BRITISH MUSEUM (NATURAL HISTORY). BRITISH ANTARCTIC (“TERRA NOVA”) EXPEDITION, 1910. NATURAL HISTORY REPORT. ZOOLOGY. VOL. VI, No. 2. Pp. 25-268. PROTOZOA. PART II.—FORAMINIFERA. BY EDWARD HERON-ALLEN, F.R.S., AND ARTHUR EARLAND, F.R.M.S. WITH ONE FIGURE IN THE TEXT AND EIGHT PLATES. LONDON : PRINTED BY ORDER OF THE TRUSTEES OF THE BRITISH MUSEUM. Sold by Longmans, Green & Go., 39, Paternoster Row, E.O. 4 ; B. Quaritch, Ltd., 11, Gratum Street, New Boud Street, W. 1 ; Dulau & Co., Ltd., 34--36, Margaret Street, Cavendish Square, W. 1 ; The Oxford University Press, Amen Corner, E.C. 4 ; AND AT THE British Museum (Natural History), Cromwell Hoad, SAN', 7, 1922. [Ali rights reserved.] 1 .1 » 11 v; [Issued 2bth March, 1922.j 25 PROTOZOA. PART II.-FORAMINIFERA. BY EDWARD ARTHUR EARLAND, F.R.M.S. WITH ONE FIGURE IN THE TEXT AND EIGHT PLATES. CONTENTS. PACK I. Introduction. (i) Material ....... 26 (ii) Literature ...... 28 (iii) Arctic and Antarctic Types of Foraminifera 30 (iv) Pelagic Foraminifera .... 34 (v) Classification adopted in tias Report . . 38 (vi) New Species and Varieties Recorded . 43 II. List of Stations. (i) Official—Harmer and Lillie .... 44 (ii) Geographical—Heron-Allen and Earland . . 44 III. Description of the Material Examined . • • 45 IV. Species and Varieties Recorded .... 58 V. Appendix. A. —Official Station 308 “Off Tierra del Fuego ” . 230 B. —“ Royal Sound, Kerguelen Island ” . . . 231 C. —Official Station 42, “ Off Rio de Janeiro ” . . 232 D. —Material collected by the “ Discovery,” 1901-4 . 233 E. — Wagnerella borealis, Meresch. .... 235 F. —On the Colouration of Polytrema miniaceum (Pallas) 247 VI. List of Works Referred to in the Report . . 236 Index ......... 249 E VOL. VI TERUA NOVA' EXPEDITION. % t I. INTRODUCTION. (i) BATFILIAL. Skvex rea is have passed since we received from the present Director of the I I Natural History Museum iittv-lmir tubes and forty-three iars of varvina; sizes * * » 0 * containing material for examination and report. The work has necessarily been retarded by the War. but Iais resulted in the recording of more thaii 050 species and yarieties of Foraminifera, forty-six of which are new to Science. 1 Before the Expedition started we were consulted as to the best methods of collecting, and gladlv nave tile results of many years* experience, but circumstances appear to have rendered impossible any serious attempt at the collection of fora m in itérons material. Apart from the tidies containing the "cores" of soundings (which are of little value from a faunistic point of view), and a few sands from the New Zealand coastal arca. the material received consisted principale' of sandv débris, evidently the residuum from gatherings of assorted Benthos, and usuallv '* preserved iii formalin, thaii which no more unsatisfactory medium for the "preservation* of Foraminifera can be imagined. Now sueli débris is nf eoume foraminilerons. otten rich, but as compared with speciallv collected material it is extremely difficult to clean, and the results are often fragmentarv. Neither tina* nor trouble was spared iii the tedious processes involved, but from the point ol view ul specialists we can only view tile results as a tantalising sketch ol tile possibilities which would have attended upon an ample supplv of pn tperlv c< >1 looted Anta ret io material. The method oi preparation employed was to wash the material on graded sieves raii ainu dowii to silk gauze of 1(H) meshes to tile linear incii. The material thus retained i< rei erred to in the subsequent report as " Sieves.