03 1 M THE LIBRARY OF THE UNIVERSITY OF CALIFORNIA PRESENTED BY PROF. CHARLES A. KOFOID AND MRS. PRUDENCE W. KOFOID >f L\U XXVIII. Description o/'Parkeria lan of growth, was inconsistent with the facts disclosed by a careful study of the organization of the five typical Genera described in my first and second Memoirs : since it had the effect of bringing together Genera whose strongly marked physiological differences required that they should be separated by the widest possible interval; whilst it ranked under different Orders generic types which exhibit the closest physiological relationship. And I then laid it down as a fundamental principle, " that physiological conformity in the condition of each individual segment, as indicated by the structure of its shelly investment, is a character of primary importance; whilst the plan of groiuth, that is, the mode of increase in the number of chambers, is a character of subordinate importance." And in the " Concluding Summary " appended to Part IV. (Philosophical Transactions, 1860, p. 569), I further expanded this doctrine, by showing that all the types which I had described might be ranged in two parallel Series : one of them cha- racterized by that peculiar texture of the Shell which had been appropriately designated PorceUanous by Professor W. C. WILLIAMSON*; whilst the shell-substance of the other has the texture which had been described by the same excellent observer, with equal appropriateness, as Hyaline or Vitreous. A third type of Shell-structure had been noticed by Professor W. C. WILLIAMSON (loc. cit.) under the designation Arenaceous ; the shell being mainly formed, not by a calcareous exudation from the sarcode-body of the animal, but by the aggregation of particles of sand obtained from without, the cement by which these are attached together being all that the animal supplies. These differences in the character of the Shell were regarded by Professor W. as indicative of " physiolo- gical differences in the living sarcode, or secreting animal substance, that have at least fl 3 ,fl*', and opening above into the successive interspaces int 1 , int 2 , int 3 ; at rp, r'p' are seen the radial processes by which these interspaces are bounded ; and at t\ t 2 are seen two of the radial tubes laid open longitudinally. Magnified 70 diameters. Fig. 2. Portion of the preceding enlarged to 250 diameters, to show the arrangement of the component sand-grains. LOFTUSIA. 21. The extraordinary nature of the remains of FORAMINIFERA discovered within the past few years in the Palaeozoic rocks of Canada, has in many ways affected previously received views concerning the testaceous RHIZOPODA. In no respect is this so manifest as in the increased importance accorded to the whole group, on account of the size of its newly added members. On the first separation of the Foraminifera from the Mollusca, mi- nuteness was regarded as a distinctive character of the suborder ; and though it was found necessary to place the Nummulites in a systematic series, which consisted otherwise of microscopical organisms, they were looked upon as exceptional, in point both of mag- nitude and of complexity of structure. The discovery in recent times of specimens be- longing to larger types, such as those dredged off the coast of Borneo by Sir EDWARD BELCHER, and subsequently described by Dr. CARPEXTER under the generic name Cyclo- clypeus*, scarcely excited sufficient attention to affect the general idea that the group was composed of animals of insignificant dimensions ; and it was not until the announce- ment and description by Dr. DAWSON, of Montreal, in 1864, of Eozoon Canadense, that the views of Naturalists became modified as to the size attainable by a class of animals of so simple an organization. It is perhaps not too much to say that the controversy respecting the Protozoic, or at least the Animal, origin and characters of the remains of Eozoon, though eventually centering in questions of minute structure, would never have arisen at all, but for doubts initiated by the dimensions of the fossil. To those who have made the lower forms of animals their special study, the peculiar arrangement of the calcareous shelly layers on an acervuline plan of growth, already well observed in other types of Foraminifera, whilst it accounts for the irregular and asymmetrical external * Philosophical Transactions, 1856, p. 547. 5 F2 740 DR. W. B. CARPENTER AND MR. H. B. BRADY ON contour of the Canadian fossil, equally explains the indefinite extension of the shell- masses. 22. The addition of Eeceptaculites to the list of probable FORAMINIFERA, and the sugges- tion that Stromatopora, Archceocyathus, and some other obscure fossils, hitherto regarded as SPONGES in the absence of any very accurate knowledge of their structure, may find their nearest allies in the same category, are indications of a field of research from which great results may be anticipated. At the present moment, therefore, any investigations tending to throw light on what may properly be termed the gigantic types of Foramini- fera have greatly enhanced interest. 23. Amongst the fossils collected by the late WILLIAM KENNETT LOFTUS, during his Archaeological and Geological researches near the line of the Turko-Persian Frontier *, were certain somewhat obscure bodies, oval or fusiform in shape, and occurring in suffi- cient abundance to give a special character to the rock in which they were imbedded. As they bore a general resemblance to some forms of Alveolina, a well-known genus of Foraminifera, from which, indeed, they seemed to differ in point of size rather than in any structural peculiarities revealed by a cursory examination, they were assigned by their discoverer to that genus ; and, having attracted but little subsequent attention, have been left by Palaeontologists in the same position. 24. In Mr. LOFTUS'S memoirf these fossils are spoken of as specimens " of a gigantic species of Alveolina 3 inches in length;" but no further mention is made of them. Messrs. W. K. PARKER and 1. RUPERT JONES, in one of their earlier papers on the " Nomencla- ture of the Foraminifera" J, make a passing allusion to them. Amongst their notes on the fossil forms of Alveolina, especially those of the Nummulitic Period, they say, " The largest we have seen was collected in Persia by the late Mr. W. K. LOFTUS, and is three inches long and an inch and a half in diameter." The two sentences quoted appear to comprise all that has hitherto been written on the subject of the present paper. 