Vol. 37, No. 1-2
ST ANISLA W SKOMPSKI
The dasycladacean nature
acta gaalaglca pOlenlca
Warszawa 1987
of Late Paleozoic palaeoberesellid algae
ABSTRACT; Tthe .ex<.'etPtionailly well preserved moulds of problematic microfoSlSiIs from the tribe PalaeobeireseJ.lae MAMET & ROUX 1974, were found m the Lower Carboniferous !iimestOlIles in the wblm COIall Basin and in the Suidetes in Poland.
The mOll'pholugy of moulds allowed to state adalSYclaJdaicealn a:ffinill;y of the gI'Ollp,
which previ'OlUSl.y !Was assigned not only to the calC8lI'eousalgae, but also to the foraminifers and even to, the "'iPOnges. The ~tian of the dasycl8ida,cean nature of the tribe Paw.eobel'eselJleae evidences IpI'ecisely their role as a paneoe[wrur.OIIlmenta~
indicator, what is exempldfied by the Waw'sortian buRdups in Western Europe.
INTRODUCTION
The tubular benthic microorganisms, preserved in the form o·f cal- careous ,perforated tubes, and internally segmented by transverse annular partitions, have composed .one of the moot frequent bi.oti.c components in the shall'Ow water aJI'eas of the Devonian
srnd
Mississippiaa:JI seas. In the fo~al sense 'Of taxonomy this g1'oup of lIl'icrofossils is cl,assified as the tribe Palaeobereselleae MAMET & ROUX 1974. Informally, it has been named the palaeoberesellid or thekamaen,id group. All the hitherto made observations were based on investigations of thin sections, and they allowed to note a great internal differentiation in this group, 'what was supported hy the ahundance of species and genera,viz. Anthraco- porellopsis . MASLOV1956, Kamaena ANTROPOV 1967, ExvotanselZa ELLIOTT 1970, Pseudokamaena MAMET 1972, Kamaenella MAMET &ROUX 1974, PaZaeoberesella MAMET & ROUX 1974, Parakamaena MAMET & ROUX 1974, Subkamaena BERCHENKO 1981, Brazhnikovia BERCHENKO 1981, StyZaelZa BERCHENKO 1981, TrianguZinelZa MAMET
&PREAT 1985,and (connected with :them) Cribrokamaena BRENCKLE
1~85 and Crassikamaena BRENCKLE 1985. Despite of larger aoo larger amount .of papers concerning taxonomical affiliation of the palaeo-
ber~selli9s, this problem is still unsolved both in general, either animal
STANISl.AW SKOMPSKI
or plant attribution, and in a mOTe detailed sense of theiT taxonomical position.
The material presented in this paper and comiJng from the Lower CaT'boniferous deposits of Poland, affords new, probably decisive axgu-.
ments which indicate an algal nature of the palaeoberesellids. The here- after described specimens were pr,epared primarily from the Caribo- niferous. Limestone beds of the Dublin Ooal Basin (SE Poland) and, partly, from those exposed at Srebrna G6ra (Bardo Mts, Sudetes). The CarbolIl- iferous Limestone beds of the Lublin Goal Basin are lithostratigraphicaUy included into the parallic, Limestone~C1aystone Series (see PORZYCKI 1980), which is characterized hy the Yoreda'le type o,f cyolic sedimenta- tion, and whi,ch chronostratigraphically corresponds to the uppermost . Visean.
An extraordinary state of preservation of the algae in these deposits allowed the author (SKOMPSKI 1981, 1984) to revise a few genera, among others Calcifolium and Kulikia. The sedimentary enviTOO1ment of the Lower Carboniferous algal limestones and paleontological charac- teristics of their a,}gal 'content were also presented by the author in other earlier papers (SKOMPSKI 1985, 1986).
