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- 26 - :ELOOLOGIC R1iLATICI:SHIPS 07 LAIR FOR~`,-I~IF7r:Ph lay Ear:L H. 1ly-ers Ho ~k~ rats .: ~ri.ne Station or Stanford University Pacif ic Gro~re, Cal if ornia i!~uT had speculated upon the Bite activities of the larger For~ifera but since the largest of tipsy are Groove extinct, it ~ s necessary that con- c~usio,~s be based upon the observed activities of sneer species nor-; CLi~r- i,~g in the shallot; wet=-s of troops ca' regions that are red ated to or re- stumble those found in older marine sed=.lellts. Ii: corlside'^ir,~ this problem, one ,~ght ask: Hoers numerous -are these larger Forar~ni~?~:~.., and ho;-. widely ,.-e they distributed dux~inc~, ~a geologic period? IT.~at -was the life spar? At -hut rate did their reproduce? Mat Areas the nature of the flood? has growth congruous or fiscoi~tii~uous and fishy? fore rzoo~::rho: Me essenti al to respiration and. the el:im~nation of nitrogenous waste? In this dis- cussion observed act~vi+.,ies and preparations of livid Fora"~ni~era v;~i-~] be described and methods explaii~d that might contribute -GO a better u~der- suandinc~ of these arid others p:-oblemse FOOD The Fora~=nifera utilize as food most `~u:6e bland and anoraks organisms and much dctril;al organic Trial toured both in the watered a:;:cl" on the bol;tom but are unable to capture ally or;~anism capable of vigorous mo~reme~t. The nature of the :Cood o~ the species is refed to the size and position of l;he pores and apertures ~ the test. In most J?oraminifviiag a ~-~;e aperture, costive orgar~isriLs of considorab] ~ size arc ill~,~ested. The nature of the, food mar be readily determined from cytoplasm~c Accuses observed i;: paraffin sections. T~no~orus, Cr~rpsina, arid crobab1:r m;~3T of -she ]ar=er _. ~ ~ O Fo::;~arn~ ife-ra that voters pro~rided~:ith grouts of pores but no larger aperatures, probably utilized bacteria mad other flute soft bodicc1 o-rga.~.~isr~.~s, or ashen larger `or~ranisrrs Acre uti:tized they revere digested ~iYithin a mass of ectoplasm that collected at one side of the test. These and other large card Hate- fo~amini~ ers When living,, in ax aquariurr1 make no attempt to capture minute arthropods nor have we ~ outed the remains of these or other large or~,a:~isr~s e;rtar~gled in the pseudopodic of preserved material although feeding e~Y- peri~Lents prove that l;l:aey readily accept Id digest 7oCy.r~eans of ferments in the pseudopodic the soft parts of dead attire orga~isrns. The pseudopodia of Operculina, _teroste~ina, Al~reoline]]a, arid other relatively large species axes as finely attenuated as' arid no oilier than th.ose of a large Qu;nquelo- cu~i~a., and it is so:-~e:~hat su-rprisi;~: to see the tests or these species mov- it up the Vertical walls of an aquarium b:r means of such delicate strands of protoplasm. In heteroste~ina and Coercu:Li:~a,, no food o-r,~anisr.s ~;e-~e ~_ , . observed in the c~rtoplas~ as was also the case in T:nocorus and G~r?:sina., while in Al~reoDine31a and ~.!~ar.=inopora n~r~utc. Soft bodied o~.=anisr:~s aid 'ire

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b.,~oker~ frustule s of diatoms are co~r::aon in the cut oplasm contained In tile distal chambers . A1 veolin~1 la and 1.1ar~inopora are ~ror~cious feeders ~ i -'JJe - ar~ =.o judge from the mass of organic d~tritu.~. the:. usual3~r covers the avpertura~ face of the tests even in mater: removed flora the s.ea. The amount of food required by densely populated beds of these larger Forar^~n~fera could. so far as he knoll, culy be prociuced within the Photic zone. FAT:S OF GRo7~fi;4T ACID - Ll 77 SPA`N Frorn cu:Lturd;L e~er~ents and statistical data of populations in the sell' it '>as found that a full dater was ~equir~3~ -to produce a single chandler o ~ E] phidiura' Fi~otalia, and even Miller species of Discorl::>is ~ ancl that ~ .A ~ ~.~^c~m five to seven daters of grazing ard metabolic activities passed before the ~1~^ chamber 1,ra.s added . A' ~ trough the ternger.atllre ~ n tropical - maters terms to accelerate the rate of r.