CHAPTER IX.-GEOGRAPHICAL DISTRIBUTION.
226. Universal Marine Distribution.-Radiolaria occur in all the seas of the world, in all climatic zones and at all depths. Probably under normal conditions they always float freely in the water, whether their usual position be at the surface (pelagic), or at a certain depth (zonarial), or near to the bottom of the sea (abyssal). This appears both from numerous direct observations, as well as from conclusions which may be drawn from their organisation (and especially their promorphobogy) regarding their floating life (compare §§ 40-50, 219, 220). Hitherto no observation has been recorded, which justifies the assumption that Radiolaria live anywhere upon the bottom of the sea (on stones, Algae, or other firm substances), either sessile or creeping. They perform the latter action, however, when they fall accidentally upon a firm basis or are accidentally placed upon it, but they seem normally always to float freely in the water with pseudopodia radiating in all directions. Active free-swimming movements are only met with in the case of the flagellate zoospores (§ 142). The development of Radiolaria in large masses is very remarkable (see note A), and in many parts of the ocean is so great that they play an important part in the economy of marine life, especially as food for other pelagic and abyssal animals (see note B). Medium salinity of the water seems to be most favourable to their development in masses, although it is not unknown in seas of high and low salinity (see note C). There are no Radiolaria in fresh water (see note D).
A. The development of Radiolaria takes place in many parts of the ocean in astonishingly large masses on the surface, in different strata, and near the bottom. The Collodaria (and especially the Sphaerozoida) often cover the surface of the sea in millions, and form a shining layer, phosphor escent in the dark like the Noctilucae, as I observed in 1859 in the Strait of Messina, in 1866 at the Canaries, and in 1881 in the Indian Ocean. Similar masses of Sphaerozoum and Acanthometron were seen by Johannes Müller on the French and Ligurian coasts (L. N. 12), and John Murray found another in the Gulf Stream, off the Faerde Islands, from the surface to a depth of 600 fathoms ; considerable masses of large PHAEODARIA live there also.
B. The alimentary canal of Medusae, Salpae, Crustacea, Pteropoda, and many other pelagic animals is a rich field for the discovery of Radiolaria, and many of the species hereinafter described are from such sources. Fossil coprolites too (e.g., those from the Jura) often contain many Polycystina.
C. Some ACANTHARIA (Aacanthometra) and PHAEODARIA (species of Mesocena and Dictyocha)live in the Baltic ; I found their skeletons in the alimentary canal of Aurelia, Ascidians and Copepods.
D. The so-called "fresh-water Radiolaria," which have been described by Focke, Greeff, Grenacher and others, are all Heliozoa, without either central capsule or calymma.
227. Local Distribution.-As regards their local distribution and its boundaries the Radiolaria show in general the same relations as other pelagic animals. Since they are only to a very slight extent, if at all, capable of active horizontal locomotion, the dispersion of the different species from their point of development (or "centre of creation") is dependent upon oceanic currents, the play of winds and waves and all the accidental causes which influence the transport of pelagic animals in general. These passive migrations are here, however, as always, of the greatest significance, and bring about the wide distribution of individual species in a far higher degree than any active wanderings could do. Any one who has ever followed a stream of pelagic animals for hours and seen how millions of creatures closely packed together are in a short time carried along for miles by such a current, will be in no danger of underestimating the enormous importance of marine currents in the passive migration of the fauna of the sea. Such constant currents may, however, be recognised both near the bottom of the sea and at various depths, as well as at the surface, and are therefore of just as much significance for the abyssal and zonarial as for the pelagic Radiolaria. It is easy to explain by this means how it is that so many animals of this class (probably indeed the great majority) have a wide range of distribution. The number of cosmopolitan species which live in the Pacific, Atlantic and Indian Oceans is already relatively large. In each of these three great ocean basins, too, many species show a wide distribution. On the other hand, there are very many species which are hitherto known only from one locality, and probably many small local faunas exist, characterised by the special development of particular groups. The observations which we at present possess are too incomplete, and the rich material of the Challenger is too incompletely worked out, to enable any definite conclusions to be drawn regarding the local distribution of Radiolaria.
The statements made in the systematic portion of this Report regarding the distribution of the Challenger Radiolaria are very incomplete. In most cases only one locality is mentioned, and that is the station (§ 240) in the preparations or bottom deposit from which I first found the species in question. Afterwards I often found the same species again in one or more additional stations (not seldom in numerous preparations both from the Pacific and Atlantic), without the possibility of adding them to the habitat recorded under the description. The necessary accurate determination and identification of the species (measuring the different dimensions, counting the pores, &c.), would have occupied too much time, and the writing of this extensive Report would have lasted not ten but twenty or thirty years.
