietly extended on the side, breathing as if after exertion. Meantime the eardiac impacts have been altering in character; at first their rate is increased, the area over which the impact is felt becomes larger and the force of the impact greater ; later, the impacts become irregular in time and force, and as the irregularity becomes more marked, they become very feeble, until finally they are impalpable. ‘Thereafter, general muscular spasms occur, and in some cases are very violent; the pupils, which have previously contracted slightly, now dilate rapidly, and the surface of the eyeballs becomes insensitive ; and urine may be expelled during the spasms before death occurs. After death, the pupils rapidly contract and then slowly dilate. If the heart be at once exposed, it may be found to be contracting feebly or to be quite motionless ; and it may still respond imperfectly to mechanical stimulation. The left ventricle may be firmly contracted and pale, in which case the right ventricle is softer and much larger, while the auricles are both large and dark. The ventricular muscle is acid in reaction to litmus paper soon after death. The skeletal muscles respond to electrical stimulation if tested immediately after death, and sometimes the response is better to direct stimulation than to stimulation through their nerves. The skeletal muscles become acid to litmus soon after the heart muscle. Only in one rabbit was an excessive salivary secretion noticed, and in a few experiments slight muscular twitchings were seen in the muscles of the neck and cheek. Feces and urine were not passed in the majority of the experiments. The rectal temperature was not materially affected till about the time of death, when it fell rapidly. Summary of General Effects in Cats. With similar lethal doses, cats display almost the same symptoms as rabbits. The condition of the heart after death and the reaction of the heart and skeletal muscles to litmus paper are the same. The respirations appear to be slowed, but a symptom which is very marked in cats, with sub-lethal as with lethal doses, makes it difficult to observe the effects of the poison on the respiration and the circulation. This symptom is vomiting. It appears soon after the injection. In fatal cases, it continues almost without interruption until death supervenes, and in non-fatal cases until a short time before the total disappearance of symptoms. 364 SIR THOMAS R. FRASER AND MR ALISTER T. MACKENZIE ON From the foregoing experiments, it is evident that S. sarmentosus acts upon the cardiac and the skeletal muscles; there is also some evidence of an action upon respiration and upon parts of the cerebro-spinal nervous system. In only one case was an increase of salivary secretion observed, and it is noteworthy that fibrillary twitches of the skeletal and cardiac muscles are not conspicuous symptoms in poisoning with this species of Strophanthus. Gaping movements of the mouth and rubbing of the mouth with the anterior extremities were not observed in the experiments on the general effects, but such movements did occur in other experiments where about fifteen times the minimum-lethal dose was administered subcutaneously. C. ACTION ON THE CEREBRO-SPINAL NERVOUS SYSTEM. (a) Brean and Spinal Cord. In the experiments performed to determine the general effects of S. sarmentosus in frogs, rats, rabbits and cats, restlessness and, in many cases, an increase of alertness are evinced a short time after the administration, and, still later, the general condition of the animal is one of quietude, while in rats there occurs distinct drowsiness. In cats, vomiting, which is a conspicuous symptom, does not begin until twenty or more minutes after subcutaneous injection. No experiments, however, were made with the object of ascertaining how far these and other symptoms might be due to a direct action on the brain. In experiments performed to ascertain the action of the extract on the spinal cord, the brain of a frog was destroyed anteriorly to a line joining the posterior margins of the palpebral apertures, and thereafter the iliac, femoral and epigastrico-vesical (HcKER) arteries of one posterior extremity were ligatured. The arrest of the circulation was ascertained by examining the web under the microscope. For electrical stimulation, a Du Bois Reymond’s apparatus and a single bichromate cell were used. With the secondary coil at 120 mm., the current was just felt on placing the electrodes on the tip of the tongue. When a dose of the extract not much exceeding the minimum-lethal dose was administered subcutaneously, the effects on the nervous system were found to be too slight to admit of accurate observations being made. Much larger doses were there- fore given. Experiment XILV.—Half an hour before the injection of 0°003 gram of extract subcutaneously into the left flank of a male frog (Rana esculenta) weighing 60 grams (= 0°05 gram per kilogram, or fourteen times the minimum-lethal dose), the vessels of the right posterior extremity were ligatured, and both sciatic nerves were exposed. The reflexes were found to be active, and the frog jumped well when irritated. Ten minutes after the injection, stimulation of the web of either foot with the secondary coil at 120 mm. caused the irritated foot to be slowly drawn away, and stimulation over the spinal cord STROPHANTHUS SARMENTOSUS: ITS PHARMACOLOGICAL ACTION. 365 just above the urostyle, with the secondary coil at 100 mm., caused the frog to jump. Twenty minutes after the injection, the attitude of the frog was still normal; the right (protected) foot was withdrawn when its web was stimulated with the secondary coil at 60 mm., and the left (unprotected) foot was withdrawn when its web was stimulated at 100 mm.; and the frog jumped away when the skin over the cord, just above the proximal end of the urostyle, was stimulated with the coil at 80 mm. ‘Twenty-five minutes after the injection, the heart was exposed and was seen to be completcly motionless, with the ventricle in systole. Thirty minutes after the injection, the frog was lying flaccidly on the thorax and abdomen; stimulation of the right (protected) web with the secondary coil at 60 mm., and stimulation of the unprotected left web at 120 mm., caused crossed reflexes; and both posterior extremities were rapidly and fully extended when the skin over the spinal cord just above the proximal end of the urostyle was stimulated with the secondary coil at 80 mm. Thirty-five minutes after the injection, direct stimulation of the right (protected) sciatic with the secondary coil at 370 mm. caused a contraction of the right gastrocnemius, and stimulation of the left (unprotected) sciatic at 370 mm. caused a similar contraction of the left gastro- enemius. Forty-five minutes after the injection, stimulation of the skin just above the proximal end of the urostyle and over the spinal cord with the secondary coil at 110 mm. caused rapid extension of both posterior extremities, the left (unprotected) being more forcibly extended than the right. Fifty minutes after the injection, similar contractions of their respective gastrocnemii were obtained by direct stimulation of the right (protected) sciatic with the secondary coil at 480 mm., and of the left (unpro- tected) sciatic at 410 mm., and a crossed reflex was obtained on stimulating the right sciatic at 350 mm. and the left sciatic at 300 mm. Fifty-five minutes after the injection, no reflex could be obtained by the stimulation of either foot with the secondary coil at zero, while stimulation over the lower part of the spinal cord at 70 mm. caused rapid extension of both limbs, and at 80 mm. caused slight extension of the left (unprotected) limb only. Two hours after injection galvanic stimulation of either foot with the secondary coil at zero produced no reflex; on stimulation over the spinal cord just above the urostyle with the secondary coil at 80 mm., both gastro- cnemii moved slightly ; and at zero mm., a more distinct contraction of the gastrocnemii occurred, but no movement of the limbs resulted in either instance. Two hours 5 minutes after the injection, similar contractions of the gastrocnemii were elicited by stimulating the right sciatic nerve at 330 mm. and the left sciatic nerve at 370 mm., but no reflex movements were produced on stimulating either sciatic directly with the secondary coil at zero. Two hours 50 minutes after the injection, no movements of any kind resulted from galvanic stimulation of either foot at zero, or of the skin over the spinal cord at zero. Three hours after injection, equal movements of their respective gastrocnemii were produced by galvanic stimulation of the right sciatic nerve with the secondary coil at 120 mm. and of the left sciatic nerve at 140 mm.,'and at this time | sections of the right (protected) thigh muscles were feebly acid to litmus paper, whereas TRANS. ROY. SOC. EDIN., VOL. XLVII. PART II. (NO. 15). 