GEOL. CROAT. 49/1 I · 79 3 Figs. !O Tabs. 22 PIs. ZAGREB 1996 

Taxonomic Review of Some Barremian and Aptian Calcareous Algae 
(Dasycladales) from the Dinaric and Adriatic Karst Regions of Croatia 

Branko SOKAC 

Key words: Calcareo us algae (Dasycladales), Bio­
strati graphy, Taxonomy, Lower Cretaceous, Dina­
ric-Ad ri atic karst, C roatia. 

Abstract 
Selected species of the followi ng dasyclad genera, deriving frO Ill 

Barremian ,lt1d Aptian depos its of the Ad ri alic and Dinaric karst, are 
elaborated: Salpingoporello, Acroporclla, ILcfilloporella, Clypeil1(f 
and Pralltr/olle/fa. A new genus, /Jirijerel!a, is establi shed. Descrip­
tions of (he following Salpillgoporelfa species arc compl e ted with 
newly establi shed characteristic fealu res and i llust rat ed with appropri­
ate sections: SalpillgolJOrella Il/Ilehibert~ii, S. melitae, S. IIispall/ca, S . 
lUISi, S. IIriadonasi, and S. dil/arica. In addit ion, the follow ing new 

spec ies are dcscribed: S. l'caIiCO.W, S. hera/dica, and S. po/ygol/alis. 
Apine/fa jafre:oi. the type species of the ge nu s Apillella, is considered 
to be a younge r synonym of Salp illgoporella hispallim . ACfilloporella 
fJodolh·a and Acroporella radoicicae are commcnted upon. ProWl" 

lonella dalli/ul"ae is ill ust rated. Sel/ipo,.el/a da/matiea and Likallella 
pejOl'il:(le have been transferred to Pratl/rlol/ella. Because the pro­
bl emati c lIalicory/le /lerae has a lso been transfe rrcd to the ge nu s 

Prafllr/ollc/fa (with it 's inadequate diagnosis and description), a new 
diag nosis and more comple te description o f th is species is give n. 
Clypeilla? solkGni is illustra ted by Barremian specimens and Ciypeino 
gigalllea n.sp. from the Lower Barremian of Mljet Is land is described. 
Bes ide Ihe type-s pecies. ririferella SpiIlOS(I, Ihe species hi therto 
labelled Clypeilla somoli(·(/, or Similidypeilla somalica, has also been 
trans ferred [ 0 [he new gerllLS Pinferella. 

Klju cnc rijeci: vapllellacke alge (Dasycladales), b io­
slrati grafija, taksonomija, donja krcda, d inaridski ­
jadranski krs, HrvalSka. 

Sazetak 
Iz bogatog matcrijala fosi lnih vapnenackih alga (Dasycl adalcs) 

priku pljenih u naslagama barema i apia d inarskog i jadranskog krsa 
obradenc su ncke vrste rodovu: Sa/pillgoporella, Acroporclla. AClino­
pore/hi, Clypcina i Prall/rio /l e lia, a usposta vlje n je rod Piri/"erella . 
Kod vee pOlnalih vrsta roda Stilpillgoporel/a: S. I/lItclllhergii, S . //Ie/i­
/(le, S. hi.\·/Jallica, S. hasi, S. lIr1adQ/w~·i i S. dil/arica, LI Z ilustrativne 
fo tografije presjeka razlici tih orjentacija izvrsene su dopune opisa i 
navcdene novo uocenc znacajke. Opisane su nove vrste ovog roda: S. 
I'errucosa, S. heraldica i S. po/ygollolis. Rod i vrsta Apillella jt/Fewi 
smatra se mlad im sino nim om S. hispallictl. Komen[irane su vrste 
Acrilloporella podo/i eo i Acropore lla radoicicat' . Od vrsta roda 
Prar/lrlollelfa ilu strirana je P. dalli/ovae. Rod u su pribrojene Sel/i­
poreI/o da/matica i Likallelfo pejol'icae, a s obzirolll <la je osporcna 
fia licorYllc nerae prebacena 1I rod Praf/lrlollel/a s neadekvu[nom 
dijagnozom i opisom, to j e dana nova dijagnoza i dopuna op isa ove 
vrste. Uz vrs!u Clypcilla? so/kalli prikazanu baremsk im primjcrc ima 

opisana je Clypeillo gigal/rca n.sp. iz donj eg bnrema otoka Mlj e lll. 
Novom rodu Phi/ereI/o, LIZ ti psku vrslu Pir!fcrcfa .Wi/rusa, pribraja se 
i vrsta 1'. sOllla/ica do sada nejasne tak sonOlll skc klasifi kacije, 
prethodno oznncenn kao Clypeil/a sO/1/a/ica odnosno Simi/iclypeillll 
somalh.·a. 

Contents: 

I. INTROOUCfION .. . ....................... ......... .................... 2 

2. SYSTEMATIC DESCRI PTIONS ................ . . ....... .. ... ............. 2 

Sa/pillgoporclla IIIlIeh/hergi (LORENZ) PIA in TRA UTH ....... 2 

Salpillgopo/"el/a lIIeliw e RADO IC IC .......... ..... ........ 3 

Safpillgoporel/a Itispollico CONR AD & GRABNER .. .... ..... ... ..4 

Sa/pingopore/la \'ernu:osa n.sp .. .. . ............ 5 

Salpillgopo/"ella lierafdica n.sp .. .. _ ............... ...... ...... ...... ..... 8 

Salpillgoporel/a polygol/alis n.sp ... .. .. .... .................. ... ........... ~ 

Sa/pillgoporella hasi CONRAD et al 

Sa/pillgoporella IIr/adol1a.ti CONR AD CI al 

....... .... ............ 13 

... .. ............... 15 

S(lfpillgoporella dil/orico RADOle lC ......... .. .......................... . 17 

Pmlltrlollella nerae (DRAGASTAN el al.) BUCUR ....... ........ 19 

Prall/dollella da/lllmica (SO KAC & VELl C:) n.comb ............. 21 

Prmllriollella pejOl'icae (RA DO ICIC) n.comb ........................ 23 

Acroporella /"{Idoicicae (PRAT UR LON) PRATU RLO N 

& RA DOICIC. . . ..... ............................ .. .... 24 

Clypeil/(/? so/kalli CONRAD & RADO JC IC .. ... ... ..... .. ... .. ... ... . 25 

Clypeilla gigalllea n.sp ........... ....... .. .............. . 

AClilloporella podoJica (ALTH) CONRAD CI a l 

..26 

.... 27 

Pin/ad/a 5pillosa n.gen .. n.sp ............................... ....... ... .. ... ... 28 

Piriferd/a somalica (CON RAD et al.) Il .comb ...... ........ .......... 30 

3. REFERENCES ............................................................. .............. 31 

PLATES I-XX II .. ..36 

Inst itute of Geology, Saehsova 2, P.O. Box 268, HR-J()(X)() Zagreb, Croal ia. 



2 

1. INTRODUCTION 

Long- lasting investi ga tions of Mesozoic deposit s 
car ri ed ou t by the author and nume rou s colleagues in 
the Dinaric and Adriatic karst regions of Croatia have 
res ult ed, alllong othe r things, in an abundant collection 
of foss il material. Among this, calcareous algae (Oasy­
cladalcs) rank as, pe rhaps, Ihe foremost const ituent of 
the foss il assemblages in gene ral , with regard to both 
thei r divers ity and abundance of specimens, in parti cu­
lar strHtigraphic hori zons. 

The analysis of th is rnate rial enabled the definition 
of precise stratigraphic positi-on and/or range for many 

of the me ntioned species, owing to the pos ition of 
algal -bearing samples in the st ratimctrically logged sec­
tion s, as well as an swering a Ilumbe r of taxonomic 
qucs tion s owing to the abundance and good preserva­
tion of the material , whic h can be illustrated by appro­
priate sections . In the Mediterranean realm , the foss il 
Dasyc ladales have been the subject of num c rous 
palaeontologic and biostratigraphic inves tigations, pre­
sented in numerous pape rs by differe nt authors. How­
ever, the primary descriptions are o ft en based on scarce 
and/o r poorly preserved mate rial (w hi c h somet im cs 
shows diffcrent fealurcs and thus implying heterogene­
ity), accompanied by few and inadcquate illustrations -
sometimes even one section of a fragment with blurred 
c haracteri s tics has been lIsed as the basis for establish­
ing a new species o r the alleged rev is ion of an cxis ting 
one . Such bad prac ti ce has resulted in inadeq uate 
knowledge about particular taxa, un sound establi shing 
of taxa, a ll eged c harac teri stic fea tures resu ltin g from 
the subjective inte rpretation of the author, unjus tified 
inc rease in the nu mber of ne w taxa, and frequent trans­
fe rring of individual species to anothe r genus without 
sound arguments ("n. com b. "). Besides, several species , 
or groups or species, of the same genus (e.g., Sa/pil/go­

parella) may show simiiarl y shaped ramifica tion s 
(branc hes) which can produce sec tions that may appear 
s imilar o r almost identical for two or three species, 
leadin g to co nfu sion in identifica ti on by less experi­
enced workers. Thi s implies that the original descrip­
tions ofte n lack adequate illust rations; whereas many 
are accompani ed by too fe w characte ris tic sections, 
some of the sec tions illustrated arc 110t species-charac ­
te ri sti c and many belong to two or more species (t his 
may depend sometimes on very s li ght d ifferences in the 
plane, or "depth", of the section). The most obviolls 
e rro rs occur w he n the identifi catio n of a species is 
based 011 - o r, at le as t, res ult s fro m - it s s trati graphic 
positi on. An even more seriolls problem lies in the 
intraspec ific variation which can be more or less pro­
nounced and can affect the shape of the branches (most 
frequently, the shape o f the pores at their dis tal e nd), 
numbe r of branches per whorl , and - most frequ entl y -
the range o r dimensions. All thi s produces the dile mma 
of whi ch differences should be treated as intraspeci fi c 
va riati ons and whi ch a re spccies-characteristic. Some­
times, admilled ly, we are dealing wi th continuous 

Geologia CrO;l lica 49!J 

ranges or forms whose end members, howeve r, clo 
show clear differences. From a pragmati c point of view, 
proble matica l ident ificatio ns and eonrusio n of taxa 
have serio LI S conseque nces with regard to the ir s trati ­
graphic use fulness and rel iability, though, in my opin ­
ion, these parameters s ti ll seem to stay quite firmly , at 

least on a local to regional scale. Of course. st ratig raph­
ic ranges are always subjec t to change, but the new data 
should be based on indisputable identification and pre­
cise a nd well documented evidence of stratigraphi c 
position. Ait that has bee n mentioned above represent s 
but a smail fraction of the unsolved questions in the 
study o f fossil algae, a ll of which concentrate on, or can 

ultimate ly be reduced to, the establi shin g of val id and 
unequi vocally adopted criteria for identification . 

Perhaps the tone of thi s paper may sometimes sOllnd 
polemic or unduly critical. My inte ntion was, howeve r, 
to clarify the taxonomy by comple ting the hithe rto exi­

s ting descrip ti ons of sOllle taxa, to support my cla ims 
by unequivocal sections (thercCore the somewhat larger 
Ilum ber of illustrations - a fact that was obviously miss­
ing in Illany earlier pape rs), to correct some incorrect 
interpre tations, to describe some taxa in the manne r I 
consider correct, to de limit the ir s trati grnphie ranges in 
the Dinaridi c and Adriati c karst areas, and to expla in 
some of my reflcctions on the validity or the new taxa 
and the criteria of the ir establish ing and recogn it ion. 

2. SYSTEMATIC DESCRIPTIONS 

Genus Sa/pingoporc/la PIA in TRAUTH, 19J8 

The generic diagnosis by PIA (in TRAUTH , 1918) 
has been comple tcd by CON RAD ( 1969) and summa· 
rized by SC HINDLER & CONRAD ( 1994). To thi s, 
the followin g addition should be added: Species with 
branched thallus, or showing a te ndency toward branc h­
ing, <:Ire also included in the genus. 

Salp;ngoporella IIlllch/bergi; (LORENZ, 1902) 
PIA in TRAUTH, 1918 

PI. I, Figs. 1· 13; PI. XX, Pi gs. 48 , 58 

Selected synonym y: 

1902 Dip/opora !I1i;h1bergii n.sp. · LORENZ. p. 52· 
54, Figs. 3-6 , 'I Fi g. 7 

191 8 Sa/pingoporella l1Iuehlbergii (LORENZ) l1.gel1., 
n.comb.· PIA in TRA UTH , p. 2 11 ·213, Fig. 4a 

1978 Sa/pillgoporel/o mllch/berg ii (LORENZ) 
8UCUR, p. 95, PI. 8, Fig. 5 

1978 Sa/pillgoporel/a IIIlIeh/berg ii (LORENZ) 
DRAGASTA N, 8UCUR & DEMETER, p. 22· 
23, PI. 2, ?Fig. 4 ; PI. 8, Fig. I pars; PI. 9, Pig. 3, 
non Figs. 1,4, 'IFig. 2; PI. II , Fig. 4 pars 

1978 So/pillgoporel/a mllchlberg ii (LORENZ) 
CON RAD & PEY8ERNES, p. 69·83, PI. 2 , 
Fig. 10 



lobe: Taxonomic Review or Somc B~lrrcmian and Aptiall Calcareou~ Algae ... 

1978 Salpingopore//a mllehlbergii (LORENZ) 
YELlC & SOKAC, p. 3 19-334, PI. 2, Fig. 2 

1979 Salpingopore/la IIlllehlbergii (LORENZ) - PEY­
BERNES & CONRAD, p. 743 -751, PI. J, Figs. 
6-7 

1980-1981 Salpingopore//a I1lllelilbergii (LORENZ) -
BUCUR, p. 55, PI. 1, Figs. 7-8, non Fig. 6; PI. 2, 
?Figs. 1-2 

1984 Salpingopore//a /1/llelilbergii (LORENZ) -
LUPERTO-SINNI & MASSE, p. 341, PI. 38, 
Figs. 3-4 

1985 Salpingopore//a mllehlbergii (LORENZ) 
B UCUR, p. 83, PI. 4, Fig. 2 

1989 Salpingopore//a I1lllehlbergii (LORENZ) 
DRAGASTAN, p. 13-14, PI. 5, ?Fig. 5, Fig. 6, 
non Fig. 7 

1989 Salpingopore//a mllelilbergii (LORENZ) 
CONRAD & MASSE, p. 283, PI. 2, Fig. 6 

1993 Salpingopore//a mllelilbergii (LORENZ) 
BUCUR, I. COCIUBA & M. COCIUBA, p. 34-
38, PI. 1, Figs. 9,11-14 

1993 Salpingopore//a mllelilbergii (LORENZ) - SOT­
AK & MISiK, p. 403, PI. 10, Figs . 1-6 

1994 Salpingopore//a mllelilbergii (LORENZ) - BO­
DROGl, BONA & LOBlTZER, p. 233-248, PI. 
1, ?Fig. 8; PI. 16, non Fig. 6; PI. J7, Fig. 8 pars 

This is a frequent ly mentioned and figured species. 
The emended description and the designation of a neo­
type was given by CONRAD ( 1970). In the descrip­
tion s of thi s species the phloiophorous shape of the 
ramifications has been emphasized: tubular in the prox­
imal part and funnel -shaped at the d istal end. The distal 
widenings are supposed 1O be mutually touch ing outside 
the calcareous envelope. The shape of the distal wide­
ning depends, according to CONRAD (1970), on the h 

and w values; specimens with slightly flattened ramifi ­
cations, such as figured by PRATURLON & RADO­
lCIC (1967, Figs. la-d,S), have also been includ ed 
within the specics. 

Thc spec imens studied and illust rated in this paper 
(PI. 1, Figs. 1-1 3), which I consider to be typical reprc­
senlatives of the species, show a comparatively long 
proximal tubular part of the branches, which retains 
about an equal diameter in both longi tudinal and trans­
vcrse sections until it attains the outer surface of the 
calcareou s envelope. The comparatively high h va lue 
between the alternately arranged whorls, as well as the 
small number of ramifica tions per whorls (w), gives the 
appearance of widely spaced ramifications in deeper 
tangential secti ons. The outer (dis tal ) part of the 
branches, where they abruptly widen, is rarely pre­
served, and where present, appears more square-shaped 
than rhombic. The differences with other contempora­
neous Salpillgoporella species in the Barremian depo­
sits appear morc visually pronounced than expected 
from simplified and generalized descriptions and com­
parison of individual dimensional values. Among nUffi-

3 

crous sections obtained from several localities in both 
the Adriatic (including the islands) and Dinaric karst 
regions, two sections have becn studied which suggest 
the existcnce of a branching thallus (PI. I, Fig. 6). 

According to the observations gathered up to thc 
present, the first occurrence of this species in the Adri­
atic and Dinaridic karst regions is exceptionally and 
rarely assoc iated wit h Campanclulla cGpuensis (DE 
CASTRO), the lattcr being generally considered to 
have a strat igraph ic range correspond ing to the Upper 
Hauteri vian and Lower Barremian. In this case too, S. 
muehlberg!i is accompanied by nUlllerous dasyclads 
characteristic of the Barremian algal assemblage. Its 
upper limit (last occurrence) is also the first occurrence 
of Palorbitolina lenticularis (BLUMENBACI-I). Thus, 
taking into account both the position of aU samples with 
this species in the analyzed (logged) sections and the 
assoc iated algal assemblage, the stratigraph ic range of 
S. muchlbergii in the Adriatic and Dinaridic karst 
regions is considered to correspond to the Barremian. 

Salpingoporella melitae RADOICIC, 1967 

PI. II, Figs. 6-13; PI. III, Figs. 1-2; PI. XIX, Fig. 7; PI. 
XX, Figs. 2B, 3B . 

Selected synonymy: 

1967 Salpingoporella ",elitae n.sp.- RADOIC1C, p. 
121 -126, Figs. 1-2, PI. 1-4 

1975 Salpingoporeila melitae RAD01CIC DRA ­
GASTAN, p. 61, PI. 86, Fig. I 

1976 Salpingoporella melitae R ADOICIC - CON­
RAD & PEYBERNES, p. 191, Figs . 13 b, h 

1978 Salpingoporella melitae R ADOICIC - DRA ­
GASTAN, BUCUR & DEMETER, p. 23 -25, PI. 
7, Fig. 3; PI. 12, Figs. 1-2; PI. 14, ?Fig. 3. 

1978 Salpingopore/la melitae RADOICIC - YELIC & 
SOKAC, p. 319, PI. I, Figs. 8-10 

1980 1981 Salpillgoporella melitae RADOICIC -
BUCUR, p. 53-55, PI. 1, Figs. 1-5 

1984 Salpingoporella melitae RADOICIC - LUPE­
RTO-SIN NI & MASSE, p. 34 1, PI. 38, Pigs. 2, 
7-8 

1987 Salpingoporella melitae RADOICIC - RADO­
ICIC, PI. 1, Fig. 6 pars; non Fig. 7, Fig. 8 

1989 Salpingoporella melitae RADOICIC - CON­
RAD & MASSE, p. 283, PI. 2, ?Fig. 12 

1989 Salpingoporella me/itae RADOICIC - DRAGA­
STAN, p. 13, PI. 5, ?Fig. 3, Fig. 4 

1992 Salpingoporella me/itae RADOICIC - MANCI­
NELLI, p. 9, 13, PI. 4, Figs. 2, 12, nOll Fig. 11 

1993 Salpillgoporella melitae RADOICH': - BUCUR, 
I. COCIUBA & M. COCIUBA, p. 35-38, PI. 1, 
Fig. 16 

1994 Salpingoporella melitae RADOICIC - BODRO­
GI, BONA & LOBITZER, p. 233-288, PI. 1, 
Fig. 6, non Fig. 7; PI. 17, Fig. 11 



4 

The suffi ciently detailed original descripti on (RAD­
OleIC , 1967) clearly defines this species as a rather 

large-s ized alga, with a spacious axia l cavity and dense­

ly arranged alternating whorls of ram ifi cations. The 
author emphasized the funnel-shaped pores with a com­

parativel y long tubular proximal part (PI. II , Figs. 7, 
13). The di s ta l part o r the ramificati ons is abruptly 

widened and transversally nattcllcd , i. e ., horizontally 

elongated, produc in g an e longated-rhombic shape (PI. 
fl , Fig. 7; PI. lIl, Fig. I), Deeper tan gen tial sections, 

culting the proximal part or the branches, show widely 

spaced rounded pores in alternating arrangement 

(whorls arc not tightly compressed) , whereas the slml ­
lower tangenti a l section s show the abrup tl y widened 

pores, mutually touc hing on the surface. T ypical, well­
d eve loped , spec im e ns clearly show the ir rhombic 

appearance in shallow tangential sec ti ons cu tting 
through the di stal part or the branches. The corners of 

the more or less e longated rhombs are clear and sharply 

de lineated (PI. II , Fig. 7; PI. Ill, Fig. I), but inwards the 
corners becomc 1110rc blurred and the rims more round­

ed. Because this species incl udes specimens with highly 
va ri able dimensions (D, d), the above mentioned char­

acter isti c features should be used as the primary criteri­

on for a safe and re liable spec ies identification. 

Howe ver, thc dis tal ends of the ramifications ean be 

va rioll s ly shaped and/or de rormed. Bes ide the rather 

typica l sections (PI. III , Fi gs. 1,2), speci mens can be 
found that show a more o r less pronounced dive rgence 

in Ihe distal shape of the pores (PI. 11, Fig. 9), in which 

case thcy appear to re present an inte rm ediate form 
bet wcen S. !lIe fila e and some othcr species or that genus 

(perhaps S. hasi or S. IIr/adanasi). Thu s we are faced 

with the dilemma of how much of divergence should be 

regarded as bei ng w ithi n the range va ri a tion of the 

spec ies and a lte rnati ve ly, which diffe re nces should be 

trea ted , taken togethe r, as sufficie nt fo r the establish­
ment of a new taxon. 

S. /J/c/irae occurs as the most abundant species, both 
s ingly and in assemblages with S. nllleh/bergii, S. 

lJiakol'cllsis, S. ilispanico, S. dinariccl , Pratllr/onello 
dOlli/ovae, AClilloporella podalien, C/ypeina? so/kalli, 

Clypeilla sp. and ot her a lgal and, morc ra rely, fo ra­
miniferal species. According to our observations in the 

Adriatic and Dinari c kars t regions, its firs t occurrence 
co in c ides with the las t occurrence of Ca /J/pane/i/lla 

caplIcnsis DE CASTRO. The laLter species being char­

acte ri s ti c of the Uppcr Hauterivian - Lower Barremian, 
this would the re fore correspond to the Lower Barremi­

an. The las t occurre nce of S. melilae is just be ll ow the 
firs t occurrcnce o f Pa/orbilolina lelllicu/aris (BLU­
MENBACH), as is a lso the case with S. "Il/ ehlbergii. 
Only one sample, derived from Ml. Ve lika Kapela, con­

tain s S. melilae associa ted with P. lel/tiel/laris, which 
re liably defines its Barremian age. 

Geologia Croatica 49! l 

Salpillgoporella hispallica CONRAD & GRABNER , 
1975 

PI. V, Figs. 1-8, part. Figs. 9-11 

Se lected svnonymy: 

1975 Sa/pingoporella hispanica l1. sp.- CONR A D & 
GRABNER, p. 30-39, Figs. 2-6 

1979 Sa/pingoporella hisponica CONRAD & GRAB­
NER - PEYBERNES & CONRAD, p. 946-748, 
PI. 2, Fig. I: ?S. cf. hispanica; PI. 2, Fig. 2 

1980- 1981 So/pil/gopo/'e l/o hispal/ica CONRAD & 
GRABNER - BUCUR, p. 53-54, PI. 2, Figs. 3-4 

1986 Apil/ella hispanico (CONRAD & GRABNER, 
1975) n.eomb. - GRANIER, M ICHAUD & FO­
URCADE, p. 804-806. 110t fig ured 

1986 Apil/el/a jaJJ/'ezo i n. sp. - GRANIER , M ICHA­
UD & FOURCAD E, p. 804-807, Fig. 2, PI. I , 
Figs. a-j; PI. 2 , Figs. a-j 

1991 Sa/pingoporel/a alT. S. hisponica CON RAD & 
GRABNER - KUSS & CON RA D, p. 876 , Figs. 
4, 14 

1992 Salpillgoporella itispallica CONRAD & GRAB­
NER - MANCINELLI , p. 9,12-13, PI. 3, Fi gs. 
1-5 

1993 Sa/pil/goporella hispallica CONRAD & GRAB­
NER- BODROGI , CONRAD & LOI3ITZER , p. 
64-66, PI. 2, rigs. 7- 10 

1994 Salpillgoporello itispanica CONRAD & GRAB­
NER - BODROGI , B6NA & LOBlTZER , PI. 6 , 
Figs. 1-2 

In their original description, CONRAD & GRABN­
ER ( 1975) emphasized the sma il number (w=4) of 
short , shallow calc ified ramifications, arranged in alle r­

nating consecllti ve whorls, with a calcareous ske le ton 
consis ti ng of thill-wa ll ed , hyaline, calcite. The recon­
st ruct ion (CONRAD & GRABNER , 1975, Fi g. 7) 
shows the connecting pores be tween the ramifi ca tions 
and the axia l cavity to be s ituated in thc middl e of the 
proximal part orlhe ramifications, which means that the 
ramifications regularly swell in both upward and down­
ward directions and have a short and thinl y calcifie d 
partit ion be tween each and every ra mifi cation and a 
s li ght te rminal bulbou s swc lling. This has le d the 
au thors to interpret Ihe di stal ends or the ramifi cation s 
in shallow tangential sec tions as square-shaped, which 
howe ver should be regarded as thc personal inte rpreta ­
tion. Moreover, it was just Ihis differencc that was used 
by GRANIER et al. ( 1986) as the basic dille renee for 
es tabli shing their ncw gen us a nd species Apille//a jaf­
Jrezoi GRA lIER et a1. , in whic h the connect ing pore is 
shifted downward and the ramification itself is ve rtica l­
ly flatt ened at its distal end and has a hexagonal co n­
tour. 

The spec imens from the Barremian of Cres Is land 
that are illus trated in the present pape r (PI. Y, Figs. 1-8) 
ancl labe lled Salpillgopore/la Itispallico do in fa c t, by 



Sobc: Taxonomic Revicw of Somc 13arrernian and Aplian Calcareous Algae ... 

their key characters, stro ngly diverge from CONRAD 
& GRABNER's (1975, Fig. 7) reconstruction. Instead, 
they show the connecting pore shifted downward (PI. 
Y, Figs. 3, 6, 8, 11) and a hexagonal contour of the dis ­
tal pore openings (PI. Y, Fig. 2), those being the charac­
teristics that make them identical to the specimens 
described as Apil/clla jaffrczoi. This, however, shows 
that recons tructions, no matter how nice they may 
appear (and how welcome, in certain cases, they really 
are), represent nothing more than the authors' interpre­
tation and thus may be misleading in certain cases. 
Some minima l dilTerences between the Kimmeridgian 
specimens from Mexico and our Barremian speci mens 
do exist; firstly the extreme dimension values, though 
these values, for the most part of their ranges, do over­
lap; secondly, there is a minor differencc in the widen­
ing or the axia l cavity between the branch-bearing 
whorls, which is more pronounced in the Kimmeridgian 
specimens (GRANIER et aI., 1986, PI. I, Fig. a; PI. 2, 
Fig. b) than in some of the Barremian specimens (PI. Y, 
Fig. 3). However, this fea ture is not a regular character­
istic: in some specimens it is missing and therefore can­
not be accepted as a valid criterion for species-specific 
differentiation. On the contrary , the above mcntioned 
list of identical features seems to confirm the identity 
bet ween the Kimmeridgian specimens from Mexico 
and the Barremian specimens of the Mediterranean 
realm , indicating that Apinella jaJfrezoi shou ld be con­
sidered a younger synonym of Sa/pillgoporclla lIispani­
ca . Herewith, the latter species acquires an extended 
stratigraphic range. 