25. A portion of Mr. LOFTUS'S geological collection was presented, some time after his decease, to the Museum of the Natural History Society in Newcastle ; and finding amongst other things a number of examples of this supposed Alveolina, I asked, and readily obtained, permission of the Committee to make such preparations from them as might be requisite for the elucidation of their structure. 26. A very slight examination by means of transparent sections convinced me that, not- withstanding a general similarity in external contour, the internal structure was distinct in many important characters from either of the previously known genera of fusiform FORAMINIFERA. In Alveolina the shell-wall is opaque, homogeneous, and Porcellanous ; in * Mr. LOFTUS'S collections were made in the years 1849-52, during tlie progress of a Joint Commission ap- pointed by the English, Russian, Turkish, and Persian Governments for the demarkation of the Turko-Persian Frontier. t " On the Geology of Portions of the Turko-Persian Frontier and Districts adjoining," by WILLIAM KENNETT LOFTUS, Esq., F.G.S., in Quart. Journ. Geol. Soc. Lond., August 1855, vol. xi. ; foot-note, p. 285. J Annals and Mag. Nat. Hist., Ser. 3, vol. v. (1860), p. 182. TWO GIGANTIC TYPES OF ARENACEOUS FORAMUSTIFERA. 741 Fusulina it is Hyaline and perforate ; whilst in the specimens under consideration it was found to be of distinctly granular texture, resembling the built-up 'tests' of some of the smoother Arenaceous types. The obvious conclusion was that these singular fossils were widely separated in organization from their supposed congeners, and that they belonged to a new type, which probably bore a similar relation to Alveolina and Fusulina, that Trochammina (incerta) bears to Cornuspira and Splrillina. At the suggestion of my friend and colleague Professor T. RUPERT JONES, I propose the generic term LOFTUSIA for the type, thereby to associate with it the name of its discoverer, my lamented pre- decessor as Secretary to the Natural History Society of Northumberland and Durham. 27. External Characters. Most if not all of the specimens of Loftusia that have been brought to this country, bear evidence of having formed part of a hard, compact, Lime- stone rock, from which they have been separated with the utmost difficulty. Indeed the process of mineralization in the animal remains, seems to have gone on simultane- ously with changes in the physical character of the calcareous marl of which the matrix was originally composed ; and the whole has been converted into a uniform subcrystal- line mass, resembling some of the "fossil-marbles" of our Carboniferous system, and capable, like them, of receiving a high polish. The rock is traversed by irregular veins of white crystalline carbonate of lime, very similar to the material that has displaced the sarcode in the chambers and cellular portions of the shells. A piece of the lime- stone with the fossils in situ in the Newcastle Collection (Plate LXXVII. fig. 1) shows the condition in which they are found. It has apparently been long exposed to the action of weather, and is thereby a good deal roughened ; but still it shows how large a proportion of the rock is composed of Organic Remains, chiefly those of Loftusia ; and the course which the fracture has taken, right through the fossils at whatever angle they happened to lie, without deviating to follow either their periphery or any of their structural lines, indicates the determined adhesion which exists between them and the matrix. The appearance of the specimens that have been roughly separated on the spot, testifies to the same fact ; for scarcely any of them show an exterior surface that can be regarded as satisfactorily representing the shell during the life of the animal. The general ex- ternal features, however, are readily made out ; and we are in no worse position in re- spect to this, than we were with the analogous genus Fusulina, which until a year or two ago was only known from the sections of Limestone in which it occurred ; yet the recent discovery of specimens in a free state has done little beyond confirming the accu- racy of the conclusions previously arrived at*. 28. In shape the specimens are all oblong or oval ; but they vary considerably in their proportionate dimensions. Many of the longer ones taper almost to a point at either extremity (Plate LXXVII. fig. 2); whilst a stouter variety (fig. 3) exhibits obtuse * [Mr. BEADY here refers to the results I have recently obtained from the examination of specimens of Futu- Una kindly transmitted to me by Mr. C. A. WHITE, of Iowa, U.S. ; which results conclusively establish the cor- rectness of the opinion I had founded on the study of less perfectly preserved specimens, that Fusulina belongs to the Vitreous or perforated series, instead of ranking with Alveolina (as was supposed by Messrs. PARKER and T. UUPERT JONES) in the Porcellanous or imperforate series. W. B. C.] 742 DE. W. B. CAEPEKTEE AND ME. H. B. BEADT OX rounded ends and a much shorter conjugate axis. The two largest perfect examples that have come under my notice* have about the same weight (three ounces) ; and their dimensions represent fairly, and in by no means an extreme light, the difference in pro- portion alluded to. Their measurements are respectively 3^ inches by 1 inch, and 2j inches by 1|- inch. In other words, the proportions between the axis and the transverse diameter are in the long form as 65 to 20, in the short thick variety as 36 to 20. In both of these specimens the transverse section is circular ; and others of the same form have proportionate dimensions ranging between the two. There are, however, some few in which the transverse section is not circular but lenticular (bi-convex). These are of smaller size, and resemble an almond in general contour (fig. 4) ; but whether the peculiarity is the result of compression, or is due to an inequilateral plan of growth, is not easily determined. I am inclined to attribute it, for reasons which will presently be given, to the former cause ; the more so as there is no ground for specific or even varietal distinction in the structure of the interior. It may be remarked in pass- ing, that a similar or even wider range of variation in external form exists in the iso- morphous genus Aheolina. In that group may be found every gradation in shape, from a nearly perfect sphere to a spindle with pointed ends, having a length four or five times as great as its thickness through the centre. Exceptional specimens of Alveolina, analogous to the compressed examples of Loftusia above referred to, are occasionally though rarely to be met with in Tertiary limestones. 