55 366 SIR THOMAS R. FRASER AND MR ALISTER T. MACKENZIE ON those of the left (unprotected) thigh were strongly acid. Three hours 15 minutes after injection, galvanic stimulation of the right (protected) sciatic nerve with the secondary coil at 100 mm. caused a contraction of the right gastrocnemius, whereas galvanic stimulation of the left (unprotected) sciatic nerve at zero caused no movement of its gastrocnemius. Three hours 18 minutes after injection, the gastrocnemii were exposed ; the right (protected) gastrocnemius was of a reddish colour; the left was quite pale and firmer than the right. Three hours 20 minutes after injection, the right (protected) gastrocnemius contracted when directly stimulated with the secondary coil at 100 mm., while the left (unprotected) gastrocnemius did not contract when directly stimulated with the secondary coil at zero. Three hours 45 minutes after the injection, galvanic stimulation of the right (protected) sciatic nerve with the secondary coil at 80 mm. caused a contraction of the right gastrocnemius, while galvanic stimulation of the left (unprotected) sciatic nerve even at zero caused no movement of its gastrocnemius ; and when directly stimulated, the right (protected) gastrocnemius contracted with the secondary coil at 30 mm., but the left (unprotected) gastrocnemius did not contract even with the secondary coil at zero. Nine hours 45 minutes after the injection, galvanic stimulation of either sciatic nerve or of either gastrocnemius at zero gave no result ; sections of both gastrocnemii were very acid to litmus, and there was general rigor present throughout the body, though least marked in the right (protected) limb. In this and other similar experiments, the spinal reflex disappeared within 1 hour after the injection of the extract, both in the protected and in the unprotected sides. In the above experiment, the heart’s ventricle was finally paralysed 30 minutes before the spinal reflex had disappeared: Experiments in which the heart’s movements and the blood circulation are stopped by ligaturing the base of the heart, show that the reflex function of the spinal cord persists for at least 25 hours after complete arrest of the heart. It seems probable, therefore, that the relatively early disappearance of the spinal reflex is due to a direct action of S. sarmentosus on the cord itself. The conductivity of the motor nerves is not destroyed until several hours after the administration of the extract. Three hours after the injection, a cut surface of the muscles of the unprotected thigh was strongly acid, whereas a section of the muscles of the protected thigh was only feebly acid in reaction. (b) Sensory Nerves. In order to investigate the effects of the extract on afferent nerves, large frogs (Rana esculenta) were used. Their brains were destroyed down to the level of a line joining the posterior margins of the eyelids, and the experiments consisted in observing after what interval of time the pelvic extremities were withdrawn from a solution of 1 in 500 sulphuric acid in water—before and after the immersion of one foot in a solution of the extract of S. sarmentosus. Immediately on withdrawal the foot was washed. Unless otherwise stated, the limbs were in every case immersed in the acid STROPHANTHUS SARMENTOSUS: ITS PHARMACOLOGICAL ACTION. 367 solution up to the same point, viz. the level of the ankle-joint, and the left foot was placed in the acid after the right had been withdrawn. The following experiment illustrates the results obtained :— Experiment XLVI.—S. sarmentosus extract, 1 in 500. The left foot of a large frog was immersed for 5 minutes in a solution of the extract in distilled water, 1 part in 500, to a point just above the level of the ankle- joint; and during the same time, the right foot was placed in distilled water to the same level. Before immersion, the time which elapsed before withdrawal of either foot from the acid was 5 seconds. This was tested on several occasions. One minute after the immersion, the unpoisoned foot was withdrawn in 8 seconds and the poisoned foot in 7 seconds; 8 minutes after immersion, the unpoisoned foot was withdrawn in 17 seconds and the poisoned one in 7 seconds; 20 minutes after the immersion, the unpoisoned foot was withdrawn in 15 seconds and the poisoned foot in 9 seconds. The left foot was again immersed in the poison, this time for 10 minutes, the right foot being meantime retained in distilled water. One minute after this immersion, the unpoisoned foot was withdrawn from the acid solution in 22 seconds and the poisoned in 18 seconds. Six minutes after immersion, the unpoisoned foot was withdrawn in 16 seconds and the poisoned foot in 18 seconds; 1 hour 40 minutes after the immersion, the unpoisoned foot was withdrawn in 23 seconds and the poisoned one in 26 seconds. The left foot was again immersed in the poison, this time for 30 minutes; and meantime the right foot was kept in distilled water. Three minutes after this immersion, the unpoisoned foot was withdrawn from the acid solution in 20 seconds and the poisoned foot in 47 seconds; 10 minutes after the immersion, the unpoisoned foot was withdrawn in 15 seconds and the poisoned foot was not withdrawn in 2 minutes; 25 minutes after the immersion, the unpoisoned foot was withdrawn in 15 seconds and the poisoned one was not withdrawn in 3 minutes; 30 minutes after the immersion, the unpoisoned foot was withdrawn in 10 seconds, whereas the poisoned foot was not withdrawn in 5 minutes. In the next observation, 40 minutes after the last immersion, the poisoned foot was purposely placed in the acid solution to a point above the level of the contact with the poison ; it was withdrawn in 28 seconds, while the unpoisoned foot was with- drawn in 17 seconds; 50 minutes after the immersion, the unpoisoned foot was withdrawn in 18 seconds and the poisoned foot in 60 seconds—only the poisoned portion was in contact with the acid solution ; 55 minutes after the immersion, the unpoisoned foot was withdrawn in 16 seconds and the poisoned one in 80 seconds; 70 minutes after the immersion, the unpoisoned foot was withdrawn in 15 seconds and the poisoned foot in 25 seconds; 90 minutes after the immersion, the unpoisoned foot was withdrawn in 15 seconds and the poisoned foot in 30 seconds; 2 hours after immersion the reaction time was 16 seconds for the unpoisoned foot and 30 seconds for the poisoned foot ; 4 hours 10 minutes after the immersion, the times were 28 seconds for the unpoisoned foot and 40 seconds for the poisoned foot; and 21 hours after immersion, each foot was withdrawn from the acid solution in 45 seconds, 368 SIR THOMAS R. FRASER AND MR ALISTER T. MACKENZIE ON Results of a similar kind were obtained with more dilute solutions (e.g. 1 in 1000), but longer contact with the extract was required to produce them. Further evidence of the action of this extract on sensory nerves was obtained by applying a solution of it to the cornea of the rabbit. The next experiment typifies these results. Experiment XLVIT.—It was found that a light touch with the point of a blunt stylette applied to the centre of the cornea of either eyeball of a rabbit caused the animal to close that eye rapidly and momentarily. The transverse diameter of each pupil was 8 mm. Of a solution of the extract of S. sarmentosus in distilled water (1 to 500) one-tenth of a cubic centimetre was placed on the right eyeball. In 3 minutes, the reflex in each eyeball was equal and unchanged. In 8 minutes, the reflex in both eyeballs was still equal and unaltered, and both pupils measured 7 mm. in transverse diameter. In 15 minutes, no reflex movement occurred when the right eyeball was lightly touched with the stylette, but the left eyeball was as sensitive as before; the right pupil was 7 mm. in transverse diameter and the left was 6 mm. In 20 minutes, the right cornea could be depressed with the stylette without any reflex movement following; the reflex in the left eyeball was acute and the transverse diameter of each pupil was 6 mm. In 30 minutes, the point of the stylette could be drawn lightly almost half-way across the right corneal surface without eliciting a reflex ; less gentle application caused partial closing of the eyelids; on the left side the corneal reflex was acute; the right pupil was 9 mm. and the left pupil 8 mm. in transverse diameter. In 