An summary, the establishment of the genus Apillel­
la , based on the reduced degree of calcification and the 
numbe r and shape of thc ramifications - bei ng broadly 
elongated above a short and eccentrically situated con­
necting pore in Apine//a - does not seem justified for 
the following reasons: 

1) The apparently reduced degree of calcifica tion can 
directly depend on the more pronounced swelling of 
the ramifications, partieu[arly if that swelling occurs 
abruptly from the very base; 

2) The superficial calcification may only be an artefact 
of preservation and a consequence of the external 
destruc tion of the thin partitions between the ramifi­
cations (visible in the illustrations of GRANIER et 
ai., 1986, where the partitions between the ramifica­
tions sometimes appear longer and sometimes 
almost completely reduced, i.e. destroyed by sec­

ondary processes); 

3) The reduced number of ramifications can be a direct 
consequence of their shape and mode of wideni ng, 
whether from their very base or more distally, and 
of the degree of transverse compression (horizontal 
f'lattening). There arc a number of Sa/piflgoporclla 
species with a varying number of ramificat ions, e.g., 
5-11 in S. fIIuchlbergii, 8-12 in S. af/nulata, 5-7 in S. 
katzcn', 5-7 in S. lIasi, ctc. Tn general, those Salpin ­
goporella species which have transversally com-

5 

pressed (= horizonta lly Ilattened) ramifications do 
also have a smaller number of ramifications. There­
fore, it follows that the delimiting number of ramifi­
cations for the genera Apinclla and Salpingoporella 
would be 4 and 5, respectively. 

4) The shape of the ramifications - broadly elongated 
above the short , narrow, and eccentr ically si tu ated 
connecting pore - is only more strongly pronounced 
in S . hispanica, while also present - though more 
slight ly developed - in S . katzeri. This latter species, 
however, was not included in the genus Apinella. 
Also, in the original description of S. katzeri there is 
no mention of the branches being strongly swollen 
upward and the connecting pore remaining at the 
level of the lower ramification rim (PI. X, Figs. 10-
12). The eccent rically s ituated connecti ng pore also 
occurs in the genus Trip/oporella and accordi ng to 

my personal observations is also present in Palaeo­
dasycladus medilerranclls PIA, which, however, 
does not question the generic and species-specific 
attribution of the mentioned forms. Instead , it is an 
indication of the variation within a genus and, even, 
within a species. 

An conclusion, it may be said that the lack of char­
ac terist ic featu res that would unequivocally differenti­
ate Apinclla from Sa/pingoporclla indicates that Apirlel­
la should be cons idered to be a younger synonym of 
Sa/pingopore//a, as has already been clearly stated by 
BODROGI et a l. (1993). The large geographic distance 
between Mexican and European fin ds, as well as the 
cons iderable difference in their stratigraphic positlon 
(Mexico: Kimmcridgian; Europe: Barremian), do not 
alone seem to justify the establishment of a new taxon 
(there is also the possibility of migrations , common 
ancestor, etc.). 

Stratigraphic position: In spite of the largc number 
of samples collected from Jurassic and Cretaceous beds 
from all over the Dinaric karst area in Croatia, the finds 
of this species remain rarc. It has been unequivocally 
idenlified at only two localions: in the Barremian a lgal 
assembJage on the Mljet Island and in Barremian depo­
sits of the Cres Island, where it is associated with S. 
hasi (spec im ens illustrated in PI. V, rigs. I-II). As a 
resuit of it s rare occurrence, the definitive establ ish­
ment of its stratigraphic range remains open, though the 
two hitherto known localities both are of Barremian 

age. 

SalpillgofJorella verrllcosa n.sp. 

PI. VI, Figs. 1-15 

Origin of the name: The species has been named 
aIter the wart - like protrusions on the outer surface of 
the calcareous envclope which mark the terminations of 
the distal ends of the ramifications (Latin verruca = 

wart). 
Type locality: Island of Mljet, road-cut on the road 

Sobra Babino Polj e, a bou l 2 km from Sobra 
(17'36'OO"E and 42°44'02"N, Fig. I ). 



6 

'\:2) 
o 

rig. [ Topographic map of a pari of the Mljcl Island. The algal­
bearing localities arc marked as follows: A) Clypeillo gi,~mll('a 
n.sp., S(lfpingo{Jort'!{(l po/ygollofis 11.Sp.: H) Sa/pillgoporella 
1'('I"I"II('osa 11.51'., Piriferella ,lpil1osa ll.gcn., n.sp. 

Type stratum: Well-bedded , light coloured, biotur­
bated algal wt\c.:kcstoncs with casts and moulds of green 

algae and gastropods. The environment of sedimenta­

tion is defined as within peritidal zone, wit h sporadic 

cmcrsions into the vadose zone. 

lIo!otype: Slightly oblique, a lmost longitudinal, 

sect ion rigured in PI. VI, Fig. 4, slide Mg-81/8. All 
slides containing the material or the new species arc 

stored at the Institute of Geology. Zagreb. 

Dimensions in mm: 

Maximum observed length (L) 

Outer thallus diameter (D) 

Inner thallus diameter (d) 

The diD relationship 

Distance between whorls (b) 

Length of the branches (I) 

Maximum ramification diameter (I') 

Number of branches pcr whorl (w) 

Inclinalion of branches «x) 

total range 

2.04 
0.38-0.72 
0.20-0.32 

34-59% 
0.14·0.24 
0 .1 2-0.20 
0.06-0.10 

6-8 

eca 15° 

Geologia Cro:lIica 49/1 

~ __ ~====~2 ____ ~3km 

Diagnosis: Sa/pillgoporel/a w ith rare, alternating, 

phloiophorolls ramirications with bulging (convex) out­

er surface. covered wi th a thin calcareolls sheet. 

Description: The cylindrical calcareous envelope is 

composed llP of coarse-grained, blocky, light yellowish 

calcite. The outer surface is covered by alternating 

wart-like bulges, which represent the distal ends of the 

ramifications. The inner surface of the axial cavity is 

s tra ight and sharply delineated. T he axial cavity most 

most frequently 

0.44-0.64 
0.24·0.30 
40-50% 

Table 1 Dimensions of Salpillgopo/"ef(a 

l'e/TIlCosa n.sp. 



Sokac: Taxonomic Review of Some Barremian and Aptian Calcareous Algae ... 

frequently comprises 40-50% of the total diameter, 
depending on whether it is measured at the level of the 
bulges or in between, where the outer surface appears 
slightly depressed. 

Simple, undivided, phloiophorous branches commu­
nicate with the axial cavity by means of short and tiny 
connecti ng pores (channels) and broaden slightly and 
continuously from their very base toward their distal 
ends. The branches reach the surface and end in wart­
likc protrusions that are covered by thin calcareous 
membranes. The calcareous envelope appears to be 
attenuated outward, which along with the ramification, 
produces a star-shaped transverse section at the level of 
whorl s (PI. VI, Fig. 14). The ramifications are directed 
slightly upward, at an angle of about 15°. 

Similarities and differences: Salpingoporella ver­
rucosa belongs to the group of Salpingoporella species, 
in which the ramifications are neither 10ngitudinaUy nor 
transversally compressed, giving in tangent ial sections 
a rounded pore regardless of where it is cut (i.e., in both 
deep and shallow tangential sections). It differs from 
other species of this group by the coarse-grained mosa­
ic calci te of the calcareous envelope, a very small num­
ber or ramifications per whorl, and a comparatively 
large distance between the whorls, the latter features 
giving the impress ion of pores being situated few and 
far between on the outer surface. The end in gs of the 
pores on the outer surface make bulging, wart likc pro­
trusions, which give this alga its characteristic appear­
ance. 

Stratigra phic position: Salpingoporelfa verrucosa 
n.sp. occurs in an interval of well-bedded fossiliferous 
mudstones and fossiliferous peloid and oncoidal wacke­
stones with several intercalations of intraformational, 
marl-cemented breccia which test ifies to repeated shal­
lowing and local emers ion (Fig. 2). That interval com­
prises about 30-40 m or stra ta beneath the algal-bearing 
leve l and about 30 m above it, and corresponds, as a 
whole, to depos ition in restricted shoals with frcqucnt 
oscillations and excursions into very shallow subtidal 
and intertidal zones. These strala contain Saipingo­
parella mueh/bergii (LORENZ), S. aff. eemi RADO~ 
rcrc, S. melitae RAD01C1C, S. hispauica CONRAD & 
GRABNER, Sa/pingoporella dil/ariea RADOlCIC, 
Salpingopore/la sp. ex gr. genevensis-hasi, AcrojJorella 
radoic/cae PRATURLON, Pralurlol/ella dani/ovae 
(RADOICIC), P. ncrae n.sp., Piriferella spinosa n.sp., 
and AClil/oporc/la podolica (AL TI-T), most of which 
occur in the same sample with S. vent/cosa n.sp. Based 
on the above mentioned alga l assemblage, and the 
supcrposilional location of the sample in the higher part 
of the column , which has well-defined boundaries (Fig. 
2), the age of the S. vcrr/lcosa bearing beds at the type 
locality may be establ ished as Upper Barremian. 

Lithologically, the entire Barremian sequence is 
characterized by tex tural and structural features which 
arc. according to T1SLJAR (1986), typical of rhythm ic 
sedimentation ranging from gradua l shallowing to 

g~ VAvl 

~ 
~ 
B 

-- Mlj- 81 

- Mlj-78 

Mlj -73 
Mlj-71 

- Mlj-65 

I 
ZOm 

10m 

Om 

7 

2 

3 

• 

Fig. 2 Detailed lithofacies co l LImn of the type- locality of C/ypeillo 
gigallfea n.sp., So/pillgoporef{o po/ygollotis n.sp., Sa/pillgoporef­
fa ve/"mcosa n.sp. and Pirifeu:!la spillosa n.gcn., n.sp. Legend 
(facies types after TISLJAR & VELIe, 1991): I ) f-acies of 
suprat ida l and/or temporary emersions: soft pebble conglomer­
ates and cll1crsion breccias - facics type K-l; 2) Facies of inter­
tidal 10 shallow subtidal limestones: LLH st!"Omatoliles, fenestral 
mudslones to pellelal wackestones - facies type K-2: 3) Facies of 
lagoons and restricted shoals : mudstone/wackestone and onko­
lites - facies type K-3 : 4) Facies of late diagenetic dolomites -
fac ies type K-5: Mlj -63 - Mlj -83: sample designation. 

restricted shoal or from the subtidal to the lower inter­
tidal and the vadose zone, respectively. In vertical 
sequence, th is is manifested by the transi tion from a 
homogeneous micrite/pelmicritc into fenestral micrite/ 
pelmicrite with eroded upper bedding planes and/or 
vadose features, and sometimes even tidal channels. 



, 
Sa/pillgoporella heraldica n.sp. 

PI. VTI, Figs. 1- 18 

Origin of the name: In tangentia l sections, th is alga 
shows an alte rnation of dark and light squares, reminis­
cent of the Croatian coal of arms. 

T ype locality: Island of Korcuia , road-cut loward 
Prizba, about 50 m from the cross-road in Bma Village 
( 16'5 1 '30"E and 42°54 '29"N, Fig. 3). 

Type stratum: Well -bedded brown al gal -pe lo idal 
g ra in stone. The sample was collected during a fi e ld 
reconnaissance campaign and had been taken at ran­
dom. T herefore no detailed lithological analysis can be 
given. However, acco rding to the gene ral geologica l 
his to ry o f the a rea, the age can be in fe rred as be ing 
transiti ona l Barrem ian-Aptian, and the environment as 
a peri tida l-tidal bar. 

Holot ypc: T angel11ial section figured in PI. VII , Fig. 
5, slide Br-3 A/28. All slides containing the new species 
afC kept at the Institute of Geology, Zagreb. 

Diagnosis: A Sa lpillgoporella with large ax ial cav i­
ty and fu nnel-shaped , alternatin g ramifi ca tions, whi ch 

o 
o 

Fig.3 Topographic Illap o ra part or the Korcula tsland . Locality with 
SalfJillgofJordla heraldica n.sp. and Pmrur!onella lIeme (DRA ­
GA$TAN CI al.. t978) is marked with aITOW. 

Geologia Croalica 49/1 

in tangential sec tions appear square-shaped , producing 
the characteristic chequered appearance. 

Description: A comparat ively small d asyc lad alga 
Wilh a cyl indrical, sometimes gently c Ulved Lhall us. The 
calcarcous cnve lope, similar to many othe r spec ies of 
this genus, is composed up of ye llow ish, crys talline cal­
ciLe. The axial cavity is borde red by a smooth surface, 
pe rforated by comparati vely large pores at the begin­
ning of the ramifi cations . The oul e r surface also 
appears to be smooth, but the oute r termina tions of the 
ramifications arc possibly slightly convex . 

The ram if ications are simpl e , undivided , arranged 
into regular alternating whorls. They are orientated per­
pendicul ar to the axial cavilY and appear sho rt , which 
may be a consequence of a destroyed oute r surfacc. The 
ram ificatio ns s lart with a compara tively w ide pore in 
re lation to Ihe LOLaI size - and widen regularly and uni­
fo rmly toward the dis tal e nd (PI. VII , Figs. 16- 18), 
re ta ining at rhe same time a square contour, regardless 
of the depth of thc tangent ial section (PI. VII, Figs. 3, 5-
6, 10). The alternation in shall ow tangent ia l sections of 
approxi mate ly equal dark squares o r the pores and light , 
calci tic squares of the ske leton produces the charac tc r-

Korc~ 
C> O ~ 

~O ____ -===== ____ -"3km 



Sokai:: Taxonomic Review of Some Barremian and Apfian Calcareolls Algae ... 

Dimensions in mm: 

Outcr diameter (D) 

Inncr diameter (d) 

TIle dID relationship (di D) 

Distance between consecut ive whorl s (h) 

Lcngth of rami f ica tions (I) 

Diametcr of ramificat ions (p) 

Number of ramifications per whorl (w) 

total range 

0.35-0.68 
0.25-0.39 
50-70% 

0.04-0.05 
0.09-0.1 2 

0.05-0.09 
11-14 

iSlic chessboard palte rn (or the Croatian coat of amls). 
This appearance is s trengthened by the fac t that the 
da rk squares of the porcs a lmost touch onc anothe r at 

thei r corncrs, which emphasizes both the diagonal and 
the quadrangul ar pattern (PI. Vll , ri gs. 3-5). This fact 
was the reason for choosing a tangent ial section as the 
holotype. 

Similarities and differences: Salpingoporella 
heraldicci n.sp. is clearly distinguished from other Salp­
ingoporella species, i.c. from those in which the distal 
parts of the ramificat ions a rc e ithe r longitudinally or 
transversely compressed. If the dis tal ends of the pores 
arc compressed, they produce a hexago nal , rhombic, 
rec tangular, or more rare ly, quadrangul ar (square) pat ­
tern on thc outer surface. In those species in which the 
pores in cortica l secti ons appear square-shaped , the 
pores in deeper tangential sections appear more or less 
rounded , as d is tinc t fro m S. heraldica, in which the 

pores re tai n the ir square shape regardless o f the 
whethe r we arc deal ing wi th shallower or deepe r tan­
gential sections. By its general dimensions, S. heraldica 
belongs to the grou p of rathe r small Sa lpingoporcllae. 
By its s ize (dimensions), the di D re lat ionship , and the 
num ber of branches per whorl , S. heraldica comes quite 
c lose, or even overlaps, with the corresponding values 
f"o r S. dina rica. However, it clearly diffe rs from the lat­
ter by the coarse-grained calci te in the calcareous enve­
lope, square po re open ing in the tan gcnti al sec tion s, 
pronounced diagonal paltern in pore arrangement, etc. 
Other species from the sa me group, i.e. S. is/riana, S. 
johnsoni, S. selli i e tc., arc charac te ri zed by phloio­
phorous ramifications Ihal do not show e ithe r longitudi­
na l o r transverse compression. The diffe rences with 
regard to these species concern the d imensions, number 
and shape of branches, e tc. (see Table 3). 

Concerning the question of how far the dimensional 
va lues (measured e lements) and thei r proportions 
should be taken int o conside ration as signifi cant crite­
ria , the original data shou ld be considered first, provid­
ed the spec ies is recogni zed as va lid. Late r data often 
relate to incorrec t identifica tions and are often addi tion­
al ly biased because of the small num ber of measured 
sections (whose spec ific, and sometimes even generic, 
allribution is frequent ly problematical), which renders 
any statistical inte rpre tation questionable. 

Stratigraphic position: Salpingopore/la heraldica 
n.sp. has been found in a random sample , which makes 

most frequently 

0.54-0.64 
0.34-0.39 

50-60% 
0.05 

9 

Tab le 2 Dimensions of Salf1ill~{)fJo/"(!lla 
heraldica n. sp . 

a more precise definiti on o f its s trati graphi c pos ition 
and environmental sell ing imposs ible. The sampl e in 
qucst ion shows some characte ri sti c features which incli ­
cate a perit ida l-t idal bar e nvironillent , as s imilar fea­
tures occur in samples collected from logged sections, 
in the same or st rati g raphica ll y close hori zons, in the 
study area. The same sample a lso contains the follow­
ing mi c rofoss ils: Triplopore //a bacilli/onl/is SOKAC 
(the same sample is also the " type specimen"), 1'. mar­
sicollo PRATURLON, T. IIrogieliiformis CON RAD & 
PEYB ERNES, and Praturlol1ella I/erae n. sp. The 
occurrence of rare Palorbitolina lenticularis (BLU­
MENBACH) in the immedi ately overlying deposi ts 
indicates a terminal Barremian or, possibly, basa l Apt­
ian age. 

Salpillgoporella polygollalis n.sp. 

PI. III , Figs. 3- 10; PI. IV, Figs. 1-17 

1994 Sa/pingoporel/a nr/adanosi CONRAD, PEY­
BERNES & R ADOICIC - BODROGL B6NA 
& LOBITZER, PI. 17, Fig. 4. 

Orj~in of the name: The name of the species is 
der ived from the irregular, po lygona l pores as they 
appear in shallow tangential sections. 

Type locality: Island of Mljcl, road-clit al the Sabra 
- Babino Polje road, aboul 2. 15 kill before Sabra, 150 III 
away from the locality of S. I'errllcosa (17°36'00"E and 
42°44'02"N, Fig. I). 

Type slralum: Well-bedded, lighl-coloured , biotur­
bated alga l wackestones with casts and moulds of dasy­
c lad al gae and gas tropods (Fig. 2) . Bioturbations arc 
fill ed wit h pell o ida l grai ns tone and small benthi c 
fo raminife ra. The sedi menta ry e nvironment may be 
defined as shallow subtida l. 

Hololype: Obliquc seclion figured in PI. III , Fi g. 3, 
slide Mlj.- 63/68. All slides 0 1' thi s specics arc kepI al 
Ihe Institule of Geology, Zagreb. In 101al, about 25 dif­
ferently oriented sections have been examined. 

Dia~nosis: A Salpingoporella wi th large, a lte rna t­
ing. ramifications, that are funne l-shaped from the vely 
base, di sta lly thickened and hori zontall y flatt ened, 
which show an irregular, rhombic-hexagonal to c llipti ­
cally polygonal paltern , with an indication 0 1" di s tal 
splillin g. The calcareo us wall is composed o/" ve ry 
coarse-grain ed, heavily weathered and recrysta lli zed 
calcite. 



0; 

SPECIES 0 d dID h p P pro~;on.1 P ,li,!al a w 

SalpingoporeUa adriatica (GUS IC, 1966) 1.36-2.62 0.26-0.50 0.30-0.35 0.50-2.20 0.20-0.40 20-22 

S. a/J/llI!ata CAROZZl, 1953 0.30-0.64 0.10-0.25 0.15-0.20 8-20 

S.ambica ELLIOT, 1968 0.31-0.73 0.21 -0.47 55-66% 8-10 

S. arllmael/sis OKLA, 1995 2.1~2.8 1.6- 1.8 0.20-0.5 0.30-0.5 0.20-0.27 20 

S.? atlantica (JOHNSON, 1968) 0.71 0.36-0.375 0.050-0.075 0.025-0.017 0.042-0.055 45 0 

S. biokoviellsis SOKAC & VELIe, 1979 0.55 -0.74 0.12-0.22 0.20-0.34 
• 0.44-0.74 0.10-0.25 16-476% 0.18-0.32 0.22-0.25 0.10-0.32 7-R 

S. bl/cllri DRAGASTAN, 1989 0.95-1.30 0.25 0.050 0.20 0.10 to-20" 14-16 

S. cemi (RADO ](:: IC , 1968) 1.8-1.6 0.44-0.52 0.38-0.48 0.16-0.24 20-25 

S. circcassa FARINACC I & RADO IC [C, 1991 0.165-0.38 0.06-0.14 0.20-0.5 0.30-0.5 0.20-0.27 20 

S. croatica SOKAC, 1992 0.34-0.92 0.14-0.38 0. 12-0.16 0.08 -0.12 8-14 
frequently 0.42-0.52 0.24-0.28 0.12-0.14 0.08-0.10 12 

S. dinorica RADO ICIC, 1959 0.24-0.56 0.128-0.40 0.048 8- 10(7) , 
0.07 -0.48 0.05-0.38 4 1-47% O.oJ-0.06 0.04 0.08-0.12 7-12 

S. enayi BERN IER, 1984 0.867-0.950 0.325 -0.425 38-44% 0.25 -0.275 0.10-0. 12 30? 

S. etalfolli BERNIER, 1984 0.325 -0.450 0. 1-0.137 28-30% 0 .125-0. 150 0.05-0.062 70-800 20? 

S.? cxilis (DRAGASTAN, 1971) 0.6-0.8 0.2-0.3 0.025 -0.030 

S.jlucgcli DRAGASTAN, 1978 0.54-1.20 0.25-0.56 0.080-0.25 0.025 -0.030 0.090-0.15 10- 12 

=- Mcgaporella pucgcli (DRAGAST AN) 1989 0.60- 1. 10 0.30-0.56 ? 0.025-0.030 0.10-0.15 12 

S. gencvcl1sis (CONRAD, 1969) 0.6-1.0 0.27-05 I 45 -51 % 0.09-0.11 6? 7-9 

S. grudii (RADO ICIC, 1963) 0 .128-0.48 0.064-0.240 0 .032-0.1 20 12-20 

S. hasi CONRAD et aI., 1976 0.2- 1.12 0.1 -0.72 0 .06-0. 13 5-7 , 
0.34-0 .82 0.20-0.54 4 1-77% 0.04-0.08 0.12-0.14 0.20-0.24 5-7 

S. heraldica n.sp. 0.35-0.68 0.25 -0.39 50-70% 0.04-0.05 0.09-0.12 0 .05-0.09 11 - 14 co 
n 
0 
C S. hispa/Jica CONRAD & GRABNER, 1975 0.25 -0.46 0.1 6-0.36 66-77% 0. 15-0.26 4 n~ , 

0.24-0.44 0.17 -0.30 64-84% 0.12-0.14 0 .14-0.28 4-5 n a 
S. humilis (BYSTR ICKY, 1967) 0.633-1. 183 0.283-0.750 42-68% 

~ 

0.033 -0.050 0.033-0.058 7-12 g 
~ 

'" 



~ 

S. istriano (GUS IC, 1966) 0.25 -0.74 0.14-0.41 0.03 -0_04 0_06-0.105 0.012-0.05 16-28 ~ 
S.jolmsolli(DRAGASTAN, [971) 0_20-0.31 0.10-0.15 0_033-0_045 0 .05-0_07 0.02-0.04 14-20 

;:;' 
0 

Q 
S. km::eri CONRAD & RADO ICI(::, 1978 0_26-0_80 0.10-0.40 0.08-0_16 5-7 

, 
o· 

" S. me/flae RADOle,,:, 1967 
0 

0.48-1.255 0.32-0.960 161-24 < 
('i" 

* 0_64-1.68 0.36-1.14 50-68% 0.05-0.08 0.17 -0.34 0.19-0.28 14-20 
, ., 

S.lIIilOl'GnOl'iri RADOIC1C, 1978 
~ 

0.070-0.240 0.035-0.1 12 0.040-0 .075 9·15 9 , 
0 

5· 11 " S. lIIuehlhergii (LORENZ, 1902) 0.2 -0.85 0.1-0.3 0.07 -0.18 
~ , 

0.28-0.86 0.10-0.54 26-62% 0.08-0.1 3 0.10-0.20 0.10-0. 18 7-8 
::;" 
0 

S.pirilliac KARAS & RAD01CIC. 1991 ?0.180 0.064-0.160 0.064-0.090 10-1 4 " 1l 
0. 240-0.530 > 

frequen tl y 0.350-0.400 " ;:;-, 
S. potn/fills; BueUR , 1985 1.10-2.300.11-0.28 10-24% 0.22-0.28 0.56-1.00 0.040-0_078 0_16-0.31 25-35° 7-9 Q 

IT 
" S. polygollalis n.sp. 0.82 -1 .48 0.35-0.74 44-55% 0_14-0.19 0.22-0.35 0_06-0_09 0. 14-0.34 10-12 ~ 

S. po/saki SOKAC & JELASKA, 1991 
> 

0.48-0.72 0.12-0.24 25-30% 0_08-0_11 0.20-0.28 0. 12-0.24 20° 9-12 rf 
0-

S. pratllriOlli (DRAGASTA:-.J, 1971) 1.2-4.0 0.6-1.6 0.25 -0.90 0.07 -0.030 20-40 

S. pygmaea (PI A, 1925) 0.43-0_72 0_17-0.26 0.02 0_09 16-19 

S. rob/ISla SOKAC, 1993 0.77 - 1.44 0.20-0.48 27·30% 0.12-0.24 0.28-0.48 0.15-0.28 8- 11 

frequently 0_86-1.1 0 0.24-0.30 0. 15-0.20 0.30-0.40 0.20-0.24 8-9 

S. se//ii (CRESCENT!, [959) 0.24-0.44 0.06-0.16 0.018 0.016-0.02 25-27 

S. steil/hauseri CONRAD e t a [. , [973 0.26-0.33 35-44% 0.08-0_1 1 45° 7-8 

S. sturi (BYSTRICKY, 1967) 1.611-3.444 0.888-2.222 50-67& 2.27-4.49 0.055 -0 . 166 167-32? 

S . tosaensis (YABE & TOYAMA, 1949) 0.71 0 .41 0 .05 0 .025 50 

S. wrgida (RADOI C ](:, 1965) 0 .57-2.1 0.22-0.72 0.064-0_ 13 0.19-0.64 0_032-0_080 0_09-0_19 12-24 

S. IIrladal1axi CONRAD ct. <11., 1977 0.65 -1.1 4 0_17-0.62 33-73% 0_015-0_13 4-8 

* 0.20-0.72 0_10-0.38 30-60% 0_08-0_10 0_11 -0.28 0.14-0.24 5-76 

S. /lbaiydhi RADO IC U:, 1979 0.24-0.65 0.08-0.32 0_050-0_096 0.Q40-0.090 8-14 

S. verticil/ala (SOKAC & NIK LER, 1973) 0.45 - 1. 18 0_2 1-0.56 0.04-0.105 0.15-0.44 0_02-0.05 0.05-0_12 22-30 

S. )'c/T/lCO.m n.sp. 0.38-0.72 0.20-0.32 34-59% 0.14-0.24 0.12-0.20 0.06-0_10 6-8 

Table 3 Comparison of the dimens ion s of genus Sa/pil/goport:!la species. * - Complemented by own data. 