29. As might be expected from the nature of the matrix, the exact condition of the exterior of the test, in respect to inequalities of the surface, ornamentation, or markings determined by structural peculiarities, is not very readily learnt. One or two of the specimens which appear to have been enucleated with less disturbance of the superficial layers than the rest, have a series of tolerably regular furrows, nearly equidistant, tra- versing the shell from end to end, somewhat resembling the uncut portion of that figured in Plate LXXVIIL, though more uniform. These depressions are not so sharply defined nor so deeply excavated as similar lines in the smaller Alveolince, neither have they the same structural significance. It seems probable that they indicate only alternating periods of more and less vigorous growth, or that they are dependent on external circumstances. Arenaceous Foraminifera generally show but little tendency to the surface-ornamenta- tion common in the Vitreous and Porcellaneous groups ; and there is no reason to sup- pose that, in the living condition, Loftusia differed materially in superficial texture from Trochammina. Apart from the longitudinal riblets before described, the exterior may have been either quite smooth or slightly granular, according to the nature of the sand of which it happened to be built up, and the proportion of calcareous material which formed the cement. 30. Internal Structure. Although, in general contour, Loftusia most closely resembles * A specimen in the Museum of the Geological Society, London, must originally have been somewhat larger than either of these ; but, as it has had sections cut from it for microscopic examination, its exact dimensions cannot now be obtained. TWO GIGANTIC TYPES OF AEENACEOUS FOEAMINIFEKA. 743 the two genera already mentioned, Alveolina and Fusulina, its structural relations may be best understood by reference to the lower members of the Rotalian series, such as Planorbulina or Discorbina, or to the still simpler type Inwlutina. This relationship, though at first sight it may appear far-fetched, becomes obvious upon a comparison of their various sections. It is only necessary to imagine one of the simple Rotalians thickened and drawn out at the umbilici, so that what was before a convex disk should become a cylinder tapering more or less towards the ends ; and the analogy is at once apparent. The transverse section of a body so constructed would correspond to the horizontal section of the original disk ; and the ideal diagram B would represent equally well the arrangement of the principal chambers in the Rotalian types and in Loftusia, as seen in section. At the same time, the lines indicating the septa have in some respects a different significance in different cases. In theory the test of Loftusia may be said to consist primarily of a continuous lamina coiled upon itself, like a scroll con- stricted at the ends. The space enclosed by this 'primary lamina' is divided into chambers by longitudinal septa. The septa are of ' secondary ' growth ; that is to say, they are not continuous with the principal wall or ' spiral lamina,' but are rather offshoots from it. The chambers separated by the septa are long and very narrow, and extend from one end of the shell to the other. The septa are not perpendicular to the ' spiral lamina ' as in Alveolina. but very oblique ; and they often take also a more or less oblique direction longitudinally. The longitudinal section is in this way somewhat confused, and less to be relied upon than the transverse, in its bearing upon the form and relation of the various parts. The chambers are further divided by numerous irregular extensions of the secondary or septal system, which it may be convenient to regard as subordinate to the rest. These 'tertiary' ingrowths are generally at right angles to the septa, or nearly so. The diagrams A, B, and C will make somewhat clearer the general plan of structure ; but it must be borne in mind that they are purely ideal, and drawn without reference to scale. The first of the three, A, may be supposed to represent a transverse section of the spiral lamina, or 'primary' wall; B shows, in addition, the 'secondary' system to which the septa belong ; whilst C indicates roughly the subdivision of chambers by the 'tertiary' ingrowths. A. B. C. 31. We must now consider these various structures seriatim, in relation to the appear- 744 DE. W. B. CAEPENTEE AND ME. H. B. BEADT ON ances presented by the actual specimens. The process of infiltration, which has in every instance extended to every part of the organism, has obscured its most important peculiarities. Fortunately in the discovery of Parkeria, and in the results of Dr. CARPEN- TER'S researches on this new and most interesting type of FORAMINIFERA, we find a clue to the reading of several portions that would otherwise have remained unintelligible. Many of the specimens of Parkeria are completely infiltrated with a subcrystalline mineral, very similar in physical characters to that occurring in the chambers of Loftusia. But there are others, which, either from the nature of the matrix, or from the compact texture of the peripheral layer of the test, contain no such deposit ; but remain, as nearly as can be judged, in the state of a dead and empty recent shell. In addition to these, a few examples of Parkeria have been met with partially infiltrated ; only their exterior layers having been consolidated by mineral deposit. Each of these three sets of speci- mens has added its quota to the facts upon which the elucidation of a somewhat com- plex organization depends ; and each has a distinct value in the study of Loftusia. By comparing the appearance of corresponding portions of the infiltrated and uninfiltrated test in Parkeria, reliable data are obtained from which to estimate the condition of Loftusia prior to the filling up of its cavities with inorganic matter. Constant compa- rison with the less altered type has been found needful, in order to demonstrate the organization of the more obscure form ; and for the opportunity of pursuing the subject in this way I am indebted to the kindness of Dr. CARPENTER. 