~ctabo:Lic active ties, our: evic~ence so., ts that the rate of gro,`-rth in the tropics does not exceed that observed in the spring and ea:::~ly sun~.rner it tile h~,her latitudes and is more closely related to the amount of food available than to the te~i~pe.rature of the '.iate.r. Ire those species in which the tee-., consists of annular series of c'i~amber~ets the rate of growth Ray be accelerated as -,,:cS the case in Smiles and Amph~sorvs. lichen growth is interrupted due to a s~lorta:,e of .) food, as during the ~.~ter in the middle arid hillier Jatitucies, -~he fe~v CM=:n5CrS added are short and nary compared Oh those Ned -.~.ile food is abu,ndant. From variations in the size of l;he cn=,::r~s in an othe~`i~se o.~derly series -A is USUG117 possible ~ 0 :~eco`~'ni~Jo ;~vidua" s Chat have s.l-rvLved the -rater or hairs lived for More than one :rear. i;!) most tropical regions there are tic periods each Mar when good is Roy- a.'~undarlt as C-' result o' -kind incluced turbulence, and curds Rich Ox the irnpo~rerished surface :~!-~ith deeper nut:^ient-;^ich Diatom. Evidence or: Aces -. observed in Oocrculina c=d other soccic~s t~-~en near the ,.~es~ern end of the Java Sea Scar . Strait ooenda' but `-~as less Riders in matcr~al from. t'~e contra: portion of this sea era the food supply is more nearly co:~star~t cat al' ~ ir.,es. cargo indi~r~dua:Ls of Operculina often contain a hundred caners a"nd it is probable that these have lived for at :L~vast too ~ cars. The 1 if e sepals of T~noporus must be equally long since orgy one i3: many hu:1c~reds o~ Living ;~-crosphe;ric individual s is provided -I'd th brood chambers ~}~dicatin~, it they had arrived at The r~.a~xi!num ~ro-~'r.lh s-ta:= ix~ forger foramir~fers are indeed rare. RUSTS OF REPRODUCT1:OT`T Evidence of brood chambers The number of progeny asexua~l..: produced by a .~crospheric ~orarnin~fer varies .'itr~ the size of the individual. Small species including S731~i:L1i-I99 Pat,el1i~1a, and Cornus,oira seldom produce more than =~O]Q i to 12 while larger species of Discorbis, E:Lpl~idium, An~histe~ina, and Ca;Lca.^ina produce as . . many as 225. The number of p-.cogeny asexually produced is l==ted bar the number of masses of protoplasm containing ~ stipple :~uc2Leus that result then the protoplasm of the r~icrospheric 3ndi~ridual Zeros multiple ~is~-,ior In most species these monoilucJeate masses ap?3~0~ate in size the prolocu~u~n of Neal ospheric tests except, as ~ species of DiScOrcis, vi'n`cre there Is

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- 2S - a co`~i:~.ete bread of the septa between the chambers of the microspheric test during repx~o:'uct;ion and the juveniles are provided -with torso or even 'e chambers before leaving the parent test, or, as in the Peneroplidae, Revere tile juveniles are provided :;ith a curcu~nambie~t cha^^iber aid, as in Tinoporus, where the ea-~:Liest juveniles consist of three unusual :Ly :Large chambers While stil:L retained in the brood chambers in which they are produced. Should one wish to know the probable number of progeny asexually produced b:r one of the ~ arger species of For=~.inifera he -out ct dine the approximate volume o~ severa:t m~crosphe~ic tests and cli~rid.e this by the approximate volume of the ~roloculum and the --elated chamber or charnbors. This could be casi:Ly accomplished when ~e]1 preserved tests are available in Itch the changers have not become fitted with minc~ra:L salts by first infiltrating, the tests ~it': paraff:~n and then treating them with acid. After the weig~nt of the pa-raflin casts has been deteri~inec:, it ~.'ould be possible to remove ancl-~;ei=~ that portion of the cast resting the . ~ pr3:norc .1um. NUC~LOCYTOPLAS~IC R2L^TIO>IS-tILPS In many species r.=crospheric tests are farther than Sloe Exospheric. Ti.