Vol.
32,No.
1·2STANiSl.AW SKOMPSKI
acbl ,,8010glca polonica
Warszawa
t
982The nature
and~ystematicposition · of the microfossils.
Glohochaete a/pina Lombard, 1945
ABSTRACT: Well preserved aoinif.orm assoc.iatioos of G,obochaete alpina Lom- bard discovered wi·thin Upper V,isean limestones of the IJublin. Coal Basin, eastea:n
p~ allow to =recognize thereJatd.OlII9bi.p of these m1crofossils to the present- -day green algae of the :family Chl.ora9pbaeraceae. The revdsion of the whole ge- nus Globochaete is unde.rttaken, and the species Nostocites vesiculosa Ma610v is
regarded as a subieclive synonymy of Globochaete alpina Lombard.
INTRODUCTION
The microfossil Globochaete. alpina Lombard, 19~5, commonly ·inter- preted as a calcified zoospore of green algae, has long been described from primarily Mesozoic and Tertiary carbonates developed in both pelagic and extreme shallow marine enviroiunentS (cf. Misik 1959, 1979;
Lefeld & Radwailski 1960; Zawidzka 1972; Borza 1975; Fliigel 1978).
A few reports concern .also Paleo:lOic occurrences in the Silurian of Al- geria, . Devonian . of Morocco and Dinantian of France (Durand Delga
1956), Devonian of Czechoslovakia (Biely & KuIlmanova 1979), Silu.rian through Carboniferous of the Soviet Union (Linetskaya & Muromtseva 1973, .Muromtseva. 1979), Carboniferous of Afghanistan (Vachard 1980), and Fermian of Mexico (Tellez-Giron .& Trejo 1974).
The specimens discussed in this paper -come from the Upper Visean limestones with Calcifolium pierced. by the borehole Podedw6rze 2 in the-north-eastern part of the Lub~in Coal Basin, eastern Poland; lo.ca-' tion of the borehole and -stratigraphic setting of the limestones are pre.,;
sented elsewhere (Skompski 1980, Skompski & Sobon-Podg6rska 1980).
Ac"kn.owledgements. The AUthor is greatly ·indebted to a-ofeesar M. S~u~
wski, ~l1titute of Geol9g:y, U~lversity of Warsaw, and to Docent. M. Batko, Insti- tu.te of Botany of· the same University, far dilscussion end crl.t.ical ;readdng·"of the
manusar.ipt. .
48 .s. SKOKPSKI
MORPHOLOGICAL· TYPES OF GLOBOCHAETE ALPINA LOMBARD
Lombard (1945) definied Globochaete as heMispherical or spherical lIs, preserved in diverse stages of divislon,single or joined in linear associations, commonly affixed epiphytically to concavo-convex frag- ments of algae, or of other undetermined organisms. The cells are fi- lled with radial calcite· which effects iil a black cross under the crossed nicols. The size of the cells ranges widely from 4()p. to 10~, or even to 20(41. as noted by the subsequent authors. A lengthy, but not compre- hensive discription offered by Lombard (1945) was supplemented by sketch-drawings of Globochaete in the stage of partition and germina- tion, in the mature epiphyticstage,.and. in the final senile stage. These sketches are eVidently simplified When ··compared to the earlier study (Lombard 1937) which presented Globo£haete as "Organism Dn. The partition of the unicells was w~l1 illustrated by Colom (1955). Both Lombardand Colom did not present any associated forms with a lar- ger nuinber of· unicells, and such very structures were recognized later, e.g. by MiSik (1959, 1979), Zawidzka (1972), Borza (1975) and recently, under the name Globochaete gregaria, by Schafer & Senowbari-Daryan (1980). A variety of the associated forms are the spherical specimens composed of well arrariged Globochaete, and illustrated by Tellez-Giron
& Trejo (1974), who doubted about the zoospore ~ature of Globochaete.
The investigated Visean specimens from the Lublin· Coal Basin repre- sent mostly associations of a greater number of Globochaete. and called here the aciniform associations which varY into the spherical, . envelo- ped forms (empty inside), circular in sections (PI. 2, Figs 1-2), and globular forms with their interior filled (PI. 1,. }i'ig. 2; probably also PI. .1, Fig. 1). Moreover, recorded are the linear associations (PI. ·3, Fig.
1), whereas the unicells and epiphytic cells. are absent.