** while the finest mud. passing thnandi the silk gauze, and subsequently separated bv means of a filter-paper, is described as " Residues. It niav be noted here that we received no material whatever from five of * the " Antarctic Benthos Stations in the Official List, Dut we have received facilities for examining some of the Sponges collected at these Stations for sessile species. A1 that three pus of material were entirelv unlocated and some others bore <1 > r 1 1 no Stall'*11 mind>ors. wen* successful! v by a comparison of the t k fauna contained. Our researches have confronted us with several notable lacunae. Conspicuous bv their absence, for example, are Miliolina Incoenis, Keramosphaera murrayi. Hyper- Ujnm nri truitti is. ( \i laie t tia infula. Anomalina ammonoides, and Pulvinulina mcnanlii. ali ui which have been recorded from similar by Chapman or bv IVarcev. A st 1 iking leal ul e o| our records is the occurrence of arenaceous isomorphs I vaimm forms hitherto known only as porcellanuus or hyaline. These have ( ) FORAMINIFERA—HERON-ALLEN ANI) EAKE AND. afforded an opportunity of adumbrating in Section V. of this fuiroduH ion a lm »l't scientific classification of the Order, especially in regard to the family Fai uolidae. The occurrence of sueli arenaceous isomorphs might have iiirii reasonai>lv anticipated in the Antarctic. The substitution of an adventitious test for « *n<‘ composed of calcium carbonate is no doubt a physiological process due to tile low temperature of the animal's environment. It has long been observed that tropical organisms secrete calcium carbonate in great abundance, while benthic and Arctic organisms are characterised by thin, delicate shells, and the experiments of Murray and Irvine proved that the reactions involved in the fixation of CaC03 were retarded bv cold and accelerated bv heat. see p. III.) The fossil or sub-fossil material received from Stations 2. :>, and 0 presents some difficulty. The distinctive nature of the material does not appear to have been recognised when it was obtained from the dredge, and no records were made of its occurrence and relative abundance. The material from tine lse Station: therefore consists of : (1) recent muds and sands recognisable as such (2) more or less disintegrated lumps of the deposit in question ; (3) a mixture of the two in which it is not alwavs possible to discriminate between the constituents. Some of tile harder lumps of this deposit have been submitted to various authorities who, as a nile. decline to commit themselves to anv definite statement as to its age. The general consensus of opinion is, that the deposit is probable of no great antiquity but a recent sea-bottom conso bv tin solution and subsequent re-deposition of OaCO., in sea water. Mr. A. W. Waters, on the other hand, after preparation and examination of the material, is inclined to attribute a Tertiary origin to the deposit, on the strength of the occurrence therein of certain Polyzoa {Cellaria amiastilnba. Verticillata, 'a, fossa) known only as Tertia rv fossils. He compares the Polvzoan fauna with certain lieds in Victoria attributed to tile Miocene or Oligocène. The matter lies outside our province and can only be settled by a further examination of suitable material. We will only observe that it the deposit is of the age suggested, we should have anticipated tile occurrence of similarly typical Foraminifera. With the exception of a single specimen of Xuimmriites found at an early stage of our work, anterior to the receipt of Mr. Waters's report, no distinctive fossils occur among the Foraminifera.* The species listed from the hard deposit are to ali intents and purposes identical with the recent forms at the same Stations, and anv differences are principally due to development, which is usually more noticeable in the fossii than in the recent * tW luci rejected tias Nlllumulltv without hesitation as dae to extraneous ruvuiusfanees, although \\v felt that ali possible precautions had heen adopted in rhadina the material, bat in view of Mr. Waters's hvimislv a considerable valae as evidence 111 support of (lis theone report its oiTiirroin1, if accepted, has o “TEIIIIA NOVA" EXPEDITION. s One point only appears Io us to he definitely proved by the com specimens paratively low percentage of mineral grains, viz. : that the deposit was laid dowii ime. and under conditions, when the shore-line was at a far greater distance at a t from tile Stations in ouestion than it is to-day. (ii) LITERATURE. The multitudinous and world-wide material collected l>v T Orbigny prior to ” Tableau Méthodique," included none from Antarctic the (late of remous. Freycinet and Duperrey. from whom lie received much material, did C."' , 1 * net cruise 8. o! Tasmania. The literature of our sub mav be said to commence with