12 Geologia Croatica 49/1 

Dimensions in mm: 

Outer di,unctcr (D) 

Inner diameter (d) 

The dID relationship 

Distance between whorls (h) 

Length of the ramifications (I) 

Proximal diameter of the ramifications (Ppm) 

Distal diameter of the ramifications (Pt/is) 

Number of ramificLltiotls per whorl (w) 

0.82- t ,48 
0 .35-0.74 

44-55% 

0.14-0.19 
0.22-0.35 

0.06-0.09 

O. t4-0.34 

10- 12 Table 4 Dimensions of Safpillgoporella polygonalis 
n.sp. 

Description: The massive calcareous skeleton of 
the cylindrical thallus of the new species is composed 
up of light grey, very coarse-grained, and variously 
rec rystallized calcite. The outer surface of the calcare­
OliS envelope is always damaged, partly destroyed or 
dissolved, and subsequently recrystallized. That the 
wall was originally comparatively th ick is indicated by 
rare ly preserved, outwardly projecting, thin partitions 
between the distal ends of the ramifications (PI. lIT, Fig. 
3; PI. IV, Figs. 2, II, 13, 16-17). Axial cavity - mea­
sured in the few most complete specimens - occupies 
only about 30-35% of the 10la l diameter, while more 
often il appears to be about 50% due to the secondary 
destruction of the wall. It is delimited by a smooth 
plane, which in section appears as a clear and sharp reg­
ular line (PI. Ill, Figs. 6, 9). In some sect ions, however, 
the inne r surface may be more or less destroyed, too 
(PI. HI, r:igs. 8, 10), as indicated by the appearance of 
the larger pores of the ramifications. 

Simple large (massive) ramifications arc arranged 
alternately in the consecutive whorls. The general shape 
of the branches is irregular and variable. Stal1ing from a 
smaller or larger connecting pore, they slightly widen in 
the firs t half of their length, after which they abruptly 
swell (PI. III, Figs. 3, 6; PI. IV, Figs. 6, 10), producing a 
funne l- or bowl-shaped distal part (PI. IV, Figs. 14-16). 
Tn some specimens, the distal swelling is equally well 
vi s ible in both longitudinal and equatorial sect ions. 
However, more often the ramifications afe more or less 
flattened and elongated horizontally (i .e., by vertical 
compression; PI. III , Figs. 4, 7, 10), which gives the 
characteristic irregular and variable pattern in shallow 
tangential sections. The shape of the distal opening of 
the pores varies from approximately hexagonal-rhom ­
bic (Pl.lIl , Fig. 3) to elliptical-oval (PI. TIT, Figs. 4, JO). 
In the latter case, the slight tendency towards division 
of the stretched parts of the primary ramificat ions may 
be noticeable. The question of whether the ramifica­
tions actually divide distally, as suggested by some sec­
tions (PI. TIT, Fig. 4; PI. IV, Figs. 3, 5) , must remain 
open, for the time being. At their very end the branches 
are closed by a dome-shaped, thin, probably calcareous 
membrane, as suggested in a few sections (PI. IV , Fig. 
16) in which thin , mutually separated partitions of the 
adjacent ramifications are prescrvcd, which probably do 
not touch at the ends. The branches arc perpendicular or 

subperpendicular to the main stem, in some specimens 
being slightly directed upwards. 

The value of D is probably somewhat larger, 
because the outer surface was partly destroyed in all the 
sect ions observed. 

In some sections (P I. III, Figs. 4, 7) small hem i­
spherical recesses in the distal parts of the ramifi ca­
tions' calcareous envelope can be seen. These arc simi­
lar to, but less well pronounced then those observed in 
S. genevensis in which this feature was supposed to be 
caused by the imprint of slightly calcified cysts. In S. 
po/ygollalis , however, this supposition can neither be 
confirmed nor definitely disproved, though the imprint 
of slightly calcified cysts into a calcareous wall seem, 
in this casc at least, unlikely. Unequivocal remains of 
even slightly calcified cysts have not been recognized, 
however. 

Similarities and differences: Sa/pillgoporel/a 
fJolygonalis n.sp. can be, in general , ascribed to that 
group of the Sa/pillgoporella species in which the rami­
fications at their distal ends are horizontally flattened, 
regardless of what shape they eventually assume -
hexagonal, rhombic, quadrangular, or, as in this case, 
polygonal. The new spec ies is most similar to S. 
melitae, which, indeed, does show a broad variation 
range, from typical forms (PI. IT , Figs. 6- 13; PI. Ill, 
Figs. 1-2) to the such forms that, as the tangential sec­
lions cutting through their calcareous walls become 
progressively deeper, gradually begin to show features 
intermediate to some other species, e.g. S. hasi or S. 
gel/evensis. Therefore the new species appears most 
similar to the above mentioned species in the deepest 
tangent ial sections (i.e. those cutting the calcareous 
envelope in its most internal part) , where the pores of 
all the species concerned appear similar, regarding both 
their arrangement and density. Going outwards, the dif­
fere nces between those species and S. polygonalis 
become progressively more and more pronounced, 
being most clearly expressed in cort ical (i.e., shallow­
est) tangential sections cu lling through the extreme dis ­
tal pa rt of the ramifications, which in S. hasi and S. 
gellevensis have a quadrangular, in S. urladanasi a 
more or less deformed quadrangular shape (PI. IX, Figs. 
2, 4-5; PI. X, Figs. 3-4, 6), and in S. biokovellsis a 
square shape, whereas in S. melitae the pores appear 
clearly rhombic and horizontally (transversally) rIat-



Sokac: Taxonomic Review of Some Barremian :lJ1d Aptian Calcareous Algae ... 

tened. In S. po/ygoJ/afis, however, the entrance pores of 
thc branches have a larger diameter than in S. rneLifae 
and the branches irregularly widen di stally, which 
makes them somewhat similar to those in S. genevellsis, 
but with differently shaped distal ends. In S. hasi the 
entrance pore is followed by a very short channel, 
which is, in turn, foll owed by a bowl-shaped widening, 
whereas in S. polygollaIis n.sp . the swe lling of the 
branches is irregular, sometimcs even more pronounced 
(but still uneven) in the distal half of their length. Salpi­
IIgoporel/a polygonalis clearly differs from S. katzeri, 
in which the bra nches also swell very rapidly, often 
being more swollen toward the upper side (PI. X, Figs. 
10-1 2), with thin partitions between the branches and 
the more or less pronounced hexagonal to rhombic pat­
tern al the d istal ends. In S. hispanica, the skeleton is 
much weakly calcified, the number of branches per 
whorl is smaller, and the branches also have a more or 
less pronounced rhombic or hexagonal shape at their 
ends. Tn S. dillarica, the fibrous calcite, consi derably 
smaller dimensions, the characte ristic arrangement of 
tiny pores in proximal (deeper) tangential sections, and 
rhombic or completely flattened pores at their distal 
ends make clear differences to S. po/ygonalis. The dif­
ferences to numerous other species of this genus, in 
which thc distal ends of the pores are compressed, i. e. 
flattened vertica ll y, or the branches widen regularly 
toward thcir distal ends, giving the marc or less round­
ed pores at the outer surface, arc clear and need not be 
discussed at length . 

Stratigraphic position: SalpillgoporcI/a polygo­
Ilalis n.sp. has been found in a sample containing a rich 
algal assemb lage, consisting of numerous Salpingo­
porcI/a l7Iclifac, S. l11ueIJlhergii, intermed iat e forms 
between S. melirae and S. hasi, and less numerous S. 
liispanica, Cfypcil/a? so/kalli, Prafl/rlol/cIIa danifovae, 
P. pejovicae, and numerous unidentified dasyclad frag­
ments. The algae arc accompanied by the foraminifera 
Call/panelillla capucnsis (DE CASTRO) , MaYllcina 
hl/lgariea HAUG et a1., Vercorsella searsel/ai (DE 
CASTRO), and olhers. About 20 m above thi s sample, 
in a continuolls sequence, Sa/pingopore/la biokovellsis 
and S. /lr/adanasi have been found . The abovc men ­
tioned dasyclad assemblage and the position of the 
algal-bearing sample within the stratigraph ic column at 
the type-locality suggest an Early Barremian age of S. 
po/ygonalis. Though the above mcntioned foraminiferal 
species occur more frequently in the Upper Hauteriv­
ian, they also range into the Lowcr Barrcmian and thus 
do not contradict the proposed Early Barremian age. 

Sa/pillguporella hasi CONRAD, RADOrCIC & 
REY,1976 

PI. 11, Figs. I ~5; PI. VJJJ, Fi gs. I ~ I 0 

Selected synonymy: 

1976 Sa/pillgopore//a hasi n.sp.- CONRAD, RADO~ 

ICIC & REY, p. 99~ I 04, Fig. 2, PI. I , Figs. 1-10 

13 

? I 986 Sa/pingopore//a aff. hispanica CONRAD & 
GRABNER ~ RADOICIC, Pl. 5, Fig. I 

?1991 Sa/pillgopore//a hasi CONRAD, RADOTCIC & 
REY - KUSS & CONRAD, p. 876, Fig. 4/17 

1992 Sa/pingapore//a hasi CONRAD, RADOTCIC & 
REY - MANCINELLI, p. 9, I I - I 2, PI. 3, ?Fig. 
15, non Fig. 16 

1994 Sa/pingopore//a hasi CONRAD, RADOICIC & 
REY - BODROGI, BONA & LOBITZER, p. 
233-243, PI. 17, Fig. 2 

? J 994 Sa/pillgopore//a hasi CONRAD, RADOICIC & 
REY - SCHINDLER & CONRAD, p. 8 I, PI. 6, 
Fig. 14 

Diagnosis: Dasyclad alga with cy lindri cal thallus 
and with phloiophorous branches which are, after a 
short connecting channel, uniformly and clearly bowJ­
shaped (widened) from their very base. At their distal 
ends the branches are mutually compressed and hori ­
zontally Ilaltened, producing a regular pattern of rectan­
gu lar, horizontally elongated, pores. Calcareous parti­
tions between the neighbouring branches as well as 
betwecn the branches of two consecutive whorls, are 
rather thi ck proximally and taper distally, passing into 
thin walls which, in some sections, appear as spines. 

Salpingoporel/a hasi CONRAD et al. was originally 
described from the Upper Albian and Middle Cenoman­
ian of Kosovo, Serbia, and from Portugal on the basis 
of several, rather atypica l, poorly preserved sections 
and some heavily recrystallized fragments, which, 
when scrupu lo us ly compared with each other, show 
some differences. This suggests that the descrip ti on 
includes material belonging to different forms, or, alter­
nati vely, that the species is characterized by the pro­
nounced variability of some of it s characteristic fca­
tures. This is perhaps thc reason that this species is sel­
dom mentioned and even less frequently illustrated. Its 
age has been most frequently restricted to thc Albian­
Cenomanian. The inadequately defined diffcrcnces 
between the strongly similar features of the related S. 
gencvensis - S. fIGsi - S. ur/adallasi species, lead to fre­
quent misident ifications in which Barremian specimens 
are mostly identificd as S. genevensis or sometimes S. 
hispanico (RADOICIC, 1986, Pt. V, Fig. I) or just 
related to S. hasi (CONRAD & PEYBERNES , 1976, 
Fig. 13 ci , e) and Cenomanian species are without rcser­
vations assigned to S. hasi. 

In the original description of S. hasi (CONRAD et 
al., 1976) , the authors mention the possibility of the 
consecutive whorls not always being in mutual contact 
but , instead , the incomple te calcification between the 
whorls produces a more or less deeply incised wcdge­
shaped dcpression. This feature is regarded as being 
species-specific. To my mind, however, this feature is 
doubtful, i f present at all (i.e. in somc sections it seems 
to be either lacking, or unobservab le; e.g. CONRAD et 
at., 1976, Pt. 1, Figs. 4, S, 7 - Iransverse, slightl y 
oblique section), in others, express ly mentioned, sec-



14 

lions il is unclear (op. eil. , Fig. 2, PI. I , Figs. 1,6, 10), 
and even in the reconstructions (op. CiL , Fig . I) it is not 
c learly defined. Therefore as this feat ure is nol men ­
tioned in the d iagnos is but only in the ciescription, the 
above mentio ned di agnosis of thi s pape r essentially 
agrees with the original one . However, des pite the 
ques tionable value of this feature, the validity of S. hasi 
as an independent species is beyond question. 

The randomly collected sample RC- \ from Barremi ­
an deposit s or the e rcs Island yielded numerou s, well­
preserved sections that , when compared wi th some of 
Ihe publ ished ma te rial of S. hasi (CONRAD e t aI. , 
1976, PI. I) and despite some new ly d iscovered mor­
pholog ic details, can be unequi vocall y assigned to S. 
/ws i. The new ly d iscovered charac te ris tic are poss ibly 
the result o f be ller prese rvation of thi s mate ria l and, as 
such, are inadequate for the establi shment o f" a new tax­
on. They do however, make possibl e minor additions to 
the description o f S. hasi in order to better differentiate 
S. /wsi from S. gene\lensis and S. IIr /adanasi, which, in 
some sections, may indeed appear very similar to be 
almost ind is tinguishable from S. /wsi. 

The compara tively la rge, horizo nta lly fl attened , 
ramifications in S. hasi show the following specific fea­
ture, visible in tran sverse secti ons: foll owin g a short 
connecting pore, or channel (exceptionally the channels 
ent e ring the axial cavity can be sli ghtl y lon ger), the 
ram ifi cations abruptly and unifollnly widen, forming a 
bow l- shaped (PI. II , Figs. 2 -4) o r funn el-shaped (PI. 
VIIl , Figs. 8, 10) swell ing toward the exte ri or and at the 
same time rctaining the ir original shape, with no signi f­
ican t deformations (PI. VIII , Figs. 2-3). The transvcrse­
ly cut branches (i. e ., in longitudinal and/or tangential 
section s) fully corres pond to the reconstruc tion givcn 
by CONRA D c t al. ( 1977, Fig . 4, sec ti on A- A ') for S. 
Ifr/adall(fsi . The calca reous walls between the ramifica­
tions of Ihe same whorl are th ick onl y in thei r proximal 
pa rts, but d ue 10 the abrupt swelling of the ram ifica­
tions, the wall s soon became very thin partitions (PI. 
VIIT , F igs. 7, 9), whi ch in beller preserved spec imens 
appear as spines (PI. VII I, Fig. 7). Thi s is never obse ­
rved in S. gellel'eflsis or S. IIr/adal/osi, and this is also 
the main diagnostic d ifference from S. gel/evensis : in S. 
/wsi the ramifica tions widen abruptly and uni follllly in 
the hori zontal pl ane with thinl y ca lc ined pa rt itions 
between the ir d iswl ends, whereas in S. gellcvclIsis the 
ramifications arc more robust, more irregul ar and with a 
la rger co nn ectin g po re, but arc al so hori zontally nat~ 
lened . Tr we compare some previously publi shed sec­
tions of S. gel/el'el/ sis (e.g., CONRAD, 1969, Figs. 1-9; 
DR AGASTA N, 1989, PI. 6, Fi gs. 1-6) with the trans­
verse oblique and long itudinal sections o f S. //(Is i in Ihis 
s tudy (PI. 11 , Fi gs. 1-5; PI. VIII , Figs. 1- 10), the above 
mentioned diffe rences become c lear enough to justify 
the validity of S. hasi . In tangential sec ti ons, however, 
the diffe rences are nOI so well identifiable and therefore 
such sections cannot be used for spec ies- level id cnti fi~ 

cation betwee n S . /w si and S. gCl/e ve nsis . The differ­
ences with S. IIr/adanasi , in wh ich the branches, in the ir 

Geologi:l Croatica 4911 

proximal pan, also often appear very simi lar, or som e­
tim es even identical, to those of S. hasi , will be di s­
cussed with S. IIrladal/asi. 

In connection with S. hasi , le t us brielly di sc Li ss the 
re lationship between S. has; and S. carpalliica DRA­
GASTAN (1969), with reference to some partial simi­
la rities in longitudinal section s be tween the two 
species . This should be done in spile of the fac t that S. 
carparhica probabl y should be considered in valid o r, at 
least, doubtful, being es tabli shed 011 poorly preserved 
and heavily recrystalli zed mate rial. CONRAD ( 1970) 
firs t mentioned that the spcc ime ns fi gured by DRA ­
GASTAN (1969, PI. I , Figs, 1-6) probably be long to S. 
/11l/ell/bergii (LORE_ Z), which was accepted by BAS ­
SO ULLET e t al. ( 1978). Cert a in sect ion s f igured by 
DRAGASTAN (1 969, PI. I , ri gs. 1-2 and possibl y Fig. 
4 ) poss ibly do belong to S . I1Il1ehlberg ii , wh ereas th e 
sections in PI. 1, Figs. 3, 5-6 arc inadequate for correc t 
identifi cation owing to poor prescrvation and heavy 
recrys tallization. For th is reason and because Fi g. 2 was 
designated as the holotype by Dragas tan, il is clear that 
S . carparlJ ica cannot be acccptcd as a va lid spec ies. 
Despite Ih is, S. carpalhica is continuously mClllioned in 
Rumani an papers. DRAG AST AN ( 1975, p. 62, PI. 8 1, 
r ig. 3 and PI. 82, Figs . I , 3) reconsidered S. carpat/lica 
and introduced two secti ons from the sam e slide as 
hypotypes . However, no new data were given because 
the fi gured sections consist o f full y recrys ta llized spcci­
mens that do not allow greate r insight int o species-spe­
cific features, which should have fac ili tated the identifi ­
cation o f the new spec ies. This a lso applics to !<ll er 
illu strati ons figured by DRAGA STAN ct 01. ( J978, PI. 
8, Fig. 2) and DRAGASTAN (197 8, Fig. 21'; PI. 4, Fi g. 
9), res pectively. In DRAGASTAN ( 1978), th e re is a 
contradi cti on betwecn a fig ured sec tion (PI. 4, Fig. 9), 
whi ch shows horizontall y compressed and rIatt ened 
branches, and the reco nstruc tion (p. 11 2, F ig. 20 in 
wh ieh the branches are shown as be ing compressed lat­
e rall y ( i.e., elongated along the growth ax is o f Ihalus) . 
The renewed description by DRAGASTAN ( 1989, p. 
14- 15) gives no suppor! ror the validity of thi s species. 
The illustrated specimcns (PI. 5 , Fi gs . 8- l3) inc lude a 
great dcal of the already published mate ria l o f problem­
aLical va lue. inc lud ing the ho]otype (PI. 5, r ig. 9 = PI. I , 
Fi g. 2 io DRAG ASTAN, 1969), whi ch CO NR A D 
(1970) a lready identified as belong ing to S. f1/ uelIlbergii 
- an opinion with which the <l ulhor fully agrees. As thi s 
is the only transve rse secti on of S. cmpalli ica whi ch 
uncquivoca lly shows 5 ramifi cations in a whorl , the 
s tatement of S. carpar/Jic(f hav in g 4 (5) branches pe r 
who rl (DRAGAST AN, 1969, 1989) al so appears 
inva lid. The remaining sections (DRAGASTAN, 1989, 
PI. 5, Figs. I O~ 12) arc too poorly preserved for adequate 
comparison, part icularl y wit h the section fi gured in PI. 
5, Fi g. 13, which belon gs to an unidentifi ed form , but 
difrerent from that (or those) fi gured in PI. 5, fi gs. 10-
12. According to all that has been said above - particu­
larl y with regard to the incorrectly dcsignated hololype 
(DRAG ASTAN , 1969, PI. I , Fi g. 6 = DR AGASTAN , 



Sokai": Taxonomil: Review of Some l3arrerniilll aile! Aptian Calcarcou~ Algnc .. 

1989, PI. 5, Fig. 8) - the question remains which char­
acteristics arc relevant for distinguishing S. carpathica 
from other closely related and/or similar species. 

In conclusion, it may be said that S. carpalhica docs 
not seem to fulfil the requirements for being accepted 
as a valid species. More generally, the ever increas ing 
uncritical acccptance of new species or new combina­
tions, which are not adcqua tely documented, disturbs 
and practically (though not formally) invalidates regu­
lar taxonomy and leads to confusing and overburdened 
synonymy. Of course, mi stakes are normal in any activ­
ity, including research, and in future research they are 
normally corrected. However, the problem arises when 
the alleged correcti on of mistakes is carried out on the 
same (inadequate) material, without new discoveries, 
and we should ask ou rselves the following question: 
which cri teria were used by the author in the original 
description and which in a later revision. Therefore, as 
aforementioned and in order to avoid future confusion, 
it is strongly suggested and recommended that the 
establishing of a new taxon, or the revision of an exist­
ing one, of fossil DasycladaJcs should be accompanied 
not only by illustration of the Iype specimen but also of 
several, variously oriented, sect ions which should clear­
ly show the essential distinguishing features of the new 
taxons and, thus, complement the information obtained 
from th e type spec imen (or section, respectively). In 
this way, the subjective, often specu lative, interpreta­
lion of the au thor should be successfully eliminated. 

SalpillgoporeUa llrladallasi CONRAD, PEYBERNES 
& RADOICrC, 1977 

PI. IX, Figs. 1-13; PI. X, r igs . J-9 

Selected synonvmv: 

1977 Sa/pillgopore/la Ifr/adallasi n.sp. - CONRAD, 
PEYBERNES & RADOICIC , p. 73-82, PI. I , 
Figs. 1-7 

1979 SalpiJ/goporel/a urladaJ/asi CONRAD, PEY ­
BERNES & RADOICrC - PE YBERNES & 
CONRAD, p. 746-748, PI. 2, Fig. 3 

1986 Apillclla llrladallasi (CONRAD, PEYBERNES 
& RADOICIC) n.comb. - GRANIER, MICHA­
UD & FOURCADE, p. 801 -807 

1989 /-Iellsollella urladallasi n.comb.(CONRAD PEY­
BERNES & RADOICIC) - MASSE, p. 281 

1993 Salpillgoporclla urladallasi CONRAD, PEY­
BERNES & RADOICIC - BODROGl , CON­
RAD & LOBITZER, p. 66-67, PI. 3, r ig. I 

1994 Salpillgoporclla urladallasi CONRAD, PEY­
BERNES & RADOICIC - BODROGI, BONA 
& LOBlTZER, p. 233-248, PI. 10, Fig. 8; PI. 17, 
non Fig. 4 

From the original description of this species (CON­
RAD et ai. , 1977), the following characteri sti cs con­
tai ned in the diagnosi s should be emphasized: Phloio­
phorous ramifications arranged in dense whorl s, trallS-

versa ll y compressed, giving a honeycomb-like pattern, 
communicate with the axial cavity th rough narrow, 
round pore. Calcareous partitions between the branches 
in a whorl are bent toward the periphery in the same 
direction , but in ailernat ing directions in consecutive 
whorls. Distal swelling of the branches is interpreted as 
resulting from the imprin ts of cysts. The calcareous 
envelope is bu il t of spath ic calcite which extends to the 
axial cavity, which is lined by a thin dark layer COIll ­

posed of small, radially arranged crystal s . In the 
description, the opposite direction of the inclination or 
partitions in the adjacent whorls is mentioned , and 
emph asized in the reconstruction (CONRAD et aI., 
1977, Fig. 4C), though it is not clearly vis ible in the 
holotypc (op. cit., PI. I, Fig. I). In the description, there 
is also the fo llowing statement: the calcareous envelope 
fills the enti re space between the ramifications, though 
a specimen from Spain is mentioned, in which the 
cnvelopes of the branches do not touch at their ends. 
However, regardless of what is stated in the diagnosis, 
the reconstruction presented with S. hasi CONRAD et 
al. (1976, Fig. 1) and S. lfrladallasi CONRAD el al. 
(1977, Fig. 4) show a remarkable similari ty in their lon­
gitudinal sections, whereas the transverse sect ions arc 
too simplified (schemati c), being obviously based on 
one set of transverse sections in slides whereas another 
set, different from the first set, have been neglected, 
c.g . the reconstruction of the transverse sect ion of S. 
hasi (CONRAD et aI., 1976, Fig. 4) seellls to be based 
on the transverse section figured in PI. I, Fi g. 2, where­
as another transverse section , which g ives a different 
picture (PI. 1, Fig . 7) seems to be neglected; however, it 
is sim ilar to the section A-A' in the reconstruction or S. 
1I1'Iadallasi (CONRAD ct aI., 1977, rig. 4). However, 
the above mentioned transverse section (CONRAD et 
aI., 1976, PI. 1, Fig. 2) possibly belongs to S. melitae 
RADOICIC. Nevertheless, in spite of considerablc sim­
ilarities between the two species and frequent dilemmas 
concerning their identification, the val idity of the two 
spec ies is confirmed by the available material, where 
both species are abundantly represented and lheir dif­
fere nces become evident. Numerous, well-preserved 
sections ascribed to S. hasi (PI. II , Figs . 1-5; PI. III , 
Figs. 1-10) and S. IIriadanasi, respectively (PI. IX, Figs. 
1-13; PL X, Figs . 1-9) make possible the visualisation 
and definition of some features essential ror their dis­
crimination , which were not mentioned in the respec­
tive original descriptions. 

The caleareous envelope of S. u/"Iadanasi, in con­
trast to that of S. hasi, is built of coarse-grained, yel­
lowi sh calcite, identical in colour as that observed in S. 
dinarica (these two species are associated in the materi­
al deriving from Ml. Dinara and thus can be directly 
compared). In S. urladanasi, however, th e texture of 
calcite is clearly blocky, i.e. mosaic-like, whereas in S. 
dinarica it is rad ial -fibrous. As already stated in the 
original diagnosis (CONRAD ct aI., 1977), some speci­
mens of S. urladanasi may show a very thin, sometimes 
somewh at darker layer lining the axial cavity (P I. IX, 



16 

FigS . I, 3). In present material it is d ifficull to tell 
whether that layer is composed of mosaic-like or radial 
crys tals. This feat ure has already been the subject of 
discussion (CONRAD & V AROL, 1990; SIMMONS et 
aI. , 199 1) and was also mentioned by BODROGI ct al. 
(I993). Such peculiarities in the structure of the skele­
ton of S. urladanasi may, among other th ings, serve as 
additional distin gui shing features for separati ng thi s 
species from S. hasi 0 11 the one hand and from S. dinar­
iea on the othe r. The origin of this feature, however, 
remains open for further investigation. 