~,~en this is -the case ii~egalospheric individuals have out a she nucleus while the i~crospheric has 40 or rive depending upon the species. This however, is not awaits the case for in Operculina and IJeteroste~i.na tests of the +.wo generations are about equal in size and the much ear relationship is star to tha1; o'!ose~,^>red in species in which there is ~ difference in the size of m~crospheric and megaJospheric individuals. In T~noporus the too types of indi~ricl.uals are approx=aate~y equal in size uteri] ==ediate:Ly prior to reproduction when large brood ch~abe^-s are added to the m~crospneric tests. In this s~?ec~es5 the nucleus Or megalospheric individuals is re- ta~ecl by one of '~he three 1 arise chambers that constitute the juvenarium until immediately prior to ga~netogenesis. In liar~ii~opora and Alveo~inella the tests of both generations contain many hundreds o~ nuc:Lei, and in the fore genus ~negalosphe-~ic ~ndi~ricinals reproduce either sexually or asexual- ly. tinc~eoc~op:l~sm~c relationships similar to those described probably exists in extinct species in which ted morphology and. the earliest chambers resembled those of recent species. T,',T, A TORAH BE :ETA T The natural habitat of a ~ora~iniT^er is often su~;ested by Char moo pho:Logical char; acl;er~stics G: the tee,,. Since -within -6he intertidal zone only those species that are firr::Ly attached to the substratum or 1 ~, tre on reef leveed c al: sur~'i~re the turbulent surf g one of she poorest ??1 aces to 'ook for fora~ninifors is the leering potion of a coral reef. The majority of species living in Balloons are r;~l=tive~y smal' SO bill receive only pass=" covenant. Tinopo~us is found on Eileen roving immediately below the intertidal zone and is attached to the weed by masses or strands of protoplasm that have a diameter Maul co that of the spine -Mom which ~ t arises. Usually there are several points of ?~ttacument, and when weed covered with this species a;~o preserved in ~ o~~ma:tin the :foramini:l~ers are

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- 2S - not easily cTislod~ed t:y st~akii~ '~he contaii~. B_u~_-;=s,:~a protected i;..:itI: croci: longer spitzes are fou.r~d on weed :ro~.~i~ at, de'`~t'~s of from 20 to 60 med;ers, and is often 4;,a`~e;.: or a firm sandy bout or: at some distance from patches of need. May species of Calcarina are provided 'pith spies ,out .,Lese live on reed in the intertidal zone Mere thence is a mode~a-' su.-~f o Jot al1 species 3?ro~rided. -.;.ith spews -e cn weed since total i a ou~chel.~= is you only Cal the softest mud bottoms, but; in this case the , soirees are exceedir~l~r ion`; aced alel~der and probably serve to keep l;h<= test from sett:Lin~ into the mucT. A:Lveolinella anc3 keteroste~ina ar.e found on firm sandy mud bottoms ally Arc mos-L nur.~erous rlea-r pickle of reed >,r~'he}~r t He ju~reniles may be fallen in considerable nubs. `~!la.;~inooora is usually i'~.l~y- a=tachec~ bar animal cement and a nest-like mass of' organic debris to -,,.e`~6s.' dead. corals, shells, or s.i~iLar objects in the shall-water bo~.aerii~-, a heel' vie the smallest s:eci.es of' Sorites could ~phisorus are Powell on wand growing in the ir~^tidal zone. The large much :~atter~ed i;esLs of Oper~lina are -sound. on (,he sol ~ .~uc3 bottoms to which it is pa-rt~cularl~y~ wail adapted, wit; Hetei~oste;-~ina :.;~7ith an ecua.J7~.~r Barge test; but; pro. u;,.:hiJica:L 41~SS _ , _ 1 A 1 ~ is better ad.ai~ted to the ~i.rm saris or sandy mud bot+~o~-':hc-~ core its natural substratum. These observations suggest that most Forc~Linifora prided wraith Tar,, p~anospiral, 1u.sifo:^ln, or discoidal tests are laptop to Tilers sandy off sar~dy-mud bottoms within the sub:Liticral zone at deaths t2~c,.t do not e~ceecl toc~.t fit :!:hich ~hotos:=thotic organisms rbri~re and arel.:os~ ':um.erous in areas adjacent to reefs where coral, calcareous algae, Cal encrustin~.,^ bryo~oans or all three are ab.undar~. Strom, currents are req,.~ired Also ~ra~.~spo~=t these 1?.rge tests a-~i.~d under Teresa cond",tio~s one often :Cind.s t~om-~.~~ix~d fifth coarse detrital Beef material i.ncludinr.; broken cor'?~3 or even gr~-vi`~l frog.: previous horizons. 1,;\r~er;~ C~3.r, e~t action has lured -the ._~iS7 at seems more :liXely that the ~ irler sediments have been removecT ::a4~.e:^ 1;~= the' the test s have been t ~anspo Voted . DEI]SI'1~! 