228. Horizontal Distribution.-From the extensive collections of the Challenger and from the other collections which have furnished a welcome supplement to them, it appears that Radiolaria are distributed throughout all seas without distinction of zones and physical conditions, even though these latter may be the cause of differences in their qualitative and quantitative development. In the case of the Radiolaria as well as of many other classes of animals, the law holds good that the richest development of forms and the greatest number of species occurs between the tropics, whilst the frigid zones (both Arctic and Antarctic) exhibit great masses of individuals, but relatively few genera and species (see note A). In the Challenger collection the greatest abundance of species of Radiolaria is exhibited by those preparations which were collected at low latitudes in the immediate neighbourhood of the equator ; this is true both of the Atlantic (Stations 346 to 349) and of the Pacific (Stations 266 to 274) ; in the former the richest of all is Station 347 (lat. 0° 15w' S.), in the latter Station 271 (lat. 0° 33' 5.) (see note B). From the tropics the abundance of species seems to diminish regularly towards the poles, and more rapidly in the northern than in the southern hemisphere ; the latter also appears, considered as a whole, to possess more species than the former. A limit to the life of the Radiolaria towards the poles has not yet been found ; the expeditions towards the North Pole (see note C), like those towards the South (see note D), have obtained bottom-deposits and ice enclosures which contained Radiolaria ; in some of the most northerly and most southerly positions which were reached the number of Radiolaria enclosed in the ice was relatively great.
A. The greater abundance of Radiolaria in the tropical seas is probably to be explained by the more favourable conditions of existence, and in particular the larger quantity of nutritive material (especially of decayed animals) and not by the higher temperature of the surface, for at depths of from 2000 to 3000 fathoms where the abyssal Radiolaria live, the temperature is but little above the freezing point or even below it (compare the bottom temperatures in the list of Challenger Stations, § 240).
B. Station 271 of the Challenger Expedition, situated almost on the equator in the Mid Pacific (lat. 0° 33' 5.), exceeds all other parts of the earth, hitherto known, in respect of its wealth in Radiolaria, and this is true of the pelagic as well as of the zonarial and abyssal forms. In the Station List the deposit at this point is stated to be "Globigerina ooze" ; but after the calcareous matter has been removed by means of acid, the purest Radiolarian ooze remains, rich in varied and remarkable species. More than one hundred new species have been described from this Station alone.
C. Regarding the Arctic Radiolaria compare the contributions of Ehrenberg (L. N. 24, pp. 138, 139, 195) and Brady on the English North Polar Expedition, 1875-76 (Ann. and Mag. Nat. Hist., 1878, vol. 1. pp. 425, 437).
D. Regarding the Antarctic Radiolaria, compare § 230, note A, and
Ehrenberg, Mikrogeobogie (L. N. 6, Taf. xxxv., A.), also L. N. 24, pp.
229. Fauna of the Pacific Ocean.-From the splendid discoveries of the Challenger, and the supplementary observations obtained from other sources, the Pacific seems to be the ocean basin which is richest both quantitatively and qualitatively in Radiolarian life, excelling both the Indian and Atlantic Oceans in this respect. It may be assumed with great probability that by far the largest port.ion of the Pacific has a depth of between 2000 and 3000 fathoms, and that its bottom is covered either with Radiolarian ooze (§ 237) or with a red clay (§ 239), which contains many SPUMELLARIA and NASSELLARIA, and has probably been derived for a great part from broken down and metamorphosed Radiolarian ooze (see note A). Pure Radiolarian ooze was found by the Challenger eastwards in the Central Pacific (over a wide area between bat. 1 2° N. and 12° S., Stations 265 to 274), and also westwards in the latitude of the Philippines, twenty degrees to the east of them (between lat. 5° N. and 15° N.). The great abundance of Radiolaria present in the neighbourhood of the Philippines and in the Sunda Sea was already known from other investigations (note B). The red clay also, which covers a great part of the bottom of the North Pacific, and which was obtained of very constant composition by the Challenger between bat. 35° N. and 38° N., from Japan to the meridian of Honolulu (from long. 144° E. to 156° W.), is so pre-eminently rich in Radiolaria that it often approaches in composition the Radiolarian ooze, and has probably been derived from it. The track of the Challenger through the tropical and northern parts of the Pacific describes nearly three sides of a rectangle, which includes about half of the enormous Pacific basin, and from this as well as from other supplementary observations it may with great probability be concluded that by far the largest part of the bed of the Pacific (at least three-fourths) is covered either with Radiolarian ooze or with red clay, which contains a larger or smaller amount of the remains of Radiolaria. With this agrees also the important fact that the numerous preparations of pelagic materials and collections of pelagic animals, which were collected by the Challenger in the Pacific, almost always indicate a corresponding amount of Radiolarian life on the surface. This is true in particular also of the South Pacific, between lat. 33° S. and 40° S. (from long. 133° W. to 73° W., Stations 287 to 301) ; the surface of this southern region and the different bathymetrical zones were rich in new and peculiar species of Radiolaria.
A. Many specimens of bottom-deposits from the Pacific, which are entered in the Challenger lists either as "red clay" or "Globigerina ooze," contain larger or smaller quantities of Radiolaria, and the number of different species of SPUMELLARIA and NASSELLARIA which they contain is often so great that the deposit might have been almost as appropriately termed "Radiolarian ooze," e.g., Stations 241 to 245, and 270, 271 (compare §§ 236-239).
