Berliner geowiss. Abh (E) 9 253-281 8 Abb., 4 Taf. Berlin 1993 
DO'G" "2$.' d __ t 
Palaeobiological Reconstructions of selected BIOGENE SEDtMENUnON 
Sphinctozoan Sponges from the Cassian Beds (Lower Carnian) 
of the Dolomites (Northern Italy) 
• 
S. MÜLLER-WII.LE & J. RElTNER 
PalaeobiologicaJ reconstruction of selecled Sphinctozoan Sponges from the Cassian Beds (Lower Camian) ofthe Dolomites (Northem 
IlaIy).- Slaffan MÜLLER-WILLE & Joachim REITNER. Berliner geowiss. Abh. (E) 9: 253-281; . 
Abstrad: PalaeobiologicaJ models of four selecled species of sphinctozoan coraIIine sponges from the Cassian Ileds (Lower Camian, 
Dolomites) have been established using, among othels, luminescence tecbniques. The latter has beeil Sllccesfully to wimate the 
organie content and diagenetic history ofthe skeletons. Recent in"estigations yield the differentiation ofthree steps in the secretion ofthe 
skeleton in coralline sponges, according to which skelelal elements can be classified. The identification of these elements rendels 
infoiiuation on the way ofsecretion ofthe skeleton, the relative position ofthe soft tissue, and the function ofthe skeleton. Two 
basic type.s of sphinctozoan organisation can be distinguished: a matrix type, where a rigid framework, secreted in an organo-spicular 
matrix peneti"ates the soft t.issue (as in suomatoporoid coralline sponges) and a cortex type, where the skeleton is secreted by a speziali'WI 
layer (cortex) surrolmding the sponge body. These organizational types bear no phylogenetic implication. 
Es wurden paläobiologische Modelle von vier ausgewählten Arten sphinctozoider Schwälluue alls den Cassianer 
Schichten (Unteres Kam, Dolomiten) unter Zuhilfenalime von I ~lmineszenZll\ethoden eisIeIlt. Diese konnten mit Erfolg zur Abschätzung 
des Gehalts an organischen Stoffen und der diagenetischen Geschichte der Basalskelette eingesetzt werden. Neuere Untersuchungen 
erlauben die von drei Schritten in der Genese des Basalskeletts der corallinen Schwämme, die zu einer Klassifikation der 
Skelettelemente herangezogen werden können. Eine Identifikation der jeweiligen Elemente entsprechend dieser Klassifikation liefert 
Infoiinationen über die Art der Sekretion des Basalskeletts, die relative Position des Lebendgewebes und die Funktion des Skeletts. Zwei 
gnmdlegende Organisalionstypen können innelhalb der Sphinctozoen unterschieden werden: ein Matrix-Typ, bei dem das in 
einer organo-spikulären Matrix sekretiert wird, die den Weichkölper durchzieht (ähnlich den stromatoporoiden "CoraIline Spongien") 
lind ein Cortex-Typ, bei dem die Sekretion des Basalskeletts in einer spezialisierten, subdeiioalen Schicht (Cortex), die den Weichkölper 
umgibt, stattfindet Diese Organisationstypen haben keine Implikationen. 
Adnsl of the authors: Dipl.-Geol. S. Müller-Wille & Priv.-Doz. Dr. J. Reitner, Institut fllr Paläontologie, Freie Univelsität Berlin, 
MalteseIstr. 74-100, HallS D, 0-12249 
• 
1 IntroductioD 
The Sphinctozoa erected by S1E 1882 
are a mainly fossil group of sponges with an ara-
gonitic or Mg calcitic basal skeleton, which is built 
up by chambers. Though the sponge affinities of 
this taxon have been known since long time, due to 
occasional findings of scleres in some representa-
tives, it remained an enigmatic group up to the dis-
covery of the recent species Vaceletia crypta. This 
can be explained by the fact that the sphinctozoans 
represent a highly polyphyletic taxon. As most 
workers had asSllmed monophyly for this sponge 
group on the ground of the similarities in skeletal 
organization, they encountered conflicting 
characters, leading to great di fficulties in solving 
questions conccming the biology and phylogeny of 
these animals. Examples are the controversial 
discussions about sclere mineralogy (S1EIN-
1882, RAUFF 1914) or soft tissue 
organization (RAUFF 1914, SEll..ACHER 1962). 
the sphinctozoan cora11ine sponges 
play an important role as reefbuilders in Pemuan 
and Triassic times (OlT 1967a, SENOWBARI-
DARYAN & RIGBY 1988), an understanding of 
their biology is crucial for facial analysis of bio-
helms of this time. Since the (re)-discovery of 
coralline sponges in the beginning of the sixties an 
actualistic approach to the palaeobiological re-
construction of coralline sponges has been possible. 