32. The texture of the ' test ' has been stated to be Arenaceous, that is, built up of sand- grains held together by a structureless calcareous cement. It is only necessary to refer to it in this place, in so far as it affects the relation of the walls to the sarcode-cavities. The granules vary considerably in size, but are comparatively much larger than those from which the investment of Parkeria is formed ; hence in the transparent sections of Loftusia the texture does not appear so homogeneous, nor is the outline of the laby- rinthic ingrowths so well defined ; indeed it is obvious that in the recent condition the interior surfaces could not have had the same smooth finish that is to be observed in the open portions (' interspaces ') of Parkeria. 33. After careful examination of a large number of sections of Loftusia, made on the median line in both a transverse and a longitudinal direction, I find no indication of the existence of a central cavity, or anything resembling the large primordial chamber which is so usual a character amongst the FORAMINIFERA. The tendency to fill up the interior of the chambers with shelly ingrowths, which leads to some of the most striking pecu- liarities of its organization, is manifest from the very earliest period of life. We have no very young specimens from which to study the condition of the test before it assumes the form and habit of maturity, except such rare instances as may be found in micro- scopical sections of the limestone matrix, and these are far from satisfactory. One section of the rock, figured at Plate LXXVIII. fig. 5, shows, however, amongst other minute fossils, the transverse section of what I have little doubt is a very young Loftusia(a). The whole is about -fa in. in diameter, and shows the space enclosed by the first turn of the spiral lamina, and about half the circuit of the layer immediately surrounding it. TWO GIGANTIC TYPES OF ARENACEOUS FOEAMINIFEBA. 745 The spiral lamina itself has much the same dense opacity as is presented by older indi- viduals ; and the interior has, so far as can be made out, a loose, arenaceous texture, which, with a further deposit of cementing material, might be expected to give the sort of struc- ture we find at the axis of the mature shell. But in the absence of evidence from speci- mens in the intermediate stages, much importance cannot be attached to the characters of a single individual, the original features of which may have been greatly altered in the process of fossilization. 34. In the fully grown examples, the first circuit of the ' spiral lamina' encloses a space of variable dimensions, in some cases measuring from -^ to -^5- inch in diameter at the centre, and from i to J inch in length, more or less ; and the revolutions succeed each other with tolerable regularity at intervals of -$ to -j inch. From twelve to twenty revolutions are usually found in an adult specimen ; but twenty-five have been noticed in one instance, and doubtless even a larger number may occasionally be met with. 35. The ' spiral lamina' or ' primary skeleton,' as it may be regarded, is composed of almost impalpable calcareous grains closely cemented. It is imperforate, and not more than from - t Q Q to -g-^y of an inch in thickness. From this extreme tenuity it necessarily depends for support upon accessory structures. 36. The space enclosed by its first revolution constituting the central axis (Plate LXXX. fig. 2, c a) is occupied by a mass of shell-substance somewhat resembling in general features a piece of fine sponge, but not quite uniform in its structure. Quite in the centre it assumes the form of a network of irregular anastomosing tubes, with the in- terspaces filled in with shell-substance to a greater or less extent ; but nearer the primary lamina, the irregularly disposed growths resolve themselves into a more definite series, and take a uniform direction. The outer portion becomes in fact the commencement of a system of parallel columnar or tubular processes springing from the inner surface of the spiral lamina, and having their free ends directed inwards (Plate LXXIX. fig. 1, #, and fig. 2, a). This system of parallel tubuli may under favourable circumstances be traced throughout the course of the spiral lamina, except when interrupted by the occur- rence of septa ; and it forms, as well shown in the figures just referred to, a sort of lining to its inner surface. The two structures are in such close juxtaposition that they appear continuous ; and their physiological distinctness is only noticed in exceptional places, when a minute portion of the sarcode appears to have become entangled between them, leaving for a short distance the primary lamina free from the accessory skeleton (Plate LXXIX. fig. 2, b, and Plate LXXX. fig. 3, 1). The nature and extent of the labyrinthic portion of the layers may be best understood from a longitudinal section taken on a line very near the periphery, as shown in Plate LXXIX. fig. 1 ; for whilst a central section presents all the layers in the same aspect, one taken near the exterior bears a different relation to each consecutive layer. Thus we have about the centre of the figure at c the transverse section of the parallel processes just described as lining the spiral lamina ; whilst at a is seen the lateral aspect of the same. Such a section leaves no doubt as to the tubular character of even the more compact portions of the labyrinthic system. MDCCCLXIX. 5 G 746 DB. W. B. CAEPENTEE AND ME. H. B. BEADY ON 37. The septa, which divide the space enclosed by the spiral lamina into chambers, are directly connected with the labyrinthic system, and form a part of the accessory skeleton. There is no continuation of the primary lamina as an imperforate facing to them, nor is there any analogous investing organ. The septa are therefore entirely secondary, and are merely extensions of the labyrinthic system, at regular intervals ; taking a very oblique direction, and terminating on the outer surface of the preceding whorl. The end of the shell, where, from the greater depth of the layers, and the gradual thinning out of the chambers, the septa form a prominent feature, shows most clearly their can- cellated structure and the sort of connexion that exists amongst them. A magnified drawing of this region (Plate LXXIX. fig. 3) bears a strong general resemblance to those portions of Parkeria in which the 'radial tubes' are largely developed, though differing in several essential characters. Not only are the cancellated structures of the septa connected, but there is free communication between the adjacent chambers of the same layer. In other words, whilst the spiral lamina is imperforate, the septa have numerous perforations which allow the passage of sarcode-stolons. 