B. Pacific Radiolarian ooze was first obtained by Lieutenant Brooke (May 11, 1859) between the Philippines and Marianne Islands, from a depth of 3300 fathoms (lat. 18° 3' N., long. 129° 11' E.). Ehrenberg, who first described it, found seventy-nine different species of Polycystina in it, and reported "that their quantity and the number of different forms increased with the depth" (Monatsber. d. k. preuss. Akad. d. Wiss. Berlin, 1860, pp. 466, 588, 766).
230. Fauna of the Indian Ocean.-As regards its Radiolarian fauna the Indian Ocean is the least known of the three great basins. Still the few limited spots, regarding which investigations are forthcoming, indicate a very rich development of Radiolarian life. Probably it approaches more nearly the fauna of the Pacific than that of the Atlantic, both as regards the abundance and the morphological characters of its species. The researches of the Challenger are very limited and incomplete as regards the Indian Ocean, for the expedition only just touched upon this great ocean basin (2000 to 3000 fathoms deep) at its two extremities (westwards at the Cape of Good Hope and eastwards at Tasmania), its course lying for the most part south of lat. 45° S. and extending beyond lat. 65° S. (from Station 149 to 158, south of lat. 50° S.). It is true that this portion of the South Indian Ocean was shown to contain Radiolaria everywhere, but these were more plentiful in individuals than in species. Only from Station 156 to Station 159 (between lat. 62° and 47° S., and long. 95° and 130° E.) was the bottom, which consisted partly of Diatom ooze and partly of Gbobigerina ooze, richer in species (see note A). The gaps left by the Challenger in the investigation of the Indian Ocean, have, however, been to some extent filled from other sources. As early as 1859 the English "Cyclops" expedition had shown that the bottom of the Indian Ocean to the east of Zanzibar (lat. 9° 37' S., bong. 61° 33' W.) is covered with pure Radiolarian ooze (see note B). Also since the Tertiary rocks of the Nicobar Islands are for the most part of the same composition, and since a great abundance of Radiolaria has been shown to be present both in the east part of the ocean, between the Cocos Islands and the Sunda Archipelago (see note C), and in the northern part or Arabian Sea between Socotra and Ceylon (see note D) ; it may be assumed with great probability that the greater part of the basin of the Indian Ocean, like that of the Pacific, is covered either with Radiolarian ooze or with the characteristic red clay. With this agrees the richness of the surface of the Indian Ocean in Radiolaria of the most various groups, which has been more extensively demonstrated.
A. The Radiolarian fauna collected by the Challenger on the voyage from the Cape to Melbourne, shows in part, namely, from Station 156 to Station 158, very peculiar and characteristic composition ; in particular, the Diatom ooze of Station 157 passes over in great part into a Radiolarian ooze, mainly composed of Sphaerellaria. This is worthy of a more thorough investigation than I was able, owing to lack of material and time, to give it.
B. The remarkably pure Radiolarian ooze of Zanzibar, discovered by Ehrenberg
in 1859, was the earliest known recent example of that deposit. It was
brought up by Captain Pullen of the English man-of-war "Cyclops," from
a depth of 2200 fathoms, between Zanzibar and the Seychelles, and "under
a magnifying power of 300 diameters, showed at the first glance a mass
of almost pure Polycystina, such as no sample of a deep-sea deposit has
hitherto shown. It is very noticeable that in the whole of this mass of
living forms, no calcareous shells are to be seen " (Ehrenberg,
L. N. 24, pp. 148, 149).
C. For the most important material from the Indian Ocean, I am indebted to Captain Heinrich Rabbe of Bremen, who during many voyages in the Indian Ocean, in his ship "Joseph Haydn," made numerous collections in different localities with the tow-net and the trawl, and admirably preserved the rich collections thus made. The greatest abundance of Radiolaria was found in those obtained to the east of Madagascar, and next in those from the neighbourhood of the Cocos Islands. I take this opportunity of expressing my thanks to Captain Rabbe for the liberality with which he placed all this valuable material at my disposal.
D. On my voyage from Aden to Bombay, and thence to Ceylon (1881), and especially on my return journey from Ceylon, between the Maldive Islands and Socotra (1882), I carried on a number of experiments with a surface net, which yielded a rich fauna of pelagic animals, and among them many new species of Radiolaria, for observation. On several nights when the smooth surface of the Indian Ocean, unrippled by any wind, shone with the most lovely phosphorescent light, I drew up water from the surface with a bucket, and obtained a rich booty. A number of other new species of Radiolaria from very various parts of the Indian Ocean I obtained from the alimentary canal of pelagic animals, such as Medusae, Salpae, Crustacea, &c. Although the total number of Radiolaria known to me from the Indian Ocean is much less than from the Atlantic and Pacific, there are several new genera and numerous species among them, which show that a careful study of this fauna will be of wide interest.