40 minutes, each pupil was 7 mm. and the state of the corneal reflexes was as last described. In 50 minutes, the right cornea could be gently touched and even slightly depressed by the stylette without causing reflex closing of the eyelids; the corneal reflex in the left eyeball was acute; both pupils measured 8 mm. in transverse diameter, and the anzesthesia seemed to be passing off. In 2 hours 10 minutes, the right cornea was less sensitive than the left, whose reflex was acute ; and the transverse diameter of the right pupil was 8 mm., and of the left 7 mm. In 3 hours after the application of the extract, the right corneal reflex was sluggish and the left was still acute; and both pupils measured 8 mm. in transverse diameter. In 4 hours, the right cornea was insensitive to very light touches only, but less so than it had been an hour earlier, while the left cornea was acutely sensitive ; and both pupils were 7 mm. in transverse diameter. In 5 hours, the right cornea could still be touched very lightly without a reflex following, but heavier touches caused reflex closing of the eyelids ; the left corneal reflex was acute; and each pupil was 9 mm. in transverse diameter. In 7 hours, the lightest touch on the right cornea with the stylette caused a partial closure of the eyelids, but the left cornea “was still more sensitive, and each pupil measured 9 mm. In 23 hours after the application of the extract, each pupil measured 8 mm. in transverse diameter; the corneal reflex had reappeared in each eyeball, but still very light touches with the stylette on the right cornea failed to elicit a reflex. The only evidence of irritation was the occurrence of several quickly STROPHANTHUS SARMENTOSUS: ITS PHARMACOLOGICAL ACTION. 369 succeeding closures of the eyelids immediately after the application of the solution of the extract. From these experiments it appears that the extract is capable of abolishing the sensibility of afferent nerves for a considerable time, and of diminishing it for several hours longer. Subsequently, there is complete recovery of the function of these nerves. Slight dilatation of the pupil accompanies the aneesthesia of the surface of the eyeball. (c) Motor Nerves. Among the general effects following the administration of lethal doses of the extract, it was noted repeatedly, especially in the rabbit, that soon after death galvanic stimulation of motor nerves failed to cause contraction of the muscles they supplied, while the muscles themselves, when stimulated directly, still responded to moderately strong galvanic stimuli. The experiment already described (Experiment XLV.) in connection with the examination of the effects upon the spinal cord shows, in addition, some of the effects of S. sarmentosus upon motor nerves. The results of that experiment are as follows:—The heart was finally arrested within 25 minutes of the injection; the spinal reflex had disappeared within 1 hour of injection ; galvanic stimulation over the spinal cord produced extension of the lower extremities within 2 hours, but had no effect after 3 hours; galvanic stimulation of the sciatic nerve within 3 hours was effective on the unprotected side, and within 34 hours it was quite ineffective; on the protected side similar stimulation pro- duced contraction of the gastrocnemius 3? hours after the injection ; direct galvanic stimulation of the gastrocnemius caused contraction of the muscle on either side 3 hours 15 minutes after the injection ; and failed to cause contraction on the unprotected side in 8 hours 20 minutes, but still caused contraction on the protected side after 4 hours. It is evident that, on the side exposed to the action of Strophanthus, the motor nerves conduct impulses after the spinal reflex has disappeared, but that galvanic stimulation of the poisoned nerves fails to produce contraction of their muscles while the poisoned muscles themselves still respond to direct galvanic stimuli; and, further, it is seen that those motor nerves which are protected from the effects of the extract preserve the power, when electrically stimulated, to cause their muscles (also protected) to contract, not only for some time after stimulation of the unprotected nerves has become in- effective, but even for a considerable time after the poisoned muscles have ceased to respond to galvanic stimuli. To investigate the action on motor nerves further, a series of experiments was performed with nerve-muscle preparations consisting of the frog’s gastrocnemius with a long piece of the sciatic nerve attached. The nerve-trunk of one preparation was placed in a suitable vessel with the muscle of a second preparation, while the muscle of the first preparation and the nerve of the second occupied a contiguous vessel. One of these vessels contained Ringer’s solution, and the other a solution of the extract in Ringer's solution. By means of a single bichromate cell and a Du Bois Reymond’s 370 SIR THOMAS R. FRASER AND MR ALISTER T. MACKENZIE ON induction apparatus, the minimum stimulus required to cause contraction of each muscle (a) through its nerve and (6) on direct stimulation of the muscle was noted repeatedly before and after immersion, as in the following experiment. Only single-break shocks were used. Experiment XLVIII.—Immediately after pithing a frog (Rana temporaria) weigh- ing 50 grams, two nerve-muscle preparations were made. In the following table these preparations are designated A and B. ‘The nerve of A and the muscle of B were each immersed in 3 cc. of a 1 in 500 solution of extract in Ringer's solution. Before immersion the normals were recorded. Minimum Single-break Shock required. Muscles. Nerves. Interval. Notes, A B B. A mm, mm, mm. mm, Before Poisoning. 10 minutes . ; 190 200 500 500 Dols 5 . | 200 200 500 500 Control | Poisoned] Control | Poisoned} Muscle B and nerve A were immersed in ah ien a solution of the extract (1 in 500) in After Poisoning. Ringer’s solution. 3 minutes 220 280 500 490 ie ‘ . | 250 270 500 500 1 eae ; . | 240 270 500 500 iy Se : . | 250 290 500 500 20s : . | 240 260 500 500 2), : . | 240 260 500 500 BOly gy : A | ee 240 420 410 Alay ae ; . | 180 170 380 350 ca ; ae 70) 120 310 310 Stimulation caused fibrillary twitches in the poisoned muscle, ADs 53 : .| 140 150 320 290 Spontaneous fibrillary twitches occurred in the poisoned muscle. BO 35 3 ; 120 110 310 280 DON ; : 110 110 280 260 Spontaneous twitches, coarse in character, occurred in poisoned muscle. (0) ' 5 | EG 110 280 260 op , ; 130 130 260 300 KOe . a 130 120 320 350 Twitches have ceased. (ae : - 140 140 280 290 Do. sO; ; i 160 130 360 300 Do. Sima; . : 170 140 330 320 Do. 50 , : 160 150 310 340 oe : : 180 120 370 480 100) > 5 : ; 190 110 Zero 270 105. ; a ey A0) 120 Zero 200 PLO US : . | 200 110 30 190 Lb ; : 200 120 Zero 140 Twitches have ceased. 120 __i,, : ; 200 130 40 130 NWA or ; He220 120 30 150 130 15 , S| PALO) 120 30 140 140) ys x | 200 120 50 160 STROPHANTHUS SARMENTOSUS: ITS PHARMACOLOGICAL ACTION. 371 Expervment XLVIII.—continued. Minimum Single-break Shock required. Muscles, Nerves. Interval. Notes. A B B. A mm, mm. mm, mm. | Ajter Poisoning. 150 minutes . : 190 110 60 200 155 S,, i : 190 120 50 180 160 ,, : 190 90 30 170 iO” 5 ; : 140 80 0 150 80> ,, ‘ ; 140 80 0 160 0) : 2 130 80 0 160 200 _=s=*é~“4 : : 120 80 0 150 BOs «55 ; 0 160 90 0 150 220 =~, ; : 140 80 0 150 230 =o, : : 140 70 0 150 240, é ; 110 70 0 0 245—C,, 5 ; 90 60 0 6 200) ;, ; : 80 40 0 0 260 =, ; ‘ 70 60 0 0 ZOe : ; 70 60 0 0 7ike\ Va 4 ; 70 60 0 0 340 =, : . |. 100 80 0 0 380) 5; : : 110 70 0 0 425 a, : : 90 60 0 0 30 hrs, 15 min... 70 0 0 0 34 ,, 45 ,, 5 60 0 0 0 Both muscles are distinctly acid to litmus | paper. The above experiment shows that, where a muscle and its nerve-ends are acted upon by S. sarmentosus, the response of the muscle to direct electrical stimuli is soon diminished in comparison with the response elicited from a non-poisoned muscle, and disappears completely some hours before the non-poisoned muscle ceases to respond to direct stimulation. Further, it isseen that electrical stimulation of the nerve-trunk of the poisoned muscle fails to cause any contraction in its muscle within three hours of poisoning, and causes only a modified response within two hours; while, on the other hand, electrical stimulation of the nerve-trunk of a non-poisoned muscle elicits a contraction of its muscle for a considerable time after the poisoned muscle has ceased to respond to stimulation of its nerve-trunk. As in experiments in which the extract was subcutaneously administered, the poisoned muscle responds to direct electrical stimula- tion of its fibres for several hours after stimulation of its nerve-trunk is ineffective. The production of fibrillary twitches is specially evident in experiments of this kind (Experiment XLVIII.). The next experiment was made in order to determine whether these twitches result from an action on the nerve-ends in the muscle or from a direct action upon the muscle-fibres. 