38. But neither the cancellated structure immediately lining the primary lamina, nor its septal developments, can be distinguished in any very definite manner from further ingrowths of subordinate importance, which to a greater or less extent occupy the inte- rior of the chambers. These ' tertiary ' extensions are of very irregular contour ; and being usually built up of the coarsest particles, are less easily made out. 39. Thus whilst the ' primary skeleton,' or what may be regarded as such, is simple and easily understood, the accessory structures are of somewhat complex character, and present appearances very diverse in different specimens, not only from the variable extent to which they are developed, but also in their disposition and texture. Some specimens show scarcely any traces of the accessory skeleton, beyond that already indicated as lining the spiral lamina and forming the septa ; whilst there are others in which the sarcode-cavities are to a great degree filled up by its extension into their interior ; but in either case some portions of the superadded structure in each chamber are prolonged, until they rest upon its floor. In those specimens which have their cavities least filled up, the ingrowths take the form of tubular columns, which traverse the chambers in a radial direction (i. e. perpendicular to the spiral lamina), terminating either on the septum of the previous chamber, or on the exterior wall of the preceding whorl of chambers. In others they are more massive and irregular, and appear to be arranged so as to subdivide the chambers in an incomplete manner into chamberlets. When this latter condition exists, the intersection of the chambers does not take place at regular intervals ; but the wide central portion of each is left comparatively open, and the ingrowths increase in frequency as the sides thin out. Transverse sections of the larger specimens present to the naked eye an appearance as of dark spots set at very regular intervals along the spiral band, which on examination are found to indicate the central larger chamberlets in the successive chambers; that is, the wider portions have the fewest intersecting shelly growths. The smaller chamberlets at each side are hardly to TWO GIGANTIC TYPES OF ARENACEOUS rORAMES'IFERA. 747 be distinguished, unless magnified, in the general labyrinthic system. This distinction of large and small subdivisions is one of degree only, and is not nearly so striking under the microscope as might have been expected from the appearances presented to the unassisted eye ; but it nevertheless does exist, and is a means to an end. 40. The 'accessory skeleton' in Loftusia may be regarded as the homologue of the labyrinthic lamellae in Parkeria ; although there are important differences, the precise significance of which it is impossible to explain with our present limited knowledge of the relation of the two types. The most remarkable of these is that in Loftusia the labyrinthic portions take their rise from the inner surface of the primary spiral lamina, and are directed inwards, that is, towards the central axis ; whilst in Parkeria the order is reversed, the corresponding structures presenting their free ends towards the periphery. Again, the ' radial tubes ' which complicate the labyrinthic system in Parkeria, have no precise analogy to any portion of the accessory skeleton of Loftusia ; the nearest approach to anything of the sort being the lines of tubular communication between the septa of the individual layers, at the ends of the shell, where the layer is thickest ; but here, as in other parts, the spiral lamina cuts off direct communication between the layers. The office fulfilled by the accessory skeleton in Loftusia is, I conceive, simply that of a support to the primary lamina, imparting the necessary solidity to the organism. The subdivision of the chambers into chamberlets seems to be an accidental circumstance, and has but little bearing on the general economy of the animal. 41. In considering its fitness for this purpose, the various external relations of the organism must be borne in mind. It is manifest that the delicate calcareous lamina, described as the ' primary wall,' would, both from its contour and extreme tenuity, be utterly insufficient of itself to protect a mass of sarcode three inches long and one inch in diameter, or to impart that rigidity which, judging from other examples, is necessary to animals of its class. 42. The layer immediately within the 'primary wall' adds greatly to its strength, not only from the additional thickness it imparts, but also from the connexion its septal pro- longations establish between the successive whorls. This portion, however, does not represent a solid mass ; and the septal portions are further weakened by irregular perfo- rations for the stoloniferous tubes connecting the sarcode of the adjacent chambers. The columnar extensions of the shell-substance provide direct vertical support ; and their distribution on the plan described is that likely to ensure the maximum of strength combined with economy of material. A longitudinal section whose direction nearly coincides with the long axis of the chambers (Plate LXXX. fig. 4), shows the primary walls as parallel lines, and the septa (s, s) as slightly oblique bands diverging in the most gradual way, and eventually connecting one wall with the other. In the exaggerated view so obtained, the columnar supports (ts, ts) appear at regular intervals; and throughout the long narrow ends of the chambers they are very close and massive in proportion to the space left for sarcode. As the chamber widens, they diminish in fre- quency and proportionate dimensions. 