231. Fauna of the Atlantic Ocean.-The Atlantic Ocean in all parts, of which the pelagic fauna has been examined, has shown the same constant presence of Radiolaria, and in certain parts of its abyssal deposits a larger or smaller quantity of different types belonging to this class ; on the whole, however, its Radiolarian fauna is inferior to that of the Pacific, and probably also to that of the Indian Ocean, both in quantity and quality. Pure Radiolarian ooze, such as is so extensively found on the floor of the Pacific, and in certain places in that of the Indian Ocean, has not yet been found in the Atlantic (see § 237). The red clay, too, of the deep Atlantic does not seem to be so rich in Radiolaria as that of the Pacific ; nevertheless, the number of species peculiar to the Atlantic is very large, and at certain points the abundance of species as well as of individuals seems to be scarcely less than in the Pacific. This is especially true of the eastern equatorial zone not far from Sierra Leone, Stations 347 to 352 (see note A) ; also of the South Atlantic between Buenos Ayres and Tristan da Cunha, Stations 324, 325, 331 to 333 (see note B) ; and, lastly, in the North Atlantic in the Gulf Stream and near the Canary Islands (see note C). The fauna of the latter agrees for the most part with that of the Mediterranean (see note D). In addition to the material collected by the Challenger, other deep-sea investigations have furnished bottom-deposits from different parts of the ocean, which have proved very rich in Radiolaria (see note E). Furthermore, since the island of Barbados consists for the most part of fossil Radiolarian ooze, it is very probable that at certain parts of the tropical Atlantic true Radiolarian ooze, like that of the Pacific and Indian Oceans, will eventually be found in depths between 2000 and 3000 fathoms, perhaps over a considerable area.
A. The tropical zone of the eastern Atlantic seems to be especially rich in peculiar Radiolaria of different species. This is shown by numerous preparations from the surface, and from various depths (between lat. 3° S. and 11° N., and long. 14° W. to 18° W.), which were made towards the end of the cruise. Unfortunately no bottom-deposits were obtained from the most important stations (except Nos. 346 and 347, depths 2350 and 2250 fathoms) in this region ; at these the deposit was a Globigerina ooze containing numerous different species of Radiolaria.
B. In the South Atlantic, between Buenos Ayres and Tristan da Cunha (between lat. 35° S. and 43° S., long. 8° W. and 57° W.) there appears to be a long stretch covered partly with Globigerina ooze (Stations 331 to 334), or red clay (Stations 329, 330), partly with blue mud (Stations 318 to 328), which contains not only large masses of individuals but numerous peculiar species of SPUMELLARIA and NASSELLARIA. The preparations from the surface-takings of this region are also rich in these, as well as in peculiar PHAEODARIA.
C. The northern part of the Atlantic appears on the whole to be inferior to the tropical and southern portions as regards its richness in Radiolaria, and from the western half more especially, only few species are known. From my researches at Lanzerote in 1866-67, it appears that the pelagic fauna of the Canary Islands is very rich in them, as is also the Gulf Stream in the neigh bourhood of the Faeröe Channel, according to the investigations of John Murray (see his Report on the "Knight-Errant" Expedition, Proc. Roy. Soc. Edin., vol. xi., 1882).
D. The Radiolaria of the Mediterranean are of special interest, because almost all our knowledge of these organisms in the living conditions and of their vital functions has been derived from investigations conducted on its shores. Johannes Müller laid the foundation of this knowledge by his investigations at Messina, and on the Ligurian and French coasts at Nice, Cette, and St. Tropez (L. N. 10). The many new Radiolaria which I described in my Monograph (L. N. 16, 1862), were for the most part taken at Messina, the place which possesses a richer pelagic fauna than any other, so far as is yet known, in the Mediterranean. Other new species I found afterwards at Villafranca near Nice, in 1864 (L. N. 19), at Portofino near Genoa (1880), at Corfu (1877), and at other points on the coast. In Messina also, Richard Hertwig collected the material for his valuable treatise on the Organisation of the Radiolaria (L. N. 33), after he had previously made investigations into their histology at Ajaccio in Corsica (L. N. 26). Lastly, at Naples, Cienkowski (L. N. 22) and Karl Brandt (L. N. 38, 39, 52) carried out their important investigations into the reproduction and symbiosis of the Radiolaria. With respect to the character of its Radiolaria, the Mediterranean fauna is to be regarded as a special province of the North Atlantic.
E. Among the smaller contributions which have been made towards our knowledge of the Atlantic Radiolarian fauna, the communications of Ehrenberg on the deposits obtained in sounding for the Atlantic cable, and on the Mexican Gulf Stream near Florida, deserve special mention (L. N. 24, pp. 138, 139-145).
232. Vertical Distribution.-The most important general result of the discoveries of the Challenger, as regards the vertical or bathymetrical distribution of the Radiolaria, is the interesting fact that numerous species of this class are found living at the most various depths of the sea, and that certain species are limited to particular bathymetrical zones, i.e., are adapted to the conditions which obtain there. In this respect three different Radiolarian faunas may be distinguished, which may be shortly termed "pelagic," "zonarial," and "abyssal." The pelagic Radiolaria swim at the surface, and when they sink (e.g., in a stormy sea), only descend to a small depth, probably not more than from 20 to 30 fathoms (§ 233). The complicated conditions of existence created by the keen struggle for existence at the surface of the sea, give rise to the formation of very numerous pelagic species, especially of Porulosa (SPUMELLARIA and ACANTHARIA). The abyssal Radiolaria are very different from those just mentioned; they live at the bottom of the deep-sea, not resting upon nor attached to it, but probably floating at a little distance above it, and are adapted to the conditions of existence which obtain there (§ 235). Here the Osculosa (NASSELLARIA and PHAEODARIA) seem to predominate. The zonarial Radiolaria live floating at various depths between the pelagic and abyssal species (§ 234). In their morphological characters they gradually approach the pelagic forms upwards and the abyssal downwards.