a paper bv Ehrenberg to the Berlin Academy in 1844. dealing with material collected bv Dr. J. Hooker uii the cruise of tile "Erebus" and ‘‘Terror,” under A volume of 100 pp. was issued containing instructions to Sir James .o * observers and collectors upon this voyage, the data and material collected to lie worked out at home after the ! etium of tile Expedition.! According to Ehrenberg it was lie who caused Humboldt to press for tile collection of micro organisms and to prescribe the methods of collection. We are told that in January 1841, lying becalmed, Boss put over a dredge which brought up rock fragments and ” a surprising profusion of animal life." Dr. J. Hooker collected forty packets of soundings from (‘ape Horn to Victoria Land, and three jars of water which were sent to Uermany. and we read that forms collected in 184*2 near Victoria Land were still almost fresii when thev readied (Terinanv in 1844- < «/ Hooker and Ehrenberg both worked on tile material, and the conclusions arrived * J1 at by Ehrenberg were, shortly, that the relations of organic life were identical 4 at the North and South Roles, that tile surface lavers (" Pancake ice "i were crowded with life, and that the? supposition that organisms cannot live below 100 fins, had become untenable.! At the time Ehrenberg wrote, only three packets, between 63° and 70° 8. (190-270 fms.) had been examined; what became of the rest is not known to us. On p. 188 he records, as found by Hooker, in floating (" Pancake ”) ice, from 78° IO' 8., 162c \V„ four E<mamillifera ; of three of these lie gives a nebulous diagnosis on pp. 207-8. the other is simply recorded as " alina:' The t H ' three diagnosed forms are; (1) Grani nwsto)mnn divergens, \v he subsequently Ehrenberg. " \ orlautige Xaehrieht ülht das kleinste Lobeli im Weltnieer, am Siidpol, limi iii i Io i' Mocivs Tiefon. Mmiatsbor. Borlinor Ale. Wbs.. is 14, pp. 182-207. t Tlio reader b T*'fi'i]t’il to Dr. 11. JL Mills work *' The Siege of tlie South Poli* " (London, 1905, pp. -52 M2li.) Tho " Erebus ’ and " Terror " started from England Sept. 30, 1839, and were within the Antarrtie < Ui<*L■ (8. ul (iii 3ii S. hit.) between dan. 1 and March 1, 1841, Jan. 1 and March 6, 1842, and .Marrii 1 and Man ii 11. isis. Flaw returned to Emdand in September, 1843. Î An abbreviated translation of Ehrenberg s paper was published iii Ann. Mag. Xat. Hist, xiv, 1844, pp. hi;1 -PI. " Ihi Mierosropic Lift* in the Ocean at tho South Polo and at Considerable Depths." F< >UAM I XIFKRA -HEPO >N -ALLEX AX I ) F AH LA X I >. figured (E. 1854. M. xxxv. A. Fig. xxii, 22)* and recorded as " (Jattai iiat f dicen/cns and 1844." Sherboni suggests Bulimina, and it is probable referable to that genus, of the B. pupoides type. Ehrenberg aiso records the same species (on p. 192) from Gulf Erebus and Terror, 63c 40' 8., 55° W. (270 fms.).t (2) /? m antarctica, and (3) /?. e>Wb. He gives no figures. but from the diagnosis they appear to be starved forms of B. beccarii (Linn.). And then came what J)r. Miii calls "the generation of averted interest/' Xo other exploration of importance, and certainly no collection of material took place in Antarctic waters until the vovage of the "Challenger" in 1874, whose "furthest south" was 66'40'8., 7S°30'E.. 8. of Kerguelen Island. But the "Challenger" made the first systematic series of O %■ soundings in the Antarctic, and the statement was made that here the Globigerina ooze (or Bed Clay in deeper water) merges into Diatom ooze, and then into terrigenous Blue Mud. (Cf. Dr. Harvey Pirie/s Map in B. 1913, 8.N.A.) The j)rineipal works dealing with the Foraminifera of our areas are, for the Xew Zealand area. Chapman 1905, G.B. I ; 1909, S.X.Z. ; and Cushman 1919, R.F.N.Z. : the latter paper being founded merely upon four type-slides, sent to the author bv our late friend Mr. B. L. Meataver. of Wellington, X.Z.. containing * » ' ~ O specimens from the "Poor Knight's" Islands (35° 30' 8., 174° 49/ E.) ; it therefore lacks the authority of Chapman's papers, which were the outcome of research upon ample material. For the Antarctic area, the works available to the student are Pearceyi papei* on the Foraminifera of the Scottish Antarctic Expedition. 