748 DE. W. B. CAEPENTEB AND ME. H. B. BEADT ON 43. The tubular condition of the principal part of the secondary skeleton has been in- ferred from the appearances presented by portions of exceptional specimens in which the infiltration has least obscured the structure. But in the absence of these, there would have been sufficient evidence to be gained from a close comparison with corresponding appearances in Parkeria, to demonstrate the general tubularity of the labyrinthic system. This character is foreshadowed in a group of Foraminifera of much simpler type. I have elsewhere* described the occurrence \r\ Ellipsoidina (an interesting genus of Fora- minifera discovered by Professor SEGUEXZA in the Miocene clays of Sicily) of a line of tubular columns, w"hose only ostensible office is to support a series of chambers which otherwise would have but little connexion with each other. There is a strong reason why the accessory skeleton which forms so large a proportion of the entire bulk of the shell in Loftusia should be built up on a plan that would ensure the greatest strength with the least weight. The habit of Foraminifera is to live on the surface of the sand or mud at the bottom of the sea, and recent shells taken from a position entirely beneath the top of the mud are dead and empty ; in point of fact the animal dies if it is buried in the sand. It is clear from the nature of the limestone matrix, that the floor of the sea in which Loftusia lived was a very fine calcareous mud, soft and oozy. Now the specific gravity of the material of which the skeleton is built is about 2-7; and sarcode itself may be regarded as but little heavier than water ; so that if, as may be supposed, the hollow sinuses were occupied by sarcode, it would materially alter the relation between the specific gravity of the animal and the element in which it lived ; that is to say, the mass comprising the shell and the sarcode would be of much lower specific gravity than would be the case were the skeleton solid : hence the animal would be correspondingly better fitted to preserve its natural habitat. It is not certain, however, that the laby- rinthic sinuses were occupied by sarcode, or even that the ends projecting into the sarcode-cavities were open ; and it is still possible that they may have fulfilled some distinct functional purpose. In the absence of evidence on this point, it is needless to dwell upon it ; but it is within the range of possibility that the cancellated structure may during life have formed a sort of water-system, or perhaps may even have been filled in part with air. These are but surmises that have presented themselves during the investigation ; but if either condition existed, it would further reduce the general specific gravity. 44. Physical and Chemical Relations. The condition of this fossil is very unfavourable for the determination of the elementary physical characters of the original organism, owing to the completeness of the mineralizing process to which it has been subjected. The sand of which the test is formed is entirely Calcareous ; and its identity in chemical composition with the mineral substance occupying the sarcode-cavities renders it impos- sible to separate or distinguish the two by means of reagents. We learn also from the study of the different condition in which specimens of ParJceria have been found, that the infiltration of a substance having the same chemical composition as the test has a * Annals and Magazine of Natural History, 4 ser. vol. i. p. 333, pi. xiiL TWO GIGANTIC TYPES OF AKENACEOUS FORAMINIFEKA. 749 much greater effect in obliterating its structural characters, and even in obscuring their outlines, than the same process when a different material is concerned. Thus the Calcareous test of Parkeria becomes almost devoid of character when its cavities are filled with a subcrystalline calcareous mineral ; whilst a specimen having its chambers occupied by silex has lost none of its distinctive characters. Chemical analysis of the infiltrated fossil (Loftusia) shows that at least 99 per cent, consists of Carbonate of Calcium, the remaining 1 per cent, being chiefly siliceous matter, a composition repre- senting equally well an average sample of the limestone matrix. The test, therefore, is built up of Calcareous sand-grains, incorporated by a cement of carbonate of calcium. But although the selective power which seems traceable in some Arenaceous Forami- nifcra, enabling them to choose certain sand-grains in preference to others*, has no exercise in respect to the chemical nature of the constituents of the test, there is still something of the same sort observable in relation to the size and distribution of the par- ticles which go to its formation. Thus whilst the septa and the looser portions of the labyrinthic structures are coarsely arenaceous, the spiral lamina is composed of exceed- ingly minute particles. The presence of a number of specimens of the smaller species of Foraminifera imbedded by accident with the sand in which they were living, and now forming a portion of the fabric (Plate LXXX. fig. 3, f,f\ renders it comparatively easy to estimate the size of the sand-grains. The largest that could be satisfactorily measured was about y^ of an inch in diameter ; but they seldom attain more than one half this size. Specimens of the same species abounding in the Limestone matrix run to much larger dimensions. One of the almond-shaped specimens of Loftusia which appears to have had its walls somewhat disintegrated by pressure or otherwise, presents its constituent granules in a very uniform condition, both as to their general appearance and their dimensions, and in this instance the average diameter is about ^^ of an inch. In the more compact portions of the labyrinthic system, the granules are smaller; and as they approach the spiral lamina they become still more minute. In the thin layer which constitutes the lamina itself, a magnifying-power of 500 diameters (the highest I have been able to use with advantage on any section yet prepared) shows the ultimate structure only as a uniform, densely packed mass of particles, individually too small for even approximate measurement. The transparent sections (Plate LXXIX. figs. 4 & 5) cross regions in which the constituent granules are exceptionally small and uniform, and show well their close setting in the compact portions (sg, s(j). 45. The variable external appearance of such of the fossils as have been exposed through the unequal action of the weather, especially on the fractured surfaces is pro- * The question of selection of materials amongst Foraminifcra with composite tests is one to which my attention was drawn in a recent chemical examination of subarenaceous Hil'iola; (Quinqutloculina ayylutinans) obtained by Mr. JEFFREYS from deep water in the Hebrides. Their tests were clearly formed of sand-grains and cement. They occurred in siliceous sand, having but a very small percentage of calcareous matter derived from the debris of Molluscan and other shells ; yet they were entirely soluble in very weak acid, leaving scarcely a microscopic trace of silica. 