372 SIR THOMAS R. FRASER AND MR ALISTER T. MACKENZIE ON Experiment XLIX.—Two male frogs, each weighing 45 grams, were selected. The — Rana temporaria was used, as it had been noted in the course of the investigation that the muscles of this species showed more marked fibrillary twitches than those of Rana esculenta. One of the frogs received by subcutaneous injection into the dorsal lymph-sac 0°3 ¢.e, of a1 percent. solution of curara. Half an hour afterwards, this frog was paralysed. Both frogs were then pithed, and three nerve-muscle preparations were made of the gastrocnemii and sciatic nerves. The preparations from the curarised frog were designated A and B, and that from the non-curarised frog, C. Preparation A was immersed in 4 ¢.c. of Ringer’s solution containing 0°001 gram of curara to prolong the paralysis of its nerve-ends. Preparation B was simultaneously immersed in 4 c.c. of a solution of 1 part of the extract of S. sarmentosus in 650 parts of Ringer’s solution, also containing 0'001 gram of curara. Preparation C was immersed at the same time in 4 cc. of a solution of 1 part of extract in 650 parts of Ringer’s solution, and this preparation, therefore, was not subjected to the action of curara. The following table gives the results of the experiment :— in mm. at which a Single-break Shock was effective. | Maximum Position of the Secondary Coil / | | LENE Muscles. Nerves. pituess A B C. A B C. mm. mm mm. mm mm. mm | After Immersion in Minutes. 5 0 0 | 430 The curarised muscles, A and B, did not respond 10 170 | 200 | 200 to break shocks even at zero, but responded ‘ s es a = feebly to make shocks. 15 bet SGP Se 0 0 | 420 se eee Bt po 0 0 4 60 { A and B do not respond to make shocks now. 30 260.) 230),) QUO) ssee Mi seteaali feze 35 fe ae hc 0 0 | 420 | No fibrillary twitches in A, B, or C. | 40 ZAQ- BAO 5), BION SRN eee ieg | 41 ore | veins | cgmes | gmee | eee) bean | pOntaneous fibrillary, twitches ini C | 45 wee ves A en MOU OLS |e oeD Do. do. 46 baa ores 1» | oe |. |... | Coarse and frequent fibrillary twitches in C; none in A or B, 50 260) LOO 2On | cae A eee. nee Dor do. D5 vig OW Chee OES 0 O | 410 57 Pe dow (|, omar ore geal le ... | Spontaneous fibrillary twitches occur in C, but they are less strong. 60 310 | 290 | 300 | ... | ... | ... | A few feeble fibrillary twitches in C after stimu- lation ; none in A or B. 65 EPP Vier lon 0 0 | 400 | Feeble fibrillary twitches in C after stimulation. 67 dcee | paral eee : . ... | Spontaneous feeble fibrillary twitches in C; none in A or B, STROPHANTHUS SARMENTOSUS: ITS PHARMACOLOGICAL ACTION. 373 Interval. After Immersion in Minutes. 70 75 80 82 85 90 93 95 100 103 105 110 115 116 120 125 130 135 140 142 145 150 155 Maximum Position of the Secondary Coil in mm, at which a Single-break Shock was effective. Experiment XLIX.—continued. Muscles. Nerves. Nou A. B. C, A, B. C. mm mm. mm, mm mm. mm, 330 | 330 | 300 | ... a ... | Strong fibrillary twitches in C after stimulation. Foe aiiliezicsualliasss 0 ‘0 | 380 | A few fibrillary twitches in C only. 240 | 210 | 220 ‘ Do. do. 1 Bes ... | Spontaneous strong fibrillary twitches in C. may ean eee 0 O | 360 220 | 170 | 180 ... | Slight fibrillary twitches in C after stimulation. : 506 Be “ts Do. without stimulation. ae oe or 0 0 | 410 Do. after stimulation. 250 | 160 | 180 | oa ase oro ce sla Do. without stimulation. noes nll ene ete 0 0 | 400 Do. FeV) UU ar OM (eral sea al ase 0 0 | 400 Slight fibrillary twitches in C without stimula- tion ; none ever in A or B. PASI Nc) aN -8 (0 ae ee Pees || ane 0 0 | 380 | Very slight fibrillary twitches in C. ' BAO ZANE TSO vcs 1 |) otal ae. oe Sa set 0 0 | 350 Do. do. 250 | 170 | 180 . | No fibrillary twitches in A, B, or C. ae SCA leet teen liter emer ur ciot: do. in€. > ae ar uae 0 0 | 400 | No do. in A, B, or C. POO LOOM TSO cee st) aes, | ght do. in C. BNI A fla) lt 0 0 | 340 | Slight do. in C, MSO P20 POO messi csc | eae oe timate 0 0 | 320 | Veryslight do. in C. 180| 160%; 70, |. < ... | No fibrillary twitches in A, B, or C. 0 0 0 | C does not respond to stimulation of its nerve; no fibrillary twitches. 200 | 110) 80 No fibrillary twitches in A, B, or C. Muscle C is pale and rigid. Twenty-two hours after immersion, muscle C did not respond to strong electrical stimuli applied directly to the muscle; after the same interval of time, muscle A contracted slightly when directly stimulated with the secondary coil at 60 mm., and muscle B contracted when directly stimulated with the secondary coil at 40 mm. Accordingly, the muscles whose motor nerve-ends were paralysed by curara did not exhibit fibrillary twitches when poisoned by S. sarmentosus. It is by an action on the motor nerve-ends in muscle, therefore, that the fibrillary twitches are produced. Very soon after the fibrillary twitches cease, stimulation of the motor nerve-trunk fails to cause contraction of its muscle. TRANS, ROY. SOC, EDIN., VOL. XLVII, PART II. (NO. 15). 56 374 SIR THOMAS R. FRASER AND MR ALISTER T. MACKENZIE ON D. Errecots oN SKELETAL MUSCLES. Among the general effects following the injection of S. sarmentosus, there were noted early slight rigidity of the muscles, the appearance of fibrillary twitches in them, and subsequently enfeeblement followed by paralysis of the muscles; and soon after death, the muscles were observed to be pale, rigid, non-contractile and acid in reaction. Experiment XLV. showed that muscles unprotected from the action of S. sarmentosus were pale, rigid and acid in reaction within 3 hours; and in 84 hours they failed to respond to direct electrical stimulation. On the other hand, muscles protected from the extract’ responded to direct electrical stimulation for 24 hours, and did not become rigid within that time. Several experiments were made in order to obtain graphic records of some of the changes produced in muscle. For this purpose, the procedure was to make two muscle preparations with the gastrocnemii of a frog in which the muscles were not separated from the femur, and retaining the tendo Achillis and the portion of the femur to which — the muscle is attached. Hach preparation was placed in a small glass cylinder, closed at its lower end by a cork. Through this cork passed a piece of stout platinum wire, whose upper end was hooked so as to fix the attached portion of the femur; while the other end was connected with one pole of an induction coil. The tendo Achillis of the muscle was attached to a light lever writing on a smoked surface. To complete the electrical circuit, a piece of fine platinum wire was hooked into the tendo Achillis, and its free end was connected with the other pole of the induction apparatus. Single- break shocks from a Du Bois Reymond’s apparatus and a single Daniell’s cell were used as stimuli, and the wires from the two muscles were so connected that each stimulus passed simultaneously through both muscles, and in the same direction along each muscle. The muscles were immersed in Ringer’s solution. One muscle was used as a control throughout the experiment, and the solution surrounding the other was afterwards replaced by a solution of extract in Ringer’s solution. The position of the secondary coil is indicated on the tracings. It was found that the muscles of Rana temporaria were more affected by the extract than those of Rana esculenta. Ezxpervment L.—Effect on the muscles of Rana temporaria of extract of S. sar- mentosus dissolved in Ringer’s solution (1 in 1000) (Plate II.). Muscle A is the control and muscle B the poisoned muscle. Before poisoning, normal curves were taken with the secondary coil at 100 mm., as at 12.23 p.m. (fig. 1). 12.25, 2 c.c, of solution of S. sarmentosus extract (1 in 1000) substituted for the Ringer’s solution of muscle B. 12.50. No change (fig. 2). 12.52. Within the last 2 minutes the lever of B had risen owing to shortening of the muscle. The rise was gradual and there were no visible fibrillary twitches. 12.55. Spontaneous fibrillary twitches of B were visible, and the lever was jerked up by some of them. After stimulation at 100 mm. and contraction of the muscle (fig. 3), these fibrillary twitches became very coarse, STROPHANTHUS SARMENTOSUS: ITS PHARMACOLOGICAL ACTION. 