The outer surface is covered with protrusions of 
ramifications wi th their own envelopes extending from 
thc middle of their length to the distal end (PI. IX , Figs. 
10, 12- 13; PI. X, Figs. 2, 3 pars,S). Thi s is the reason 
for the occurrence of wedge-like incisions in the skele­
ton and mutuall y unconnected, deformed quadrangular 
patterns in superficial tangen tial (con ical) sections 
resulting from individual, mutually separatcd ramifica­
tions from the same whorl or from adjacent whorls (PI. 
IX, Fi gs . 11 - 13). These features are ment ioned by 
CONRAD et a!. (1976) as being characteri stic o f S. 
hasi, but they could not be observed in the avail able 
materi al, nor do they seem to be visible in the original 
illustrations. Furthermore, the available material seems 
to exclude the presence of such features in S. hasi (PI. 
VlIl , Figs. 2-3, 6A). Ahernatively, in S. urladanasi the 
separate calcareous envelopes of adjacent branches 
clearly show the wedge-shaped intrusion s between the 
branches that somet im es penetrate deep into the prox i­
mal part of the skeleton (PI. IX, Figs. 12-13). In cases 
when the envelopes in S. urladanasi are in mutual con­
tact, the thickness of the partitions between the ram ifi­
cations doubles (PI. IX, Figs. 2-4; PI. X, Figs . 4 pars, 
6), which is not the case in S. hasi. The fu sed envelopes 
disappear deeper with in the calcit ic mass between the 
proximal part of the ramifications (PI. IX, Figs. 2, 7-8), 
and thus the difference between S. hasi and S. urla­
dallasi also disappears. In S. lIrladanasi, the horizontal­
ly flattened branches may be deformed dista lly to vari­
ous extents, which can be seen in transverse and tan­
gential sec tions. Thus, in transverse sections, the 
branches may be abruptly and uniformly swollen, form­
ing a bowl-shape widening following the short and nar­
row pore (channel), similar to S. hasi (PI. X, Figs. 8-9), 
or they may be irregularly swollen with one side more 
inflated than the other (PI. IX, Fig. II pars; PI. X, Fig. 
IA), or they may be fu nnel-shaped, with gradual 
widening. Some times, in spec imens with irregularly 
swollen ramifications, the channels that connect the 
ramifications with the axial cavity are directed obliq ue­
ly to the ax ial cavity (in the horizontal plane, i.e. in 
transverse section), i.e. they are not perpendicular to the 
inner surface (PI. X, Fig. 8). A particularly pronounced 
deformation of the branches, emphasized in the original 
description (CONRAD et. a/., 1977) as a primary 
species-spec ific characteristic, concerns the shape of 
their distal ends in tangential sections. This shape in S. 
urlaclanasi can vary from a comparatively regular rec-

Gcologi~ Croalica 49/1 

tang le (PI. IX, Fig. 5) to a rectang le squeezed in the 
middle like a bow-tic (PI. IX, Fig. 4 ), sometimes with 
more strongly stre tched corners on one side, remi nis­
cent of the longitudinal section of a femur (PI. IX, Fig. 
6; PI. X, Fig. 4). The part itions between the branches of 
the same whorl may be vertical to subvertical (PI. IX, 
Fig. 5) in less deformed branches, or oblique, inclined 
to the same side. This can also be the case in the adja­
cent whorls, or else, as has been stated in the original 
description, the inclination can be reversed in adjacent 
whorls (PI. X, Fig. 4 pars). The intensity of deformation 
of the ramification' s distal ends varies from individual 
\0 individual and even in the same individual. The vary­
in g exten t of deformation of the ram ifi cati ons in S. 
urladanasi proves it to be of secondary origin and thus 
neither this feature can not always be used as absolutely 
reliable [or speci fi c determination. Therefore, it is not 
considered likely that Ihe distal swellings of ram ifica­
tions are due to imprints of cysts, or rather, there are no 
indications for such an interpretation. 

Regardless of the original descriptions of S. hasi 
and S. urladanasi and the accom panyin g reconstruc­
tions, all data based on the abundant avai lable material 
suggest that the difference s between the two spec ies 
can be specified as follows: In S. hasi the ramifications 
are more or less regular and in transverse sections (= in 
hori zontal plane), after a shon channel, abruptly and 
regularly widen into a bowl - or funnel-like Shape. The 
calcitic mass between the prox imal part of the ramifica­
tions within a whorl abruptly thins di stally, givi ng, in 
better preserved specimens, the appearance of spines. 
Such dis tally th in partitions make it impossible to 
accommodate uncalcified ho ll ow spaces (recesses) 
between the ramifications and thus prevent the individ­
ualization of rami fications at their outer ends. O n the 
contrary, in S. urlaclanasi the calcit.ic mass between the 
ramifications of the same whorl also becomes thinner, 
but halfway out (or in the outer third) , and sometimes 
even earli er, it sp lits into two separate ca lcareous 
envelopes of the dista lly individualized branches. As a 
consequence of this , there are uncalcified wedge­
shaped recesses which som etimes may reach deeply 
into the proximal part of the skeleton (PI. IX, Figs. 12-
13). In S. hasi, the part itions between the ramifications 
of the same whorl and of the adjacent whorls are regu­
lar and in the outer tangential section hal f as thick as in 
S. urladanasi, in whi ch they arc often separated from 
each other with deep uncalcified , wedge-shaped inser­
tions. The partitions between the ramifications in S. 
hasi being this thin, make it impossible for the ramifi­
cations LO bulge out on the outer surface. Therefore the 
ramifications were probably covered by an uncalcified 
membrane, in contrast to S. lIrladanasi in which the 
protruding ramifi cations are evidently covered wit h a 
calcareous envelope composed of the same calcite as 
the proximal part of the skeleton (PI. X, Fig. 2). 

Stratigraphic position: In spite of the fact that 
unequivocal finds of S. urladanasi - as defined in this 
paper - are not very numerous, it s total ra nge in the 



SOkiIC: Taxonomic Review of Somc Barremian and Aptian Calcareous Algae ... 

region or the Dinaric Karst in Croatia can now be reli­
ably detennined due to its lowest (oldest) occurrence on 
the island of Mljet and its highest (youngest) occur­
rence on Mt. Dinara. On Mljet, S. urladanasi has been 
found associated with Salpingoporel/a biokovcnsis, in a 
sample situated 20 m above the rich algal assemblage 
also containing Campanellll/a capuensis (DE CAS ­
TRO), which indicatcs the Lower Barremian age (see 
the description of Clypeina gigantea). On the other 
hand, on Mt. Dinara , the sample with S. urladal/asi is 
situated 25 m above the assemblage consisting of Salp­
ingoporella dinarica RADOICIC, Acropore//a radoici­
cae PRATURLON, Orbitolina (Mesorbitolina) milluta 
DOUGLASS, Sabal/dia minllla (HOFKER), Nezzazata 
simp/ex OMARA, Deharina hahouJlercnsis FOUR­
CADE, and Vercorse//a scarsel/ai (DE CASTRO) 
(foraminiferal identifications by I. VELlC). About 15 m 
above the bed with S. lIrladanasi, the first Orbifo/ina 
(Mesorbilo/illa) lexalla (ROMER) occur, and pass 
upward into a typical Albian microfossil assemblage. 
Therefore the total range of S. urladanasi, in lhe region 
of the Dinaric and Adriatic Karst , can be defined, for 
the time being, as Barremian-Aptian. 

Salpillgoporella dillarica RADOICIC 1959 

PI. XI, Figs. 1-10; PI. XII , Figs. 1-13; PI. Xlll, 
Figs. 1-8; PI. XIV, Figs. 1-9 

Selected svnonymy: 

1959 Sa/pil/gopore//o dil/arica n.sp. - RADOICIC, p. 
33-42, PI. 3-5 

1960 HellSol/e/la cylindrica n.gen., n.sp. - ELLIOT, p. 
229-230, PI. 8, Fig. I 

1982 Hel/sol/e//a - LUPERTO-S INNI & MASSE, p. 
870, Figs. 12 ct-e 

1983 Sa/pillgoporel/a dinarica RADOICIC - CON­
RAD, PEYBERNES & MASSE, p. 93-94, PI. 6, 
Figs. 6 pars, 12 pars 

1984 /-/ellsollel/a sp. (gr. dinarica) - LUPERTO-SIN­
NI & MASSE, p. 340, PI. 33, Figs. 7-8 

1985 /-Iensol/el/a dinarica (RADOICIC) - MASSE, p. 
719 

1987 Salpingoporella dillarica RADOICIC - RADO­
lCIC, p. 73-79, PI. I , Figs. 5 pars, 6 pars 

1989 Salpillgopore//a dinarica RADOICIC - DRA­
GASTAN, p. 13, PI. 5, Figs. 1-2. 

1989 Hensone//a dinarica (RADOICIC) - MASSE, p. 
281, not figured 

1989 Hellsonel/a dil/arica (RADOICIC) - MASSE & 
LUPERTO-SINNI, p. 32, not figured 

1990 Sa/pingoporel/o dillarico RADOICIC , 1959 
(alias /-Iel/sonella cylindrica ELLIOTT, 1960, 
a lias Hensollclla dinarica) - CONRAD & 
V AROL, p. 206, Fig. 9 

1991 Sa/pingoporel/a dinarica RADOICIC - KUSS & 
CONRAD, p. 874 - 876, Fig. 4.15 

17 

1991 Hellsonel/o dinarica (RADOICIC) - SIM­
MONS, EMERY & PICKARD, p. 955-959, Fig. 
J A-C 

1992 Sa/pingoporel/a dinorica RADOICIC 
MANCINELLI, p. 9, 13, Pl. I, Fig. 10 

1993 /-Iellsollelia gr. dinarica (RADOICIC) - LUPER­
TO-SINNI & MASSE, p. 300, PI. 4, Figs. 8-10 

1994 Sa/pingoporel/a dinarica RADOICIC , 1959 -
SCHINDLER & CONRAD, p. 77, PI. 2, Figs. 9-
II; PI. 4, Fig. 16 

This is a very well-known species, which occurs 
frequently and abundantly in the areas surrounding the 
Mediterranean and the Middle East (i.e. the southern 
Tethyan margin). It was originally described as a small 
alga with cylindrical thallus, rather broad axial cavity, 
and comparatively thin walls (RADOICIC, 1959). The 
simple, undivided, phloiophorolls ramifications are 
characteristically arranged into a quincunx pattern, ancl 
are horizontally flattened at the distal ends, where the 
porcs assume an elongated-rhombic patlem. A species­
specific characteri stic is that the calcarcous envelope 
consists of radial-fibrous calcite and is lined on the 
inside by a thin dark layer of microcrystallinc calcite. 
These two characterist ic features were usee! by 
ELLIOTT ( 1960) for the establishment of his new 
genus and species Hensonella cylindrica (Problcmatica, 
Annelida?). Later, however, ELLIOTT (1968) consid­
ered it to be a younger synonym of Sa/pingopore/la 
dinarica. Since then, many authors have discllssed the 
species (e.g., LUPERTO-SINNJ & MASSE, 1982, 
1984; CONRAD & V AROL, 1990; SIMMONS et al., 
1991; SCHINDLER & CONRAD, 1994), mainly due to 
the two peculiar features mentioned above, which were 
co nsidered uncharacteristic of the Dasycladales. SIM­
MONS et al. (1991) have summarized the previous dis­
cll ssions ancl, basing their conclusions on the results 
obtaincd by cathodoluminiscence and microprobe 
analysis, have concluded that the calcareous envelope 
of S. dinarica was composed of primary low-Mg calcite 
and thalthis was its original structure, no mailer how 
highly unusual this was for the dasyclads and how dif­
ferent from the other Salpingoporclla spccies. For this 
reason they ascribed this species to Hensonella , as has 
previously becn done by LUPERTO-SINNI & MASSE 
(1982, 1984). The truth is, that almost all sections of 
this form show a more or less pronounced radial­
fibrous calcitic wall structure. This structure is in gener­
al, better displayed in specimens in which the pores of 
th e ramificat ion s arc only slightly visible (see, for 
instance, ELLIOTT, 1960, PI. 8, Fig. I; SCHINDLER 
& CONRAD, 1994, PI. 2, Figs . 9- 1 I and the sections in 
the present paper, e.g., PI. XI, Fig. I). However, this is 
not a rulc without exception, though the reverse is also 
often true (P I. XI, Figs. 4-6; PI. XII, Figs. 6-7). In the 
majority of the observed spccimens, thc axial cavity 
and the surrounding space are filled with coarse-grained 
blocky calcite cement or the surrounding sediment (PI. 
XI, Figs. 6, 8, 10; PI. XII, Fig. 1). 



18 

However, in spite of the facL Ih al the wa ll s of S. 
dinaric."o arc almost always composed or radial-fibrous, 
yellowish calcite, the re arc a few exceptions to thi s ru le, 
too. Conside r, for instance, sections fi gured in Pl. XII, 
Fi gs. 11 , 13, in which the fibrou s calcite is partly or ful ­
ly replaced by microcrystalline ca lcite identical 10 tha t 
of the surrounding sediment. The oute r and inner sur­
faces of the ca lcareous envelope a rc marked by a thi n 
layer of yellowi sh ca lc ite. Thi s phenomenon should 
probably be conside red as a consequence and result of 
diagene tic processes . A lso, the re arc a few sections in 
which the radial - fibrou s ca lc ite g rows upo n the thin , 
dark , microcrys ta lli ne layer ins ide the axia l cav ity, fill ­
ing it pa rt ly (PI. XTl , Figs. 1, 5) or wholly (PI. Xli , Figs. 
8 pars, \9). The same rad ia l-fibrous calc ite also occurs, 
though more rare ly, on the oule r surface. In such cases, 
too, it is separated from the calcareou s wall itself by a 
thin dark layer probably of microcryst,:lIline calcite (P I. 
XlI , Figs. 5, 10). The developmcnt of thi s radial -fibrou s 
calcite , which is certa inly of secondary or igin and syn­
ch ronous wi th the blocky cal ci te ceme nt in the ax ial 
cavi ty, may be thought of as a conseque nce of o ver­
growth ( i.e . syntax ia l rim cement ) . This indicates that 
during diage nes is the axia l cav it y may be partly or 
wholly fill ed with either (a) surrounding li me mud (PI. 
XI, Fig. 7), (b) secondary blocky calcite cement , or (e), 
more rare ly, radial -fibrou s calcite (P I. XIII , ri gs. 5, 9-
10). EvenlUally, both within the ax ial cavity and on the 
ex te rnal s ur face o f the ske le ton , synchronously and in 
the sam e e nvironme nt , blocky (granula r) and radia l­
fi brous calc ite deve lops, and thus the q ues tio n ari ses 
why in somc cases there is an ove rgrowth (if indeed it 
is), and in others there is not. 

The thin dark inne r layer has been noti ceci by all 
worke rs who have dea lt with thi s spec ies, beg inning 
wit h the original descr ip tion by R ADOI C IC: (1959). 
ELLlOlT ( 1968) s tressed lhe pri mary organic na lurc of 
thi s layer and S IM MONS e t a l. ( 1991 ) w lled Ihal it 
could be interpre tcd as a primary organic membrane, 
about 0.012-0.018 mm thick, that lines the calcareous 
e nve lope on the inside. C ON RAD & VAROL (l990) 
a lso consi der this layer in S. dillaric{( to be of primary 
o ri gi n. In the available material (P I. XIl , rigs. 1-2, 5 , 
LO; PI. X][] , Figs. 1-4), two types of dark mi eroerys­
tallinc laye r can be di s tinguis hed , which di ffe r as a 
result of the time and origi n of the ir growth . T he firs t 
type may be more o r less clearly vi s ible but is a lways 
prcscnt , lining the inne r surface of the calcareous wa ll 
and being in imm ed iate contact with the radial -fibrous 
ca lc ite of the wal l. The primary origin of this laye r and 
its function during the life of the al ga, as assumed by 
CONRAD & VAROL ( 1990: " in vivo"), is confirmed 
by the ex iste nce o f the pores in the layer, wh ich are vis­
ible in tangential sections (PI. XIIJ , Figs. 1-2). The sec­
ond type inc ludes the ve ry dark m ic rocrys talline layer 
that may be unevenly developed (i.e. be ing of variolls 
thickness) and whi le lacking in some specimens, in olh­
ers it may be more pronounced than the firs t type . This 
layer origi nated during ca rl y diagenesis, probab ly as a 

Geologi" Croalic:, 49/1 

result of biological ac tivity of mi crobial organi sm s or 
by accretion of organic muc us on the a lga l s kel e tal 
rema ins (thi s is the third or the fourth possibdit y men­
tioned by CONRAD & VAROL, 1990) . It can develop 
both on the outside and the ins ide of thc ca lcareous 
e nve lope; in the firs t case it occurs as a th ill layer (of 
va riable th ickness) eit he r wholl y ' ·enc rusti ng" the 
skeleta l remain , or only in part ; it is usuall y onl y deve l­
oped on some specimens wi thin a sample . The second 
case, i.c. when thi s laye r is developed on the ins ide of 
the calcareous envelope, is shown in PI. XII , Figs. [ -2, 

5 , 10. Sometimes the ou te r laye r visib ly dest roys the 
outer surface of the calcareous e nvelope and pene trates 
deeply into the rad ia l-fibrous calci te, reaching, in some 
cases, Ihe ax ial cav ity (PI. XII , Fig. 2). It may be c learly 
and sharply de limited toward the surrou nding sedime nt 
(PI. XII, Figs. 1-2,5), or it may pass gradually i11l0 the 
surrounding scci iment , [arming a darkened zone around 
the a lga l ske le lon (PI. Xl , Figs. 8, 10). When developed 
in the ax ia l cavi ty, it can partly cover the pr imary da rk 
microcrystalline layer (PI. X II, Fig. 10). lt is more fre­
que nt ly developed ins ide the ax ial cav ity , e ithe r as an 
unin te rrupted zone (PI. XI I, Fig. I) or fragme ntaril y (PI. 
Xli , Fig. 5), if the ax ial cav ity has already bee n partly 
fill ed, in which case it int errupts the growth of the sec­
ond gene ration calcite ceIlle nt within the axia l cav ily 
(PI. XIf , Fig. 5) , be it rad ia l- fi brous or b locky. In thi s 
case, Ihc narrow zone o r the primary dark mi c rocrys­
ta ll ine laye r is sharply de lineatcd as a reg ular oval 

(depe ndi ng on the o bl iqu ity of the section), which is 
li gh ter than the irregular seconda ry m icroc ryst a ll ine 
zo ne and darkc r than th e li ght calc it e of the ske le ton 
and the first generation or the secondary calci te cement 
(radia l- fibrous or blocky), which always fill s the centra l 
part o f the ax ial cavity (PI. X II, Fig. L). 

Amo ng num erous samples contain ing S. dinarica, 
the re a rc on ly two samples (bo th Aptian in age, one 
from the Dugi otok Is land and the other fro m the Mali 
Losinj Is land) , thaI , in addi ti o n 10 typical spec im ens, 
a lso contain a few sections which show the tende ncy of 
a branching th allu s and , eve n more rare ly, some sec­
ti ons in whi ch the branching of the tha llus is c lear ly 
deve loped, s im ilarly to some recent gene ra ( V A LET, 
1968). T he tende ncy of the branching tha llus is di s­
played by longi tudina l spli tt ing of the axial cav ity and 
can be seen in transverse and obliq ue sections wit h 
more or less pronounced extensions of the inner surface 
into a fold di rccted toward the centre of the ax ial cav ity. 
The initia l formation of th is fold appears, in some sec­
tions, like a denticle (PI. XIV, Fi g. 5). If Ihe ex te nsion 
of the fo ld is Illore pronounced and proceeds frol11 both 
oppos ing s ides, it leads to a part ia l (PI. Xl!! , F ig. 6) or 
comple te splitting of the axial cavi ty illlo two para lle l 
(P I. XI II , Figs . 6-7) or s lightly divergent (PI. XIV, Figs. 
3, 6-7) tubular cavities which are, still , connected by a 
common calcareous enve lope (PI. XIV , Figs. 1,7); that 
is, the indepcndent branches a re not ye t full y form ed. 
Precise ly thi s fact led to the supposit ion of two spec i­
mens tightly pressed toge ther, in which the ca lcareous 



'obi;: Taxonomic Review of Some lbrrcmi:m and Aptian Calcareou.'; Algae ... 

~nve lopcs along the contac t were supposed to be partly 
)J" completely dissolved. Howcver, this is refu tcd by thc 
~x i s t e ncc of secti on s tha t show the s in g le calcareous 
~nvelope, in which the charac teri stic arrangement of the 
Jo res (well visi ble in tangential sec tions) was re tained 
. md which, by its mere d iamete r, eanllot accommodate 
. wo individ uals (PI. X lii , Fig. 3; PI. XIV, Fig. 8). The 
longi tudinal sec tion fi gured in PI. X III , Fig. 5, clearl y 
, hows the branc hing of an initially tubul ar tha i us, 
whi ch continu es to grow with two slightly divergen t, 
ind e pe ndcn t branc hes, eac h wit h it s own ca lcare ous 
:;nve lopc. 

In sp ite of two unique features, the thallus of th is 
alga is composed of low-Mg calc ite (STMMONS e t a I. , 
199 1) and the re is a thin dark inne r layer of m icroclYs­
ta lline ca lcite (w hi c h is al so rare in othe r dasyclad 
spec ies), 1 am , for the time being, not inclined to 
chan ge it s taxonomic c lass ification , i. c. 10 remove it 
from the genus SafpiJ/gopo/'clfa. If we accept the taxo­
nom ic c mcndation , thi s would mean givi ng grea te r 
in1po rLancc to a new paramete r - the primary calciti c 
mineralogy of the skele to n - w hose ph ys io logic rol e 
and biolog ic s ignificance are far from clear, and rcjcc t­
ing, in turn , the clear morphologic c rit e ria that have 
se rved us well in the hitherto es tablished class ification 
of the dasyc lad algae (shape and arrangement or ramifi­
ca ti ons, etc.). These morpholog ic: criteria are striking ly 
s imilar to those in othe r Sa/pillgoporcl/a spec ies . 
Besides, the re is the new ly di scovered featu re of the 

branching thallus, obviously not so nue in this species, 
and tent atively indi ca ted even in the type spec ies , S. 
/III/eh/bergii (PI. I , Fig . 6). 

The stratigraphic ran ge, geographi cal di stribution, 
and e nvi ronm ental p refere nces of this species have 
been frcquently commented upon, as is ev idenced by an 
exceptionally abundant synonymy (BASSOU LLET et 
aI., 1978, and th is pape r). Tn the Dinaric and Adri atic 
Karst areas it occurs in va rious fac ies types of the shal­
low wate r carbonates , frequen tl y assoc iated wi th other 
ca lcareous al gae, benthic foraminifera (both " large" and 
"sm a ll "), corals, bivalves, etc . Therefore , co ntra ry to 
what has been stated by S IMMONS et al. (199 1), in my 
opinion it cannot bc cons idered as be ing charac teri stic 
o f particular k inds of e nvironme nts. It occurs mos t 
abundant ly in mi c riti c limes tones of protected Uppe r 
Barrem ian and Apt ian lagoons. Its oldest occurrence in 
inves tigated area is in the Valanginian of wes tern Istria 
(PI. Xl , Fig. 1), where in a continuous lithostratigraphic 
succession , it occ urs not far above the first occurrences 
of Epifllosfopora cekic i RADOICIC and Trip/opo/'el/a 
lIeocollliell sis RADOIC I<:. which arc cons ide red to be 
mos t common in the Lower and Middl e Valang inian. 
Thus, the to tal s tratigraphic range of S. dinoric(l in the 
Adr iatic and Dinarid ic kars t areas can be inte rpre ted as 
from the Middle (or more likely, Upper) Valanginian , 
up to and including the Albian. Its lower boundary thus 
coincides with the data g iven by CONR AD & V AROL 
( 1990) and SCHINDLER & CONRAD (1994). 

19 

T here is s till anothe r phe nome non whi ch see ms 
worth me ntioning in connec ti on w ith this alga: the 
comparativc ly frequen t occurrence of eharaeeans asso­
ciated with S. dillarica or, very abu nd antly, in laye rs 
immediate ly be low or above the beds wi th S. dinarica . 
In western Is lria , in thc M iddle-Uppe r Valanginian , S . 
dinarico is associated wi th a peculiar type of characcan 
(PI. X IV, Figs. 10-13), w hi ch has so rar not been 
observed in the younger (Barremian-Aptian) deposits 01" 
that are a. Maybe such characean s ca n be caut ioll s ly 
used as an indi cation of the Valang inian age of the oth­
erwi se ba rre n deposits , at leas t in a geographi ca ll y 
rcstric ted area. 

Genus Pralllrlollella BARATTOLO, 1978 

Praillriollella lIerae (DRA GASTAN, BUCUR & 
DEMETER, 1978) BUeUR, 1993 

PI. XV, Fi gs. 1- 19 

? 1978 Ha licorYlle nerae n. sp. DRAGASTA N, 
BUeU R & DEMETER, p. 28, PI. 12, Figs. 5-6 

1993 Prawrlonella lIerae (DR AGAST AN, I3 UeU R 
& DEMETER) n.com b. - BUeUR, p. 103, PI. 8, 
Fig. I, ?Figs . 2- 12 

1993 Uk,," ell,,? sp. - MASS E, p. 3 16, PI. 2, Fi g. 8 

1993 I'r{///frlonell" dallilovae (RADOICIC) - SOTAK 
& MISiK, p. 403 , PI. 6, Fig. 8 

This species, first described as Holicoryne /Ierae by 

DRAGASTA N e L al. (1978) was later transfe rred to 
Prawrlonella by BUeUR ( 1993). In the original 
description or 1-lalicorYl/e lI erae (DRAGASTAN et aI., 
1978), the a uthors described oval -ellipsoid bodies, g iv­
ing the ir d imensions (length , w idth , number and diame­
ter of cysts), whic h represcn ted an inadequate descrip­
Li on of these frag me nts . The re ro re MIS IK ( 1987) con­
s ide re d thei r assignment to Ha/icoryne as doubtful. 
Among alilhe sections figured by BUeUR (1993, PI. 8, 
Figs. 1- 12), only one sec tion of a segme nt (op.e it. , PI. 8, 
Fig. 1) is c lear enough, where as other sections (O p. CiL , 
PI. 8, F igs. 2- 12) cannot be ide ntifi ed with certainly 
with Ihe section figured on his PI. 8, Fig. 1. Moreover, 
the diffe re nce is also v is ible whe n the dime ns io ns of" 
cys ts in DRAGASTAN et al. ( 1978) are compared w ith 
the va lues of "p" (both o n di s ta l and prox im al e nd s) 
given by BUeUR ( 1993) . In spite o r some visual s imi­
larity be tween the sec tions fi g ured by BueUR ( 1993, 
PI. 8, Fi gs . 3- 12) a nd those in the present paper (PI. 
XV, Figs. 1-19) , only a prob le mati c al compari son is 
poss ible and the present author is not. convinced that we 
are deali ng wi th the same foss il form . The question is, 
among othe r things, how can 95 % of the sections fi g­
ured by BUeUR ( 1993) represenl tangential (?) sections 
th rough isolated segments, whereas almost 100% of 
section s studied here (PI. XV , Figs. 1- 19) show Lhe 
more or less complete a lgal skeleton , regard less of the 
ori entation. However, independent of these considera­
tions, the assignment of Halicoryne J/erae (DRAGAS -



20 

c is, or s light ly below the whorts, appears s ligh tly nar­
rowe r. P. ncrae is al so distinctly smalle r than P. jol'­
dallica K USS & CONRAD, [rom which il also di ffers 
by the shape of the ramification s and the number of 
rows of ramifications within a ring. P. ha!lll1ll1r/ai 

(RADOI C IC), which has rcccmly been ass igned 10 Fa/­
solikallella (SCHLAGINTWE IT, 1990) and was or ig i­
nal ly desc ribed [1'0111 heav ily recrysta llized mate rial 
(RADOIC IC, 1975 , Pi s. 1-2), shows a s imilar heighl of 
calc ified rings (segments), according 10 KUSS & CON­
RAD ( 199 1), this howe ver being th e sale similarit y. 
The c1i1lcrcncc is most ev ident in the di D values, whic h 
arc distinc tl y larger in P. lIerac. 