The hitherto preSented descriptions and illustrations of Globochaete, as well as the· investigated material allows to distinguish the following types of morphologies (see Text-fig. 1) observable in the thin sections of Globochaete:
A --.;;unIcellar type: all Itbe. ~e· forms, and cells c:luriDg partitioll, usually into 2 or 4;
B - linear aBsoolatioo.: all the chain-Wte (":rosa.ry-~"; "cbapeklt" of Lorn-
baIrd, 1945) groups IWbich might 'ha.ve developed due to ;ooe-dim~l parii- tdan of the cells;
C - epiphYltic celis: all the :&iJngle or .grOuped oel:l:s attached to or~ r~
commonly .of the algal origin. .AB rtbese cells are usua1ly ~ed to the Clltl-
cave Bide. oftbe epiphyse (foe exceptQons see Pl 3, Fig. 2; and Bromrlmann 19155, Text-fig. 60); Zawidzka ·(1972) suggested their nature as the remnants of vegetati'Y'e cells 'WIith affbc.ed ~res;
D ~ aclndfarm. aSl!lOOia.tiQIl: the cella f§rouped into ihe grapes, the ~ ~Velo- pe at W'hicb maybe the only one preserved. .
NATURE OF GLOBOCHAETE
Additionally, as the fifth type, the endophytic cells are to be dist~
guished, viz. those presented by Linetskaya & Muromtseva (1973), who recognized Globochaete inside the thallus of the alga Beresella; an ob- scure photo does not allow to verify that record.
The above "section" -types correspond to the four morphological types of the. occurrence of Globochaete. It should however be noted that a two-dimensional image from thin sections.is not always recognizable to a certairity as to its three-dimensional shape.
SYNONYMY OF GWBOCHAETE ALPINA LOMBARD
Within the genus Globochaete. the five species have hitherto been recognized, viz. G. alpina Lombard. 1945; G. tatrica Radwanski, 1968;
. G. Bpinosa Dragastan, 1971; G. h'l"onica Borza, 1975; and G. gregaria Schiifer . & . Senowbari~Daryan, 1980. The last three of these species should eVidently be revised, as the criteria of their distinction· are eit- her unclear or simply wrong. Highly similar to some Globochaete are also the microfossils Nostocites vesic'Ulosa Maslov discussed beneath.
The species Globochaete tatTica Radwai)skzi, 1968, described for the first time from .the Rihaetian· of the Tetra Mts (R,adwanslm 1968), has subsequently been
\~~ . /
\ \ rBJ!
Jet \ /
\ /
\ It
\ /
\
Fdg· 1. ~ogim1 types of Globochaete alJrina LambeM
A - s.i.Dgle cella (after Colom 1955), B - lIirneaI' assooie4'Lon of cells (aftef' MdAik
1969, Linetskaya & Muromtseva i973), C - epiphytic cells (after Bronnimann 1955, Bnn.et 1956, Mui .. k 195&), D ~ acinifQl'm asaoaiation of. cells: solid 10rm (at left)
,and lfIJperficlal farm (at right)
"
50 S. SKOMPSKI
acCepted and recQI"ded bath :from the Rhaeman of
the
Tatra Mts and of· the Mps (Flilgel 1972, 1978; Gazd*koi 1974; Sc.hllfer & Senawbari-Darya.n 1980), as wellaB fJrom the sllratigraphdca1Jywide ranging deposits (CanWm through Neocomiao) of Slovakia (BGrza 1975, WAik: 1979).
The species Globochaete spinosa Dragastan, 1971, recognized in the Jurasstc- -Cretaceous deposits of Romania, does oot display the ~equested ~ic features, and was akea.dy denied by MiAik (1979).
A
_time
diagnostic of Gl.obochaete hronicd Bor.za, 1975, was a large, conical depressioo. filled with micrite :inside the cell. However, ocly in some sections this structure is reoogni.rz;able as a depression; in. most caBe6, the holotype ·incJu- diDg it d.s a darit spot .inside the cell Such a spot varydng in size (see PI. 3, Fig.6), is also chairacter,iBtic ~ G. atpina and may even a:ppeai- at the edge of the cell, as illustrated by Barza (1975, PLl, Fdg. 8). C'loniSequently, there al"e no speci- fic featua-es distinguishing the species hronica whicb therefore is rejected. lt is to n.ote othat the taxonomic value of the dall"k spot is doubtful, as this tStructure,
ns size, shape, .and location within the cells are .supposedly 1"t!S11WDg anay from fossilization and diagenesis, similarly as in some bluegreen algae (see Golubie &
HofmamJ. 1976). .