375 12.59. Coarse spontaneous fibrillary twitches were occurring in B. These began at irregular intervals. lp.m. After stimulation at 100 mm. and contraction of the muscles, the lever of B fell lower and very coarse fibrillary twitches occurred, jerking it up again. When it had returned to its former level, the fibrillary twitches disappeared. 2 p.m. Spontaneous fibrillary twitches were present in B. They were much fainter than before. After stimulation at 100 mm. (fig. 4) and contraction, about twenty very coarse fibrillary twitches occurred in B, jerking the lever upwards. 2.18. There were no visible fibrillary twitches in B. 2.20. There were no fibrillary twitches before or after stimulation. 2.25. The lever of B has returned to its original abscissa (fig. 5). 2.35. No fibrillary twitches before or after stimulation. 3.25. Do. (fig. 6). 10 a.m. next day. Secondary coil at zero. A contracts better than B. Experiment LI.—Effect on the muscles of Rana temporaria of S. sarmentosus extract in Ringer’s solution (1 in 10,000) (Plate III.). were taken with the secondary coil at 120 mm., as at 12.2 p.m. (fig. 1). Before poisoning, normal curves Muscle A is the control and muscle B the poisoned muscle. 12.7 p.m. 2 c.c. of solution of S. sarmentosus extract (1 in 10,000) substituted for the Ringer’s solution of muscle B, 1.2. Up to this time, no fibrillary twitches were seen. The observations were interrupted until 2 p.m. 2p.m. The lever of muscle B has been raised, and fine spontaneous fibrillary twitches were present in the muscle. 2.22. Before stimulation of muscle B, no fibrillary twitches were visible, but after contraction and relaxation of the muscle the lever was lower than before contraction (fig. 3), and very coarse fibrillary twitches appeared, jerking the lever upwards to its former position. 2.25. Fibrillary twitches were still present in B, but were very feeble. 2.27. After stimulation of the muscle B, the fibrillary twitches became very coarse. 2.36. Coarse fibrillary twitches were occurring in B. 2.42. Stimulation with secondary coil at 120 mm. On relaxation the lever of B was lower than before contraction (fig. 4). 2.44. Very coarse fibrillary twitches occurred in B, jerking the lever back to its higher position again. 3p.m. Both A and B were stimulated twenty times in rapid succession. The fibrillary twitches which followed in B were recorded on a slowly moving surface (fig. 5). 3.17. After contraction and relaxation fine fibrillary twitches occurred in muscle B. 4,26. Do. (fig. 9). 5.35 p.m. No fibrillary twitches occurred in B before or after stimulation. 10 a.m. Muscle A still contracts well when stimulated, muscle B does not respond to electrical stimulation. ‘From these and other similar experiments it appears that immersion of a skeletal muscle in a solution of extract of S. swrmentosus is followed some little time after- wards by shortening of the muscle and consequent elevation of the lever (see Plate II. figs. 3 and 4, and Plate III. figs. 3 and 4). This shortening occurs very rapidly with strong solutions, and it is not at first accompanied by visible fibrillary twitches. On relaxing after stimulation, the muscle becomes more elongated for a time, but does not reach the original abscissa. This elongation is soon overcome by coarse fibrillary 376 SIR THOMAS R. FRASER AND MR ALISTER T. MACKENZIE ON twitches which shorten the muscle once more. Gradually, the shortening of the muscle passes off, and usually it had entirely disappeared when spontaneous fibrillary twitches had ceased (Plate II. figs. 5 to 7). The jerking of the lever has been referred to above. After the initial shortening of the muscle occurs, the fibrillary twitches to which this is due are very noticeable. These soon become very coarse and incessant, and thereafter become less frequent and finer, until they disappear entirely. Even after spontaneous fibrillary twitches have disappeared, coarse fibrillary twitches may be caused for a time by stimulation of the muscle, and, at any time during which fibrillary twitches are present, they become more coarse and more frequent after contraction and relaxation of the muscle. Such twitches are seen in Plate III. fig. 5. After the cessation of spontaneous fibrillary twitches, changes in the muscle curve appear. Usually the first change is that the poisoned muscle requires stronger stimuli to cause contraction. When it contracts, its curve is not so high as that of the non-poisoned muscle ; the rise of the lever is more gradual, the summit of the curve is more rounded, and the fall of the lever much slower than in the case of the control, as in Plate II. fig. 7 and Plate III. figs. 7 and 9. Finally, the poisoned muscle fails to respond to strong electrical stimuli long before the non-poisoned muscle does so. E. Errects oN THE CIRCULATION. (a) Heart. From the earliest experiments, both in cold-blooded and in warm-blooded animals, it was obvious that S. sarmentosus has an action upon the skeletal and cardiac muscles in comparison with which its other effects are of secondary importance. The heart-muscle seemed to be particularly sensitive. Observation of the cardiac impacts revealed that these were at first increased in rate, in force and in extent; later, they became irregular in rate and in force; and just before death they were feeble, irregular and difficult to determine. When the heart was exposed immediately after death, it was usually found that the heart’s ventricle was motionless, pale and firmly contracted ; it did not respond to mechanical or electrical stimuli, direct or indirect, and soon became acid in reaction. In the case of warm-blooded animals the left ventricle of the heart showed these characteristics more distinctly than did the right. Experiments to determine more exactly the extent and the nature of this action upon the heart will now be considered. The following experiment shows the eflects upon the frog’s heart of a dose of the extract, subeutaneously injected, which caused profound general effects, but from which the frog would probably have recovered :— Experiment LII.—Sixty-eight hours 20 minutes after receiving into the dorsal lymph-sac a dose of extract equal to 0°0031 gram per kilogram, the frog (Rana esculenta) lay on the abdomen and thorax with the limbs loosely flexed; the throat — STROPHANTHUS SARMENTOSUS: ITS PHARMACOLOGICAL ACTION. 377 respirations were 20 per 10 seconds, the flank respirations 3 or 4 per 10 seconds; the conjunctival reflex was acute; the pupils were dilated; the frog was unable to flex a hind-limb which had been passively extended; it made feeble and unsuccessful efforts to recover the prone position when laid on its back; and the cardiac impacts could be counted. Sixty-eight hours 25 minutes after the injection, the frog was decapitated and the heart exposed. ‘The heart was contracting regularly seven times in 60 seconds ; the diastolic phase was long, and all the chambers were very large and dark during diastole ; systole was rapid and complete, the ventricle becoming very small and pale, and then rapidly dilating ; a portion near the apex was the last to contract in systole. Five minutes after decapitation, three complete contractions of the heart occurred in 30 seconds, and then the heart paused in diastole for 40 seconds. After this pause, five good and regular contractions occurred in 60 seconds. Fifteen minutes after decapitation, seven contractions occurred in 60 seconds; the ventricle became very pale in systole (which occupied less than 2 seconds), and was large and dark in diastole (which occupied 6 or 7 seconds). Forty-five minutes after decapitation, five con- tractions occurred in 60 seconds, and the systolic and diastolic phases were as last noted. One hour 5 minutes after decapitation, seven cardiac contractions occurred in 60 seconds and at irregular intervals. Systole was performed completely and rapidly. Three hours 8 minutes after decapitation, six contractions occurred in 60 seconds; the first of these was succeeded by a pause in diastole for 40 seconds, the other five occurred in the next 20 seconds; the systolic phase was rapid and complete, but the diastolic phase predominated. Three hours 45 minutes after decapitation, four con- tractions occurred in 60 seconds and at irregular intervals. Four hours 30 minutes after decapitation, four contractions occurred in 45 seconds, followed. by a pause in diastole of 25 seconds, and systole was now less complete. Four hours 50 minutes after decapitation, five contractions occurred in 30 seconds; dark ‘“‘ pouchings” usually appeared at the apex and the base of the ventricle in systole and moved from one part of the cavity to another; and at times dark rings ran over its surface from base to apex ; and during these vermicular movements of