A great visual si mi la rity which, in some sec tions, 
amount s to an impression of total likeness, ex is ts 
between P . lIerae and Carbon iferous Windsopore/la 
radiala MAMET & RUDLOFF. Comparison belwccn 
the illu s tra ted sections of W. radiata CMAMET & 
RUDLOFF, 1972, PI. 3, Figs. 18-25) and P. lIerae (PI. 
XV , Fi gs . 1- J 9) reveals an id enl ieal ske letal form, 
arrangeme nl of ramifica tions wit hi n the calc ified rings 
and thc ir direction 0 1" growth dive rg ing bot h upward 
and downward from the hori zonta l plane, a nd al so 
s light widcnin g of the cent ral cavity at the leve ls of 
whorls, all these features being essentia lly the same as 
in P. /lerae. The dillcrcnces concern the much small er 
dim ens ions of W. radiota and the pattern of ram ifica­
tions, which, accord ing to MAMET & RU DLOFF 
( J972), are grouped into bundles of four issui ng from a 
common basis , or constitu te a bunch of a ramified pri­
mary branch, rcspecl ivcly (sec MAMI2T & RUDLOf'f', 
1972, Fi g. 3). Thi s last s tatement, however, remains 
specu lative, w ith rega rd to the author 's assertion that 
the calci fi cation of the ramification's e nvelope occurs 
no sooner than from the ir departure point, and thus the 
exis tence of common departure point , i.e. the primary 
ramifica tion , is made doubtful. Thi s doubt is further 
s tre ngt he ned by secti ons fi gured by MAME'T & 
R UDLOFf' ( 1972, PI. 3, Fig. 18, Fig. 22: lefl sidc o f Ihc 
uppe r ring, Pi g. 23: le ft s ide of the lower ring), whe re 
one gets the impression o f rami fications issu ing sepa­
rately from the axial cavity , s imilarly as can be seen in 
some sections of P. salcmitana fi gu red by BARATTO­
LO (1978, e .g. PI. 5, Figs. 10, 12, 14; PI. 7, Figs. 1,3, 
4, e tc.). However, being uncertain does not necessarily 
mean tha t the inte rpre ta ti on of Maill et and Rudloff is 
impossible; there fore the object ivisati on of this poss ible 
and s ignificant d iffe re nce be tween Windsoporella and 
Prallf l"loJ/clla should be a matter for furt her resea rch on 
the type-species of Wi/ld.mporel!a. If, on the other hand 
the main characte ristics of Windsoporella are proven to 
be ide ntical with those of the tale r established PraHI/·­
IOllel/a, the quest ion of the validi ty of the genus Pratl/l ·­
/onella ar ises. However, the large time span separating 
two taxa with identical c haracteristic fea tures, suggests 
agrccmcnl wilh KOCI-IANSKY -DEVIDE & GUSIC 
(1971 ) who mentioned severa l ins tances of the repeated 
occurre nces of s imilar and/or (almost) iden tical fo rms 
and other c haracteristics in the course o f evolution of 

Geologia CrO;llica 49/1 

the Dasycl ada[es without implying the ir generic unity , 
which would a lso seem a li ke ly ex planation fo r the 
lVilldsoporellalPratlirlollella s itua tion, im plying tha t 
the two genera do not belong to the same li neage. Thus 
Prall/rlol/ella would remain a va lid genus. 

Stratigraphic position: The remains of this a lga 
have been found in a randomly collected sample during 
geologica l reconnaissance of the central part of Korcula 
Island. Therefore it is impossible to assess its full strati­
graph ic posit ion and its age can only be determi ned on 
the bas is of the accompanying mic ro fossils, which 
include Trip/opore//a bacililormis SOKAC, T. marsi­
COI/O PRATURLON , Sa/pillgoporel/o hera/dim n.sp. , 
and Cylilldroporella ivallovici (SOKAC) . In addition , 
th e algal-bearing sam ple lies immediate ly (probab [y 
about 2-3 m) be low the first find s of Pa/orbitolillll 
ICllticlflaris (BL UM EN BACH), sugges ting that the 
stratigraphic posit ion o f the algal-bearing sample is top 
Barrem ian or maybe Lower Aptian. 

Pmturlonella dallllatica (SO K AC & VELle, 1978) 
n.co mb. 

1978 Sel/iporel/a da/lIlatico n.s p. - SOKAC & 
VELIC, p. 23 l -234, Pis. 6-9. 

1982 Ukallel/a? dani/ovae R ADOIC IC (p .parlc anel 
all. dalli/ovae) - CONRAD , p. 2. 

Cons ide r ing its generic attribution and taxonomic 

valid it y, Ihis species was conside red by CONRAD 
(1982) to be a you nger synonym o /" Prali.lr/olle /lo 
dani/ovo e (RADOICIC) or, alleasl, parlly s imilar 10 il. 
The moS! rcccnl quolalion (SCJ-JJND LER & CONRAD, 
1994) (ll so inc ludes the possibilit y of ass igni ng S. daf­
marica to Pratllrfone /!a. Our illu s tra ted mate rial 
(SOKA C & VELIC, 1978, Pis. 6-9) shows some hilhcr-
10 insufficient ly e mphasized features which make it 
poss ible to d ist ingu ish thi s fo rm from P. donifol'ae and 
to va lidate it taxonom icall y as Pratlfr/Olle//a dalmatica 
(SOKAC & VELIC) n.comb. As can be seen from 
Table 6, the d iffe rences betwee n P. dOl/i lo vae and P. 
da/matica include the larger height or the rings, large r 
variat ion range of the dID ratio , smaller and more var i­
ab le ramification length, and possib ly smaller number 
of ramirica tion rows in a ring in P. dal/ilovac. Another 
difference , which has not been adequate ly emphasized 
to date , a nd cannot be observed from nume rical va lues 
a lone, concerns the shape of the ramifi cation s: in P. 
dallilo vae they are c learly fractured at their dista l e nd 
(RADOICIC, 1968, PI. I, Figs. 2-3; PI. 2, Figs. I , 3; PI. 
3, Figs. 2-4; PI. 4, Fig. 4; PI. 5, Fig. 4), which is nol vi s­
ible in P. do/matico (SOKAC & VELIC, 1978, Pi s. 6-
9). The latter has clearly phlo iophorous branches. grad­
ually widening from the base toward their d istal e nd. 
Besides, in P. dOllilo\'ae the li ghtly compressed ramifi ­
ca ti ons diverge both upward and downward from the 
horizonlal plane (RADO ICIC, 1968, PI. I , Figs. I -3; PI. 
3, Fig. 3; PI. 6 , Figs. 1,5 and PI. XVI, Fi gs. 13-14 in 
th is pape r) , w hereas in P. do/matico the ram if ications 



~ 
0 
~ 

Prall/rlol/ella Prall/rionella Prt/l/lr!oneffa Prall/r/Olle/ta Pra/urlollella Pralurlollella K, 
dal1ifol'ae safemilalla pejOl'icae adriarica jordallica neme ;: 

" (RA DO[C[C, [968 ) BARATTOLO, [978 (RADO[C [C, [969) (SOKAC &VELlC, 1978) (KU SS & CON RAD, [991 ) (DRAGASTA N CI aI., [978) ~ 
Il.comb. n.comb. 3 

" " 0 
< 

I-Ieighl of segments H • 0.95-1.45 • Hs 0.5-1.4 • 0,75-0,80 • 0.7-1.2 0.24-0.48 o· • 
Diameter of thallus in the level of segments 0 • 1.10-2.40 • Ds 1.3-2.2 • 1.2-3 .04 • 1.60-2.25 • 1.48-2.00 • 0,36 

.. 
~ • 0.44-0.92 • '" Diameter of thallu s between segments D' • 0.40-0.75 '-:> 0.48-0,62 • 0.55-0.84 ~ 
3 

Diameter of central cavi ty d • d; 0.6-1.0 • 0.29-0.72 • 0.52-0.79 • 0.18 
:;j' -• 

O. [9-0.40 0 
~ 

> 
Relat ion inner/ouler diameter dID 0 15-35% "'~ 24-38 % ¢ 22-32 % • 35-40% 32-48% " g' 

Distance between whorls or groups of whorls h • 0.40-0.72 0.29-0.64 Q 
g-
o 

Height of thallus between two segments h ' • 1,45- 1.60 • Hi 0.2·1.0 • 0.37,0.43 ~ 
• 0.079-0.095 

> 
Length of ramifications I 0 0.33-0,73 ¢ 0.58-0,68 • 0,69-0 .82 • 0.52-0.62 ;, 

0. 14-0.35 ~ 

Diameter of rami fications (distal) P(di"~I) • 0. 16-0 .30 • 0,19-0.32 • 0, 18-0.25 • 0.20-0.59 • 0.031-0.047 
1-0.40 0.05-0.13 

Diameter of ramifications (proximal) p (P. O\ ,) 0 30-35 JI • 35-70}1 • 0.012 

0.025·0.07 

Number of ramifications per segment • 60- 100 • 50-70('80) • 30-52 r:> 75· 120 • cca.40 eca.50 

Number of hori zonTal row s per segment 0 min. 2 2 3-4 • min. 4 3 

-l 
" cr 

- :.., ('") 0 - -
• "'\ 0 • ~ ~ 0' -. ~ '" 

v a~:"o:;:;iC\ 

; ~ g ng ~9: r ~· ~3 ~ 9. 
Q ;:3 ~3 ~3~§ ~~g~ 
I';;;;l = ;:!. :;:o"1:l ~.!! ~ :::: :::: 5'- ~ :::: 
c: ~ ~ » [ -g. 2i . ~ 3 5' ~ ~ ~. 
2 a· ~ 0 Co. ~ ~ ~ ~ ~ 2. ~ 
'" ;7 g 'D ~ -, (5 g' § ~. £, 

0'-0 0 "" - - ~ 

~ r~~ : HIB 
o en tr. '0- "' -

~ xc [ 9· ~ ~ ~. ~ 0: 



22 

are, in general, obliquely directed upward by varying 
angles (si milar as in P. salernitalla) and ohen bent like 
an arch toward the exteri or (SOKAC & VELlC, 1978, 
PI. 6, Figs. 1-2; P I. 7, Figs. 1-3), so that sometimes only 
the dis tal ends of the lowest row of ramifications can be 
situated below the horizontal plane, i.c. below the level 
of the lowes t row o f branches (SOKAC & VELlC, 
1978, PI. 6, Fig. 2 pars; PI. 8, Fig. I pars). The d iffer­
ences in the direction of ramification growth result in 
th e differen t shape of the ca lcareous rings-segments, 
which is ball -shaped in P. dani/ovae (sec reconstruction 
in BARATTOLO, 1978, Fig. 18), in contrast to P. dal­
mGlica where it can be envisioned as being rather bow l­
shaped. All the above rncnlioncd diffe rences seem 10 be 
sufficientl y clear \0 support the va lidity or p. 
da/matica. 

The strat igraph ic pos it ion of thi s species corre ~ 

spo nds approximate ly to thc Barremian s tage, though 
the small number of find s requires furthe r research. 

Pralurlollella pejovicae (RADOICIC, 1969) n.comb. 

PI. XV II, Figs. 8- 14 

Selected synonvmv: 

1969 Clypeilla pejovici n.sp. - RADOIC IC, p. 7 1-84, 
Pigs. J ~3 

1978 Likallella pejovicae (RADO IC IC) n.comb. -
BASSOULLET et aI., p. 145, PI. 5, Fi gs. 1-2 

1978 Selliporella pejovicae (RADO IC IC) n. comb. -
SOKAC & VELlC, p. 231 , not fi gured 

1986 Clypeilla pejovicae RADOIC IC - RADOl C IC:, 
p. 54, not fig ured 

1993 Likallella pejovicae (R ADOIC IC) - BUeUR, p. 
103, not figured 

1993 Falsolikallella pejovicae (RADOIC IC) - DRA­
GASTAN & BUeUR, p. 17 

In the original descript ion of Clypeilla pejovicae it 
was emphasized that thi s species has whorls consisting 
of two alternating rows of ramificat ions, which grad ual ~ 

ly and regu larly widen toward their arched distal ends 
(RADOICIC, 1969, Fi g. la-c, Fig. 2g, i, Fig. 3a-d). Two 
alterna tive possibil ities have been put forward fo r the 
ex istence of two rows of branches : firs t, that the fertile 
ramifications form two consecutive whorls , and second , 
that we are dealing with olle whorl with numerous ram­
ifications, arranged into two rows, which was cons id­
ered by RADOICIC (1969) to be a more likely al te rna­
tive. Based on the second possibility - two rows of 
branches with in a whorl - BASSOULLET et al. (1978, 
p. 145) assigned thi s spec ies to Likallella as Likalle/la 
pejovicae (RADOICIC) n.comb. , bas ing their decis ion 
on what was then generally thought to represent bas ic 
fea tures of the genus Likane/la. Radoicic later (RADO~ 

Ie le, J 986) maintain s it s assignmcnt to Clypeilla and 
thinks the re are no reasons to transfe r it to Likal/e/la, 
though she docs not give any argument to support her 
opinion. Rare fi nds o f this species, which a re, mo re-

Geo!ogia Croatica 49fl 

over, limited to the Dinaric Karst region, scem to be the 
reason for the general neg lect of thi s species and it s 
exclus ion from com para tive treatmc nts. In the es tab ~ 

lishment of Prafllrlollella, BARATTO LO ( 1978) 
ass igned to the new gen us Likanella? danilova e 
RADOIClC. but fail s LO mention Likanella pejovicae, 
in spite of the simil a rity between the two spec ies. 
KUSS & CONRAD ( 199 1) also fai l to mention L. 
pejovicae in their comparative table that accompanies 
the description of Pra!urlonella jordallica. All this indi ­
cates the fac t tha t P. pejovicae is considered, by the 
majority of workcrs, as being inadequately known and 
of unclear systematic pos it ion. 

The main charac teristics of this species can be sum­
marized as follows: phloiophorolls, c lub-shaped ramifi ­
cations; a rranged into two rows within a whorl (as in P. 
salemilalla; BARATTOLO, 1978, F ig. 13), or, maybe, 
two tightl y compressed whorls (as in P. danilovae; 
BARA TTOLO, 1978, r ig. 18); the ir arrangement along 
the tha llus in widely spaced, clearly pronoun ced seg­
ments; s ligh t downward bending of the l ower~row ~ ram­

ifications and s li ght upward inclina tion of the upper­
ro w ramificat ions. If a ll these a re propcrly assessed, 
thi s spec ies should undoubtcdly be assigned to the 
genus Pratlfr/Olle/la. Even more surpri si ng, therefore, is 
its proposal as Falsolikallella pejovicae (RADOIC IC) 
by DRAGASTAN & BUeUR (1993, p. 17), which was 
done wi thout any re levant argumentation. 

Thc comparison o f P. pejovicae w ith Clypeilla 0110-

gasli RADOICIC shows some interesting facts. Sadly, 
in the ori ginal description, C. onogosfi was illustra ted 
with only two poorly preserved sections (RADOIC IC, 
1986, p. 54-55, PI. 3, Figs. 1-2). Moreover, if one com­
pares the two descrip tion s - RADOI C IC (1969) for C. 
pejovicae and RADOI C IC (1986) for C. ollOROSli -
some ident ical features may be noti ced. In both spec ies 
the branches are rather swollen from the ir vcry base, 
tubular and slightly widened dis tally, and wit h smooth ~ 

Iy rounded distal ends. In P. pejovicae, the branches a re 
arranged in (wo rows, whereas C. ollogosfi is sa id to 
havc individual branches si tuated c ither above or below 
the whorl leve l as a conseque nce of too littl e space 
around the comparatively narrow axia l cavi ty to accom­
modate the branches int o one level. The Sllille reason­
ing, however, applies to P. pejovicae. In both species, 
the ramifica tion walls are weakly ca lcificd and, a lso in 
both spec ies, the branches are more c learly individual ­
ized toward the outer pe riphe ry of the skeleton (com­
pare PI. XVII, Figs. 8- 10 in the present paper w it h 
RADOI CIC, 1986, PI. 3, Fig. I). 

According to the original descrip tio ns the cl i ffe r ~ 

cnces in dimensions seem to be somew hat c lea re r, 
though th is should not be considered a dec isive feature, 
particularl y because the dimensions fo r C. ollogosfi 
apparently refer to on ly one speci men (sect ion ). Th e 
purely vi sual impress ion of two rows ramificati ons 
within a whorl in P. pejovicae and the tendency to 
atta in s uch an arrangement in C. ollogosf i d oes not 
seem re li ab le enoug h for the di sti nction o f these two 



Sakai:: Taxonomic Review of Some llarremian and Aptian Calcareous Algae ... 

species, which raises the question of the validity of C. 
ollogosti. Doubt is further strengthened by the fact that 
C. ol1ogosti (two specimens?) was derived from the 
same sample as P. pejovicae, Actinoporella podotica 
(ALTH), Sa/pingopore//a cOlli (RADOICIC), S. me/i­
tae RADOICIC, S. ex gr. geJlevensis, and P. danilovae. 
The same algal association has been identified in the 
analyzed sample from the Mljet Island (M1j -63). Beside 
the illustrated and many other sections of P. pejovicae 
(PI. XVII, Figs. 8-14), sections visually similar to C. 
onogosti have been observed, which may represent 
variet ies or atypical sections of P. pejovicae. However, 
for the time being, it is only necessary to draw attention 
to the problem of their delimitation or their identity , 
respectively, despite the inclination towards the latter 
solution. The definitive answer to this question must 
await until a more thorough analysis of more numerous 
and better preserved sections from the type- locality of 
C. onOROs[i is done. 

Based on this study and the ascertained stratigraphic 
levels, the tota l stratigraphic range o[ P. pejovicae cor­
responds, so far, to the Hauterivian and Barremian 
stages. 

Remarks on the species assigned to the genus 
Pratllr/ol1ella: Recently GRANIER & BERTHOU 
(1994) have established the new genus Milanovicella, 
with the most important distinguishing character bcing 
the two-rowed whorl s of branches arranged at equal 
distances along the thallus. Now, disregarding (for the 
moment) the genera Likal/ella and Draconisella, the 
following questions remain: to which genus - Pralllr ­
IOllclla or Milallovicella - should the species described 
above be assigned? The statement of a species having 
two-rowed or one-rowed whorls of branches seems [or 
the largest part to be hypothetical, (assu med on the 
basis of tangential sections, but ones that are too shal ­
low, i.e. do not show the siwation near the main stem) 
and is therefore a reflection of the author's interpreta­
tion rather than the facts. Consider the example of P. 
IIcrae, which will show, r hope , how prob lematic the 
question is. Judging by tangential sections alone (PI. 
XV, Figs. 5,11), P. nerac can be assumed to have two­
fold or three-fold whorls. Othcr scctions, however (PI. 
XV, Figs. 15, J8-19), show that at the same level of 
each of thc transverse section prescnted, two to three 
rings of pores of branches can be distinguished, cut at 
different distances from the axial cavi ty. The inner ring 
with numerous channelets cutting through the cylinder 
wall (PI. XV, Fig. 18) and the inner ring of very tiny 
pores nearcr to the axial cavity (PI. XV, Fig. 19) show 
that the number of pores is twice, or even more , thc 
number of large pores that rcpresent the branches of the 
more distant circle. In contrast to the aforement ioned 
tangential sections, this suggest one row, or at most 
two(?) densely packed rows , or branches per whorl. 
Because of all that - i.c., the not firmly es tabl ished 
number of rows o[ branchcs per whorl - the ge nus 
PratIfflonella, seems, for the time being, suffi cient to 

2] 

accommodate all related species with similar character­
istics. 

Genus Acroporella (PRATURLON, 1964) 
PRATURLON & RADOICIC, 1974 

Acroporella rado;cicae (PRA TURLON, 1964) 
PRATURLON & RADOICIC, 1974 

PI. XVIII , Figs. 1-20 

Selected synonymy: 

1964 Acroporella radoicici n.gen. , n.sp. - PRATUR­
LON, p. 177-179, Figs. 8-11 

1974 Acroporella radoicicae PRATURLON 
PRATURLON & RADOICTC, p. 17-20, Fig. la­
b, Fig. 2a-d, Fig. 3a-d 

1984 Acroporella radoicicae PRA TURLON 
LUPERTO-S INNT & MASSE, p. 339, PI. 33, 
Figs. 1-2 

1992 Acroporella radoicicae PRATURLON 
MANCINELLI, p. 9-13, PI. 6, Figs. l-5 

non 1993 Acroporella radoicicae PRATURLON 
LUPERTO-S INNI & MASSE, p. 300, PI. 4, 
Figs. 11-12 

This species was originally dcscribed by PRATUR­
LON (1964) as a cyl ind rical a lga wit h unsegmented 
skeleto n, characterized by simple, undivided, acro­
phorous ramifications, situated obliquely, and with an 
alternating arrangement in consecutive whorls. The 
ramificat ions are directed upward between of 15-4SO 
from the hori zontal plane. Later, PR A TURLON & 
RADOIClC (1974) revised the generic diagnosis, 
adding that long primary ramifications show incipient 
branching - " in clusters of button-like sccondary twigs" 
- at the distal ends. However, compari son between the 
section illustrated In the origina l description 
(PRATURLON, 1964, Figs. 8-11) and those illustrated 
in the revisio n (PRATURLON & RADOTCTC , 1974, 
Figs. 2-3) shows some additional differences: the cal ­
careous skeleton of the originally depicted specimens 
shows coarse-grained sparite, which is also the case in 
sections illustrated by LUPERTO -SINNT & MASSE 
(1984, PI. 33, Figs. 1-2), MANCINELLI (1992, PI. 6, 
Figs. 1-5), and in the present paper (PI. XVIII, Figs. 1-
20), but it is not visible in the sections illustrated by 
PRATURLON & RADOICIC ([974). Further, the sec­
tions illustra ted by PRATURLON & RADOTCIC 
(1974, Figs. 2-3) show an almost ideally rcgular alter­
nating arrangement of branches in consecutive whorl s 
with constant distance bctween the whorls, whereas this 
fcature is not so clear by far, nor so constant in the orig­
inally described sections (PRA TURLON, 1964, Figs. 8-
9), or in the sections presented by the above mentioned 
authors, ancl, particularly, in the sections figured in the 
present paper (PI. XVIII , Figs. 1-5,9, 13), which clear­
ly show not only irregularly alternating arrangement of 
branches (PI. XVIII , Fig. 2) but also variable distance 
of whorls. Another difference concerns the shape of the 



24 

TAN c1 al.) into the genus Pratur{onclla involved the 
transference of an inadequate holotypc (regardless of 
the statement in Article 7.2 of the ICB N that the 
nomcnclatorial type, i.e. holotypc, is not necessarily the 
most Iypical or representative clement of a taxon) and, 
consequently and morc important, an inadequate diag­
nosis. However because the above mentioned Article 

legalizes such a transfer and accepts the thus transferred 
taxon as valid, the necessity of a new diagnosis and a 
new description in accordance with real characteristics 
of that fonn appears imminent. 

Diagnosis: Small dasyclad al ga with cylindrical cal­
careous enve lope consist ing of ring-shaped segments 
bearing phloiophoroLls ramifications alternating with 
weakly calcified intervals of the thallus envelope, with­
out ramifications. Densely packed ramifications arc 
tightl y compressed within thin calcareous rings, result­
ing in their inclination into different directions. Inner 

surface of the central cavity is straight to slightly undu­
lated, slightly widcncd in thc central parts of the 
branches-bearing ca lcareous rings. 

Description: The ncw specics is bascd on the analy­
sis or numerous section s of fragments, among which 
several include longitudinal or tangential sections cut­
ting more than five branch-bearing rings. The existence 

of these rings is a consequence of calcium carbonate 
infillings of space left among the ramifications. The 

ramifications are arranged along the thallus in succes­
sive inlervals (P I. XV, Figs. 2A, 4,14). 

The calcareous envel ope bet ween the ramifications 
bcaring rings (segments) are very thin and weakly cal­
cified, so that the skeleton is easily breakable. This is 
probably the main rcason that the remains of this alga 
are mostly represented by single rings and only rare ly 
by skeletal fragments containing 3-4 rin gs . The inner 
surface is delineated by a st raight to sl ightly undulated 
wal!. The centra l cavity is s li ght ly enlarged at regular 
intervals correspond ing to the levels where the ramifi­
cations grow out, forming the calcified rings (segments) 
(PI. XV, Figs. 9- 10). 

The undivided, clearly phloiophorous ramifications 
seem 10 be situated in whorls consisting of 2-3 tightly 
compressed rows of ramifications (PI. XV, Figs. 5, 11 , 

Geologia Croatica 49/1 

16), which occur at regular intervals along the thallus. 
These feature s, tightly compressed whorls and a com ­
paratively large number of ramifications in a whorl, as 
well as the widening toward their distal end, give an 
alternating arrangement of ramifications in consecutive 
rings and lead to thcir inclination with respect to the 
horizontal plane. The cx istence of 2-3 densely packed 
whorls is indicatcd by thc section figured in PI. XV, 
Fig. 16, where a comparatively broad proximal end of 
the ramifications can be seen, as well as their mutual 
contact at the base and the well-visible entrance pores, 
connecting the ramifications with the central cavity, sit­
uated in two closely spaced levels. The same sit uation 
is suggestcd by the transversc scction figured in PI. XV, 
Fig. 15, which shows a regular circular arrangement of 
pore of equal-diameter, probably belonging to the same 
whorl, and , in addition, indicating that the ramifications 
of one whorl have an approximately equal inclination 
from the horizontal plane. Also the same is suggested 
by the tangential section (PI. XV, Fi g. 11) showing 
three rows of alternately arranged pores in the middle 
ring. The smaller diameter of the middle-row pores, 
compared with that of the lower- and upper-row pores, 
indicates that the middle-row pores correspond to a sec­
tion of those ramificati ons whi ch are orientcd perpen­
dicularly to the growth direction of thallus. The number 
of ramifications in a whorl seems to amount to 15-18 
and total number of ramif ica tions in a ring (segment) 
can be estimated to be about 50. 

Similarities and differences: Prall/rlol/ella I/erae 
(DRAGASTAN el al.) is Ihe smal les t species of Ihe 
genus currently described. It differs from P. dalli/ovae 
(RADOTC]C) by the dimensions of all measured cle­
mcnts and, particularly, by the shape of ramifications at 
their distal e nd, which in P. dani/ovac are more or less 
dislinctly widened (PI. XVI, Figs. 4, 13-14) or even vis­
ibly fractured. Further differences includ e s li ght but 
clearly visible widenings of thc central cavity in P. IIcr­

ae at the levels of tightly compressed whorls, which arc 
not present in P. dani/ovae. With regard to the type­
species, P. sa/erllirana BARATTOLO, P. IlCI'({C differs, 
in addition to being distinctly smaller, a lso by the 
widenillgs of the central cavity at the leve l of whorls, 
whereas ill P. salernirana the axial cavity at these lev-

Dimensions in mill: total range most frequently 

Maximum observedlcngth (L) 

Outer cli,lIneter of branch-bearing segments (D) 

Inner diameter Cd) 

The dID relationship 

Distance between two consecutive whorls (11) 

Length of ramifications (1) 

Width or ramirications, distal (P<li,tal) 

Width of ramifications, proximal (PPfOJ 

Num ber of ramifications per whorl (w) 

Number of ramifications per ring (segment) 

2.4 

0.44-0.92 

0.19-0.40 
32-4f~ % 

0.29-0.64 
0.1 4-0.34 

0.05-0. I 3 

0.025-0.07 

15- 18 

abollt50 

0.5R-0.SO 

0.20-0.30 

34-44 % 

0.40-0.60 

O. I 8-0.30 

0.05-0. 10 

0.03-0.04 

Table 5 Dimensions of Pmlll/,-
1m/ella I/C/'{/C (DRAGASTAN 
et al.). 