The species Gtobochaete gregaria SchIifer &. Senowbari-Dal"yan, 1980, was estabLiished due to "'.. diameters of single spores· between 150 and 180 microns, modus of division of the spores in lOOIlettes, and aggr~elike appearance of. nu- merous spores" (Schiifer & Senowbari-Daryan .1980, p. 100). Included into synony~
my by Schllfer & SenowbaIri-Daryan (1980) !Were the gregarious specimeDs· of Gtobochaete aZpina described by za,WiLdzka .(1972). As s.talted· above, the cell diam~ ...
tar .in Globochaete atpina i1'8Jhges from 40 to 200/l, ,the <type of a80S00iation ·is a mot- pbotype nat a specifdc :t.ea:ture, and the diverse modes o~ partitJian (the rosette-1i.ke including) appeB!' withln the same COlIliPosi.te s.pecimen. Usually,the cells situated
:nearer
to the ceDJbre 01 an acdniform association e!I'e mare densely spaced and, conotmry to the ou.ter cells, they dispJay a rather polygonal outline (see PI. 3, Flig. 5). Consequently all rtbe speaimens described as G. gregaria e!I'e to be Lnclu- ded :Into G. alpina, the same as those speCIimens ,presented correctly· tlndet" this very name by Zawl:dzka (1972).Corlcerndng tbe species Nostocites vesicu,Iosa Ma&lov, 1929, ,it was established 101" ucbed or coo.tor.ted, chain-like aggregates composed of loosely spaC'ed cells, about 45/l On diameter (see Text-fag. 2) The generic Illoatne was 'coined up by Mas-
..---_._. _._--_
..._---
t\ ~ . . . .
• . . . ., . '.~. _ r.Y.l
\_~. . ~
~ 9
3QOP.
Fig.
a.
N08tocites vesicu.1osa Maelov (atte'/' Maslov 1929)lov (1929) Ala anoounce the .relatiooshij;> Wdth~e .bl11e@reeD. algae .of the genus Nostoc, the view b~ ;rejeated by Maslovhimself dn a. later paper (MasJ.ov 1956,.
Formerly, MasJov (.1929) d:ist.ingudlshed ~ithin the
CealuS
,NoBtocites the w;o 6)pecies, viz. N. vesicu.1osa and· N. problematica~ The·latter species was· subsequently· su- ggested by Maslov (1956) to consist. of foramilndferil ·fragmems, but nevell'lthe1estt,NATURE· 'OF GLOBOCHAETE
51
it IW9S accepted and illuSltrated by Jobnson. (1963), Rdch (1974), and Bl'encJde (1977) ..
.Am. atllrdbut.ion· of .that tax~ to .either foa:aminifem OIl' to problemalti<: algae ·was
offered by 'Mamet & Roux (1978), ~ .also presented the tiJrst photographs of Nostocites: vesiCuZosa, taking them however: for an ac:inifurm ~n, nor for a JineaIr one. The sdze and type 01. the -cell arrangements of' the speoimen.s des- cdbed both by Maslov (1929, 1956, 1963); and by Mamet & Roux. (1978), clearly shows that these are the Carboniferous representatives 'of Globochaete alpina,' type Band D distinguished in this paper. The name Nostocites veBicuZosa is therefore rega.rded. by tl;he present Authoa:. as a ..subiective synonymy.
'I10 the Calrboniferous oc.cunrences of Globochaete alpina should a·lso be inclu- ded the specimens desClI1Lbed as Litostroma sp.. by Jansa, Mamet & Roux (1978, PI. 1, Fig. 11 and ? 10). Their • cell size (40.-:-80",), mode of partition, and shape of ceUs are mare similarr to Globochaete atpina (espec:iaUy to ISUch forms as i:]lus-tra- te.d !in R 3, F1gs
3
and 7 of this Paper; see also suggesi;i0D6 by'Vach8lI'd 1980) than·to any forms of the genus L*tostroma Mamay, 1959.RELATION OF GLOBOCHAETE ALPINA'l1O RECENT ALGAE
The taxonomic features important in Recent algae, are hardly reco- gnizable in Globochaete in its most advanced stages which display some resemblances' to the palmelloid and gleocystid stages of green algae.