0'r~ POPUL~A~TI3I~S So - ~ due to curry action may at ties result in co:;~,iderab:Le depo.s~ts of Fo~ amirli~era cons:Lst~g of below the out elf reef opposite ~ one or trio: ~ species. T~rned~te~ ~ danger on the Tslar:d cf Bali in the Nether- Jar,ds East Ir~es Mar,- pure deposits of rI~noporus were observed, which :`hen a pole leas thrust molto the teds, were found -to be at; Jeas' 30 inches in depth. The sand or some portions of the beach in this -region CO:~S4cso:G almost a pv.re deposit of this forrr~ and is used to construct Creaks and. roads. ~- tensive deposits of Ope-~cu~ina were also observed o:,.a the Stunatra niche of Strait Soenda in which the ma, or portion of the sed.=n~-ts was co~r~oosec Or this one form. The population per unit area of' bluest species may at tins number hundreds or were: i,housa,~ds per square Scot anci it is so~.le~iT~hat amazing that so many organisms can survive in eaters in which: the amount of i-cod is prcsur~ed to be exce~ding:L:r ~,..a:Ll, although her- recap the amomrlt of Miriam life represerl-ted by a li~rirg coral redid arid tl.~cl.t flue p3:~incipa1 source of food to the corals are all or~-.:,isrns, While fine Foram~ni.~era

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are omniverous it is s~ne:.~hat less romarliable. These ~-~d other deposits suggest conditions similar to those that must have e~istec~ there certain: f oral~inif Oral limestones Are b ring :Z orbed. . ZOOXAI.~TI=LA:E It has been presumed by sor^.e that the production of OXygol~ through photosynthesis and the uti:tizatior~ of carbon dioxide and r~itrogenous waste by zooxanthe11ae may have contributed to the success of certain :1 arger Foray nifera. Ho;ve~rer5 it wou1 d seem Chat there is a close re- latiorship between the presence of zooxa~thel~ae acne test.~rphoJogy . . .. . . for these symbiotic algae seem to occur only in species having relatively large ape - Cures. Sorites, I.iar~opora, and Penerop1is often contain so many of these pJ ant cells that they seem to occupy at J east as much Of the beset as the protoplasm of the fora~r`inife~:. Zoo~'a~thel:Lae do not occur in Tinoporus, Heteroste,~na, and Operculina Probable, because- of the smaT:l diameter of the pores and apertures when present. In QuinqueJocu1ir~a and V=rtebra1ina the apertures are Barges however, no zooxa'~the1lae are . found, for eared -~hen ingested by these omnivorous [ceders whey are expelled from the test along with diatom fr~stules, the cel1u10se :-;a11s of bits of mu:Ltice:~1 ular a1 Mae, and the occasional a~openc4`ages of an arthropod. In those species which harbor zooxanth.ellae megalo spine tic juveniles . . accinire these plaint cells as c:rtopJasmic inclusions au the tire the juveniles are produced, hide in the microspheric generation they are i:~gested together with foods since the zoospores or gaxne~;es do rot contain them. Gi-4,OLOCIC SIr~t'lIFIC~4 CE although Tiiloporus is exceeds numerous in certain, ads in the Netherlands East Inches facing the Indian Ocean, l;heir disoribut~o~`n is probably limited to a few square Wiles. A7veolillel~a, ~i.~-Gerost<~zi:~a, and BacuJo~-ypsina are mom widely -distributed- but they also inhabit ~ reJat~rely small portion of the data Sea which has an average c~;~-th of only NO meters. It is p-~esur~ed that reduce of the region is one of slow su:side}:~c~. Under these conditions a progressive shifting o~:^ -reef areas and the silt:~ng over of flat was formerly ~ filly sand or sa::dy-mud bottom Elliot greatl~y contend the apparent range of these aid other foraminifors hang similar rec~uire- ments. The larger Foraminifera are used as guides to the age of g<;;ologi.c :t'o~mat~ons and the biotic conditions that prevailed aI, the three they were deposited, but in mailing Ecologic correlations theist usefu',::ess is Kited by the fact that the conditions required for their success probably was at no three extensive, cor.~pared with that of SP=CiCS topics OF softer mud bottoms .