Sobi::: Taxonomic Review of SOllle Barremian and /\pliall Calcareous Algae ... 

branches. Sections figured by PRATURLON & 
RADOICIC (1974, Figs. a-c) show primary branches 
visibly widening toward the exterior, so that the pore 
diameter at Ihe distal end is twice the diameter at the 
proxima l end (i.e. , the branches arc club-shaped), which 
is visible neither in the originally figured sections 
(PRATURLON, 1964, Figs. 8-9, 11) nor in the present 
paper (PI. XVIII , Figs. I, 18-20). Both the original and 
the present material clearly show tubu lar branches that 
retain Ihe same diameter all along thei r length. Also, 
neither of the above mentioned sections, except those 
illustrated by PRATURLON & RADOICIC (1974, 
Figs. 2-3), show secondary ramifications . Oblique and 
tangentia l sections, sllch as those figured in the present 
paper (PI. XV III , Figs. 2, 4), that pass through !1llmer­
ous whorls and cut the branches at various distance 
from the axia l cavity (PI. XVIn, Fig. 4: the pores situat­
ed near the lower tip of the thallus arc being cut at their 
most distal parts), shou ld in all probability show the 
"clusters of button-like secondary twigs". Therc is not, 
however, the slightest sign of thcir prcsence. thereforc 
it follows, that the originally described materia l (as wcll 
as that figured by LUPERTO-SINNI & MASSE, 1984, 
MANCINELLI, 1992, and in the present paper) and thc 
"revised" material (PRATURLON & RADOICIC, 
1974) do not belong to the same species. Therefore I 
accept Acropore/fa radoicicae as it has been described 
and illustrated in thc original description (PRATUR­
LON , 1964). The materials that PRATURLON & 
RADOIC IC (1974) uscclto "revise" Ihe genus Acrop ­
orella should probably be treated in conncction with thc 
genus Trip{oporclla. 

A hitherto neglected feature of A . radoicicae, which 
is clearly visible in numerous sections (about 80) in the 
available material (which has been collected at several 
localities in the Dinaric Karst area), concerns the fis­
sures, or cracks, of the calcareous envelope. The cracks 
arc bcst visible at the ouler surface, where they are also 
at their widest , whereas they closc toward the interior 
(but can, in some cases, reach to ncar the axial cavity). 
The cracks arc irregular, forming either a mosaic- li ke, 
polygonal meshwork al the surface that encloses all the 
pores and thereby contributes to sccondary individual­
ization of the ramifications (PI. XVIII, Fig. 6), or they 
delimit largcr, irregularly shaped fields on the surface 
(Pl. XVlll , Figs. 1,2, 13), or arc scarcely pronounced at 
all (Pl. XVlll, Fig. 4). This [eature appears to be pre­
sent also in the tangential part of thc holotype section 
(PRATURLON, 1964, Fig. 8). It is probably a sec­
ondary characteristic, acquired during diagcnesis, but 
nevertheless can contribute 10 the very characteristic 
appearancc of this specics. 

The stratigraphic range of A. radoieicae can be 
defined as Barremian-Aptian, which seems to be well 
supported at numerous localities in the Outer Dinarides, 
as the stratigraphical position of each collected sample 
is well establ ished. 

25 

Genus Clypeina (MICHELIN, 1845) BASSOULLET 
et al., 1978 

Clypeina? solkani CONRAD & RADOICIC, 1972 

Pl. XIX, Figs. 1-6, 8-10 

Selected synonymy: 

1972 Clypeilla? solkani n.sp. - CONRAD & RADO­
ICIC, p. 87 -95, Figs. 1-3; Pl. I, Fig. 4; Pl. 2, 
Figs. 1-4 

? 1978 Clypeina solkani CONRAD & RADOICIC -
DRAGASTAN, p. 109-ll0, 1l3, Fig. IH-b 

1978 Clypeina? solkani CONRAD & RADOICIC -
VELIC & SOKAC, p. 317, PI. I, Figs. 2-7 

1982 Clypeilla solkani CONRAD & RADOICIC -
LUPERTO-SINNI & MASSE, p. 864, Tabl. 2 

1984 Clypeina'i solkalli CONRAD & RADOIC IC -
LUPERTO-SINNI & MASSE, p. 339, Pl. 33, 
Figs. 3, '14 

1986 Clypeina? solkalli CONRAD & RADOICIC -
SOKAC, p. 46-53, PI. 3, Figs. 1-5 

1987 Clypeina? solkani CONRAD & RADOICIC -
RADOICIC, p. 75, Pl. I, Figs. I pars, 2-3 

1992 Clypeina? solkani CONRAD & RADO IC IC -
MANCINELLI, p. 9, 13, Pl. 2, Figs. 1-3; PI. 3, 
Figs. 13-14; PI. 5, Fig. 10; PI. 7, figs. 7-9 

1993 Clypeina'I solkani CONRAD & RADOICIC, 
1972 - BODROGI, CONRAD & LOBITZER, p. 
64, PI. I, Figs. 3, 6; Pl. 2, Figs. 3, 6 

1994 Clypeina? solkani CONRAD & RADOICIC -
BODROGJ, BONA & LOBITZER, p. 233-245, 
PI. 3, Fig. 6; PI. 6, non Fig. 3, ?7, PI. 10, Figs . 
4-7 

Thi s alga, thoroughly described by its authors 
(CONRAD & RADOICIC, 1972), is characterized by 
widely spaced whorls of fertile ram ifications, open 
toward the exterior. Each whorl has the shape of a 
rather thick calcareous disc and bears 8-16 (most fre­
quen tly 10-14) ramifications. Ramifications of thc sallle 
whorl stay in mutual contact for 1/2 to 2/3 or their 
length. J\ significant remark made by the authors 
(CONRAD & RADOICIC, 1972) refers to the shape of 
the ramifications, which depends on their number with­
in a whorl. When that number is smaller, the branches 
are shorter and more strongly widencd toward thc extc­
rior, having a stocky appearance. \Vhcn therc arc more 
branches in a whorl, they are longer and ·more slender, 
compressed in their proximal half. Observations in this 
study suggest that Barremian specimens observed thus 
far generally have a smaller number (8 -12) of branches 
per whorl, which rcsults, as mcntioncd abovc, in their 
stocky, pear-shaped appearance. 

Clypeina? solkani is one of the most common 
species in the Dinaridic Karst region of Croatia. Its 
presence has bccn cstab lished in western 1st ria , along 
the coas tal region, and on the islands, at almost every 



26 Geologia Croatica 49/1 

Dimensions in mm CONRAD & RADOICIC 

1972 
Barremian 
specimens 

Outer diameter (D) 

Inner diameter (d) 

The diD relationship 

Distance between whorls (h) 

Distal diamete r of branches (p dis!) 

Proximal diameter of branches (p prox) 

Number of branches per whorl (w) 

0.52 - 1.20 

0.15 - 0.31 

8-16 

locality where Lower Cretaceous deposits are exposed. 
Its s tratigraph ic range can be defined as ranging from 
the Haute rivian to the Mid~Upper Barremian. In Hau­
terivian deposits , it is associated with Sa/pingoporella 
anflu/ata, Praturlonclla pejovicae, P. dani/avae, Humi­
ella leu rae, and numerous unidentified dasyclad 
remains. Among the foraminifera, the majority belong 
to the long-ranging forms , but Campanel/u/a capuel1sis 
(DE CASTRO) is siratigraphically imporlanl. In Bar­
rem ian deposits , C.? solkani is accompanied by a rich 
algal association composed of Salpingopore//a mueh/­
bergii, S. me/ifae, S. biokoviellsis, S. dinarica, Sa/pin­
gopore//a sp., Prafur/olle/la dani/ovae, P. pejovicae, 
ACfinoporel/a podolica, and a C/ypeina sp. In the origi­
nal description of this species, the possibil ity of its Apt­
ian age has been assumed. However, this supposit ion 
has not been proven so far, just as the associa ted fossil 

assemblage at the type-locality docs not prove this age. 
There arc some indications that C.? solkani possibly 

occurs as early as Upper Valanginian, but so far no 
uneq uivocally identifiable sect ions have been found. 
Transverse sections which arc often determined as C.? 
so/kani Illay be identical with the scct ions of a so far 
uniden tified Sa/pin[iopore//a. 

Clypeina gigantea n.sp, 

PI. XVII , Figs. 1-7 

Origin of the name: This species is the biggest 
spec ies of C/ypeina described so far. 

Type locality: Road Babino Polje - Sobra, on Ihe 
Island of Mljct; road cui about 2.15 km before Sobra, 
i.e. ISO m before the point marked by coord inates men­
tioned in the description of S. verrucosa (Fig. 1). 

Type stratum: Well-bedded, light grey, biolurbatcd 
algal wackes toncs with skletal-moldic cavities of green 
algae and gas tropods (Fig. 2). Bioturbalions arc fil led 
with peloidal grainstone and small benthic foram inifera. 
Depositional envi ronment may be defined as shallow 
sublidal. 

Holotype: Oblique section figured in PI. XVII, Fig. 
2; thin scction Mlj -63/2 1. Thin sections made from the 
topotype material are stored at the Institute of Geology, 
Zagreb. In total, eight sections of this species were used 
for the description. 

0.48 - 0.67 

0.19 - 0.28 

34 -47 % 

0.14 - 0.25 

0.10 - 0.20 

0.02 - 0.05 

8-12 

Table 7 Dimensions of Clypeina? 
solkani CONRAD & RADO­
ICIC. 

Diagnosis: Large C/ypeina with distinctly spaced, 
saucer-shaped whorls of numerous club-shaped ramifi­
cations and with deeply indented rims. 

Further research may show larger variation ranges, 
as dimensions refer 10 only 8 specimens (Table 8). 

Description: A vcry large Clypeina, whose skeleton 
is preserved as translucent, coarse-grained sparile. Cen­
tral caviLy appears regularly cylindrical between the 
whorls and with relatively thick calcareous wall. Sec­
tions across that tubular parl of the skeleton suggest the 
existence of tiny longitudinal tubulcs within the wall. 
These cannot be connected with the primary ramifica­
tions nor can they be identified with the pores on the 
corresponding parts of the skeleton in C. jurassica. In 
transve rse sections cutting the cylindrical skeleton, 
these tubules appear as tiny, irregular and sometimes 
indistinci pores (PI. XVII, Fig. 3) Ihal arc partly visible 
also in Ihe holotype (PI. XVII, Fig. 2). 

The ramifications are phloiophorous and, starling 
from a comparatively wide pore (Pi. XVII, Figs. 4, 7), 
they gradually and regularly broaden toward their distal 
end (PI. XVII, Fi gs. 2-4), sometimes somewhat more 
abruptly within the first half of their length. Each 
branch has its own calcareous envelope. In their proxi­
mal part , the envelopes, in general but not always, arc 
in mutual contact. Along their distal half, or more, the 
enve lopes of the neighbouring branches are scparated 
from each other by narrow and regular, non-calcificd, 
deep indcnlations (PI. XVII, Figs. 3-4). Rarely, Ihe cal­
CaI'cous envelope bends toward the interior at the tip of 
the branch, which suggests the ir bcing primarily closed. 
The available sect ions indicate the oblique position of 
the ramification, directed slightly upward. Terminal 
swelling of ramifications results in their being shifted 
somewhat below or above the fictitious wavy saucer 
shaped surface. Such ramifications would then appear 
as more pronou nced ribs on that wavy surface (PI. 
XVII, Fig. 6). 

Similarities and differences: Clypeina gigantea 
n.sp. belongs to the group of large and quite typical 
species of the genus. With some of its dimens ions 
(maximum D values), Clypeina jurassica may come 
rather c lose to C. giganfea, but C. jurassica shows a 
much larger variation range, perpendicular to subper­
pendicular position of branches to the main stem, yel­
lowish calcite in the skeletal structure, thin dark micri-



Sok<ll:: Taxonomic Review of Some Barremian and Aplian Cdcareou$ Algae ... 

tic lin ing around the axial cavi ty, and clearly visible 
tiny pores w ithin Ihe walls of the cylindrical sta lk 
between the whorls, which is not present in the ne w 
spec ies. By its outer diameter, C. gigantea falls within 
the variation range of the poorly known C. hallabaren­
sis Y ABE & TOYAMA, but C. giganlea has a much 
narrowe r central cavi ty and larger distal ramifica ti on 
diameter, is so much more s tron g ly calcified, and the 

ramifications are c lo ser to a horizontal position. C. 
gigal/lea is clearly larger than most Mesozoic C/ype ina 
species (c. ealinilia CAROZZT, C. inopinata FAVRE -
possibly synonymo us with C. jurassica, see BASSO­
ULLET el a I. , 1978; C. marleli EMBERGER, C. parvlI­
la CAROZZT, C.7 salkani CONRAD & RADOICIC, C. 
zlIlIIelae JAFFREZO & FOURCADE); moreover, il d if­
fers from each of the above mentioned species, taken 
individual ly, by the degree of calcification, inclination 
of branches, number of branches in a whorl , shape of 
the branches, or some other fea ture, which makes it eas­
il y distinguishable. By its much larger size, C. gigantea 
c learly differs also from the strati graphically c lose C. 
nigra CON RAD & PEYB ERNES (~ Aclinoporella 
nigra (CON RAD & PEYBERNES ) GRANIER , 1994) 
as well as from the poorly known, and probably ques­

tionable, C. OIlOgosli RADOICIC (see a lso the d isc us­
sion concerning Pratur!ollel/a pejovicae), which was 
derived from the identical Barremian algal assoc iation. 

C. gigal/tea is twice the size of C. neretvae RADOICTC 
(accordi ng to the recommendat ion of the ICBN, the 

specific namc should be "I/ere/valla"), which is, more 
over, morc wcakly calc ificd, particularly the cylindrical 
parts of the stalk between the whorls, and has more 
irregularly shaped ramifications which, in contrast to C. 
gigal/lea, become sli ghtly thinne r toward their dis tal 
end. C. giganfea is a lso clearly d istingu ishable from C.? 
o/ra wii RADOICIC, which has a much broader axial 
cavi ty, larger number of ramifications in whorl , shorter 
ramifications, and more closely spaced whorls than C. 
gigallfea. C. pastriki RADOICIC is much small er, its 
branches are shorter than the diameter of the central 

cavi ty and their number per whorl is about 1/3 of that in 
C. gigalltea . C. estavezii GRANIER (1986) was inade­
q uale ly descri bed an cl poorly illuslrated (GRANIER, 
1986), so that we could only gene rally say that it is 

Di mensions in mm: 

27 

small e r and with more widely spaced whorls propor­
lionally to its size. C . gigantea also clearly differs from 
Pa laeogene species of Clypeina, by its larger si ze, a 
rather constant diD relationship (18-20%), shape and 
length of ramifications , and somet imes also by the 
number of ramifications per whorl. 

Stratigraphic position: Clypeina giganfea n.sp. has 
been found in a sample with a rich algal association, 
composed of numerou s Salpillgoporel/a melifae, S. 
mlfch/bergii, transitional 1"01111S between S. melilae and 
S. hasi , and less nume rou s S. hispanica, Clypeina? 
solkani, Praturlonclla danilovae, P. pejovicae, and 
numerous unident ified dasyelad fragme nts. Among the 
foramin ifera, there are Campane/lu/a capuel/sis (DE 
CASTRO), Mayl/cil/a bl/Igarica LAUG cl aI., and Ver­
corsella scarsellai (DE CASTRO). Aboul 20 m above, 
in the continuou s sequence, S. biokoviensis and S. 
{trladallasi have been fo und. The wholc microfossi l 
association, as well as the posi tion of the algal -bearing 
sample in the cont inuous sequence al the type locality, 
suggest that the stratigraphic position should be defined 
as Lower Barremian. The above mentioned foraminifer­
al species do not contradict that agc, though they arc 
more common, in th is area, in the Upper Hauterivian. 

Genus Actil/oporella (GUMBEL in ALTI-l, 1881) 
CONRAD, PRATURLON & RADOICIC, 1974 

AClilluporella podolica (ALTI-l, 1878) CONRAD, 
PRATURLON & RADOICIC, 1974 

PI. XX, Figs. 1-3 ,5-13 

Select ed synonymy: 

1878 Gyraparella podalica n. sp. - ALfH, PI. 6, Figs. 
J -8, non ri g. 5 

1881 AClil/oporelia podotica (ALTH) n.gen., n.comb. 
- GUMBEL in ALTH , p. 322 ( 140) 

1974 Aelinoporella podotica (ALTH) emend. - CON­
RAD, PRATURLON & R ADOICIC , p. 1-1 5, 
Figs. 1-12 

1975 AClinoporelia podotica (ALTH) - DRAG A S­
TAN, p. 66, PI. 33, Fi g. 3; PI. 36, Fi g. 4; PI. 37, 
Figs. 1,74; PI. 38, Fig. 1; PI. 51, 7Fig. 2 pars; Pl. 
52, Figs. 2, 4; PI. 71, non Fig. 2 

OUler diameter of whorls (D) 

Outer diameter of sta lk between whorls (0 ') 

Inner diameter of thallus (d) 

The dID relationsh ip 

3. 12 - 4.35 

0.94 - 1.20 

0.54 - 0.82 
8 - 20 % 

Distance between consecu ti ve whorls (h) 

Length of rmnifications (1) 

Diameter of ramifications at distal end (pd) 

Diamclcr of ramifications allhe base (db) 
Numbe r of ramifications per whorl (w) 

0.60 - 0.72 

1.00 - 1.70 

0.28 - 0.44 

0.08 - 0. 14 
25 - 28 

Tabte 8 Dimensions of Clypcilla gigallfea n.sp. 



1978 Actilloporel/a podolica (ALTH) - DRAGAS­
TAN, SUCUR & DEMETER, p. 22-23, Pl. 4, 
?Fig. 4; PI. II, Figs. "3, 4 pars. 

1978 Actilloporel/a podolica (ALTH) - DRAGAS­
TAN, p. 11 0-115 , Figs. I c-e, PI. 3, Fig. I; PI. 5, 
Fig. 3. 

1984 Actilloporella podolica (ALTl-I) 1878 emend. 
CONRAD et aI. , 1974 - LUPERTO-SINNI & 

MASSE, p. 339, PI. 33, Fig. 9. 

1986 Actilloporella podolica (ALTH) emend. CON­
RAD et aI., 1974 - GRA NIER, p. 47-48, PI. 9, 
rigs. e-h. 

1989 Actilloporel/a podoliea (ALTI-I) DRAGAS­
TAN, p. 28, PI. 14, ?Fig . 4, non Fig. 5 

1994 Aetinopore/la Jiodolica (AL TH) - GRANIER, p. 
113-117, PI. I. Figs. 1,4,8-10; PI. 3, Figs. 5-8 

1994 Actilloporel/a Jiodoliea (ALTH) CONRAD, 
PRATURLON & RADOTCIC - SCHINDLER & 
CONRA D, p. 73-74, PI. 3, Figs. 1,3,7-8. 

AClinoporella podolica (ALTH) is a well-known 
species which has been thoroughly rcdcscribcd and 
illustrated by CONRAD et al. (1974) . Un fortunat ely, 
the authors chose as neotypc a washed-out spec imen 
which docs not allow insight into the structure and 
shape of the ramifications, and whieh are indispensable 
for the proper generic and species definition. Phloio­
phorous branches are arranged into distinctly separated 
whorls . The essential generic characteristic is the 50-

called corona, which is the swelling of branches at their 
base and which is in A . podolica equally well expressed 
on both the lower and upper sides of a branch. Howev­
er, the sections illustrated by SCHINDLER & CON­
RAD (1994, PI. 3, Figs. 7-8 ), as well as those in the 
prescnt paper (PI. XX, Figs. 1,5, 10), suggest thc possi­
bility that the corona appears to differ from how it was 
depicted in the rcconstruction of CONRAD et al. (1974, 
Fig. 5) taken ovcr also by GRANIER (1994, Fig. I), 
and, instead, seem to give the impression of stunted or 
incompletely developed second3lY ramifications which 
also show a phloiophorous shape. This proves the 
importance of good longitudinal sections for good 
species determination. Isolated rows of mutually con­
nected rings, which may partially correspond to whorls 
cut through the distal part of the branches, have fre­
quently been ascribed to this species. However, some 
similarly constructed species (c. g., certain Clypeinae) 
may show almost identical sections (also cutt ing 
through the outer parts of isolated whorls), which 
makes the identification of such doubtful sect ions rather 
risky. This, in turn, has further implications rcgarding 
the stratigraphic position, geographic di stribution, and 
ecological requirements of this species. \Nh ile single 
occurrences of A. fJodolica in the Dinaridie Karst area 
of Croatia, have been recorded in the Upper Maim, its 
common occurrence marks the Lower Cretaeeolls, up 
to, and including, the Aptian. 

Geologia Croalica 49/1 

Genus Piriferella n.gen. 

Type of the genus: Pin!erella spillosa n.gen., n.sp. 

Origin of the name: The gencrie name derives 
from lhe typ ically piriferous shape of the ramifications. 

Diagnosis: Cylindrical thallus, with thin calcareous 
envelope, bears apparently densely arranged whorls of 
large, piriferous, alternately arranged ramifications, 
each ramification growing out independently and being 
completely enveloped by its own thin calcareous enve­
lope up to its distant tip. 

Pir~ferella spinosa n.gcn., n.sp. 

PI. XXI, Figs. I-IS 

Origin of the name: The name derives from the 
outer spinose appearance of the thallus, caused by out­
wardly thinn ing ramifications. 

Type locality: Road Babino Polje - Sabra on the 
island of Mljet; road cut aboot 2 km before Sobra 
( 17°36'00" E and 42°44'02" N, Fig. I). 

Type stratum: Well-bedded, lightly coloured, 
algal-pe lletal wackestones/packstones with fenestral 
fabric and vadose crystal sill. Upper Barremian (for 
details see stratig raphi c position of Safpingoporc/la 
verrucosa and Fig. 2). Depositional environment: peri­
tidal shallowing upward cycles with black pebble brec­
cia. The algal-bearing samples mostly derive from the 
firsl, subtidal, member: algal and/or algal-pelletal 
wackestones, and rarely [rom the second, intertidal, 
member with fenestral fab ri c: fenestral algal-pelletal 
wackestones with vadose fabric. 

Holoty!'e: Oblique section figured in PI. XXI, Fig. 
6, thill-section Mlj-81/28. Paratypes (PI. XXI, Fi gs. 1-5, 
7-15) derive from the same sample. All material is 
deposited at the In stit ute of Geology, Zagreb. The 
description is based on 20 variously oriented sections 
from the same sample. 

Diagnosis: Alga with a cylindrical thallus, thin cal ­
careous envelope, bearing apparently dcnsely arranged 
whorls of large, typically piriferous, alternately 
arranged, obliquely situated, upward directed ramifica­
tions . Ramifi cat ions grow out independently, each 
within its own th in calcareous envelope, and are closed 
at their distal ends. Ramifications of the same whorl or 
of two consecutive whorls may be separated from each 
other or they may touch mutually at their maximum 
swellin gs. Secondary calcite deposits in the proximal 
region (be tween the ramification basses) give an 
impression of a thicker wall and apparently closer con­
tact between the ramifications. 

In total, 15 sections have been measured. Because 
of destruction, the outer diameter (D) may be somewhat 
larger. 

Description: A comparatively sma ll dasyclad alga, 
having a weakly calc ifi ed thallus that consists or a 
cylindrical axial cavity with a thin calcareous envelope 



Sob/:: Taxonomic Review of Some Barremian and Aptian Calcareolls Algae ... 29 

Dimensions in mm 

Outer diameter (D) 

Inner diameter (d) 

The dID relationship 

Distance between conseclltive whorls (h) 

length of ramifications (I) 

M aximum diameter of ramifications (p max) 
Diameter of the cntrance pore (p pro x) 

Number of ramifications per whorl (w) 

0.48 - 0.S7 

0.20 - 0.34 
34 - 50% 

0.10-0.t4 
0.22 - 0.34 
0.10 - 0.19 
0.04 - 0.05 

7 - 8 

(PI. XX I, F ig. 2) and bears apparently densely spaced 
whorls of undivided, typically piriferous ramifications. 
The inner surface or"the calcareous cylinder is smooth, 
sharply delimited, and perforated by alternately 
arranged ramification entrance pores. Axial cavity is 
comparativcly broad. The independently growi ng, typi­
cally pirifcrous, ramifications (i.e. proximally swollen 
and tapering distally) give a spinose outer appearance 
10 thc thallus. 

Undivided, typically piriferous branches, arranged 
into alternating whorls, grow out independly from each 
other, each ramification being completely coated by its 
own th in calcareous cnvelope (PI. XXI, Fig. 10). The 
communication between the axial cavity and the branch 
is performed through a short pore, sit uated in the mid­
dle of the ramification's base or, sometimes, below it, 
as the branch 's swelling is more pronounced upward 
(PI. XXI, Fig. I). After the en(ranee pore, each ramifi ­
cation abruptly and regularly swells and then gradually 
tapers toward its distal end. While the branches of the 
same whorl, or of two consecutive whorls, are regularly 
separated from each othcr, they often touch at their 
thickest part (PI. XXI, Fig. 6). Narrow spaces between 
the branches at their base are completely (PI. XXI, Fig. 
12) or partially (PI. XXI, Figs. 9-10) filled by sec­
ondary calcite, wh ich makes the ca lcareous envelope 
secondarily thicker. The alternating arrangement of 
branches in the consecutive whorl s (PI. XXI, Figs. 7, 
12) may not a lways be wholly regu lar, depending on 
the intensity o/" swelling of the proximal parl or a par­
ticular branch. The branches are directed sli ghtly 
upwa rds, with their distal ends more strongly bent 
toward the main stem. Though the cysts have not been 
observed, the strongly swollen branches suggest the 
cladospore type of reproduction. 

Similarities and differences: PirUerella spinosa 
n.gen. , n.sp. is distinguished by its peculiarly shaped, 
spinose thallus, wh ich is the consequence of the shape, 
arrangement, and mode of growth of the branches, and 
the degree of calcification . At thc gcneric lcvel, Pir­
iferella n.gen. clearly differs from Clypeina, the latter 
being characterized by distinctly spaced whorls, with 
indentcd rims, this being a consequence of branches 
rnerging together in their prox imal parts and being 
divided only in thei r exterior (distal) parts. In contrast, 
Pir~ferella n.gen. has more closely spaccd whorls, in 

Table 9 Dimensions of PiriJerella spinosa n.gen., n.sp. 

which the branches are allernately arranged (which has 
not been observed in Clypeina) and which may come 
into mutual contact , both within a whorl and betwecn 
the consecuti ve whorls. The same features make Pir­
(lerella different to Actinoporel/a, the latter being addi­
tionall y characterized by the ex istence of "coronas" (i.e. 
vestibule-like basal widening), after which thc ramifica­
tion shrin ks and then again widens distally (A. 

podolica). By the arrangement and dens ity of branches, 
and their independent mode of growth, all of which 
rcsults in a similar general appearance of the thallus, 
Piriferella n.gen. comes more closely to Similiclypeina, 
which is, however, characterized by Actinoporella-type 
ramifications (i.e., graduall y widening toward exterior) 
and by vertically flattened coronas. In spite of the simi­
larity regarding the arrangement and density of ramifi­
cations and their thin calcitic envelopes in H%sporel/a 
(which makes this genus sim ilar to Piri!erella), there is 
a clear difference regarding the shape of the branches: 
in Holosporella, the branches are spherical to hemi­
spherical, in co ntrast to clearly piriferous to phloio­
phorous shape in Pirijerel/a. PIA (1930) has already 
emphas ized Ihat different gcncra may have the same 
arrangement of sporangia. This, of course, may result in 
the very similar or even identical visual appearance of 
different genera in certain sections. Pinjerella spinosa 
seems to be most similar to Clypeina somalica CON­
RAD et al. (1983), which was subsequently assigncd by 
BUeUR (1993) to his new genus Similiclypeina. In 
fact , thc two species are so similar as to belong to the 
same gen us, which Icd to thc proposal of Pin/erel/a 
somatica n. comb. (sec below). Thc simi larities bctween 
the two specics concern the similar dimensions within 
the same variation ran ge. A morc pronounccd differ­
ence, according to thc original dcscription (CONRAD 
et aI., 1983), is visible only in the "h" value , which 
seems 10 be twicc as large in P. somatica as in P. spin­
osa. As a consequence of this, according to CONRAD 
et al. (1983), the consecu tive whorls in P. somalica 
never touch (though it seems tha t they do, but not as 
c losely as in P. spinosa). While in P. spinosa the 
branches grow independently from their base, their 
envelopes almost always touch at the places of maxi­
mum swell ing. FurthemlOre, P. somatica is said to have 
egg-shaped ramifications, as distinct from P. spinosa 
where the branches are typically piriferous and directcd 



JO 

morc steeply upward. Though P. somalica still seems to 
be inadequate ly known, the above mentioned differ­
e nces seem sufficic!1I 10 justify the separat ion of the 
I wo species. 