PREVIOUS OPINIONS ON THE NATURE OF PALAEOBERESELLIDS
The IspecimenS mcluded i"ecently in the u-tbeBalaeObereseHeae were primarily recQgnized as foraminif&s by EHRENBERG (1854) aII1.d vou MOLLER (1879)., The f<JO.'OOlinifeml affiliation of these miiCiOOfoosills" based on a genera[ morphologicall affinity, ;persisted a long time, ood in many de&er.ipt.ions theSe fot-ms have usually been assigned to the genera Nodosi.nelZa 8I!ld Moravammina (e.g. FOMINA1960, AIZENVERG & BRAZHNIKOVA 1966, MENNER & REI'l1LINGER 1971, TOMAS 1974). TheIit a'lgaJ. ;nature wajS suggested by MASLOV (1956) and ANTROPOV(1967), Who created the new genexe AnthracoporeZZopsis MASLOV, and Kamaena ANTRO- POV\.An essen1iial· axgumerut IDdieatillJJg their aLgal! alffilllity was the presen!Ce o;f pores oriented jpel"JpeInClicularly 00 the caIlcaJL"eows waIl1 of the 'Ilwbe. The genus AnthracoporeUopsis was 'chairalctarized by diidhdtomlously branclrlng pores, whereas the ,pores m Kamaena were ,straight ~unfOl"ltunately tlhese ~ were lOOt ccm:fJiJIimed in iH'llstrations).RIDING & JASNA (1974) olassified the genus Uralopo- reZZa KQRDE, 1950, t~her iWit'h AnthracoporeZZopsis and Kamaena. Thewem-
-maxk.ed iSepta of UraloporeZZa and a qlUesfliJonalblle porosity of the IMIM, determined these authors to negate
an
mga:I affinity of the whdle group (cf. also RIDING 1977);the classificatian of rpa1aeobereseltilds was iedit an OIPE!I!l. question, but ;some siIIni~
larlities to the foraminifers (mdiCirostlructure of the wan) were pointed.
MAMET & ROUX '(1974) made a getner'ail fl.'ev.ision of the dil9cussed group of microfoosils, 3IIlId they crealteid three new genera, Kamaenella, PalaeobereseZZa and Parakamaena, mcluded ,into the new tribe PalaeObereseLleae together witlh Kamaena,. . AnthracoporeZlopsis, ExvotariseUa and Pseudokamaena. This gfiOUIP, assigned ~h
a :doobt to the order PasyiCladlHes, was ~ as aooestars of the dasycladace8n GrOUP BereselJ.eae, !Which eJq)8lll;ded m the Upper' ~botnlferous. However, MA.MET
& ROUX 0(1974) didttlOt dispel dQubts reIQtiJng (pofI.'osiJty of. the,wd, becaUse,~
PALAEOBERESELLID ALGAE 23
some twenty of several hun~ of ErnceIlent micrographs showed ,!,rregular and
fr~entaTy pores.
The thirid 'ooncE!\Pt ,of classificatiO!ll of the ,pa·laeoberesellids aweared when TERMIER, TERIMlER & V ACHARD (1975; see also 1977) iJncluded aB. :flhe abGve indioa:ted genera to the new faanily MO'!'avammiJIlidae, which was olaJSSified within the group of the .sponges Ischy:rospotngia. The Moravam.miJn.iK:lae ;dififea:' from the foramilnifoos (a.ccol1ding to TERMIER, TERMIER & V ACHARD 1977) by: "lack of
proloculus, by the branching mode of perforations and frequent forking"; transverse
s~ta tiO!ll arr:Iid the microstructure of the wa[.1 were the main feaJtures differing them fTom the fQSSii:l algae.
The invesrtigations in ihe 80's aH.owed to distinguish ;SevemI thitherto unlmlOWlll
genera within the tribe PalaeobereseIleae, but aIllY new ideas of clalSSification did not fl!PIpeM'. V ACHARD (1980) was still includd.ng the pa'laeobere.seIUds to Algo- spoogia., and he even. e.x:panded thds thesLs by a!Ili HllustraUon of a hypothetica~
phyl'Ogenetie :relationship between the famiili.es Moravamminidoae and Aoujgaltidae.
LEES, HALLETT & HIBO (1985) ~Uipported this' IiJdea on the ba;s:is of a growth model .of the Waulsortian buildups. They a:t"gued that the both fwnilies are found together, below the photic zone, and diistribution of the mOlravammiJn.ids is lr.e-
minescent of the pattern shOWlIl by 'sponge SipilCll'~. ZADOROZHY J & YUFEREV (1980) and BERCHENKO (1981) ,stated precisely the rule.s of dIstiJn.guishdlng the paiI'tkUllar ;!ij)ecies and genera within the kibe Palaerobereselleae, and they COl1l-
silstently numbered this tribe to the order D$yclada:les. Conkary totb.em, BA:SSOULLET & al. (1979) e~luded this gIl'oup from :bhe fOSSil dasycladaooan aJgae.