1902-4 (P. 1914, 8.X.A.), in which he records 242 species and varieties, and Dr. Harvey Piriei paper on the Deep 8ea Deposits (P. 1913, 8.X.A.). which, however, records no species other than those listed in Pearcev's memoir. Our friend Mons. E. Fauré-Fremiet has contributed two papers on the French Antarctic Expedition of the " Pourquoi pas ! " (F. 1913 and 1914, F.M.A.F.). M. Fauré-Fremiet is an excellent worker, but unfortunately kis material must have been very inadequate, for he only records fourteen species. In 1914 Chapman, iii the Reports of the Shackleton Expedition (1907-9), contributed two papers, one Geological (C. 1914, E.D.R.8.) on (ui a deposit 20 feet above sea-level X. of the Drygalski Glacier, and (b) another from the slopes of Mt, Erebus, 160 feet above sea-level ; the other, Recent (C. 1914. F.O.R.8.), from soundings in the Ross Sea, in which he records sixty-six species and varieties. It is obvious that the material submitted to these authors was very inadequate, but we made, originally, a point of giving references to their the papers m synonymies of ali species and varieties recorded by us from the far more extensive material submitted to us for examination. The drastic reduction of synonymies v * For explanation of the method emploved by us. with a view t economy of space, in ali our bihlio- graphical references, see p. :i30. *f Ehrenbergi ligure of (r. <1itnyntss PI. xxi. fig. Sfi, is Bohr tun ilihifafa. 1 Iss., a quite different form- TERRA NOVA” EXPEDITION. U . ) explained on p. 5$, necessitated by the present conditions of cost of printing and us to excise this portion of our work, but the papers has paper themselves are no doubt readdv accessible to the student. It remains onlv to sav that, though as we have indicated (supra), the featherings consumed to us might have been much more exhaustive and repre sensitive thaii tliev are. the material analysed in this Report is by far the most voluminous that has been brought to this country from either New Zealand or V the Antarctic regions. (iii) ARCTIC AND ANTARCTIC TYPES OF FORAMINIFERA. It is not easv to form anv comparison between the foraminifera! fauna listed in our Report and tile tallies of species from high latitudes published by Brady (B. 1 $84, E.C., pp. 779-78-3), because, of the eight Antarctic stations figuring in © © Ids list, five are from localities on or adjacent to the Southern extremity of the a American continent, and two of the other Stations rich insular gatherings round Kerguelen Id. and Heard Id. The from remaining is "Challenger" Station 1.30 in the Southern Ocean (.5*2° 4' S.) in 1.30 fathoms. This is approximately tile latitude of our Stations 11 and 12 O Ic Nos. 208-9). and the depth is verv similar, but the nature of the Challenger material, which was organic débris from a rockv bottom, yielding onlv sixteen species of Foraminifera, forbids any direct comparison with the oozes obtained at these two " Terra Nova " Stations. Brady did not list the species derived from tin1 dt water " ( * Stations off the lee Barrier (Stn. 1.53—1,675 fins., iep Stn. 1.5.3—1.300 bus.. Stn. 1.57—1,9,50 fins.), ali of which would have afforded suitable material for a comparison with our Report. Murray in tile "Summary of Scientific Results of the Voyage of the 'Chui- * » » » j len • « % *95). vol. 1, p. 498, gives a list of forty species of Foraminifera o-er from Station 153. Idus is probably not exhaustive. It contains many interesting forms, but perhaps tile most noticeable point is the absence of nerina par]t/f<lcrnw (Fhbgd, and the presence of G. dutertrei. d'Orb. Nor is it easy to make any direct comparison with the foraminifera! fauna from Arctic latitudes given in the same table because our list includes many deep water Stations, whereas 300 fathoms represents the maximum depth in Brady's list. But a comparison of the species in Brady's Arctic and Antarctic lists with mu- own records yields some noticeable results, in some confirming Brad\'s observations and iii others disproving them. Unis Brad}' records the following species as Antarctic but absent from the Arctic : - I. Art andina tumidis. Bradv. ♦ % *2. .. .. var. inornata. Bradv. FOR AMINIFKRA -FIFROX-ALLFN A X1 > K A I : FAX I ». SI 3. Clavulina communis, d'Orbigny. 