Stratigraph ic position: PiriJerella spinosa n.gen., 
n.sp. is assoc iated wit h Sa/pillgopore/la ve,.rucosa 
n.sp. , where the di scussion or its stratigraphi c posi tion 
is g iven. 

Piriferella sOlllalica (CON RAD, P EYI3ERNES & 
MASSE, (983) n.col11u. 

PI. XXlI , Figs. 1 ·20 

Selected synonymy: 

1983 Clypeilla sOll/alica n.s p . . CON RAD , PEY· 
BERNES & MASSE, p. 93·95 , PI. 6, Fig. 1· 12 

1992 Clypeil/o? somatica CONRAD , PEYBERNES 
& MASSE · MANCINELLI , p. 9· 13, PI. 2, Figs. 

4 ·7, II 

1993 Sill/ilic/ypeilla a rr. sall/atica (CO NRAD , PEY· 
BERNES & MASSE) n. eomb . . BUCUR , p. 
106·107, PI. 3, Figs. 7, 9b, 10, 12·25; PI. 4, Figs. 
4·11. 

? 1993 Clypeil/a ? somalica (CONRAD, PEYBERNES 
& MASS E) . LUPERTO·SINNI & MASSE, p. 
300, PI. 5, Figs. 1· 2 

This species was described on comparatively scarce 
and recrys ta ll ized mate ria l and figured wit h a few COI11-

plete but damaged sections that do not show all the rel­
evant fea tures. Thi s is probably the reason that since 
[hen the species has seldom been menti oned and even 
more rarely illus trated. The sectio ns fi gured by CON­
RAD e[ al. (1983 , PI. 6, Figs. 1· 12) seem [ 0 belong [0 

he terogeneous ma te r ial , that has been united int o a 
spec ies. O ne group of re la ted secti ons incl udes the 
holo[ype and [he para[ypes (CO NRAD e[ aI. , 1983, PI. 
6, Figs. 5, 8, 12) and al so some less instructi ve sect ions. 
Another group includes sec tions from Prcstat's material 
(op. Ci L, Fi gs. 4, 10). Heav ily recrys talli zed sec tions, 
part ly belonging to the type materia l (op. CiL , ri gs. 6, 7, 
9), seem to include heterogeneous mate ria l, and Ihe sec-

Gcologiil Croalica 49/1 

lion figured in Fig. II (op. c it. ) is also probl e matical. 
'rhus, sec tions in Figs. 6 and 11 hin t at the possibi lity of 
vCl1 ically Ilallcned ramifications. Such unclear rc:lati ons 
prevent thc full ins ight into the actual characteri s ti cs of 
the s pec ies, which , in turn , makes it s dete rmination 
exceed ingly difficult (and even proble matic). The lack 
o f s te rile rami ficat ions (whic h could not be ide ntified , 
in cont ras t 10 C. j urossica), as well a s ti ghtl y spaced 
whorls (a \ least in their proximal parts) , led CONRAD 
ct al. ( 1983) to cxprcss somc doubts as to the attribution 
of thi s species to Clypeilla. The latter characte ri sti c, the 
closeness of consecu ti ve whorl s, i.e. ramifications , in 
the proximal region, w here the space was infill ed by 
calci te mass, was deemed decisive [or the attribution o f 
[his species [0 Simiticiypeilla (B UCUR, 1993). 

The genus Si111iliclypeino was es tab lished wit h S. 
cOl/ radi BUeUR as type spcc ies . It has peculiar, hori­
zontally elo ngated, (i.e., verti cal ly llattcned) swe llings 
in [he proximal part or [he branches (BUCUR , 1993, 
Fi g. 3), very s imilar to the coronas in ACfin oporello 
podol ica, the only diffe rence bei ng that in S. conradi 
these s we ll ings are hori zoll ta ll y e longated , in contrast 
to A. jJodolica w here they are vertica ll y e longated 
(CONRAD e[ aI., 1974, Fi g. 5). This feature, [he corona 
or ves ti bule-like swellings, which seem to actually rep­
resent ineompletcly deve loped secondaries (see PI. XX, 
rigs. 1, S, 10), and which is thc main diagnos tic feature 
[or di s tingui shing AClifloporello from Clypeina (e.g., 
ACfinoporella mas/ovi PR ATURLON is a ss igncd to 
Clypei llo due to the lack oj" coronas), was neglec tcd by 
B UCU R ( 1993). He based his establi shme nt or a new 
genus on only one characte ri sti c : the dense , more close­
ly spaccd whorls of ind ividua li zed ramifi cat io ns th at 
often mutua lly touch, and hence introduced a number of 
the new combinations: Similiclypeino alrmvii (RADO­
IC I(:), S. somotico (CON RAD e [ al. ), S. pi/p"olensis 
(SO KAC). The refore, Similiclypeillo would include 
fo rms with d iffere nt c haracte ri s tics: form s w it h hori ­
zontally e longated, corona- or vestibule- like swellings 
in the proximal part of the ramifications (5. cOllradi and 
S.? iusfilliani), form s with 110 Sti ch swe l1ings at a ll (S . 
a/rawli and S. somalica ), and fOfms with two types of 
secondary ram ificat ions, one growing out from the 

Dimensions in mm: Clypeil1a soma/jea Simi/jclype jllu Pirijerella 

Outcr diamcter (D) 

In ner diameter (d) 

dID relationship 
Distance between 

consecutive whorl s (h) 

Length of branches (I) 

Diameter of branches (p) 

Number of branches per whorl (w) 

CONRAD et aI., 1983 

0.47 · O. 80 
0.30 · 0.42 
36 ·59% 

0.23 
0. 16 · 0.23 

Ppm,. 0.04 
6 - 7 

aff. somolica sOlllolica 

IlUCUR,1993 thi s pnper 

0.33 . 0·54 0.34 · 0.68 
0.16 · 0.23 0.1 5 - 0.38 
30 ·46 % 38·55 % 

0.07· 0.09 0.10 · 0. 14 
0.19 · 0.23 0. 15 · O.IS 

p""O.OS·O.11 P.""O.OS - 0.1 3 
5 · 6( 11 ) 6 · 7 

Table 10 Compari son or the dimensions of Ciypeil1a soma/icll CONRAD el aI. , Similicfypeina aff. ,mll/aliea BUCUR and Piriferella .tollltllictl. 



Sobi':: Taxonomic Review of Some Barremian and Aptian Calcareous Algae ... 

cylindrical stalk of the thai us , another on the primary 
branches (5. pupnatensis) . As the basal swelling of the 
ramifications is generally agreed to be crucial for dis­
tinguishing AClil1opore/la from Clypeina, and thus 
became, by general consensus, a criterion of generic 
rank, it would be iltogical to e rect a genus that wou ld 
unite forms with and without such an important feature. 
Alternative ly, if we agreed that a feature serves as a 
generic trail for di st in guishing Clypeina from Actino­
porella, the same feature should have the same value in 
assignin g similar forms to Simi/ic/ypeina. This is the 
main reason for removing S. somalica from the genus 
5imiliclypeina and its attribution to the new genus Pir­
i/erel/a. 

In thi s paper, Piri/erel/a sOll/olieo (CONRAD et al.) 
n.comb. is presented with several sections, all of which, 
except one (PI. XXII, Fig. 11), were derived from the 
same sam pic of Barremian limestone from western 
1st ria. Th is enable get be tter insight into the morpholo­
gy of this form and provided some minor amendments 
to its description. This is a comparatively small dasy­
clad alga, whose thallus bears densely spaced whorls of 
ramifications. The shape of the ramifications was 
described as being egg-shaped. Almost completely indi­
vidualized branches are thoroughly coated, each with 
its own thin calcareous envelope (PI. XXII, Figs. 4, 6, 
9). More secondary calcite deposits infill spaces only in 
the most proximal region and thus connect the individ­
ual branches at their very base. In tangentia l sections, 
the ramifications appear more clearly separated than in 
P. spinosa, and the pores have a more or less oval to 
irregu larly elliptical shape (PI. XXI[, Figs. 2, 4, 8, 18), 
indicating that the ramifications are slightly horizontal­
ly flatt ened. Thi s is diffcrent in P. spinosa, where, in 
corresponding sections (deeper tangential sections), 
pores cut at approximately the same level , appear more 
or less regularly circular. The arrangement of branches 
in consec utive whorls is clcarly alte rnating. The 
branches are perpend icular to subperpendicular to the 
main stem (PI. XXII , Figs. 6, 9). 

Th e compari son of the dim ensions shows general 
agreement between P. spinosa and P. somalica. In P. 
soma/ica, the dis tance be tween consecu tive whorl s 
appears to bc somewhat larger in the original descrip­
tion than in later quotations. The identical p values in P. 
spinosa and P. somalica may be superficially mislead­
ing: while in P. spinosa the maximum swelling of 
branches occurs in their proximal part , in P. somalica 
the same value is attained in the dista l part. This high­
lights the main difference between the two species as 
being the shape of ramifications and their more pro­
nounced regularity in P. spinosa. In the Dinaric Karst 
area, P. somalica has so far been found at two, rather 
distant, localities : western Istria and the west side of the 
Neretva Rivcr, upstream from Mostar. Its stratigraphic 
posi tion at both localities is Barremian; it is associated 
with Sa /pingopore//a mueh/bergii, a Sa/pingopore//a 
sp. , Prafur/onella ncrae, Cylindroporella ivanovici, and 
a Cylilldroporella sp. 

31 

Acknowledgements 

T am grateful to my colleagues L. FUCEK, N. 
OSTRIC and B. KOROLTJA for thc loan of fo ssil ifer­
ous sample from the islands Dugi otok, H var, eres and 
Mali Losinj. Spec ial thanks go to I. VELTC for long­
last ing fiel d collaboration, the determ in ation of 
foraminifera, and the loan of the Cretaceo Ll s sam ples 
from Ist ria. Thanks go also to J. TTSLJAR for the sedi­
mentological analyses of the slides and the determina­
tion of th e palaeoen vironm ental sett in gs and T. 
GRGASOVIC for the help in technical preparation of 
the manuscrip t. I am especially grateful to I. GUSIC 
and to M. CONRAD for their reviews of the manu ­
script. 

3. REFERENCES 

ALTH, A. ( 1878): 0 Galicyj skich gatunkach skarn i­
eniaJych otwornic rodzaju Gyroporella Giimbel. ­
Rozprawy i sprawozdania z posiedzen Wydzialu 
matematyczno-przyrodniczego, HI Akad. Umiej w 
Krakowie, V, 7 1-112. 

ALTI-I, A. (1881): Die Versteinerungen des Nizniower 
Kalksteines. - Beitr. Pal'iont. Oster.-Ung. , I , 183-
332. Wi en. 

BARATTOLO, F. (1978): Su di una nuova dasi­
cladacea (alghe verdi) nel Paleocene dell' Appenni­
no meridionale.- Boll. soc. Natu!". Napoli, 87, 1-76. 

BASSOULLET, J.P. , BERNIER.P .. CONRAD, M.A. , 
DELOFFRE, R. & JAFFREZO , M. (1978 ): Les 
Algues Dasyciadacees du Jurassique e t du Cretace.­
Geobios, Mem. spec. 2, 1-330, Villeurbanne. 

BERNIER, P. (1984): Les Formations carbonates du 
Kimmeridgien et du Portlandien dans Ie Jura merid­
ional. Straligraphie, Micropaleontologie, Sedimen­
tologie.- Doeum. Lab. Geol. . 92/1, 1-443, Lyon. 

BODROGI, I. . CONR AD, M.A. & LOBITZER, H. 
(1993): Lower Cretaceous Dasyclada1es from the 
Villany zone, Southwest Hungary. Biogeographical 
significance.- Boll. Soc. Paleont. ltal. , Spec. Vol. I, 
59-68, Mucchi, Moelena. 

BODROGI, I. , BONA, J. & LOBITZER, H. ( 1994): 
Vergl eschende Unters uchung der Foraminifcrcn­
und Kalkalgen-Assoziationen der Urgon - Entwick­
lun g eles Schrattenkalks in Voralberg (Ostcrreich) 
und der Nagyharsany Kalkstein Formation des Vi l­
lany-Gebirges (Ungar).- Jubilaums. 20 Jahre Geol. 
Zusammenarbei t 6sterreich-Ungar, 225-283, Wien. 

BUCUR, I. (1978): Microfaciesurile calcarelor albe din 
partea nordica a masivului Piatra Craiului. - Consid­
eratii biostraligrafice, Dari de Seama al e Sedint. 
LXIV (1976-1977). 4. Stratigrafie, 89 - 105 , Bucu­
resti. 



12 

BUCUR , I. (1980-198 1): Algues calcaires du Cretace 
inr6riclif des Monts Padurea Craiuiui.- Nymphaca, 
VTII-IX , 53-68, Oradea. 

BUCUR, I. (1985): A llew dasycladacean Alga in the 
Urgonian limestones from Pacturca Cra iului 
(J\pusen i Mountains): Sa/pingoporella pafrll/iusi 
Il.sp.- Revue roumaine, Geo\. G6oph. G6ogr. , 
Geologic, 29, 8 I -84, Bucarcst. 

BUCUR, I. ( 1993) : Some new or poorly known ea l­
careO Ll S algae (Dasycladales, Gymnocodiaccac) in 
the Lower Cretaceous deposits from the Res ila­
Moldova Nona zone (Southern Carpathi ans, Roma­
nia). - Rev. [span. Micropal., 25/1, 93-126. 

BUCUR , I. , COC IUBA, I & COC IUBA, M. ( 1993): 
Microracies and microfossils in the Upper Jurassic -
Lower Cretaceous limestone in the Spulhcrn parl or 
the Paclurca Crai ului Mountains.- Rom. 1. Stratigra­
phy, 75, 33-40, Bucurcsti. 

BYSTRICKY, J. (1967) : Di e obenriad ischen Dasy­
cladazccn clef Wcs tkarpatc ll. - Geo!. sbornik, 
XV IlI /2, 177-364, Bratislava. 

CARROZ1, A. ( 1953): Decouverte du genre Salpil/go­
porella dans Ie Jurass ique superieu r dl! G rand 
Sa leve .- Arch. Sci. Genevc, Vll , 319-324. 

CONRAD, M.A. (1969): Pial/ella ~el/eI'el/sis n. sp., une 
OasycJadacec cladospore des calcaires urgoniens.­
lOcI. geol. helv., 62/2, 571-582. 

CON RAD, M. A. (1970): Barremian and lower Aptian 
Dasyc ladaccae in the area surrounding Ocncva 
(Swi tzerland).- Geol. romana, IX, 63-100, Roma. 

CON RAD, M. (1982) : Com ments o n Korkyrella and 
other new Dasycladaceae.- Rev. Palcobiol. , 1/1 , 1-3. 

CONRAD, M.A. & GRABNER , S. ( 1975): Salpil/go­
porella hispallica n.sp. une Dasyc.: ladale elu Bar­
rcmien des Pyrenees espagnolcs.- C. R. Soc. Phys. 
Il isl. nal. Geneve. N. S., lX/I -3, 30-39. 

CON RAD , M.A. & MASSE, J.P. (1989): Les alg ues 
ca lcaires des formations carbonatees de I ' ll au leriv­
iell - Barrem ien pro parte du Jura vand ois el 
nellehfHelois (Suisse). - Mcm. Soc.:. NelichaL Sc.:i. 
nal., 11 ,277-290. 

CONRA D, M.A. & PEYBERNES, B. ( 1976): Haute ri ­
vian - A lbia n Dasycladaceae [rom the urgonian 
limestones in the French and Spanish Easlern Pyre­
necs.- Oeol. romana, 15, 175- 197 , Roma. 

CONRAD. M.A., PEYBERNES, B. & RADOICIC, R. 
(1977): Salpil/Roporella urladallas; n. sp., une Dasy­
clada le du Crctace inferieur d 'Espagne el de 
Yougoslavie.- Geolog. Mediterrancenne, 4/2, 73-82. 

CON RAD , M.A., & PEYBERNES, B. (1978): Su r 
quelques Oasycladales de l ' Urgo-Aptien el u 
Pre balkan bu lgarc.- C . R. Seances Soc. Phys. I-fis t. 
nal. Geneve, 12/2-3, 69-83. 

Gcologi<l Cro3tica 49/1 

CONRAD, M.A. , PEYB ERNES, B. & MASSE, J.P. 
(1983): Clypeil/a somalica n.sp. Dasye ladale nou­
ve lle du CrelacC infcricur de la Plaque Africaine 
(Rep. de Somat ic, Ital ic mcrid ionale).- A nn. Soc. 
Geolog. du Nord, 93-95, Lille. 

CON RAD, M.A., PR ATURLON, A. & RADOI CIC, R. 
(1973): Reinstatement of Ihe genus Salpingoporella 
PIA (Dasycladaccae) followed by S. sleillhall seri 
n.sp. - C. R. Soc. Phys. Hi sl. nal. Genevc, 7/2-3, 
103- 111. 

CONRAD, M.A., PRATURLON, A. & RADOICIC, R. 
( 1974): The genus AClilloporella Guembel in AL TH 
1882, Dasyc1adaJcs, green A lgae, a revision. - Geo!. 
roman a, XIII, 1~ 15, Roma. 

CONRAD, M.A. & RADOI CIC, R. ( 1972): On Mlllli­
aia baconica Deecke (Characeac) and Clypeil/a ? 
solkolli n.sp. (Dasycladaeeae) . A case oj' homeo­
morphism in calcareous green Al gae.- C. R. Soc. 
Phys. Hist. nal. Geneve, VI/2-3, 87-95. 

CON RAD , M.A., RADOlCIC, R. & RIOY , Y. ( 1976): 
Sa/pillgoporclla hasi n. sp. , une Oasyelada le de 
l 'Albian el du C6noman icn elu Portuga l ci de 
Ju gos lavi e.- C . R. Soc. Ph ys. Hi s\. nat. Oencve, 
N.S., 11/1 -3,99-104. 

CONRAD, M.A. & RADOlCIC, R. (1978): Salpill go­
porella kafzen· n. sp. une Oasycladac6e (a lguc cal ­
caire) nou velle du Bcrriasien cl du Va langinicn de 
la region mcdi lerraneene.- Geo!. vjcsnik, 30/1,69-
72. 

CONRAD, M.A . & V AROL, B. (1990): Cylil/droporel­
la tal/rica n.sp. urges 10 rev iew di rrcrenl palterns o[ 

calci fica ti on in the Mesozoic Dasyclada lcs (green 
algae).- Arch. Se., 43/1, 193-2 14, Geneve. 

CRESCENT1, U. (1959): Sopra una nuova Mocroporel­
la (Alga calcarea) ncl Maim dell' Ii alia merid­
ionale.- Giorn . geol, 27, 195-202, Bologna. 

DRAGASTAN , O. ( 1969): Alg ues ca lca ires du Juras ­
sique sllperieur CI du Cretace inferi eur de 
ROllmanie.- Rev. Micropa!. , 12/1,53-62, Paris. 

DRAGASTA N, O . ( 197 1): New a lgae in the Jurassic 
and Lower Cretaceous in the Bicaz Va lley (Easl 
Carpathians) Romania. - Rev. espan. M ieropal. , 3, 
155-192. 

DRAGASTAN, O. (1975): Upper Ju rass ic and Lower 
C retaceous Microfacies [rom the Bicaz Valley 
Basin (East Carpalhians).- Memoires, Instil. Ocol. el 
Geophys .. 21, 1-87, Buearesl. 

DRAGASTAN, O. (1978): Microfacies de la serie cal­
carie Cretace infericur cI' Aliman (Dobrogea de 
Sud) .- Dari de Seam a ale Sed inl. , LX IV, 107-136, 
Bllcuresti. 

DRAGASTAN, O. ( 1989): Calcareous Algae (new and 
revised). Microproblcmalieae and Foraminiferida of 



Sokac: Taxonomic Review of Some Barremian ,Illd Aptian Calcareolls Algae ... 

Jurassic-Lower Cretaceous deposits from the Carpa­
thian area.- Rev. espan. Micropal., 21/1, 5-65. 

DRAGASTAN, 0, BUCUR, I. & DEMETER, I. 
(1978); Date noi privind biostratigrafia depozitelor 
barremian - albiene din partea central-estica a Zonei 
Resita - Moldova Nona (Banat), obtinute prin fora­
jul de referinata de la Sopotul Non.- Dari de Seama 
ale Sedint., LXIV /5, 17-36, Bucuresti. 

DRAGASTAN, O. & BUCUR, I. (1993): The Dasyclad 
Genus Radoicicief/a and it s representatives from 
Romania.- Rcv. cspan. Micropal., 25/2, 5-23. 

ELLIOTT, G.F. (1960): Fossil calcareous algal noras 
or the Middle East, with a note on a Cretaceous 
problematicum, Hellson e//a cylindrica, gen. et sp. 
nov.- Quart. J. geol. Soc. London, 115/3,217-232. 

ELLIOrr, G.P. ( 1968): Pcrmian to Paleocene calcare­
ous algae (Dasycladaceae) of the Middl e East.­
Bull. brit. Mus. nat I-list. (Geol.), Suppl. 4, p. I-Ill. 

FARINACCI, A. & RADOICIC, R. (199 1): Latc Juras­
sic - Early Cretaceous Oasyc ladales (green algae) 
from the Western Pontides, Turkey.- Geol. romana, 
27, 135-165, Roma. 

GRANIER, B. (1986): Algues Chlorophyceae du Juras­
sique terminal et du Cretace infe rieur eu Alicante.­
Mediterranea, Ser. Geol., 5, 5-96, Alicante. 

GRANIER, B. (1994): The genus AClilloporel/a (GUm­
bel in Alth, J.881) and its representatives. A review.-
13eitr. Palaont., 19, 113-127, Wien. 

GRANIER, B. & BERTHOU, P.-J. (1994): Milallovi­
cella mOlJlcilialla n.gen. n.sp., une Algue Oasy­
cladale nouvelle du Ponlandien de I' Algarve central 
(Portuga l).- Rev. MicropaJeont.. 37/2, 113- 121, 
Paris. 

GRANIER, B., MICHAUD, P. & FOURCADE, E. 
(1986): Apinel/a jaffrezoi n.gen. n.sp. algue Dasy­
cladacce elu kimmerielgien du Chiapas (S ud. - est e1u 
Mexique).- Geobios, 19/6,801-813, Lyon. 

GUSIC, l. (1966): Two new Dasyclad species of the 
subgenu s Pial/clio from Lower Cretaceous of 
Istria. - Geol. vjesnik, 19, 35-46. 

JOHNSON, H.J. (1968): Lower Cretaceous Algae from 
the Blake Escarpment (Atlantic Ocean) and from 
Israel. - Prof. Contr. Colorado School Min. , 5, 71 p., 
Golden. 

KARAS, N. & RADOICIC, R. (1991): Salpingoporel/a 
ph'iniae n.sp. iz barema karbonatllc platforme Par­
nasa (kontinentalna Grcka).- Oeo!. an. Balk. polLl­
os!., 55/1, 163-l73, Beograd. 

KOCI-IANSKY -DEVIDE, V. & GUSIC, I. (1971): 
Evolutions-Tendenzen der Dasycladaceen mit 
besondcrer Bcriicksichtigung neuer Funde in 
Jugoslawien.- PaJaon!. Z., 45/1 -2, 82-91. 

33 

KUSS, J. & CONRAD, M.A. (1991): Calcareous Algae 
from Cretaceous carbonates of Egypt, Sinai, and 
Southern Jordan.- Jour. Paleon!., 65/5, 869-882. 

LORENZ, M. (1902): Geologische Studien im Gren­
zgebiete zwichen hclvetiseher und ostalpinem 
Facies.- Bel'. naturf. Ges. Freiburg Br., XII, 34-62. 

LUPERTO-SINNI, E. & MASSE, J.P. (1982): Contrib~ 
uto della paleoecologia alia paleogeografia della 
parte meridionale della piattaforma apula nel Cre­
taceo inferiorc.- Geol. roman a, 21, 859-877, Roma. 

LUPERTO-SINNI, E. & MASSE, J.P. (1984): Donnees 
nouvell es sur la micropaleontologic ct la stratigra­
phie de la partie basale du "Calcare di Bazi" (Cre­
tace inferieur) dans la region des Murges (Italic 
mcridionale).- Riv. It. Palcont. Stra!., 90/3, 331 -374. 

LUPERTO-SINNI, E. & MASSE, J.P. (1993): The Ear­
ly Cretaceous DasycladaJcs from the Apulia Region 
(Southern Italy): biostratigraphic distribution and 
paleobiogeographic significance.- In: BARATTO­
LO, F., DE CASTRO, P. & PARENTE, M. (eds.): 
Studies on Fossil Benthic Algae. Boll. Soc. Paleon!. 
Ital., Spec. Vol. J, 295-309, Modena. 

MAMET, B. & RUDLOFF, B. (1972): Algues car­
bonifcrcs de la partie septentrionale de I ' Amerique 
du Nord.- Rev. Micropal. , 15/2,75- 114, Pari s. 

MANCINELLI, A. (1992): Dislribuzione delle Dasi­
c1adali (a lghc verdi) nei sedimenti del Crelaceo 
in fe ri ore in facies di piattafonna carbonatica 
dell' Appennino cenlro-meridionale (Lazio e Abruz­
zo).- Studi geol. Camerti, 12,7-29. 

MASSE, J.P. (1985): Pal eobiogeographie des Rudistes 
du domaine peri meditcrranccn a I' Aptien inferieur. ­
Bull. Soc. geol. France, liS, 715-721. 

MASSE, J.P. (J 989): Relations entre modifications 
Biologiques et phenomenes geologiques sur les 
plates - formes carbonatees du domaine perimediter­
raneen an passage Bcdou lien-gargasicn.- Geobios, 
Mem. spec. 11,279-294, Lyon . 

MASSE, J.P. (1993): Early Cretaceous Dasycladales 
biostratigraphy from Provence and adjacent regions 
(South of France, Switzerland, Spain). A reference 
for Mesogean correlations.- In: BARATTOLO, E, 
DE CASTRO, P. & PARENTE, M. (cds.): Studies 
on Fossil Benthic Algac. Boll. Soc. Paleon!. I tal., 
Spec. Vol. 1,311-324, Modena. 