The views which have hitherto been held regarding the bathymetrical or vertical distribution of the Radiolaria have been entirely altered by the magnificent discoveries of the Challenger, and especially by the important observations of Sir Wyville Thomson (L. N. 31) and John Murray (L. N. 27). These two distinguished deep-sea explorers have, as the result of their wide experience, been convinced that Radiolaria exist at all depths of the ocean, and that there are large numbers of true deep-sea species which are never found at the surface of the sea nor at slight depths (L. N. 31, vol. i. pp. 236-238 ; L. N. 27, pp. 523, 525). The result of my ten years' work upon the Challenger Radiolaria, and the comparative study of more than a thousand mountings from all depths, has only been to confirm this opinion, and I am further persuaded that it will some day be possible by the aid of suitable nets (not yet invented) to distinguish different faunistic zones in the various depths of the sea. In this connection may be mentioned the specially interesting fact that the species of Radiolaria of one and the same family present in the different depths characteristic morphological distinctions, which obviously correspond to their different physiological relations in the struggle for existence. Owing to those extensive discoveries, the representation which I gave in my Monograph (1862, L. N. 16, pp. 172-196) of the vertical distribution of the Radiolaria, and of their life in the greatest depths of the sea, has been entirely changed. Compare also Bütschli (L N. 41, p. 466).
233. The Pelagic Fauna.-The surface of the open ocean seems everywhere, at a certain distance from the coast at least, to be peopled by crowds of living Radiolaria. In the tropical zone these pelagic crowds consist of many different species, whilst in the frigid zones, on the other hand, they are made up of many individuals belonging to but few species. Most of these inhabitants of the surface may be regarded as truly pelagic species, which either remain always at the surface or descend only very slightly below it. Probably most Porulosa (both SPUMELLARIA and ACANTHARIA) belong to this group ; whilst but few Osculosa occur in it, and fewer PHAEODARIA than NASSELLARIA. In general the pelagic Radiolaria are distinguished from the abyssal by the more delicate and slender structure of their skeletons ; the pores of the lattice-shells are larger, the intervening trabeculae thinner ; the armature of spines, spathillae, anchors, &c., is more various and more highly developed. Numerous forms are to be found among the pelagic Radiolaria which have either an incomplete skeleton or none at all. When the pelagic forms leave the surface on account of unfavourable weather, they appear only to sink to slight depths (probably not below 20 or 30 fathoms). Within the limits of the same family the size of the pelagic species seems to be on an average greater than that of the related abyssal forms.
234. The Zonarial Fauna.-Between the pelagic fauna living at the surface of the open sea and the abyssal, which floats immediately over the bottom, there appears to be usually a middle fauna, which inhabits the different bathymetrical zones of the intermediate water, and hence may be shortly called the "zonarial" fauna. The different species of Radiolaria which inhabit these different strata in the same vertical column of water present differences corresponding to those of the plants composing the several zones of vegetation, which succeed each other at different heights on a mountain ; they correspond to the different conditions of existence which are presented by the different strata of water, and to which they have become adapted in the struggle for existence. The existence of such bathymetrical zones has been shown by those important, if not numerous, observations of the Challenger, in which the tow-net was used at different depths at one and the same Station. In several cases the character of the Radiolarian fauna at different depths presented characteristic differences.
For the present, and until we are better acquainted with the characters of the Radiolarian fauna at different depths, we may distinguish provisionally the following five bathymetrical zones :-(1) The pelagic zone, extending from the surface to a depth of about 25 fathoms ; (2) the pellucid zone, extending from 25 to 150 fathoms, or as far as the influence of the sunlight makes itself felt ; (3) the obscure zone, extending from 150 to 2000 fathoms, or from the depth at which sunlight disappears to that at which the influence of the water containing carbonic acid begins and the calcareous organisms vanish ; (4) the siliceous zone, extending from 2000 or 2500 to about 3000 fathoms, in which only siliceous not calcareous Rhizopoda are found, and in which the peculiar conditions of the lowest regions have not yet appeared ; (5) the abyssal zone, in which the accumulation of the oceanic deposits, and the influence of the bottom currents, create new conditions of existence. So far as our isolated and incomplete observations of the zonarial Radiolarian fauna extend, it appears that the subclass Porulosa (SPUMELLARIA and ACANTHARIA) predominates in the two upper zones, and as the depth increases is gradually replaced by the subclass Osculosa (NASSELLARIA and PHAEODARIA), so that the latter predominates in the two lowest zones. The obscure zone which lies in the middle is probably the poorest in species. In general, the morphological characters of the zonarial fauna appear to change gradually upwards into the delicate form of the pelagic and downwards into the robust constitution of the abyssal ; so also the average size of the individuals (within the limits of the same family) appears to increase upwards and decrease downwards.