4. Sagrina raphanus, Parker and Jones. ,, dimorpha, Parker and Jones, o. Nes. 1 and 3 are abundant in our Antarctic material, but the others do not occur. Again Brady comments on the fact that tile following forms th occur m Arctic, while no representatives of these genera occur in the Antarctic : 1. Verneuilina polystropha (Heuss). <>. Polymorphina lactea (W. A J.). Bigenerina nodosaria, d'Orb. .. oldoiufa, d Ori). i. 3. Spiroplecta biformis (P. & J.). 8. rotundata (Born.). 4. Polymorphina acuminata (d'Orb.) 9. Operculina ammonoides (Oreni.). o. compressa. d'Orb. But No. 1 appears in our Antarctic list with four other species of Verneuilina ; No. 3 with another species. Spiroplecta annectens (Ik A J.) ; No. 9 occurs, and although only Nos. o and 8 are represented in our list of Polvmorphinae, our Antarctic species of that genus number eight, as against Brady's live from the Arctic and none from the Antarctic. Possibly the most striking fact in the comparison of the Polymorphiuae lies iii the entire absence of P. lactea (W. A J.) from both Antarctic lists, though it figures at live out of six of the Arctic It looks therefore as though P. lactea may be regarded definitely as ►J Ul I» absent from the Antarctic. Operculina ammonoides ((iron.) occurs at several Antarctic Stations. Among Bradv's other deductions are :— I’ 1. That Uvigerina and Sagrina replace Polymorphina in Southern latitudes With this we cannot- agree for reasons already given. O « O o That (C. communis, similarly replaces - Bigenerina, which appears to be correct. That the porcellanous forms display a more varied development in the o • South thaii in the North. In support of this he lists twenty-four porcellanous species, nineteen of which are Antarctic against ten in the Arctic list. There are twenty-six porcellanous forms iii our Antarctic list, including several species not common to our own list and Brady's, so the deduction appears to be correct if the Arctic list may be regarded as fairly complete. 4. That the Northern region is much richer in— O (</) The Textularia!! types ; (b) Ihdymorphinae ; (r) Nonioninae. * As regards (u) the deduction is based on a list ul t wenty-seven 1 cxtulariidae. nineteen of which are recorded by Bradv as Antarctic and only sixteen as ‘ TERUA NOVA ' EXPEDITION. • ». > * » — Arctic. The reasoning is therefore at fault if Brady had "species" in his mind. Jf however "number of specimens " was intended we may perhaps agree, as, with certain exceptions [dae ahu a (‘ommunis. Cassidulina (various chapmani. Ehrenbergina hystrix var. ytubra. etc.) the Textulariidae occur very sparinglv in our Antarctic material. Th genus Textularia itself is represented < in oni* list hv a single deep-water form. Textularia concava var. heterostoma, horii., as against three species iii Bradv's Antarctic list and two in his Arctic list ; and other genera, where represented at ali. are usually very rare. But we have some fortv-seven species and varieties of Textulariidae on our list, south of the Antarctic hi role, so Bradv's conclusion must not be accepted without further evidence. Vs regards (b) we have alreadv stated the reasons for our disagreement with Bradv's <•<inclusion. Vs regards (r) Brad\ ivht species of ali of which figure 1-t-1s1 iii the Arctic fauna, but only four of them appear in his Antarctic list, and these arc confined fo the Patagonian localities and do not figure in the Kerguelen, Heard Island or Southern Ocean nii . Our experience is very different, for we have eight species on our Antarctic list and one of them is Nonionina Iacra. d'Orb.. which is one of the most characteristic Northern forms It figures in ali tile Arctic Stations of Bradv, but at onlv one Southern Station, off the Patagonian coast. Helice we mav regard this conclusion of Bradv s to be unfounded aiso. Bradv's tables include tiftv-three genera and 189 species, of which forty-four * *. c. • i «/ represented by 137 species, occur iii the Southern area and forty-one genera. genera, represented by III species, occur in the Northern arca ; thirtv-two