MASS E, J.P. & LUPERTO-SINNI, E. (1989): Cylill­
drofJorella Iyra'a, nouvelle espece de Dasycladale 
de I' Apticn inferieur du domaine pcrimediterraneen 
nord-occidental.- Rev. Micropal., 32/1, 30-39, Paris. 

MISfK, M. (1987): The alga Halicoryne c{[}palica n.sp. 
[rom Uppcr Triassic of Western Carpathians. - Geo­
logica earpathica, 38/3, 285-293, Bratislava. 

OKLA, M. S. (1995): Late Cretaceous Dasycladales 



34 

from the Aruma Formation in central Saudi Arabia. ­
N. l b. Geol. Palaonl. Mh., 1995, H. 8,475-487. 

PEYBERNES, B. & CON RAD, M.A. ( 1979): Les 
Algues du Cretace infcricur de Hongric.- Bull. Cen­
tr. Reeh. Exp lol".- Prod. Elf - Aquitai ne, 3/2, 743-
752,Pau. 

PIA , J. (1918): Familie: Dasycladaeeae (Endl. ) Cram. 
em.- In: TRAUTll , F. (cd.): Das Eoz,invorkommen 
bel Radstat im Pongau und se ine Bez iehun gen zu 
den Gleichailcrigcn Ablagerungen bei Kirchberg am 

Wechscl lind Wimpassing am Lci thagebirge.­
Dcnkschriftcll der mat hcm. -naturw. Klassc, 95, 171-
278, Wien. 

PIA , 1. (1925): Einigc nelle oder ungcni.igcnd bekanntc 
Siphoncac vertici llate aus clem mittelcuropi:iischcn 
Malrn.- Ann. naturhist. Museulll Wien , 38, 83-88 . 

PIA, l. (1930): Upper Triassic foss il s from the Burmo­
Siamese fronti er. A new Dasycladacca. HolosjJorel­

la siamcflsis nov.gen., nov.sp., with a description of 
the allied genus Acicff/elln Pia.- India Geo!. Survey, 
63,177-181, Calcuta. 

PRATURLON, A. ( 1964): Calcareous Algae [rom 
Jurassi c Cretaceous !imestones of Centra! Apen­
nines (Southern Lat iulll - Abruzzi). - Geo!. roman a, 
fll, l7 1-202, Roma. 

PRi\TURLON, A. & RADOICIC, R. ( 1967): Notes on 
the Dasyclad genu s Sa/pillgoporel/a Pia.- Geo!. 
romana, VI, 137- 144, Roma. 

PRATURLON, A. & RADOICIC, R. ( 1974): Emenda­
tion of Acroporella (Dasyc ladaceae).- Geo!. roma­
na, XllI , 17-20, Roma. 

RADO ICIC, R. ( 1959): Salpil/gopo/"e//a dil/a/"ica nov. 
sp. u donjok redni m sedim entima Oinarid a.- Geo!. 
glasnik, 3, 33-42, Ti tograd. 

' RADOICIC, R. ( 1963): On the new Upper Jurassic 
Dasyc1adaeea Pial/ella gn{(lii nov. gen., nov. spec .­
Geol. vcsn ik, scI". A, 20, 189-193, Beograd. 

RADOICIC, R. (1965): Pial/ella tll/"gida n.sp. from the 
Cenoman ian or the Outer Dinarids.- Geo!. vjesnik, 
18, 195-199. 

RADOICIC, R. (1967): Salpil/gopo/"ella /IIelitae spec. 
nov. iz donjokrednih scdimcnata spoljasnih Dinari ­
da.- Annal. g601. Penins. balkan. , 33, [21-126, 
Beograd. 

RADOICIC, R. ( 1968): Likal/cl/a , dal/i/ol'ae spec. 
nov. et que lques autres Dasycladacccs cretacees 
inf6rieurcs des Dinarides externes.- Vesn ik Zav. 
geol. geof. istl"., (A), 26,177- 193, Beograd. 

RADOICIC, R. ( 1969): A new lower Cretaceous Dasy­
cladacea, C/ypeil/a pejovici, and not e on some 
Clypeinae.- Geol. romana, VIII , 7 1-84, Roma. 

RADOICIC, R. (1975): On Likal/e1la i1al1l/lllldai, sp. 
nov. from the upper Cretaceous of the Tripoli area 

Geologia Croal;c:I 49/1 

(Lybia) ancl the age of strate containing Disso­
cladella all/Ill/ara (RAINERI).- Annal. goal. penins. 
ba lkan. , 39, 147- 152, Beograd. 

RADO IC IC, R. (1978): Salpil/gopo/"e//a lIIi/ol'al/ol'ic i 
n. sp. a new Dasyclad from the Cenomanian strata of 
the Dinari des and a note on the Forami nifer NUII/­

IIIlIlowllilla sp. (arr. reglliaris PHILIPPSON).­
Annal. g601. penins. balkan. , 42, 375-381, Beograd. 

RADO IC IC, R. (1979): Salpillgoporella Ilbaiyd/li, a 
new Senonian Oasyc lad and some data abou t sub­
surface biostrat igraphy of the western Iraq i deserl. ­
Bulleti n Aead. Serbc Sci. Art s, Classe Sc i. mathc­
mat. natul". 68/1 9, 97-105, Beograd. 

RADOICIC, R. (1986): 0 jednoj novoj klipei ni i nckim 
drugim dazikladaccama iz jure i krcde Crne Gore.­
Geol. glasnik, 10,53-65, Titograd . 

Ri\DOlCIC, R. (1987): A con tribution to the study of 
the stratigraphy or Hvar (followed by s imultaneous 
occurrences of the spec ies Salpillgoporella me/ilae 
and S. dil/arica).- Glas SANU, 5 1, 73-79, Beograd. 

SC HI NDLER, U. & CONRAD, M.A. ( 1994): The 
Lower Cretaceous Oasyc ladales from the North­
western Fri uli platform and their di stributi on in 
chronostrat igraph ic and cyclostratigraphi c units.­
Rev. Paleobiol., 13/ I, 59-96. 

SCHLi\GINTWEIT, F. ( 1990): Taxonomic revision or 
Likallella hallllllllda; RADOICIC 1975, dasy­
cladacea n alga from the Upper Cretaceous of the 
Northern Calcareous Alps (Gosau formation, 
Coniacian) .- Rev. Pal60biol., 9/2, 257-261 , Genevc. 

S IM MONS, M.D. , EMERY, D. & PICKARD, N.A. I-l. 
( 1991 ): Hensollella dinarica (Radoicic) an orig inal­
ly calci tic Early Cretaceous dasycladacean alga.­
Palaeontology, 34/4, 955-96 1, London. 

SOKAC, B. (1986). Ciypeilla radici n.sp. (Ca lcareous 
algae; Oasyc ladaccae) from th e Neocomian of the 
Coastal Part of the Outer Din<.ujcies.- Geol. vjesnik, 
39,43-54. 

SOKAC, B. (1992): Two New Species of Calca reous 
Algae (Dasycladaceac) from the Upper Dogger of 
Sout hern Dalmatia (Croalia).- Geol. Croatica, 45, 
15-24. 

SOKAC, B. (1993): Sa/pillgopore/la robllsta n.sp. (Ca l­
ca rcO Li S algae; Dasyc lada lcs) from the Uppe r Bar­
rem ian - Lower Apt ian of Mt. Biokovo, Croatia.­
Geol. Croatica, 46/2, 175- 180. 

SOKAC, B. & JELASKi\ , V. (199 1): Salpillgoporella 
po/saki n.sp. Calcareous Algae (Dasyc ladaccae) 
from the Cenomanian depos its of the Is land of 
Brac. - Geol. vjesnik, 44, \ -7. 

SOKAC, B. & NI KL ER, L. (1973): Calcareous algae 
from the Lower Cretaceous of the envi rons of 



Sokac: Taxonomic Review of Some Ilarremian and Aptian Calcareous A lgae ... 

Niksic, Crna Gora (Montenegro).- Pal. jugos lav. , 
13, 57 p. , Zagreb. 

SOKAC, B. & VELIe , I. (1978): Redescri ption of the 
genus Sel/ipore/la (Cal ca reou s algae; Oasy­
cladaeeae) .- Geol. vjesnik, 30/1 , 225-242. 

SOKAC, B. & vELlC, r. ( 1979): A new ealcarous alga, 
Sa/pillgoporel/a bikoviellsis n.sp. (Oasycladaeeae) 
from Lower Aplie n deposit s of MI. Biokovo, Dal­
matia (south Croatia).- Gcol. vjesnik, 31, 145-150. 

SOTAK, J. & MISiK, M. ( 1993): Jurassic and Lower 
Cretaceous dasycJacl aJean al gae from the Weste rn 
Carpathi ans.- In: BARATTOLO, r. , DE CASTRO, 
P. & PARENTE, M. (cds.): Studies on Fossil Benth­
ic Algae. Boll. Soc. Paleont. Ital. , Spec. Vol. 1,383-
404 , Modena. 

TISLJ AR, J. ( 1986): Postanak em ih oblutaka i ulomaka 
C' black pebbles") u peripli ll1atskim vapnc ilc ima 

35 

titona zapadne Istre i barema otoka MJjeLa. - Geo!. 
vjesnik , 39, 75-94. 

TISLJAR , J. & VELIe, I. ( 1991): Carbonate facies and 
depositional environments of the Jurass ic and Low­
er Cre taceous of the coastal Din arides (Croati a) .­
Geol. vjesnik, 44, 215-234. 

VALET, G. ( 1968): Contributi on a I'ctude des Dasy­
c1adales. 1. Morphogenese.- Nova Hedwigia, XVI, 
2 1-82, Lohre. 

VELIe, I. & SOKAC, B. ( 1978): Biostratigraphi c 
ana lys is of the Jurass ic and Lower Cre taceolls in the 
widcr region o r Ogulin, centra l Croatia.- Geol. vjcs­
nik, 30/1 , 309-337. 

YABE, 1-1. & TOYAMA , S. ( 1949) : New Dasy­
c1adaecac from the Jurassic Torin osu lim estone o r 
the Skawabasin Sikoku .- Proe. Jap. Acad. Tokyo, 
25/7 , 40-44. 

Manuscript received February 19, 1996. 
Revised manuscript accepted May 27, 1996. 



I, 4, 6, 10- 1 I 

2-3,5 

7-9 

12-1 3 

PLATE I 

1-13 Sa/pil/goporel/a mueh/"ergii (LORENZ) PIA in TRA UTH 

Tangential - oblique sections; Figs. 1,4,6, x25; Figs. 

Longitudinal - oblique sections, x31. 

Oblique sections, x3 1. 

Transvcrse sections, x3 1. 

10- 1 I, x3 I. 

Geologia Croa!icn 49/1 

Age and gcncral location: 

1,3,9 

4 

2,5-8,13 

10-11 

12 

Lower-Middle Barrcmi an; field scction 011 thc road cut ncar Sobra, Mlje t island (Pigs . 1-2). 

Lower Barrcmian; field scction Bravci dolae - Debelo brdo, Mt. Dinara. 

Upper Barrcmian; Velika jezero, Mljcl island. 

Upper Barremian; Mljel island (rig. I). 

Lower Barremian; Mljcl is land (Fig. 1). 



Sokai: PLATE! 37 



38 

L,3-4 

2 

5 

6 

7 

8-9 

10- 12 

13 

PLATE JL 

1-5 Salpillgoporella hm"i CONRAD, RADOICTC & REY 

Transverse sections, x57. 

Transverse slightly oblique seclion, x57. 

Rcc ri sta li zcd tangential - longitudinal section, x47. 

5- 13 Salp illgofJorella llIelifae RADOICIC 

Longitudinal - tangential section, x23. 

Tangential section, x25. 

Tangent ial - oblique sections; Fig. 8, x25; Fig. 9, x23. 

Oblique scctions, x23. 

Transverse section , xIS. 

Age and general local ion: 

\-5 Barremian; eres island. 

6, 9, I [-12 Lower Barremian; field section on the road cut ncar Sabra, Mljet island (Figs. 1-2) . 

7-8, 10, 13 Upper Barremian; Veliko jezero, Mljet island. 

Geologia Croalica 49/1 



Sob c PLATE Jl 39 



40 

1 

2 

3,5,8-9 

4 

6 

7, 10 

PLATE 111 

1-2 Sa/pil/goporel/a melitae RADOICTC 

Longitudinal - tangential sec tion , x23. 

A - Oblique sect ion, B - Longitud inal section, x L8. 

3-10 Sa/pil/goporel/a po/ygollalis Il .sp., x23 

Oblique sections; Fig. 3 - Hoiotypc. 

Tangentia l section. 

LongilUciinai - oblique sect ion. 

Oblique - tangential sect ions. 

Age and gene ral location: 

1 Upper Barremian; Velika jezcro, Mljcl is land. 

2 Barremian; Gornje Igranc, Mt. B iokovo. 

3- 10 Lowe r Barremian; field section on the road cut ncarSobra, M ljct island (Fig. 1-2), 

Geologia Cro;llica 49/1 



PLATE 1II 41 



42 Geologia Cromica 49/1 

PLATE TV 

1-17 Salpingoporella polygollalis n.sp., x23 

1-5, 7-S B, 10- 11 ,13 Oblique sections. 

6 Tangential - obl ique section. 

SA-9, 12, 14, J6 T ransverse sections. 

15, 17 T ransverse slightl y oblique sections. 

Age ancl general location: 

1- 17 Lower Barremian; field section on the road cut near Sobra, Mljet island (Figs. 1-2). 



Sokac PLATE IV 43 

, '" . 
• " •• ~ :t <' • 

\jj J '. fI" 



44 

1-2 

3-4 

5-8 

9A 

10A-llA 

9B 

lOB- liB 

PLATE V 

1-8, 9A-IIA Sa/pillgoporel/a hispanico CONRAD & GRABNER 

Tangential - iongitudjnal sections; Fig. 1, x54; Fig. 2, xS7. 

Longitudinal sect ions, x57 . 

Oblique sections, xS7. 

Tangential - longitudinal section, x36. 

Oblique sections, x57. 

9B - liB Sa/pillgoporella hasi CONRAD, RADOICIC & REY 

Longitudinal s lightly oblique section, x36. 

Oblique section s, x57. 

Age and general location: 

1- 11 Barremian; ercs island. 

Geologia Croalica 49/1 



Sokac PLATE V 45 



46 

1,5-8, 10 

2,9 

3 

4 

II 

12- 15 

PLATE VJ 

1- 15 Sa/pillgoporel/a verrucosa n. sp. 

Oblique sections; Fig. I, x38; Figs. 5, 7-8 , x57; Fi g. 6, x38; Fig. 10, x47 . 

Longi tudinal sections, x47. 

Tangent ial section, x57. 

Longitudinal slightly oblique section - Holotypc, x57 . 

Transverse slightly oblique section, x47. 

Transverse sections; Figs. 12- 13, x57; Figs. 14-15, x47. 

Age and general location: 

1- 15 Upper Barremian; field section on the road cut near Sabra, Mljet island (Fi gs . 1-2). 

Geologia Cro;lIica 49/1 



Sokac PLATE VI 47 



PLATE VlI 

1-18 Salpillgoporel/a heraldica n.sp. , x31 

1-2,4,7,9, 11- 14 Oblique seclions. 

3 

5-6, SA, 10 

15 

8B, 16- 18 

Tangential - oblique section. 

Tangential sections, Fig. 5 - Holotype. 

Transverse slight ly oblique section. 

Tran sverse secti ons. 

Age and genera l location: 

1- 18 Top Barremian or Lower Aptian; Brna village, Korcula island (Fig. 3). 

Gcologia CrO:lliC<l 49/1 



Sokm; PLATE VII 49 



50 

I 

2 

J, 6A 

4-5,68,7 

8, 10 

9 

PLATE VIII 

1- 10 Salpillgoporella llasi CONRAD, RADOICIC & REY 

Longitudinal section, x47. 

Tangential - longitudinal section, x47. 

Tangential sect ions, x57. 

Oblique sections; Figs. 4-5, x47; Figs. 6A-7, x57. 

Transvcrse sections, x57. 

Transverse slightly obliquc section , x57. 

A ge and general location: 

1- 10 Barremian; Cres island. 

Geologia Cro;ltica 4\)jl 



Sokac PLATE VIIl 



" Geologia Cro,uiC:1 49/1 

PLATE IX 

1-13 Salpillgopore/la IIrladallasi CONRAD, PEYBERNES & RADOICIC 

1, 9 Longitudinal sections, xS4. 

2,4 Tnngenliai sections; Fig. 2, x44; Fig. 4, x60. 

3,5-8, IO- IIA Oblique - tangenl ial scclions; Figs, 3, 7,10, x54; Figs, 5-6, x44; Figs, 8, IIA, x57 . 

II B, 12A - J3 Transverse seclions; Figs. liB , 13, x54; Fig. 12A, x57 . 

12B Salpill{;opol'el/a dillGrica RADOICIC- lransverse section, x57 

Age and general location: 

1- 13 Top Aptian; field section Bravci dolac - Debclo brdo, ML DinanL 



PLATE IX 



54 Gcologi:l Cro:llicn 49/1 

PLATE X 

I A Saipillgopore/la IIrladalIasi CONRAD, PEYBERNES & RADOICIC - transverse slig htly o blique 
section. 

IB 

2,6-7 

3 parI. 

4 

5 

8 part. 

9 part. 

Salpingoporel/a dinarica RADOICIC - longi tudinal sect ion, B I - oblique section, x54. 

S. IIr/adallGsi - oblique sections; Figs. 2, 7, x54; Fig. 6, x45. 

S. lIrladollasi - oblique sections, x45. 

A - S. IIrladanasi - tangential sec tion; I3 - S. d;'/CIrica - tangential section , x45. 

A - S. IIrladanasi - oblique section; B - S. dil1arica - tangential section , x57. 

S. IIrladal/asi - transverse slight ly oblique sect ion , x54. 

S. IIrladanasi - transverse section , x57 . 

10-12 Saipillgopore/la katzeri CONRAD & RADOIC1C, x22.5 

10 Longitudinal sec tion. 

I 1- 12 Obi ique sections. 

Age and general location: 

1-9 Top Aptian; field section Bravei do lae - Debelo brdo, Mt. Dinara. 

10- 12 Berr iasian; fi eld section south of the peak Stratinovac, Mt Biokovo. 



Sobe PLATE X 55 



56 

1-2 

3 

4 

5 

6-8, 10 

9 

PLATE XI 

1-10 Sa/pingopore/la dinarien RADOICIC 

Oblique seclions; IA - Charophyla; Fig. I, x60; Fig. 2, x50. 

Longitudinal sections, x60. 

Tangential section, x60. 

Longitudinal - tangential section, x60. 

Oblique sections, x60. 

A - Tangential section, B - oblique section, C - transverse section, x28. 

Age and general location: 

2, 7 

3 

4 

5-6,8, 10 

9 

Upper Valanginian; Sosici village, W lslria. 

Aptian; Bozavciea bay ncar Bozava, Dugi otok island. 

Aptian; Jesenje hill, SE Ogulin. 

Aptian; Poljana cove, Mali Losinj island. 

Upper Aptian; Velo Grablje near Stari Grad, Hvar island. 

Middle Aptian; field sect ion ncar Saranac, Mt. Biokovo. 

Geologia Croatica 4lJfl 



Sokac PLATE Xl 57 



58 

1,5,9 

2,4 

3 

6 

7 

8 

10 

I 1-13 

PLATE XU 

1-13 Sa/pillgopare/la dillarica RAD01C1C 

Oblique sections, x 150. 

Tangential - oblique sections; Fig. 2, x57; Fig. 4, x36. 

Longitudinal sect ion , x36. 

A - oblique section, B - transverse section, x44. 

Oblique section, x44. 

Transverse sect ions, x57. 

Transverse slightly oblique section, xlSO. 

Oblique sections, x57. 

Age and general location: 

2,5,8 -9, 12 Aptian; ML Velika Kapela. 

3-4, 6-7, 10-11 , 13 Upper Aptian; field section on the road cut near Sobra, Mljet island (Fig. 1). 

Geologia Croatica 49/1 



Sokac PLATE XII 59 



I 

2,4 

3 

5 

6-8 

1-4 

5-8 

PLATE XlII 

]-8 Sa/pillgoporel/a dillarica RADOICIC 

Tangential - oblique sect ion , x42. 

Oblique sections, x42. 

Tangeillial - oblique sect ion, x38. 

Longitudinal section, x34. 

Different oblique sections; Figs. 6-7, x4; Fig. 8, x34. 

Age and general location: 

Aptian; Poljana cove, Mali Losinj island. 

Aptian; Bozavcica cove ncar Bozava, Dugi otok island. 

GCOlogb Croatica 49/1 



lobe PLATE XllI 6 1 



62 

I 

2 

3-9 

10-13 

PLATE XIV 

1-9 SaipiJlgoporel/a diJlarica RADOICIC 

Oblique sect ions, x31. 

Longitudinal - tangential section, x31. 

Di fferent oblique sections; Fig. 3, x34; Fig. 4, x42; Fi gs. 5-9, x31. 

Charophytes; Fig. 10, x50; Fig. 11, x60; Fig. 12, x57; Fig. 13, x44. 

Age and general location: 

1-4,7-9 

5-6 

10- 13 

Aptian; Poijana cove, Mali Losinj island . 

Apt ian; Bo7.avCica cove near Bozava, Dugi alok island. 

Upper Valanginian; Gustinja cove, SE Rovinj , lstri a. 

Geologia CrO:lIica 49/1 



Sok:lc PLATE XIV 63 



64 

PLATE XV 

1-19 Pralllriolle/io neroe (DRAGASTAN, BUCUR & DEMETER) BUCUR, x32 

IB, 2B, 6, 9-10,16 

lA, 2A, 3-4, 7-8,12-14 

5, II 

15, 17, 19 

18 

Longitudinal sections; Fig. 2B Hololype. 

Oblique sections. 

Tangential sections. 

Transverse sections. 

Transverse s lightly oblique section. 

Age and general location: 

1-19 Top Barremian or Lower Aptian; Brna village, Korcula island (Fig. 3). 

Gl:oJogia Croatica 49/l 



PLATE XV 65 



66 

2 

3 

4-6,8,12- 15 

7,9-11 

PLAT E XV I 

I-IS Prallfr/onella d{/Ili/ovae (RADOICIC) BARATTOLO 

A - Pm/lff{Ollella danilovae - oblique sec tions, x16. 
B - Sa/pillgopore//a me/if({e - tangenti al sec tion, x 16. 
C - Pralur/ollclla pejovicae - obl ique section, x16. 
D - Sa/pingoporella sp., x 16. 

P. doni/ol'oe - differe nt sec tions, x 16. 

Longi tudina l and transverse sections, x22. 

Different oblique sections, x22. 

Transverse s li ghtl y oblique sections, x22. 

Age and general location: 

l- lS Lower Barremian; field section on the road ClI t ncar Sabra, Mljcl is land (Figs. 1-2). 

Geologia Croalica 49fl 



PLATEXYI 67 



68 

I 

2 

3A 

5-6 

7 

8 

9- 10, 13 

12 

II 

14 

PLATE XVII 

1-7 Clypeina giganlea Il. Sp. , X 18 

Transve rse sect ion. 

Oblique section , I-1olotype. 

Tral1svcrs~ slightly oblique sections. 

Tangential sections. 

Transverse section. 

8- 14 Pratllriollella {Jejovicae (RADOICIC) n.comb. , x22 

Tangential - longitudinal section. 

Oblique sections. 

Tangential ancltransversc - oblique sections. 

Tangentia l section. 

Transverse slightly oblique section. 

Age and genera l location: 

1- 14 Lower Barremian; ri eld section on the road cut near Sabra, Mljet island (rigs. 1-2) 

Geologia Croatic<l 49/1 



Sobi:: PLATE XVII 69 



7() 

1, 4-6, 9-10 

2 

3,7 -8, 11-14,16- 17 

IS 

18-20 

PLATE XVIJI 

1-20 Acroporella radoicicae (PRATURLON) PRATURLON & RADOlCIC 

Obl ique - tangential sections, x23 . 

Tangential section, x 18. 

Oblique section s; Fig. 3, x 18; Figs. 7-8, 11 - 14, 16-17, x23. 

Transverse - oblique section, x23. 

Transverse sect ions, x23. 

Geologia Cro:uica 49/1 

Age and generallocatioll: 

1-7, 9, 12-20 

8,1O- ll 

Lower Aptian; NW from the bridge on the Mreznica River, between Primislje and Ju zbasici. 

Upper Aptian; fi eld section Dravci do b e - Dcbelo brcio, Mt. Dinara. 



Sokac PLATE XVIII 71 



72 

1-2, 4 ,6 

3,5 

8 

9A, 1O 

7 

9B 

PLATE XIX 

1-6, 8-10 Clype!lIa? solkall! CONRAD & RADOICIC 

Tangential - longitudinal sections; Fig. 1, xI8; Figs. 2, 4, 6, x23. 

Longitudinal sections, x29. 

Oblique sections, x23. 

Transverse sections, x23. 

,S'a/pill[:oporclla mclitae - longitudinal section, x23. 

Sa/pil/goporel/a biokol'iclIsis - oblique section, x23. 

Age ancl gcncrallocalion: 

1-5,7-10 

6 

Barremian; Gornjc 19rane, Mt. Biokovo. 

Upper Barremian; Velika jezero, Mljc\ island. 

Geologia Croalic;1 49/1 



PLATE XIX 

" ( 
( 

",-__ 10 

73 



74 Gcologin Cro~tiCi149/1 

PLATE XX 

1~13 Act;lIoporella podolica (ALTH) CONRAD, PRATURLON & RADOlCTC, x23 

I, 2A, SA Long itudinal sections. 

3A, 6, 8- \ 0, 13 Diferclll oblique sections . 

4A,7, 11 Tangential sections. 

4C C/ypeina? solkani - longitudinal slightly oblique section. 

12 

28,38 

413,513 

Transverse section. 

Sa/pingoporc//a !/Ielirae - oblique sections. 

Sa/pillgopore//a muehlbergii - oblique sections. 

Age and general location: 

1 ~ 13 Upper Barremian; Velika jczcro, Mljet island. 



Sobc PLATE XX 75 



76 Geologia Cro<l!lca 49/1 

PLATE XX I 

1-15 PiriJerella spil10sa n.gen., n.sp.; Fig. I, x38; Figs. 2- 15, xSO 

1, 4-8,11 , 13-14 Oblique scct ions; Fig. 6 - Holotype. 

2 Longitudinal section. 

3,9, 12 Tangential sections. 

10,15 Transverse slightly oblique sections. 

Age and general location: 

1- 15 Upper Barremian; rield section on the road cut ncar Sobra, Mljcl island (Figs. 1-2). 



PLATEXXT 77 



78 

PLATE XXII 

1-20 Pirilerella somalica (CONRAD, PEYBERNES & MASSE) Il.comb., x46 

1-2,5 , 11 , 14-15, 19 Oblique sectiolls. 

3-4, [8 Tangential sections. 

6, 9-10,13 

7-8,1 2, 17 

16, 20 

Longitudinal sections. 

Tangential - oblique sections. 

Transverse sections. 

Age and general location: 

1- 10,12-20 

II 

Upper Barrem ian; sea-side 3 km N [rom Porec. 

Barremian; Raska gora, W side of Neretva river, NE Mostar. 

Geologia Cmalica 49/1 



Sokat PLATE XXII 79 



so Geologia CroatiC:l 49/1