the ventricle the auricles were very much dilated and the frog struggled feebly. Four hours 55 minutes after decapitation, the heart was beating almost regularly ; the ventricle was uniformly dark in diastole and uniformly pale in systole. Four hours 56 minutes after decapitation, the heart stopped in diastole; 30 seconds later a single powerful contraction of the ventricle occurred and was followed by immediate relaxation ; and the frog struggled feebly. Four hours 57 minutes after decapitation, another single powerful contraction of the ventricle occurred, followed by several more at intervals of 15 to 20 seconds; and each contraction was preceded by a single auricular contraction which greatly distended the ventricle. In 6 hours 25 minutes after decapitation, the heart was observed to be in diastole for 2 minutes, then ten regular and complete contractions occurred at intervals of 4 or 5 seconds, and were followed by a pause in diastole for 1 minute, succeeded by further rhythmic contractions. Seven hours 45 minutes after decapitation, 378 SIR THOMAS R. FRASER AND MR ALISTER T. MACKENZIE ON three regular and complete contractions of the heart occurred in 30 seconds. Twenty- four hours after decapitation (and 92 hours after the injection), sixteen regular and uniform contractions of the heart were seen in 55 seconds; in systole, the ventricle did not become very pale ; in the diastolic phase, only the apex of the ventricle anda portion at the base first became dark... This series of contractions was succeeded by a pause in diastole for 24 minutes; then the auricles began to contract and to distend the ventricle; at first, the ventricle responded only by superficial and imperfect contractions, but after about a dozen auricular contractions the ventricle was beating regularly and well, and then another long pause followed. Twenty- nine hours after decapitation, the heart was motionless in diastole; then the auricles began to contract and to distend the ventricle; at first the ventricle responded to this stimulation imperfectly, but by and by it resumed its contractions at the rate of 18 per minute, synchronously with the auricles, and became quite pale in systole; 4 minutes afterwards, the rate of contraction, however, became slower and diastolic pauses of irregular length occurred. General reflexes could still be elicited. Thirty- three and a half hours after decapitation, the conditions were as last noted; the heart was observed to beat well for 11 minutes at 18 per minute and then to become arrested in diastole. Thirty-six hours after decapitation, the heart was beating at 18 per minute, with occasional pauses in diastole lasting for one or more minutes. No further observations were made. In the next series of experiments the extract dissolved in Ringer's solution was applied to the outer surface of the frog’s heart, definite quantities of standard solutions being instilled within the pericardial sac. The procedure was as follows :—The brain was destroyed anterior to a line joining the posterior margins of the eyelids. The frog was then pinned down on its back and the heart was fully exposed without damaging the pericardial sac. A small portion of the sac was removed with scissors from the highest part, which was usually situated over the auriculo-ventricular junction. By means of a hypodermic syringe, from which 100 drops of Ringer measured approxi- mately 1 ¢.c., five or ten drops of one or other of the standard solutions used (= 0°05 «e. and 0°l ec.) were instilled within the pericardial sac through the artificial opening. During this process the cut edge of the sac was held with fine forceps and the upper part of the sac gently raised from the surface of the heart. After the instillation, the pericardium was not released until several contractions of the heart had distributed the solution all over its surface. These quantities of fluid were easily retained within the pericardial sac of frogs weighing 20 grams and upwards, and the heart’s movements did not cause the solution to overflow. In order to keep the tissues moist, the frog was laid on wet filter-paper and was covered by a glass funnel. In similar experiments upon frogs without the application of any poison, it was found that the heart’s movements preserved their chief characteristics for 48 hours after the heart was exposed—the contractions continued to be regular in time and to affect STROPHANTHUS SARMENTOSUS: ITS PHARMACOLOGICAL ACTION. 379 the chambers uniformly ; the ventricle was of a pale pink colour in systole and never white ; and ventricular systole continued to occupy two-thirds of the ventricular cycle. The number of contractions, however, had fallen from eight to six per 10 seconds, and the ventricle did not become quite so small in systole as at the beginning of the experiment. After 48 hours, general reflexes could still be elicited. Experiment LITI.—0:0000125 gram of extract placed within the pericardial sac of a frog (Rana temporaria). Contractions in 10 seconds, Interval. Auricles. Before Strophanthus. 12 minutes . i After Strophanthus. 10 minutes . if 20 ” 6 30 6 AO. 3; 6 ae 6 23: 5, 4 KOS 2,, 4 190 =, 4 2S ee 4 ZOO ,, 3 2D - 5, 2S: ,, Ventricle. Notes, ars Contractions regular and powerful; ventricular systole occupies two-thirds of the cardiac cycle; in diastole, the ventricle is uniformly dark red in colour, and in systole it is pink and moderately small; auricular systole and ven- tricular diastole are synchronous. 0:0000125 gram of extract in Ringer’s solution was inserted inside the pericardium. Notes as before, but the ventricle possibly becomes paler in systole. ; Contractions are regular and powerful; ventricular systole continues to occupy two-thirds of the cycle; in diastole, the ventricle appears to be darker than before the applica- tion of the extract; in systole, it is distinctly smaller and aler. Hecate regular and powerful; ventricular systole and diastole each occupy half of the cycle; the ventricle begins to dilate before auricular systole occurs. Contractions regular and powerful ; ventricular systole again occupies two-thirds of the cycle; auricular systole and ventricular diastole are again synchronous. As last noted. Contractions regular and powerful ; ventricular systole and diastole each occupy half of the cycle ; ventricular diastole is performed well and uniformly; the ventricle contracts uniformly and becomes pale pink. Ventricular diastole occupies two-thirds of the ventricular cycle; systole is powerful; ventricular diastole is in two phases, the first is a slight relaxation of the ventricle immediately after its own systole; the second corresponds with the systole of the auricles. As last noted. Regular and powerful contractions ; ventricular systole and diastole each occupy half the cycle; the ventricle is now less pale in systole. Ventricular diastole now occupies two-thirds of the cycle; at irregular intervals the heart is arrested in full diastole for periods of about 15 seconds. The heart was arrested in diastole for 15 seconds and the frog struggled violently; the ventricle became greatly dilated and then began to contract regularly. The heart is contracting regularly and powerfully. 380 SIR THOMAS R. FRASER AND MR ALISTER T. MACKENZIE ON Experiment LITT.—continued. Contractions in 10 seconds, Interval. —— Notes. Auricles, Ventricle, After Strophanthus. 283 minutes . : es se The heart paused in diastole for 5 seconds; such pauses occur spontaneously, and may also be induced by ietting the edge of the funnel fall sharply on the thighs of the frog. 5 hours : : 4 4 Regular and powerful contractions ; ventricular systole and diastole each occupy half the cycle; ventricular diastole shows one phase before and one phase accompanying auricular systole; in systole, the veutricle is well con- tracted and moderately pale. 22 hours 30 mins. . 4 4 Contractions regular, but less powerful; other notes as last recorded; in diastole, the ventricle is smaller than before, and mottled. BL ae PAD ts -(2 6 6 Regular and feeble contractions; auricles and ventricle are very pale and small ; general reflexes elicitable. DDE Gano uns oe Sc Bat The heart is motionless; the auricles are small and pale; the ventricle is small and purplish in colour; mechanical stimulation of the ventricle results in several feeble move- ments of all chambers of the heart; general reflexes can be elicited ; respiratory movements have ceased ; on section, the ventricle was empty and its cavity small ; the reaction of its muscle to litmus was doubtful—perhaps faintly acid. Experiment LIV.