235. The Abyssal Fauna.-The great majority of Radiolaria which have hitherto been observed, and which are described in the systematic portion of this Report, have been obtained from the bottom of the deep-sea, and more than half of all the species have been derived from the pure Radiolarian ooze, which forms the bed of the Central Pacific at depths of from 2000 to 4000 fathoms (§ 237). Many of these abyssal forms were brought up with the malacoma uninjured, and they show, both when mounted immediately in balsam, and when preserved in alcohol, all the soft parts almost as clearly as fresh preparations of pelagic Radiolaria. These species are to be regarded as truly abyssal, i.e., as forms which live floating only a little distance above the bottom of the deep-sea, having become adapted to the peculiar conditions of life which obtain in the lowest regions of the ocean. Probably the majority of the PHAEODARIA belong to these abyssal Radiolaria, as well as a large number of NASSELLARIA, but on the other hand, only a small number of ACANTHARIA and SPUMELLARIA are found there. A character common to these abyssal forms, and rarely found in those from the surface or from slight depths, is found in their small size and their heavy massive skeletons, in which they strikingly resemble the fossil Radiolaria of Barbados and the Nicobar Islands. The lattice-work of the shell is coarser, its trabeculae thicker and its pores smaller than in pelagic species of the same group ; also the apophyses (spines, spathillae, coronets, &c.), are much less developed than in the latter. From these true abyssal Radiolaria must be carefully distinguished those species whose empty skeletons, devoid of all soft parts, occur also in the Radiolarian ooze of the deep-sea, but are clearly only the sunken remains of dead forms, which have lived at the surface or in some of the upper zones.
236. Deposits containing Radiolaria.-The richest collection of Radiolaria is found in the deposits of ooze which form the bed of the ocean. Although the pelagic material skimmed from the surface of the sea, and the zonarial material taken by sinking the tow-net to various depths, are always more or less rich in Radiolaria, still the number of species thus obtained is, on the whole, much less than has hitherto been got merely from deep-sea deposifs. Of course the skeletons found in the mud of the ocean-bed, may belong either to the abyssal species which live there (§ 235), or to the zonarial (§ 234), or to the pelagic species (§ 233), for the siliceous skeletons of these latter sink to the bottom after their death. Almost all these remains found in the deposits belong to the siliceous "Polycystina" (SPUMELLARIA and NASSELLARIA) ; PHAEODARIA occur but sparingly, and ACANTHARIA are entirely wanting, for their acanthin skeleton readily dissolves. The abundance of Radiolaria varies greatly according to the composition and origin of the deposits. In general marine deposits may be divided into two main divisions, terrigenous and abyssal, or, more shortly, muds and oozes. The terrigenous deposits (or muds) include all those sediments which are made up for the most part of materials worn away from the coasts of continents and islands, or brought down into the sea by rivers. Their greatest extent from the coast is about 200 nautical miles. They contain varying quantities of Radiolaria, but much fewer than those of the next group. The abyssal deposits (or oozes) usually commence at a distance of from 100 to 200 nautical miles from the coast. In general they are characterised by great uniformity, corresponding to the constancy of the conditions under which they are laid down ; they may be divided into three categories, the true Radiolarian ooze (§ 237), Globigerina ooze (§ 238), and red clay (§ 239). Of these three most important deep-sea formations the first is by far the richest in Radiolaria, although the other two contain often very many siliceous shells.
The marvellous discoveries of the Challenger have thrown upon the nature
of marine deposits an entirely new light, which justifies most important
conclusions regarding the geographical distribution and geological significance
of the Radiolaria. Since Dr. John Murray and the Abbè Renard will
treat fully of these interesting relations in a forthcoming volume of the
Challenger series (Report on the Deep-Sea Deposits), it will be sufficient
here to refer to their preliminary publication already published (Narrative
of the Cruise of H.M.S. Challengers 1885, vol. ii. part ii. pp. 915-926)
also the earlier communications by John Murray (1876, L. N. 27, pp. 518-537),
and by Sir Wyville Thomson (The Atlantic, L. N. 31, vol. i. pp. 206-246).