750 DE. W. B. CAEPEXTEE AND ME. H. B. BEADY OX bably due to other causes rather than to the size of the constituent sand-grains, and chiefly to two, viz. a slight difference in the character of the subcrystalline deposit in the chambers, and the partial disintegration of the shell-structure from pressure or other dis- turbing cause, during, or it may be previous to, the process of mineralization. I have noticed that the specimens in which the exposed portions are the roughest and most granular, are those in which the structure is most confused, not merely in the labyrin- thic growths, which are naturally of coarser texture, but even in the compactly built spiral lamina, which, usually so well denned, is often scarcely traceable in such indi- viduals. 46. It has been stated that the shells of a number of the minuter forms of Foraminifera are to be found imbedded in the shelly material forming the test, their presence being due to the fact of their having been living amongst the sand in the neighbourhood of the animal whilst the process of building its skeleton was being carried on. Their identification is of considerable importance, as they afford, together with the organisms to be observed in the limestone matrix, the only data by which the depth of water wherein the animal lived, may be indicated with any degree of accuracy. In the test itself the following have been noticed : in many cases it is impossible to do more than identify the Genus, specific and varietal characters often depending on peculiarities not to be recog- nized in sections. BilocuKna ringem, Lamk. Planularia longa, Cornuel. Biloculwa contraria, D'Orb. Textularia sagittula, Defrance. Triloculina oblonga, Montagu. . Textularia variabiUs, Will. Triloculina trigonula, Lamk. Textularia pygmcea, D'Orb. Triloculina tricarinata, D'Orb. Sigencrina nodosaria, D'Orb. Quinqueloculina seminulum, Linn. Discorbina, sp. Spiroloculina planulata, D'Orb. Eotalia Beccarii, Linn. Trochammina incerta, D'Orb. The specimens seen in the limestone rock are generally of much larger size than those which have been built into the walls of Loftusia itself. Most of the foregoing varieties occur, and, in addition, the following, Spiroloculina canaliculata, Czjzek. Sulimina ovata, D'Orb. Valvulina Austriaca, D'Orb. Planorlulina ammonoides, D'Orb. Cristellaria, sp. Eotalia Schrceteriana, P. & J. Dentalina, sp. Nummulina, sp. The last named is one of the small thick forms characteristic of the lower Tertiary strata. 47. Mr. W. K. PARKER has given me much assistance in these determinations. The conclusions to be derived from them are that the rock is a lower Tertiary limestone, very similar to some of our Miliolite limestones, but richer in the small arenaceous forms ; TWO GIGANTIC TYPES OF AEENACEOUS FOBAMINIFEEA. 751 t and that the sea-bottom was a soft calcareous mud, and lay at a depth of from ninety to one hundred fathoms. A few minute fragments of Molluscan shells resembling columns of Pinna, and small pieces of fossil POLYZOA, are the only remains that are to be found of animals of higher organization than Foraminifera. 48. Zoological Relations. From the details of structure which have been adduced, the genus Loftusia would seem to find a natural place at the head of the Arenaceous series of Foraminifera, a position corresponding to Alveolina in the Porcellanous group, and Fusulina amongst the Vitreous forms. In texture it is similar to the higher Trocham- mince. Its general plan, in so far as the primary skeleton is concerned, is simple ; and there is no approach to the more complex organization found in the shell of the Num- mulite or its immediate allies. Its most striking external difference from the other members of the Arenaceous group (Parkeria of course excepted) is its size; and the chief peculiarity of its internal structure consists in the secondary shelly growths neces- sary for the support of the enlarged test. 49. Notwithstanding great diversity in the size and contour of the specimens, their dif- ference is attributable to degree of development depending on external causes, rather than to specific or varietal distinction ; and with a slight reservation in respect to the compressed specimens, I propose to place all under one species, of which the following will serve as a description. LOFTUSIA, gen. nov. Testd lilerd, regulariter rotundatd, axe elonyato ; transverse sectd orbiculari (aut len- ticulari ?} ; ex spird bene compositd, cujus orbis qidsque orbem antecedentem penitus am- plectitur, constante ; in numerosissimos loculos septis longitudinalibus partita iterum plus minusve subdivisos ; structurd arenaced ; aperturis (multis, complexis, labyrintheis?). L. PEESICA, spec. nov. L. testa elongatd, ovatd vel fusiformi ; transverse sectd orbicu- lari (vel compressd ?} ; extremitatibus obtusis aut rotundatis ; loculis multis, angustis, interne cancellatis, cancellis ad axem versis ; septis perobliguis ; aperturis (multis, compleocis, super facie ultimi loculi arcuatd sparsis ?) ; superficie Icevigatd aut suba- renaced, interdum sulcis subtilibus paribus intervallis inter se distributis in longitu- dinem sculptd. Longa, 1-5 usque ad 3'5 poll. Lata '5 usque ad l'2o poll. LOG. Persia, fossilis. Distribution. In respect to distribution but little can be said beyond what appears in Mr. LOFTUS'S Memoir (op. cit., p. 235) and the notes accompanying the specimens presented to the British Museum and the Geological Society ; but the information from these sources is deficient both in Geographical and in Geological details. In describing the geological characters of a section lying " on the direct road between Kalah Tul and Isfahan," after speaking of a confused series of gypsiferous rocks, which seem ' ; as if the bed had been shot off the side of Mererari during its sudden elevation," the author adds, " Masses of gravel-conglomerate lie in the bed of the stream and high up the slopes of 752 DE. TV. B. CAEPEXTEE AND ME. H. B. BEADY OX the mountain through which the Ab-i-Bazuft flows," and then follows this foot-note : " A few miles N.E. of this stream (but before reaching the left bank of the Kuran at Du Pulun) I procured from a hard rock of blue marly limestone a gigantic species of Alveo- lina, three inches in length." The station appended to the specimens in the British Mu- seum and the Geological Society is the " Kellapstun Pass, near Du Pulun, Bakhtiyari Mountains, Persia." Unfortunately scarcely any of the names mentioned appear on the " Sketch-map" that accompanies the paper ; and for scientific purposes the district referred to may be said to be as yet unmapped. I am indebted to Mr. KEITH JOHNSTON, of Edin- burgh, for a detailed tracing of the region, procured with some pains from unpublished German sources; from which it appears that Du Pulun is on the 32 parallel N. Lat., and that a Longitude of 50 30 E. would indicate a point halfway between it and the little mountain stream Ab-i-Baziift. The district lies between the N.E. corner of the Persian Gulf and Isfahan. 