Nev·ertheless, MAMET& ROUX (1981) strangIy a4vocaJted a position of :the ,palaeo- bereseIlLds W1it:h.i1n the green a'Igae (cf. also ROUX 1985, MAMET & PREA'r 1985), and added some new arguments :floc ·such an attriibwtion, viz. the lP:resence OIl pre-
su:mahleooo<~cles :in !l;he waH iIIl the genus PalaeobereseHa (MAMET & ROUX 1981, PI. 1, Fig. 2).
The preseO<t state of discusiSiOtn is the 'bes.t charaqi;erizeiCl by BRENCKLE (1985, p. 56), who Btated thaJt " .. .the palaeoberesellids and their descendants, the Bere- seZleae, should be united in a new family or higher taxonomic group, but left questionably within the Chlorophyta until phyletic relationships are established ... ".
Some specimens of paIaeoberesellids :trom the Lubli'll Ooa~ BaJSm. (main:ly those m thin .sections) were ;previQUiSly repoll'ted by the Qu!l;hO'!' (SKOMPSKI 1986).
The most importainthiint to'!' consideratiOlIls on the nature of thesej fossi[.s was a :ffiJnIdilIlg of I;l retirul&' rstructtm'e of !l;he pailaeober~seHid waU. TihJi.sJ feature ~
clealJ.'lly visi:ble :i:n reflected light arui it explaiJIls an effeclt of the wa1Jlporasity seen in transverse :secmaus. In coIllClusion, an aiflndty of the PalaeobereseNeae was mated (SKOMPSKI 1986, p. ~65) "to the red algae of the family Ungdarella";
ceae ... ", adthough this OjpdniOlIl wais regail'ded as "highl1l speculative in the present knowledge
.of
calcareous algae ... ".P ALAEOl3ERESELLIDS FROM THE LUBLIN COAL l;lASIN AND SUDETES
MATERIAl,. .
... The investigated specimens (see PIs 1~) have been prepared mainly
from the Upper Visean organodetritallimeStones pierced by the boteholes Orzech6w IG-2, Sawim IG-3 and Sawin IG-7 in the northeastern part of the LublinCoal Basin. In these limestones it is possible to disting),lish five microfacie$; -. ' . , . . the. . algal : . -,' ,-
M:E, ..
. -. . ... the· fQranrlnUeral ..' .. . ~ .MF,~e , crinoidMF, ", ," . ~24 STANIS;LAW SKOMPSKI
the SPiculiticMF,and the most frequently the organodetrital MF, which is composed of . mixed elements of other mkrofacies. The algal MF is characterized by the .Qccurrenceof more than 20 taxa (SKOMPSKI 1986).
among which dominate the phyl10id algae Calcifolium, aiIldrarely 'present aTe the palaeobexesellid algae, especially those of the genus Kamaena.
The l:nvestiga<ted material oorutain1s both the ~s recognized in thdn sections as well a;s thooe chemically prepared witlh. adm of 10% acetic acid. Due Ito a v~yinJg s.tage of neomorphism, some specimens dJn thin sec1li.ons diiSlptay a clear.
sparj,tic ~, while the othell'iS ~e miorosparitic O!l' paJ.1ily~tiIzed (see PI. 1 aJIlId PI. 2). A reOO'.tlSwuoction 'Of the primary, rea[ shape of the "cel!1is" (deteIl'mined by wan and partitions) . lis frequently ipOSsi:ble iIll Teflected light (PIl. 1, Figs ·lb, ab, 4b;
PI. 2, FigS 2b,5b;. see also SKOMPSKI 1986, P1. 5., F!igs 5-6; PI. 6, ~ 3-4).