In the Narrative (loc. cit., p. 916) the following table of marine
deposits is given:—
237. Radiolarian Ooze.-By Radiolarian ooze, in the strict sense of the term, are understood those oceanic deposits, the greater part of which (often more than three- quarters) is composed of the siliceous skeletons of this class. Such pure Radiolarian ooze has only been found in limited areas of the Pacific and Indian Oceans. It is most conspicuous in the Central Pacific, between lat. 12° N. and 8° S., long. 148° W. to 152° W., the depth being everywhere between 2000 and 3000 fathoms (Stations 266 to 268 and 272 to 274). In the deepest of the Challenger soundings (Station 225, 4475 fathoms) the bottom is composed of pure Radiolarian ooze, as well as at the next Station in the Western Tropical Pacific (Station 226, 2300 fathoms), the latitude varying from 12° N. to 15° N., and the longitude from 142° E. to 144° E. In the Indian Ocean also, pure Radiolarian ooze was found in the year 1859 between Zanzibar and the Seychelles, this being the first known example of it (§ 230). On the other hand, it has not yet been found in the bed of the Atlantic ; but the Tertiary formations of Barbados (Antilles, § 231) like those of the Nicobar Islands (Further India), are to be regarded as pure Radiolarian ooze in the fossil condition. Mixed Radiolarian ooze is the name given to those deposits in which the Radiolaria exceed any of the other organic constituents, although they do not make up half the total mass. To this category belong a large number of the Challenger soundings which are entered in the Station list either as red clay or Globigerina ooze. Such mixed Radiolarian ooze has been discovered (A) in the North Pacific in an elongated area of red clay extending from Station 241 to Station 245 (perhaps even from Station 238 to Station 253), that is, at least, from long. 157° E. to 175° E., between bat. 35° N. and 37° N. ; (B) in the tropical Central Pacific in the. Globigerina ooze of Stations 270 and 271. The ooze from the latter station, situated almost on the equator (bat. 0° 33' S., long. 151° 34' W.), is specially remarkable, for it has yielded more new species of SPUMELLARIA and NASSELLARIA than any other Station, not excluding even the neighbouring Stations 268, 269, and 272. Probably such mixed Radiolarian ooze is very widely distributed in the depths of the ocean, as, for example, in the South Pacific (Stations 288, 289, 300, and 302), and in the Southern Ocean (Stations 156 to 159) ; also in the South Atlantic (Stations 324, 325, 331, 332) and in the tropical Atlantic (Stations 348 to 352). When carefully purified and decalcified by acids, Radiolarian ooze appears as a fine shining white powder ; in the raw state it is yellowish or reddish, sometimes. reddish-brown or dark brown in colour, according to the quantity of oxides of iron, manganese, &c., which it contains. Calcareous skeletons (especially the tests of pelagic Foraminifera) do not occur at all or only in very minute quantities in pure Radiolarian ooze from more than 2000 fathoms, whilst specimens of mixed ooze often contain considerable quantities of them.
Pure Radiolarian ooze was first described by Dr. John Murray as regards its peculiar nature and composition under the name "Radiolarian ooze" (1876, L. N. 27, pp. 525, 526) ; compare also Sir Wyville Thomson (The Atlantic, L. N. 31, vol. i. pp. 231-238), and John Murray (Narr. Chall. Exp., L. N. 53, vol. i. pt. ii. pp. 920-926, p1. N. fig. 2). The different specimens of pure Radiolarian ooze obtained by the Challenger from the Pacific, and handed to me for investigation, are from depths of from 2250 fathoms to 4475 fathoms, and may be divided according to their composition into three different groups :-I. The Radiolarian ooze of the Western Tropical Pacific, Stations 225 and. 226, from depths of 4475 and 2300 fathoms (lat. 11° N. to 15° N., and long. 142° E. to 144° E.). II. The Radiolarian ooze of the northern half of the Central Pacific, Stations 265 to 269, from depths of 255° to 290° fathoms. III. The Radiolarian ooze of the southern half of the Central Pacific, Stations 270 to 274, from depths of 2350 to 2925 fathoms. A fourth group would be constituted by the Radiolarian ooze from the Philippines, which was brought up by Brooke in 1860 near the Marianne Islands from 3300 fathoms, and described by Ehrenberg (Monatsber. d. k. preuss. Akad. d. Wiss. Berlin, 1860, p. 765). The Diatom ooze, too, found by the Challenger in the Antarctic regions (Stations 152 to 157) is in some parts so rich in Radiolaria that it passes over into true Radiolarian ooze. Regarding the Radiolarian ooze from Zanzibar, obtained by Captain Pullen in 1859 from 2200 fathoms (§ 230), we have only the incomplete communications of Ehrenberg (L. N. 24, p. 147). A more accurate knowledge of these deposits from the Indian Ocean, and of those which we may with probability expect from the tropical eastern Atlantic, will be sure to increase very widely our knowledge of the class.
238. Globigerina Ooze.-Next to the Radiolarian ooze proper the Gbobigerina ooze is the deposit which is richest in the remains of Radiolaria. Often these are so abundant that it is doubtful to which category the specimen should be referred (e.g., Stations 270 and 271, see § 237). In fact, the two pass without any sharp boundary into each other, and both present transitions to the Diatom ooze. Next to red clay § 239), Globigerina ooze is the most widely distributed of all sediments, and forms a large part of the bed of the ocean at depths of 250 to 2900 fathoms (especially between 1000 and 2000 fathoms). It covers extensive areas at depths below 1800 fathoms, and in still deeper water is replaced by red clay. It is a fine-grained white, grey, or yellowish powder, which sometimes becomes coloured rose, red, or brown owing to the admixture of oxides of iron, and manganese. True Globigerina ooze consists for the most part of the accumulated calcareous shells of pelagic Foraminifera, principally Globigerina and Orbulina, but also Hastigerina, Pulvinulina, &c. It contains usually from 50 to 80 per cent. of calcium carbonate, the extreme values being 40 and 95 per cent. After this has been removed by acids, there remains a residue, which consists partly of the siliceous shells of Radiolaria and Diatoms, and partly of mineral particles identical with the volcanic elements of the red clay.