Commonly accepted is the opiiuon e~ressed by Lombard. (1945) that Globochaete, recognized by Lombard oniy' in unicellar and chain-like forms, has a nature of green-algal zoospores, certainly related to the
·Pr6tococcaceae group of the order Chaetophorales and the· best analogies were suggested with the zoospores of the genus Dicranochaete. Tellez- -Giron & Trejo (1974) believed however that the composite forms evi- dence a more complicated nature of the· mother· plant. On the other . hand, Linetskaya & Muromtseva' (1973) were of the opinion that Globo-
~haete were the zoospores of various algae coming from diverse facies and stratigraphic ages.
Both the investigated material (PIs 1-3) arid analyzedreferences allow to recognize that. the best· analogies of Globochaete appear not only in zoosporic but also in vegetative forms of the twoordines of the gx.een algae, namely in the Chlorococcales and in the· ChaetophOtl'ales . (systematics according to' Fritsch, 1961). Taking' into account the mo-
des of partition, cell size and 'morphologies, the best pronoUnced simila":' rities are recognizable in the family Chlorosphaeraceae (cf. Herndon 1968) of the Chaetophorales, which includes i. a. such genera as ChZo- rosphaerops£s (? . =' Chlorospha~ra)' Vischer, '1933, PlanophiZa Gerneck, 1907, and Chloroplana Hollerbach, 1936. All these genera are represen- tedby unicells or' associations' and' coenobia, sometimes' of' . thecllain- -like outline '($ee Text-fig ... 3). TIle cells, ranging in . size from a few to over 5()p." are ~mbraced by a cell wall; usually· their the best readable element is the' 'chromatophor, cup-like' or.'ovoid ih shape .. Alfthe
52 S. SKOIotPSKl
A B
~ "· ' · · I
. " 'I ..
~' .. ::: "
I '
,
I
!'
~. 3. M.Ql'pholo~ types of algae .pl tile family ChlOl"108phaemcea.e A -:- ChZoro8phaera aU.nnatis Klebs (after Ar.taa:i; taken from Frit.sch 1961), B - PZanophUa asvmmetrica Gerneck: {after Ger.neck 1907), C - ChlorQplana teTTieola Hol.lerbach (after HoUerbach 1936)
Chlorosphaeraceae reproduce either by separation or by biflagellate zoospores (the latter of less than 8p in diameter). It seems therefore doubtful that the unicells of Globochaete, the same as those in diverse stages of partition, were the zoospores (their size attains 20(}J4 in G.
alpina, and 500,... in G. tatrica). Hardly believable. is also Lombard.'s interpretation of some convexities of the cells as the germinating flagellas' (cf. Lombard 1945, Text-fig. 1).
The Chlorosphaeraceae display also iill the morphological types dis- tinguishable in Globochaete (compare ,Text-fig. 3 and PIs 1-3). The Recent representatives of .that family live either free, witho\.'!.t any relation to a substrate. or epi- and endophytically. As an example. tl1e genus ChloTo8phaeropsi8 lives as an endophyte in leaves of many water plants, and as, an epiphyte of the duckweeds.
REMARKS ON CALCIFICATION OF GLOBOCHAETE
The investigated Carboniferous Globochaete commonly occur in association with diverse calcifYing algae, e. g. with Calcifolium of the family Codiaceae in the Visean and Namurian deposits (8ee Linetskaya
& Muromtseva 1973, Skompski 1980). with DonezeUa and BeTesellil
NATUR.& OF 'GLOBOCHAETE 53
(problematic Dasycladaceae) in the other Namurian deposits (see Linets- kaya & Muromtseva 1973; Maslov 1929 in regard With Nostocites vesicu- losa). A similar situation there also exists in the Middle and Upper Jurassic stromatolitic structures (see Lefeld & Radwanski 1960, Rad- wailski & Szulczewski 1966; Szulczewski 1968). The Globochaete itself, if its' attribution' to the Chlorosphaeraceae is substantuil; did not belong to the, calcifying algae. Its preservation In a calcified, form must there- fore
be
regarded as a result of fossilization, 1; e. of diagenetic processes, ,as it was already suggested by Lombard (1945).The first stage of calcification, certainlyreaUzed in the covering and enveloping of the Globochaete grapes by a thin layer of micrite. This process was supposedly similar to that acting in present-day bluegreen algal mats, and was a result both of biochemical precipitation, and of physical trapping and binding of calcareous ooze by the mucilaginOUS cover of Globochaete cells. The thickness of the micritic layer may be estimated as about 10t.L (see PI. 1, Figs 1-2 and PI. 3, Fig. 4). The settling of this envelope became the terminatiQn of the photosynthesis and the beginning of organic decay. Under favorable conditions, for example when buried by the sediments, either the Globochaete cells or more or less complete aciniform associations, have undergone succe- ssive calcification by secretion (c/. PI. 1, Fig.' 2), or disintegrate. If t~e
latter case happened, usually only the , outermost cells remained in a calcified state (cf. PI. 2,Fig. 2; also the specimens illustrated by Te":' llez-GiroIi & Trejo 1974), whereas an emptied Globochaete-bearing sphere might suffer from contortion and collapsing (cf. PI. 2, Fig., 1)- during further advances of diagenesis.