genera, bv sixty species, being common to both areas. h of forms common to Northern and Southern waiters i rac ui might be used as evidence in favour of the theory of bipolarity by a zoologist unfamiliar with the group, but to the lhizopodist the presence or absence of a few particular species would be more convincing evidence thaii the longest list of other forms. This, for the reason that, apart from those genera and species limited in their distribution bv conditions of mean temperature (which form the characteristic shallow water foraminifera! faunas of different latitudes), the mean temperature at ali depths below' 50U fathoms ali over the w'orld (Joes not vary more thaii a lew degrees, and so would constitute no barrier to the gradual migration of Foraminifera from one pele to the other by way of the cold waiters ol the dce]i sea. Hence wre should be loth to accept the existence of any species or number of species in both Arctic and Antarctic waters as evidence in favour oi bipolarity, but should be prepared to regard the absence from either region of a species kamen to be peculiarly characteristic of the other region as definite evidence opposed to the theorv. Brady s list of Arctic Foraminifera contains at least three species peculiar to FORAMINIFERA—HEROX-ALLKX AND KAIULANI >. • » • i .h' Arctic and adjacent seas. They are Reophax arctica, Brady, known only from the Arctic, and by a few pauperata specimens from Delos (Sidebottom). Ui ppnrrepina indivisa, Parker, known from tile Cult' of St. Lawrence, the Arctic seas, and in a 4 starved form from tile Moray Firth ; Polystomella arctica, P. & J., perhaps the and abundant of ali Arctic species, a large, strong species windi most cannot be overlooked and with a Southern limit about the Moray Firth. t None of these three forms have gyer been recorded from the Southern arca. and but for the occurrence of a single specimen (broken) of Hippocrepina indivisa in our own records we should regard their complete absence as more striking thaii the occurrence of a multitude of other species which have a more or less universal distribution. It is more difficult to select similar forms from the Southern area, but there are two characteristic Antarctic forms which mar bo considered, viz. : (a Bradv, abundant in the farthest South gatherings , V m t O O This has been recorded aiso from shallow water in the tropics, but never, so fai as we know, in Northern Seas. (2) Ehrenbergina hystrix, Brady, represented abundantly in the Antarctic by our variety glabra. and otherwise known only from deep water in the Pacific. Practically the only species (so called) which could be invoked as evidence of bipolarity is G enna (Elibg.). rTn ias curiously thick-walled and nana very characteristic form is eina of Arctic Su 4 John Murray* it is a purely benthic form, its very structure our opinion, to any idea of its existence in the pelagic state. Abundant in the Arctic oozes, it becomes rarer in the North Atlantic, and. save fia- perhaps an occasional doubtful specimen, sueli as the records from the Pacific (B. 1884. F.C., p. 777). reaches its southern limit in the Faroe (.'haimei. It does not form a constituent of the bottom oozes of temperate and wami seas, as might be expected if it were either (1) Pelagic, as stated by Murray, we believe in error. (2) Benthic, but generally Yet Globigerina pachyderma (Ehbg.), occurs in the Antarctic as the typical Globigerina at ali depths. As we have stated elsewhere in our notes on the two species, the explana tion of the occurrence of this Arctic form in Antarctic oozes, and its absence in intermediate deposits, lies in tile fact that it is not a true species, but a local irina dutertrei, d'Orb., induced bv conditions of temperature. variation of f / The same gradual transition of the one tvpe into tin tin hav ii ?v u we c described in the Antarctic could be traced in the Arctic and temperate seas, and although the records of G. dutertrei are singularly lew manse it is Ö ^ * * J. Murray, ‘‘ The Oe-an ** (Lonfinn. n.fi.) 1913, p. 165 anfi '• Pelagia Foraminifera 1897. Nui Science, vol. xi, pp. 17-27. F VOL. VI.
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