Thes.ections taJngentiail. to the lSoull'face of tubes enable to show a reticuw structure of the waU(Pl. 1,· Figs 1.,.,..3). The jntell'1or I()f. tu'bets is u'S'lJ8fliy fW.led with 'bhe. same materlail as that sUl'II'oundingthe ~IlIS, but sometimes it js entirely sparltized (PI, 2, Fig. 3) I(){I." pyritized (arrowed iln P1.2, Fig. 2a). The latter specimens all'e extremely usefu~ :for mOll'iPholqgi.cal recOlllJS'tirucUOIrl.Sof the livmg pa!l"ts of the pla,n<t, becau.se dulling dissolution of li:mesione!S a fine-grained waill. of the. tube has been corroded earld.er than ,cOOIl"SJe-grained &paritic mtel'lnai mou~ds (e.g. thooe presented inPlls 3-6).; One specjmen {illUiStIl'ated ilnPL, 3, F.igs laJ..-:.1b) :iIS pa!rlticllllall'ly valuable, because it was ori~l1y composed of t,WI() parts (mouldS a;nd caJ.careous reticwate wa'l1), which have been sepa!l'aj;ed during· prepall'ation of the specimens for SEM.
In generiail, moa:e llhaJn 100 mOUl1ds were the .lSuibject of investigatiOlllS dJn SEM, but all of these specimens have been pt'epa!1'ed from saIIll!Ples which ID the 1ihiln secti~
oOrlisisted only of s.i:ngle, isOlated palaeoibelresellJil (3-5 ~mens· iIll iSta'Illdalrd thin section). Aipaa'1t from palaeobe.resellids (mairny Kamaena), the only dliiStillguish- able forms were the f·oraminiferIs (e.g. Archaedisc'Us, Howchinia, EostafelZa, Endo- th'Ura, Tetrataxis), IPhylloid Mgae (Calclfoli'Um okense), amd fil'agme.nts of Cil'iJooids.
Some moulds (PI. 3, Fig. 2 and PI. 4, Figs 1, 5) have beell1i prepared from the LqwerCarooniferOlUS limestones exposed at Srebrna G6ra in the Bardo Mts (Sudetes). These rociks are. characterized bya high&
degree of neomorphism and they represent the biomicrosparitic MFwith fragments of crinoids. tabulates, brachiopods, Sponge spiculesand single 1!nicellulalr· f,o~inifers.' The walls of rare palaeobereseUids are herein yellowish and sparitic.
MORPHOLOGY OF MOULDS
At first glance, the dasycladacean type of morphology is' clearly visible on the ilnvestigated moulds: verticils of lTamifications branch out of the central stein, andth&efore the dasycladacean terminology is used in the following description.
In general, all the 1st order ramificatiOJlS are distributed in the form of verticils, among which it is possible to distinguish three types {Text- -fig. 1): the single-row verticils, the double-Tow verticils, the oomposite verticils .
. In. the. s~e-row verticHs.; mmificaM.Ons are dd:s1lributed ID the· plane pel'- pendliCUl1:ar to the axis aJIlId in-tile equal d:irstance one from another (Pl., 6, F~ 1"":"2,
ACTA GEOLOGICA POLONICA, VOL. 37 S. SKOMPSKI, PLo 1
Palaeobelfesellid algae, Kamaena delicata ANTROPOV 1967, in thin sections, in transmitted (a) and reflected (b) light
1-3 - Longi,tudinal, 'sli,ghtIy oblique se.cti'OI1iS thrl()lugh curved :parts of tubes (retkular .,tructure of wall is visible), 1 - Ispecimen f:rom borehole Sa.win-7 (derr:>th 1083 m), X 30; 2 - e'l'1lrurged :part of Fig. 1, X 60; 3 - Slpecimen f["om Podedw6rze-2 (de:pth
584 m), X 40