Institute of Geolow of the Warsaw University,
AI. Zwirki i Wigtt.1't,i 93, , 02-089 Warszawa, Poland
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56
S. SKOMPSKI
S. SKOMPSJq
BUDOWA I POZYC.JA SYSTEMATYCZNA GLOBOCHAETE ALPINA LOMBARD
(Streszczenie)
Przedmiotem pracy jest rewizje kosmopolitycm,ycb mikroorgani:unbw Globo- chaete aZpina Lombard, dokonana w oparciu 0 material pochodulcy z g6mowizeil.s- .idcb wapiem Lubelskdego Zagl~ia W~lowego. Stan zachowanda badaneg,o, mate- lI'ialu (patrz pl. 1-3) pozwolil .na usystematyzowande datychczasznanych postaci mOldolO~C7llY.ch w' obr~bie t.ego gatunku (patrz fig. 1). P1Or6owtlanie ze wsp6kzes~
nymi formama' Die potwierdzllo:poglQ.du '0 ZIOOOP~j naturze Globochaete, lecz wskazalo na pok!rewteilstwo teIO rodzaju z postaciami weg.etatywnym.i glonbw naleZlI.£yc'h do'
roomny
Chlorosphaeraceae, (patrzftg. 3). Wysttp}Wame r6z.nych staddbW 'zachOWlBDlia pozwoJilo ponadto DB odtworZL>nie przebiegu procesu kalcyfi- k:ac;li Globochaete. PodobiensotwQ do inny.ch lropaJnych .farm glonowych umotli-' wilo 'uzoame gatunkc)w Globochaete gregaria Schiifer & Seno.wbairi-Daoryan, Globo- chaete hronica:socza
oratz NoBtocites 1)esiculosa Maslov (patrz fig. 2) za synanimy Globochaete alpina Lombard. 'ACTA GECLOGlCA POLONICA, VOl.. 32 8. SKOMPSKJ, PLo 1
1 - Aciniform association of Gtobochaete alpina Lombalt',d; tangential section 2 - Aciniform assooiat.i.on of Gtobochaete aZpina Lombar,d; transverse section
ACTA G!!Ot.oGICA POLONICA, VOL. 32 s. SKiO~ Kt, PLo 2
1 - Aciniform association of Globochaete aLpina Lombard; superficial form with
a ,thick, muJltii].ayered eJOternal cover, toro out during lM-hification; at'1l"cw.ed is the
fragment magnified in PIl. 3, F.ig. 2
2 - Aciniform association of Globochaete alpina Lombard; superficial form with one-layered external cover, di&playing both ta-aaosver-se (at Zeft) and ~ellJtial sec- tion (at bottom right)
Ar::TA OEOLOOICA POLONICA., VOL. S2 S.8K Kl, PLo 3
bnea.I') and ,ac·iniform as&OO.iatiQIls rOf Globochaete alpina Lombard
1 - Linear association (fragment ot CalclfoLLum okense at bottom left); 2 - pseudoeplphytlc fragment of that presented In PI. 2, F.1g. ·1; 3 - transverse section of small aciniform asso- ciation; 4 - fragment of that presented in PI. 1, Fig. 1, with partition of cells; 5 - frag- ment of that presented Ni Pi. Fig. 2; 6 - fragment of that presented in PI. 2, Fig. 1
(anowed are cel13 with grea,t central 5potS); 7 - transverse sec.tion of larger aciniform association