—0'00005 gram of extract placed within the pericardial sac of a frog (Rana temporaria). Contractions in 10 seconds. Interval. - — Notes. * Auricles, Ventricle. Before Strophanthus. ‘10 minutes . : 7 7 Contractions powerful and regular in time; ventricular systole occupies two-thirds of the cycle ; the diastolic move- ment of the ventricle is abrupt, the systolic is deliberate. l minute. ; 7 th As last noted. a 0:00005 gram of extract in Ringer’s solution inserted inside the pericardial sac. After Strophanthus. 2 minutes a 7 As last noted. 5 yy 6 6 Do., but ventricle is perhaps paler in systole. Sti, 6 6 Li os 6 6 20, 5 5 Contractions regular and powerful; ventricular diastole now occupies one-half of the cycle; in systole the ventricle becomes very pale; the diastolic movement of the ventricle is prolonged, and is in two phases, one before and the other accompanying auricular systole. STROPHANTHUS SARMENTOSUS: ITS PHARMACOLOGICAL ACTION. 381 Expervment LIV.—continued. Interval. After Strophanthus. 28 minutes 31 fs 34 ” 35 43 37 : 40 5 a, 42 if 43 a ee Contractions in 10 seconds, Auricles, Ventricle. Notes. Two auricular contractions precede each ventricular con- traction ; twelve auricular and six ventricular contractions occurred in 380 seconds, ventricular systole occupies 2 seconds and the ventricle becomes very small and pale ; ventricular diastole occupies 4 seconds (about), and the ventricle does not dilate very much. Twelve auricular and six ventricular contractions occurred in 30 seconds. In 20 seconds, the auricles contracted six times, and each auricular contraction distended the ventricle further, then the ventricle gave a single powerful contraction. The frog struggled ; the base of the heart is quite pale and contracted ; at each contraction of the auricles only a small area at the apex of the ventricle becomes darker in colour, and it at once pales in systole again. Three auricular and three imperfect ventricular contractions occurred in 60 seconds; the ventricle is almost completely contracted. No movement of the heart in 60 seconds. One contraction of all chambers in 60 seconds. Do. do. Eight contractions of auricles and ventricle occurred in 60 seconds; these were irregular in time and very feeble; thereafter the heart was arrested with the ventricle very small and pale; the auricles were slightly dilated; the frog breathed regularly. : The heart is motionless ; the auricles are slightly dilated and the ventricle is pale and firmly contracted; mechanical stimulation of any chamber results in a feeble movement of the auricles only; on section, the ventricular muscle appeared to give an alkaline reaction with litmus paper. Experrment LV.—0-0004 gram of extract placed within the pericardial sac of a frog (Rana temporaria). Interval. Before Strophanthus. 5 minutes . 1 3 ” Contractions in 10 seconds. Auricles. Ventricle. Notes. Contractions regular in time and powerful; movements large ; ventricular systole occupies two-thirds of the cycle. Do. do. Inserted 0:0004 gram of extract dissolved in Ringer’s solution inside the pericardial sac. TRANS. ROY. SOC, EDIN., VOL. XLVII. PART II. (NO. 15). zi BT 382 SIR THOMAS R. FRASER AND MR ALISTER T. MACKENZIE ON Experiment LV.—continued. Contractions in 10 seconds, Interval. —_ Notes. Auricles. Ventricle. After Strophanthus. 3 minutes . : 8 8 Contractions regular and powerful; in systole, the ventricle becomes much smaller and paler than formerly ; in diastole, the ventricular movement is more abrupt; the auricular movements are strong. 9 9 In systole, the condition of the ventricle is as last noted ; in diastole, the ventricle does not dilate so much; ven. tricular systole occupies two-thirds of the cycle. 1Ow gs : : 8 8 Regular contractions ; ventricular systole is as last deseribeal g in diastole, the ventricle dilates only a little and its apex scarcely expands at all; the auricles are acting well. 15 - 8 8 Do. do. Pe if 7 Do. do. 23 vhs 8 8 Do. do. 30m 5-8 8 8 Ventricular systole as last described; in diastole, the ven- | tricle now becomes very large; the auricular diastolic movement is also greater. Ciel wri; 8 8 Contractions powerful and regular in time; ventricular expansion and contraction are very large movements; in diastole, the auricles become very large and they contract powerfully. 40 ,, : ‘ 8 8 The ventricular movements are very large and quite regular in time; there is a slight tendency for the ventricle to begin contracting before its diastolic expansion is complete. tiie, Se) - T ff Movements of all chambers are very large and quite regular ; the ventricular systole occupies two-thirds of the cycle. 50 a : ; 6 6 Do. do. 52 5 ; é tes eos The auricles are contracting on their contents, but do not expel them ; the ventricle is arrested in systole. 55 ‘; 6 6 Contractions regular in time ; the auricles do not completely expel their contents in systole; they are enormous in diastole; the ventricle dilates only slightly in diastole, which occupies half of the cycle. 60) ‘ : 4 4 Contractions regular in time ; the auricles are greatly dis- tended in diastole and do not expel more than half their contents in systole; ventricular diastole occupies three- fourths of the ventricular cycle; of these, two-fourths occur before auricular systole; in systole, the ventricle is very small and pale. BD: © Sapna ; 4 4 Do. do. Gre a es : oe ae Heart stopped beating ; all chambers were in diastole for 100} seconds, then regular contractions occurred spontaneously. (Ce a Berta ge : Ans = Heart arrested for 50 seconds with the auricles fully dilated and the ventricle partly dilated ; contractions were resumed spontaneously at four per 10 seconds, the auricles contracting incompletely and the ventricle completely. GIN wore : : 3 3 oF age F ; Bac Re A long pause with all chambers in diastole. SOi eee : F She ae Pause of 40 seconds’ duration as above, and then spontaneous contractions at four per 10 seconds. 85 r i ‘ = xe Pause and recovery as in last note. 90 ”» . - 3 3 The movements had the same characters as at 60 minutes after poisoning. 110 3 ; ; 4 4 As in last note. STROPHANTHUS SARMENTOSUS: ITS PHARMACOLOGICAL ACTION, 383 Experiment LV.—continued. Contractions in 10 seconds. Interval. —— Notes. Auricles, | Ventricle. After Strophanthus, 2 hours 5 mins. . 2 2 Systole is performed fairly rapidly ; the diastolic phase is prolonged. Mee, LD 4, : 4 4 Each alternate ventricular contraction is superficial, the ‘ intervening contractions are complete. > 3 4 4 Do. do. Pei AD 4, : $5 eae Do. do. tes : : 4 4 Do. do. See LOMmMINS. 4 4 Do. In diastole, the auricles are well dilated; in systole, they expel their contents almost completely. eee tO 5, : 3 3 One of these ventricular contractions was a localised dimpling only ; the others were powerful contractions ; the auricles are contracting and expanding well. . ee ; an ue Six contractions of the auricles and ventricle occur in 30 seconds ; they are almost regular in time ; the left auricle contracts first, then the right; their contraction is rapid but incomplete; the ventricle contracts after the right auricle; it usually becomes uniformly pale in systole and very small; sometimes a wave of contraction passes over the ventricle from left to right; the systolic movements of the heart are rapid ; the diastolic phase is prolonged. HOmes OO. ,, : wae Ae | Six auricular and two ventricular contractions occur in 30 seconds; two or four auricular contractions precede each ventricular contraction; after contraction, the ventricle relaxes a little, and the successive auricular contractions distend it more and more; finally, it again contracts, and sO On. > Pe 20° 5, : ace ‘ies Heart arrested with all chambers greatly dilated. We ol ,, A site Fa Six auricular and six ventricular contractions in 30 seconds ; notes as at 9 hours 45 minutes. i, 40 ,, E Pas sie Arrest of the heart with all chambers fully dilated. From these experiments it appears that, with the smallest doses of extract used, the first change is the greater completeness of the systolic movement of the ventricle, and this occurs very early. Soon afterwards, the ventricle becomes more expanded in diastole, while the rate of the heart’s contractions becomes slower. At this stage, the contractions are regular in time, and the ventricular systole occupies two-thirds of the ventricular cycle, as in the normal heart. This condition is followed by changes in the cardiac cycle ; at first the time of the ventricular cycle becomes equally divided between systole and diastole, and afterwards ventricular diastole is the longer phase, the contrac- tions of the heart continuing to occur at recular intervals up to this point. When the diastolic phase begins to predominate, the ventricle relaxes very rapidly after contract- ing and it is already well dilated when the auricles contract. Irregular pauses in diastole soon interrupt the regularity of the cardiac contractions; these pauses occur | spontaneously, but they can be induced at times by stimulation of peripheral nerves. 