4 - Longitudill1aol section:s, specimens from borehole Sawin-3 (depth 1137 m), X 30
ACTA GEOLOGICA POLONICA, VOL. 37 S. SKOMPSKI, PLo 2
DiUerent stages of preservation of palaeoberesellid algae in thin sections,
iln transmitted (a) and reflected (b) light
I, 4-5 - Sections w.ith pan:Uy s,p.&-itized interoa'l 'Pan:ts of tube, thiln neomocphic layer is visible on some patrtiltioJ1s (FLgs 1 and 4); 2 - Section w1th partly' pyritizeid (arrowed) IinternaI pa'l't; 3 - Section with tot'a'lly .spaD:i,tized .iinteroall palrt; 1-4 - specimens from borehoJe Sawin-3 (depth 1137 m), 1 talken X 60, 2-4 tak€n X 30;
5 - fr-om Sawiln-7 fd€a>tJh 1069 ro), X 60
ACTA GEOLOGICA POLONICA, VOL. 37 S. SKOMPSKI, PL.' 3
1 - Mould of .palaeobeJreseHid a1ga, aharacterized by cOl11jposite t~ of verticil~",
(Fig. la) and fragment 0[ calcareous sleeve with retiC'U!lar structure (Fig. lb);
spec~men fTOlm bore hole Orz.eehow-2 ~de!Pth 918 m), X 125
2-4 - Mou,lds with double-Tow verticils of branches (Fig:. 3b - recOtnlStTuction of oa,lcareous 's~eeve on the mo.uld. Hg'UTed iin 3a); 2 - specilmen from Srebrna Gfua, X 150; >3 - from bOlI'ehole Orzech6w-2 (dejpth 918 m), X 100; 4 - from
Sawin-7 (!depth 1069 m), X 1100
ACTA GEOLOGIC A P'OLONICA, VOL. 37 S. SKOMPSKI, PLo 4
1-3 - MOIUlds of palae'obereseIlid aLgae, charadeo:ized by cOl!rl;posite vertidls (Fig. Ib - reconstruction of ,calcareous sleeve on the mould figurr·ed in la; Fig. k - 1.."l11ar,ged palrt of Fig. la); 1 - s;p.ecimen from Sr-e'iJIJ'na Gfua, X 60; 2-3 - specime!1ls
from b:arelhoille Orzech6w-2 ~depth 918 m), X 120
4-5 - MouLds with double-lfow vertidls of 'brau:JChes ~F.i.g. 5b - r,ec,o'!1Istrucmolll of oalcarreQUJS s1eeve); 4 - iSpecilffien from Orze·Clhow-2 (:depth 9,13 m), X 70; 5 -.
s,pedmen from Srebrna Gfu'a, X 135
ACTA GEOLOGICA POLONICA, VOL. 37 \ S. SKOMPSKI, PLo 5
Different ty,pes O'f verticils of ,pa.laeoberesellid algae: 1-3, 7 - double-r'Ow verticils, 4-5 - cOlIDpos.'ite ve:rtidhs, 6 - double-row or co.m.posj,te vertici:ls
1 - En!arged part of specimen presented in PI. 4, Fig. 4, X 140; 2 - enlarged part of specimen presented in PI. 3, Fig. 2, X 300; 3 - specimen from borehole Sawin-3 <depth 1137 m), X 200; 1 - from Orzech6w-2 (depth 913 m), X 150; 5 - enlarged part of specimen presented in PI. 3, Fig. la, X 170; 6 - ' specimen from Orzech6w-2 (depth 913 m), X 180; 7 - enlarged part
of specimen presented in PI. 3, Fig. 3a, X 130
ACTA GEOLOGICA POLONICA, VOL. 37 S. SKOMPSKI, PLo 6
1-2, 6 ~ MiOulds wit:h si~gle-row vertici·1s of bra~qh€ls: 1-2 - specimens from.
boreoole Orz.ech6w-2 (depth 918 m), X WO; 6 - from Sawilll-7 MeJPth 1069 m), X 90 3, 5, 1 - Moulds with cOIm;pOlsi<te vmkils, specimans from Orzech6W-2 (Fig. 3
from depth 918 m, X 150; Figs 5 00Jd· 7 frOlm dep<th 913 rn, X 75) 4 - MO'UI1d with double..row verticils, :fa:Qlll Orzech6w-2 ~depth 910 m), X 125
PALAEOBERESELLID ALGAE 25
6). The. iiIl!terverticil distance cO'lTesPonds' l.JiS1lHitlly 10 .the dianieterof the. central stem {PI. 6, Figs 1-2), but sCimetLTnes it lis oonskiera'bl:r smaller; (Pl 6, Fdg.6). .