50. Our knowledge of the Geological distribution of the type may be summed up in few words. Mr. LOFT us appears to regard the "blue marly limestone" as belonging to the oldest Tertiary rocks, though he does not say so very distinctly ; and the evidence of the Foraminifera imbedded in it leads pretty conclusively to the same view. The data afforded by the Microzoa are probably sufficient confirmation, in the absence of any satis- factory record of larger fossils from the same geological horizon. In conclusion I have to express the obligation I am under to my friends Mr. W. K. PARKER and Dr. CARPENTER, for the interest they have taken in the subject of the pre- sent paper; to both for suggestions derived from their large knowledge and philo- sophic views in connexion with the Protozoa generally, to the latter for light thrown upon obscure points by the study of collateral structures in Parfceria, and above all for the opportunity of constant reference to specimens of that genus, without which the history now given could not have been so far elaborated. EXPLANATION OF THE PLATES OF LOFTUSIA. PLATE LXXVII. Fig. 1. Piece of Loftusia-limestone, the surface of which has been ' weathered' by expo- sure, and the sections of the fossils thereby brought into relief (natural size). Figs. 2, 3 & 4. Loftusia Persica (natural size). Of these figs. 2 & 3 represent longer and shorter individuals of the normal form, whilst fig. 4 is the compressed variety with lenticular transverse section described in 27. Figs. 2", 3", 4" represent the transverse sections of the three specimens drawn to their natural size. Fig. 5. Section of the Limestone rock forming the matrix of Loftusia, with Foraminifera in situ. a. Young specimen of Loftusia cut transversely, showing the space enclosed TWO GIGANTIC TYPES OF AEENACEOTJS FOKAMINIFEKA. 753 by the first turn of the spiral lamina, and about half the circuit of the layer immediately surrounding it. Sections of several other Foraminifera included in the matrix are seen. PLATE LXXVIII. General view of the structure of Loftusia, showing the appearances of sections cut on different planes, and their relation to each other. Magnified about 4 diameters. PLATE LXXIX. Fig. 1. Longitudinal section of Loftusia, very near the periphery, viewed as an opake object. Magnified 45 diameters. s I. Imperforate primary skeleton or spiral lamina. a. Series of parallel, columnar, secondary shell structure, immediately lining the primary lamina. c. The same cut transversely. It must be borne in mind that this is not a central section, but a tangential one cut so near the periphery that the direction of the section differs in its relation to each successive layer, so as to illustrate fully the arrangement of the accessory skeleton. The centre, representing the innermost of the four layers of which portions are drawn, shows at c the appearance presented by the parallel columnar shelly processes (a, a) when cut across. The portion of the figure to the right exhibits a mass of the accessory skeleton formed from overlapping septa, and shows the sort of intercommunication between them. Owing to the direction of the section, or possibly to accidental causes, the spiral lamina (s I), the continuity of which is usually a prominent character, appears lost at the end of the layers to the right, though well seen in the other portions of the specimen. Fig. 2. A portion of a transverse section, s I represents the spiral lamina, and at I its distinctness from the accessory structures may be noticed. This condition may be found at points in almost every specimen. The nature of the septa, as prolongations of the series of columnar processes (a, a, a) lining the pri- mary lamina, may be easily traced. The subdivision into chamberlets, a large chamberlet (c c, c c, c c) occupying the centre of each chamber (see 39), is also apparent. Magnified 45 diameters. Fig. 3. Longitudinal section of a portion of a layer near the end of the central axis. At this point, where the layer is widest and the chambers thin out and over- lap, the intercommunication of the septa is often so regular and complete that it resembles very strongly the portions of Parkeria in which the system of radial tubes is most developed. MDCCCLXIX. 5 H 754 ON TWO GIGANTIC TYPES OF AKENACEOUS FORAMINIFEKA. Figs. 4 & 5. Transparent sections, to illustrate the nature of the subcrystalline calcareous deposit filling the chambers, and the close setting of the sand-grains (s g, s g) in the compact portions of the test. Magnified 45 diameters. PLATE LXXX. Fig. 1. General view of a transverse section of an average specimen, mounted in Canada balsam and seen as a transparent object, under a low power. Magnified 5 diameters. Fig. 2. A similar preparation from one of the compressed specimens alluded to at 28. Magnified 6 diameters. All the almond-shaped specimens are small, possibly not fully grown ; and if so, the composite test may have been less consolidated than in mature indi- viduals. This may account for their form. The arrows If, If indicate a line on which pressure at the two sides appears to have caused the fracture of many of the layers. Fig. 3. Enlarged view of a portion of the transparent section shown in fig. 1. Magni- fied 25 diameters. s I, si. Spiral lamina. J, b. Points at which the ' primary ' and the ' accessory ' skeleton are not in close approximation, and show their distinctness. f,f,f- Imbedded Foraminifera in situ. Fig. 4. Longitudinal central section of a single layer near the periphery, showing the regularity of ' tertiary ' processes subdividing the chambers. si, s I. Primary skeleton. s, s. Septa. t, s. Tertiary shelly processes affording perpendicular support. Carpenter LXXli PARKERIA. A llollirk del el hill WW.-til irn, Carpenter Phil, flwi&MDCCCLXK: /YnA- 1 JOUI1 A HolKdr del rt lilh. Carpenter fe Phil. TnuuMDCCCLSJLPLate J XW. 2. I ' - I r-p '#*" A.HoIlick del etbtk. WWesl imp- Phil. Trams. MD CCCLXDC J' ABolliokdel etM, WWrsI ,,,. Carpenter I 4 WJH.V^eoiey J,th W Weotimp. / 1W/..ZJWIS.MDCCCLXIX Plate 1 ,XXY!! 3. ' W Wf :.' Phil. TTVWS MD CCCLXK-PZote LXXM o W < H CO ck del et lilli I '/, il Trasis MDCCCLXIX / >lajte I ,XX1X A.Hollici del et lith WWe-i hil. Traun*. MDCCCLXDCPZofc I, XXX If- ^.4-ft 3s^r33i8w^ Ara.-^ 5S-*^*SiU^ *l. 735>iaBR&MfiMaiuitf ^S*^ SSO* r*i^-*.- - -.V : RETURN TO LOANPERIOD o 2/84 BERKELEY, CA 94720