384 SIR THOMAS R. FRASER AND MR ALISTER T. MACKENZIE ON Throughout the observations, the auricles contract and dilate regularly until finally the ventricle becomes permanently small and pale in systole and resists the efforts of the auricles to propel their contents into it. The auricles continue to contract after ventri- cular standstill, and they respond to mechanical stimulation after the ventricle has ceased to do so. With larger doses, the main features of the action are similar, viz. : slowing of the heart, increased extent of the systolic and diastolic movements of the ventricle, the tendency to predominance of diastole, the occurrence of diastolic pauses of irregular length and at irregular intervals, and at last permanent systole of the ventricle, which becomes very pale and rigid. The auricles are better filled and empty their contents completely in the early stages, but later the ventricle does not contract rhythmically with the auricles, and several auricular contractions precede each ventricular systole. When the final systole of the ventricle occurs, the contractions of the auricle appear to be unsuccessful in propelling the smallest portion of their contents into the ventricular cavity. In the case of the largest doses which were applied to the surface of the heart the course of events was similar in kind though different in degree. In the last experi- ment described (Experiment LV.), the observations were discontinued when the dia- stolic pause continued for 10 minutes or longer, though in other similar experiments it was found that mechanical stimulation of the heart induced contractions of all its chambers after the organ appeared to be finally arrested in diastole, and repeated irrita- tion changed the type of arrest from diastolic to systolic. In several other experiments, with large doses of extract (not described), the first noticeable change—increased completeness of the systolic movement of the ventricle— became so marked that the heart was permanently arrested in systole within 30 minutes of the application of the extract. In such cases, the ventricular cavity becomes quite obliterated and the cardiac muscle immediately becomes acid in reaction. During the experiments, the laboratory temperature was from 15° C. to 18° C. Experiments afterwards described (heart perfusion) indicate how the quantity of extract affects the nature of the changes produced in the heart. Influence of the Vagus. In order to determine if S. sarmentosus acts on the vagus, numerous experiments were performed on the lines of those just described. With a laboratory temperature of 18° to 20° C., it was found that electrical stimulation of the exposed vagus nerve in the frog caused complete arrest of the heart in extreme diastole for several seconds. Within three minutes of the application of a solution in Ringer’s solution of sulphate of atro- pine to the surface of the heart, electrical stimulation of the vagus produced no effect. When 0:000025 gram of atropine sulphate was instilled within the pericardium, the strongest stimulation (from a Du Bois Reymond’s apparatus and a single bichromate cell) of the isolated vagus nerve produced no effect on the heart during two hours, ex- cepting acceleration in some instances. After this interval, electrical stimulation of the STROPHANTHUS SARMENTOSUS : ITS PHARMACOLOGICAL ACTION. 385 vagus arrested the heart in diastole, but a second application of the same dose of atro- pine abolished the vagus-action for other two hours. When 0:00005 gram of sulphate of atropine was applied to the heart in the same way, the vagus, which had previously been ascertained to be active, failed to slow or arrest the heart, when electrically stimu- lated, during four hours; but after the lapse of four hours electrical stimulation of the nerve was followed by arrest of the heart in diastole for several seconds. Larger amounts of sulphate of atropine appeared to affect the cardiac muscle. Atropine after Strophanthus. Eapervment LVI.—0-0004 gram of extract instilled within the pericardial sac of a frog (Rana temporaria) and, when the effects developed, 0:000025 gram of sulphate of atropine instilled. Contractions in 10 seconds. Interval. Notes. Auricles. Ventricle. Before Strophanthus. 3 minutes . : 5 5 Contractions regular in time and powerful; ventricular systole occupies two-thirds of the cycle. Inserted 0:0004 gram of extract of Strophanthus dissolved in Ringer’s solution inside the pericardial sac. After Strophanthus. 4 minutes . : 6 6 All chambers are larger in diastole and all contract well in | systole. Ss. 5, : , 7 7 Do. do. ” | 10° .,, . : 7 7 Contractions regular and powerful ; ventricular systole occupies two-thirds of the cycle; auricles and ventricle are very large in diastole and contract very well in systole. AD.» : . 6 6 Do. do. ZOE w,, ¢ ; 6 6 The auricles are greatly dilated in diastole and expel only half their contents in systole; the ventricle is small and pale, and only the apex dilates at each auricular systole. 235 eae : : 6 6 The auricles fail to expel their contents and the ventricle is almost in complete systole. Bae! -55 ; : = sae 0:000025 gram of atropine sulphate in solution inserted inside pericardium. On at-55 2 ; 6 6 As last noted. 30 * : . ae sae Six auricular and six ventricular contractions in 30 seconds ; the auricles fail to expel even half of their contents ; ventricular expansion in diastole very slight. 35 A ; ; ae Se, The heart is arrested with the ventricle small and pale and the auricles large and dark ; occasional contractions of the auricles occur, followed by a feeble movement of the ventricle. 40 ” 4 4 Characters as at 33 minutes. 45 an 4 4 Do. 55 5 4 4 Do. 70 i 6 : ae ae Pause for 15 seconds with the auricles moderately dilated and the ventricle a little relaxed; thereafter the auricles expelled their contents better into the ventricle, which was less contracted. 386 SIR THOMAS R. FRASER AND MR ALISTER T. MACKENZIE ON Experiment LVI.—continued. Contractions in 10 seconds, Interval. SSS SS Notes, Auricles, Ventricle. After Strophanthus. 75 minutes . : ang ap Eleven contractions of the heart in 30 seconds, followed by a pause of 35 seconds during which the auricles were dilated and the ventricle contracted. eer. ‘ : ae sis A similar pause for 30 seconds. 82) 5 ; ; Bae ay In 30 seconds twelve cardiac contractions occurred in which ventricular systole preceded auricular systole, and then a short pause followed in each cycle; thereafter a pause of 40 seconds’ duration followed, in which the auricles were dilated and the ventricle contracted ; then the auricles gave a single spontaneous contraction, followed by contraction of the ventricle, and after that ventricular systole preceded auricular systole for ten contractions. Such series of contractions were seen several times. < 95 “4 : : sis wat Final arrest of the heart with the ventricle small and pale and the auricles large and dark. Experiment LVII,—0-0004 gram of extract instilled within the pericardial sac of a frog (Rana temporaria), and later 0000025 gram of sulphate of atropine. Contractions in 10 seconds, Interval. SSS SESE Notes, Auricles. Ventricle. Before Strophanthus. l minute. 3 6 6 Contractions powerful and regular in time ; ventricular systole occupies two-thirds of the cycle. Inserted 0:0004 gram of extract of Strophanthus in Ringer’s solution inside the pericardial sac. After Strophanthus. 5 minutes . ‘ 6 6 As last noted; perhaps the relaxation of the ventricle in diastole is less complete. LO. = : : 6 6 Ventricular systole is a little more complete ; the auricles fill and empty well. 15, . : 5 5 Contractions regularand powerful; ventricular diastole occupies one-half of the cycle and begins before auricular systole ; in systole the ventricle becomes very small and very pale. i: Sale - ; A 8s Slight pauses in diastole are occurring. LY ee ; é ate ee Five auricular and five ventricular contractions occur in 30 seconds ; the slowness is due to diastolic pauses after each relaxation of the heart. 19 Fe ; : ae Pec Three auricular and three ventricular contractionsin 30 seconds; in systole the ventricle becomes very small] and pale. 20 Sa Be? a Inserted 0:000025 gram of atropine sulphate in solution inside the pericardial sac. 22 - ; : Sa: se Four auricular and four ventricular contractions in 30 seconds ; characters as last noted. . :