In the double--row 'v~ticils(pI. 3, Figs ~; Pt 4, FigiS 4, 51; Pt .5, F~s 1~;
2-3, 6?, 7; PI., 6, Fig. 4), ramificati'OltlS are arra.ngediln two symmetrical a;nd adjoinling who!r'ls, in which partioolar 'branchelS8Il'e' generail.iy coupled (Pt 5, F.igs 2-3 and 7). Sometimes, this symmetry is not precist!
what!;
CIOIUlld' be caused.' by:a:
jpOOIr state of preservati<llll. The interveriticH distanceis
.uStiaHysiariilar· to . the width of verticils arid it is sigriificaJntly sinaJl.[er (1/3 or 1/4) than the diameter' of the centra,l stern.In the composite verticiJl:s theire M"e at iea.st two tYlPes of distribution: either ramificatiolrus are arir8lIlged iJrregularly (Pl. 3, F.ig. la; :Pl. 4, Fjg. 1; PIl. 5, Fjg. 5), or they are lSituated in a ~ adjoining wholl"ls.PartictilaX, coia'esporiding lI.'aniifiea,,;, tioru; from different whorls are lO!ften jointed 8IIld they. form smailll rolls, pazraHel to the main aX!is of' the aaga (PI. 4, Figs. 2-3;' PI. 5, Fag. 4; PI. 6, F~ 3, 5' allld 7).
The interverticil distance is al1ways distinctly sroaner than the width of verticlls.
r
:lIL. tlIl"
I I
J I
".,
~.
B' . c'
Fig. 1. Types of verlicils and calcareous sleeves
A - .si«lgle-row verlici1s, B - d9ulble-row veriticlJs, Cl - composite vertic.ils with braOOhes diStributed in the form of rolils, Cl! - COIqpOISUe verticias with in'eguRaroly lI1I1'aaJged branches; A', B', C' - hypothetical recon.structions of ca[calt'eOUS sleeves,
, ' : '. ' .: '::, ,. ~poIlidtng with parli:cUilar types of vertiClls ' .. .
26 .STANIS:t.AW SKOMPSKI
The shape .of 1st order ramifications is pretty much the same in each type of vertidls; stTaight, elongated, and cylindrical. Their length is difficult to estimate due to corrosional character of specimens; only in a few cases (PI. 3, Fig. 1 and PI. 5. Fig. 5) some ramifications have preS&ved their primary length.
The measurements of specimens change on a large scale and theyaa:-e not dependent on the type of verticils. A 'diameter of the central stettn ranges from 40 to 230 .... m. whereas the external diameter of the verticil changes from 80 to 350 .... m.
A CONSTRUCTION. I~ THE PALAEOBERESELLIDS
A comparison of informations yielded both by thin sections and by prepared specimens allows a rreconstruction of the main featmes of the palaeoberesellid structure (Text-fig. 2). ' These algae were characterized by a construction typical .of the euspondyl Dasycladaceae, 'but calcifica- tion did ;not occur around each ramification. This prrocess usually took place outside of the central stem and 0Il1 the suTface of the whole verticil.
The only e~ception seems to he the genus Exvotarisella, in wpjch calci-
s
~R2
. Fig .. 2. Scheme of
C.<m.structl9n
'of' the palaeoberesellids.i ~
calcareous
sleeve,RI .:.-..
1st' ,order mm~i~on:.2 ' -- 2nd .
orderramUieauOO
PALAEOBERESELLID ALGAE 27
ficaUon penetrated the interior of verticil and it surrounded the parti- .culM." branches. In the interver,ticil segments the calcareous sleeve was
distinctly thicker than in' parts corresponding to the verticils, but the .external diameter of tubes was constant, what in consequence gave an .effect of internal paa-titions. The calcareous sleeve was composed of wide reticular (pseudoporous) sections, homologous with verticils, and narrow, smooth sections without "pores", homologous with in1:erverticil aa:eas..
"The second-order ramifications, hypothetically reconstructed in a tTicho- phorous form (Text-fig. 2). probahly projected though reticular paa-ts of the sleeve.
PROBLEMS OF TAXONOMY
The taxonomy of any group, based only on the features trecognizable in thin sections, will always cause doubts, because the differentiation of sections is usually greater than the variety of real forms in a particular fossil. Difficulties in finding a relation between thin sections and spatial forms .of Paleozoic foraminife:rs could serve as an especially good ex- emplification of this prdblem. On account of a generally smaller number of investigations these difficulties concern also calcareous algae, what could be supported, for example, by the discovery of real morphology of the genera Calcifolium and Kulikia in the Vise'an dEWosits of the Lublin Goal Basin. In the case of the tribe Palaeobereselleae, a relation between thin secticms and prepa'red specimeru; is alsouniclirectional, i.e.
it is poss:i!ble to find the shape of sections of the calcareous sleeve on basis of internal moulds {see PI. 3, Fig. 3 and PI. 4, Figs 1, 5Y, but in- versely it is difficult.