Regarding the composition and significance of the Globigerina ooze,
see John Murray (L. N. 27, pp. 523-525, and L. N. 53, vol. i. p. 919).
Recently this author has separated from the Globigerina ooze (sensu
stricto), the Pteropod ooze, distinguished from the former by
the greater abundance of Pteropod shells and calcareous shells of larger
pelagic organism~ which it contains. It is found in moderate depths (at
most 1500 fathoms), and contains fewer Radiolaria.
239. Red Clay.-This is quantitatively the most important of all deep-sea deposits, covering by far the greatest extent of the three great ocean basins at depths greater than 2200 fathoms. It thus far surpasses in area the other deposits, both Radiolaria and Globigerina oozes, and commonly forms a still deeper layer beneath them. Probably these three deep-sea deposits together cover about three-eighths of the whole surface of the earth, that is, about as much as all the continents together, whilst only two-eighths are covered by the terrigenous deposits. Red clay is principally composed of silicate of alumina, mixed in various proportions with other finely granular substances ; its usual red color, which sometimes passes over into grey or brown, is more especially due to admixture of oxides of iron and manganese. Calcareous matter is usually entirely wanting, or present only in traces, whilst free silica is found in very variable, often considerable quantities. The chief mass of the red clay consists of volcanic ashes, pumice, fragments of lava, &c., whilst a large part of it is generally composed of shells of Radiolaria or fragments of them ; in many places the number of well-preserved skeletons contained in the red clay is very considerable, so that it passes over gradually into the Radiolarian ooze (e.g., in the North Pacific, Stations 238 to 253, see § 237). Hence it may be supposed that a large part of the red clay consists of decomposed Radiolarian ooze.
The characteristic composition and fundamental significance of the red clay in the formation of the deep-sea bed were first made known by the discoveries of the Challenger (compare John Murray, 1876, 1. N. 27, p. 527, and Narr. Chall. Exp., L. N. 53, vol. i. pt. ii. pp. 920-926, p1. N ; also Wyville Thomson, The Atlantic, L. N. 31, vol. i. pp. 226-7229). The mineral components of the red clay are for the most part of volcanic origin, due to the decomposition of pumice, lava, &c. Among the organic remains found in it, the siliceous skeletons of Radiolaria are by far the most important, and their number is often considerable. A large portion of the red clay appears to me to consist of broken down Radiolarian shells, in which a peculiar metamorphism probably has taken place. Sir Wyville Thomson was of opinion that a considerable proportion of it consisted of the remains of Globigerina ooze, the calcareous constituents of which had beeff removed by the carbon dioxide in the deep-sea water (L. N. 31, loc. cit.). Among these remains, however, the siliceous skeletons of the Radiolaria play a significant and often the most important part. Further more, John Murray has called attention to the fact that in many deep-sea deposits yellow and red insoluble particles remain, which unmistakably present the form of Radiolarian shells (L. N. 27, p. 513). At Station 303 he found "amorphous clayey matter, rounded yellow minerals, many Radiolaria-shaped " at Station 302 there was sediment " consisting almost entirely of small rounded red mineral particles ; many of these had the form of both Foiaminifera and Radiolaria ; and it seemed as if some substance had been deposited in and on these organisms." Similar transitions from well-preserved Radiolarian shells into amorphous mineral particles I have found in several other specimens of Challenger soundings, and consider them a further argument for the supposition that the Radiolaria often take an important share in the formation of the red clay.
240. List of Stations at which Radiolaria were observed on the Challenger Expedition.-The 168 Stations recorded below, in soundings or surface preparations from which I found Radiolaria, belong to the most various parts of the sea which the Challenger traversed during her voyage round the world ; they constitute about half of the (364) observing Stations contained in the official list published in the Narrative of the Cruise (Narr. Chall. Exp., vol. i. part ii. Appendix ii.).
In addition to the particulars given in the list regarding the geographical
position of the Station, depth, temperature, and composition of the bottom
deposit, I have added the result of my investiga tions as regards the relative
abundance of the Radiolaria in each. The five letters (A to E) denote the
following degrees of frequency :-A, abundant Radiolaria (Al, pure Radiolarian
ooze ; All, mixed Radiolarian ooze) ; B, very numerous Radiolaria (but
not a predominating quantity) ; C, many Radiolaria (medium quantity) ;
few Radiolaria ; E, very few Radiolaria (as they occur almost always).
In using these symbols regard has been had to abundance of the abyssal
as well as of the zonarial and pelagic forms (§ 232) ; sometimes also
the estimated number of Radiolaria has been inserted, based upon information
given by John Murray in his Preliminary Report (L. N. 27), and in the Narrative
of the Cruise (L. N. 53), as well as by Henry B. Brady in his Report on
the Foramiifera (Zool. Chall. Exp., part xxii., 1884). From Stations 348
to 352 in the Eastern Tropical Atlantic no specimens of the bottom were
obtained, but a rich pelagic Radiolarian fauna was demon strated by numerous
preparations from the surface. The depths are given in fathoms and the
temperature in degrees Fahrenheit. In the column describing the nature
of the bottom the following abbreviations are used :-