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: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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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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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 .
Representative terms from entire chapter:
brood chambers