According to MAMET & ROUX (1974), the taxonomy of the tri:be Palaeohereselleae was based mainly on the shape of "cells" and trans- verse partitians, type of porosity and on the measurements of specimens.
In the light of the aiboV'e presented data on the dasycladacean nature of discussed farms, all of'these featUJres are questionable (cf. SKOMPSKI 1986). Particular pacr:ameters oftrubes and "cells" are dependent on the plane of section, and thus they are rather accidental. The type of veriicils, which should be rega.r:ded as a principal feature of ,the palaeoberesellids, is also independent ·of the measurements and it does :not correspond with the shape of "cell" and tube. For example, the sections of the sleeve similar to Kamaena delicata Originate ,both in the case of double-row and of composite verticils. The porosity of wall, observahle in thin sections. oomesponds at moot with the less important and hypothetical 2nd order ramifications, while the 1st order branches have been left imperceptible in thin secticms. Finally, the shape of partitions is probably more dependent on the mode of preservatiOlIl and the way .of neo ...
morphism, than on the real differences in the structure of algae.
- - - -
.,
28 - STANIS:t.AW SKOMPSKI
In this situation, a dozen or so genera and a great number of species distinguished on the basis of thim sections within the tribe Palaeo- bereselleae, seem toO be a distinct overstatement. NatUTally, a great taxonomical variety of these algae is not excluded, but verification of this is possible Qnly when the specimens J)Q'epared from the Tocks are obtained. The material descrihed in this paper allO'ws. to recognize at least 3 genera with different types Qf verticils, arnd the hitherto existed descriptions of the palaeoberesellids point to the o1her next possibilities.
For example, the genus Tringulinella MAMET, 1985, is cha:racterized by a triangulatr shape of perpendicular section of the tUbe. By analogy to' the other Crurboniferous Dasycladaceae (e.g. above mentioned genus Kulikia), it seems probable that this O'rigimi.1 feature is caused by a meta- spondyl arrangement .Qf ramifications (3 clusters in each verticil). It is also probahle that in the case of a higher frequency Qf specimens, it will be possible to find statistical relation between thin sections and tmee- -dimensional specimens.
The next question refers to the pooition of the herein reconstructed Palaeobe:I"eselleae in the hitherto existing classifications. HomO'logies in the structUJre of the palaeoberesellids and the tribe Bereselleae are still veritable, in spite of new recons1n.-ucUon of the former group. They diffe!I" surely by the type Qf branches, which are th:iJnner :in the tribe Bereselleae. BASSOULLET &, al. (1979) excluded. the hoth group from the mder Dasycladales, although. HERAK & al. (1977) connected with the Bereselleae (amQng ,otherS) an important stage of evO'lution iIll tltis very order r(3rd stage, appearanc1e of ver.ticils). The Palaeobereselleae and Bereselleae are thereby similar to fQrms of such taxa as the genera Nanopora and OLigoporella, but they distinctly differ_ by their type of calcification realized arQund the verticEs. For that reaSQns it see.rn.s necessary to unit the _ both groups in the fQI'IIl of a separate tribe withim the ordeiT Dasycladales.
ECOLOGICAL IMPLICATIONS
Interpretative importanC€- of the palaeolberesellids !has oonsidelt"ahly increased by confirmation. -of the dasycladacean nature of theSe micro- fossils. Bathymetrical range of Recent Dasycladales is relatively small, as usually these algae ,occur iIll masses not deeper thaJIl 30 m, although sporadically they exist to the 90 m (FLUGEL 1985).
AbundaPJ; :remadns of the palaeoberesellid algae OCCUil' in shallow~;
-water, neaiT-subtidal environments of Devonianand Call'boniferQIUS epi';'.
platform seas, in association with the -other skeletal elements O'f similar bathymetrical limitations (cf. DREESEN & al. 198.5, HENNEBERT &
HANCE 1980, SKOMPSKI 1986, . CHUV ASHOV. & RIDING 1984). Their:
PALAEOBERESELLID ALGAE 29
value is greater in sediments significantly dominated by the Palaeo ...
bereselleae or in such deposits in which theya;re sporadic (e.g. of hyper- saline or brackish waters) .
. The presence of the palaeoberesellids (Kamaena-like forms) is also recorded from the Waulsortian build'Ups in West&n Europe regarded as deep-wat& (see LEES & al. 1977). Bathymetry of these enigmatic mud- -mounds is. still a subject of discus'Sion, but in general the init~ations of growth of thE' Waulsortian buildups tooL~ place at sUlbphotic depths, probably exceeding 300 m (LEES & al. 1985). Such ,depths are estimated
on
the basis of approximated lower limits of the calcareous algae, micri- tization, pluriloculrur forarllinifers and hyalostellid sponges. The palaeo- beresellids appeared ill the phase Band C of buildups growth, which were still connected with the subphotic zone (depth 200-300 m according to LEES & al. 1985; see also LEES & MILLER 1985, Fig. 8). Recognition of dasycladacean nature of the palaeo>beresellid algae offers a different interpretati<m. of the Waulsortian mud-mounds, which are thought to have probably originated in shallower environments.Acknowledgements
The moot ·sID<:&e th8nks aa:e expressed to Profesror A. RADW ANSKI, UniveTsilty of Warsaw, :for a critical reading ami imJ;lroVling an etail'lier
mait
of this !paper, ami to M. P ASZKOWSKI, M. Se. (Jagie1~dn!i'aal University),· who Iktiintdlly sllIPPlied the·specimens from the Sudetm. The author is also greatily indebted :to B. W &KS- MUNDZKI, M. Se., for malking C'an-efully the !Line dxaIwing fur Tm-fig. 2, and to Miss E. KLICHOWICZ, M. Se., for takdng the SEM-graphs.
The Geolqgical Survey of Poland (SMesian Branch) ikilndly aHowed the author to s~e the bo.:rebtole cores fu'om the Lublin Coal Ba,s:Ln.
Institute of Geology of the UntverBtty Of Warsaw,
Al. ~wtrkt t Wtgury 93, 02-089 Warszawa, Poland
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S. SKOMPSKI
DAZYKLAOOWATA NATURA POZNOPALEOZOICZNYCB GLONOW Z PODRODZINY PALAEOBERESELLEAE
(Streszczenie)
Plrzedmiotem pracy * jest aaializa 'budowy glonów wapi€!l'1lIlych z podrodziny paaaeo>bere,seUeae MAMET & ROUX, 1974, za'chowanych w wyjątlrowym stanie
ośródek, a wypreparowanych z do1noka!rbońSJl.tiJC1h ;wapieni Lu>be1saciego Zagłębia Węglowego oraz Sudetów. Te problematyczne mikroskamieniałości, ZI!1a!Iłe dotych-
CZaI> ze szlifów (patrz pL 1-12), zaliczane były ,me tylko do gIOIl'lów, ale il'ównież
do otwornic, a nawet gąbek. BoróWIl!B.nie morf()lQgi.i ośródek (patrz !pl. 3-6) i prze- krojów otaczających je rwapiemJ.YICIh osłonek wslkazuje wyra:bne podobieństwa bu- dowy przedstawicieli lPOCh'odziny PalaeobeIesełleae do gloOlIlów wapiennych z rzędu
Dasycladalles (patrz fig. 1-2). Sprecyzowanie taJksonamicZI!1ej pozycji pa,loobereseHi
zwiększa ich znaczen!ie ;pa:iZy analizie ś,rod<lWislka s'edymentacj'i, pow,ooując u!p. istot- ne zmi'any dotychczasowych wniosk<lwań paleobaty.metrycznych, co zaprezentowano na przY'kładzie facji Waru.1>wrtian z dolnego karbonu Eul'OIPY Zachodniej.
* Praca wykonana częściowo w ramach Cent>mlnego Plrogra'llllU Badań Podsta- wowych (problem nr 03.04). kOO1t'dynowamego ,przez Instytut Nauk Geologiczny,ch PAN.