The new information was readily taken up by 
palaeontologists working on fossil representatives 
(like ZIEGLER (1964b) for pharetronids or OlT 
(1967) for sphinctozoans), but not until the 
discovery of the recent demosponge Vaceletia 
crypta (VACELET 1977) and thorough in-
vestigations of scleres in fossil species (e.g. 
REITNER 1987b,1990) it became clear that the 
sphinctozoans are highly polyphyletic. This fact 
makes it necessary, to expect different life 
strategies in different sponge taxa with 
sphinctozoid basal skeletons. To avoid the pitfalls 
of deductions from phylogenetic assumptions four 
species have been selected from the rich and weIl 
preserved fauna of the Cassian Beds, representing 
at first glance different "end members" of sphincto-
zoan organization. Asswning that all stern from 
254 
• 
• 
i 
N 
• 
·st. C."'an 
• .. "\ 
, 20km I 
Fig. 1: The material used in this study is originated from three locations with "erratie boulders" with 
Cassian fauna (Calllian, Triassie). S = Seelandalpe (Alpe di Specie) R = Vale di Rimbianco P = 
Passo Giao 
different lineages, a palaeobiological model has 
been established for eaeh of them, in the end 
leading to the differentiation of organiVltional 
grades (sensu SIMPSON 1961). 
Therefore the species were, as far as 
possible, only assigned to higher taxa in the system 
of the Porifera by eharaeters of spieulation. 
2. Material 
The material was eollected by J. Reitner from three 
locations in the Dolomites near Cortina d'Ampezzo 
(see fig.l). They have long been famous for a very 
rieh and exeellently preserved invertebrate fauna, 
the Cassian fauna, in the Cassian Beds of lower 
Carnian age. UnfOI'tunately this fauna appears only 
in so ca1led "erratie boulders", i.e. theyare exposed 
in blocks of up to several eubie meters volume em-
bedded in an argillaceous matrix, whieh makes 
faeial and stratigraphie correlation diHicult. To 
date it secms the most probable, that they stern 
from small biostromes or biohetms whieh arose in 
shallow water at the transition between the Upper 
Cassian Dolomite, where existing basins had been 
filled with platfonn debris and the base of Dürren-
stein Dolomite, with its onset of a new trans-
gression phase, thus being of Julian age 
(austriacum-zone). Others interpreted them as 
small pateh reefs in the baek reef area or as Cipit 
boulders transported into the basins of the Upper 
Cassian dolomite (see fig.2; for further infOllnation 
see FÜRSICH & WENDT 1977, BOSSELINl 
1989, RUSSO et al. 1991). An overview over the 
sponge fauna of the Cassian Beds is given by 
DIECI et al. (1968). 
The investigated tbin sections are deposited 
at the Institut für Paläontologie at the Freie Uni-
versität Berlin. 
3. Methods 
The observations for this study were taken only 
from thin sections, using nOIIl1al as weH as pola-
rized light. Additionally, luminescenee teehniques 
were applied. These techniques have been in use in 
carbonate sedimentology only sinee a short time, so 
that some technieual and explanatory annotations 
are necessary. 
Lumineseence is the capability of certain 
minerals, to emit light when exeited by an energy 
source due to certain impurities of the erystal 
lattice. Techniques can be distinguished according 
to the exeitation souree. For this study ultraviolet 
and further short wave length light and an electron 
beam were used as energy sourees. Good general 
informations about luminescenee techniques are 
contained in MARSHALL (1988) and MACHEL et 
al. (1991). 
The luminescenee caused mainly by ultra-
violet light from a Hg-high pressure lamp 
(epitluorescence) is generally believed to be eaused 
by the exeitation of organies inclosed in carbonates 
(van GUZEL 1979). We got best results using two 
different non-UV filter sets to dete .. mine the auto-
Fig. 2: 
NORIAN 
z 
ce 
z 
a:: 
ce 
u 
z 
C 
-
:-:-:.:-:.-:-;.:.~-:.:.:-:-:-:-:.:-:-:.:-:-:-:-: . 
. '. -.-.-.'.".-.-.-.-.-.".".".-." ---"-------"-"-" 
·0"0·.-.·.·.·." J.-.-." J,",".".-." _"." .-."."_ 
.' .",".-.••.•.. -.-.-.-... -." .".".".-.".-.-.-.-." 
.. " .-."." .-." -"." ."." .".-... "." .-.".-.-.-.-.-." 
--
+ 
Upper 
CauIan 
L.I ------"---,,,,---,--,I Bain aedimen18 
255 
, 
Ralb1 Fm. 
, } ' 
Upper ca.lan Fm. 
, 
Lower CauIan Fm. 
<J 
. 
• I Wengen Fm. I 
, - , 
1) .'Ibr RUSSO 1991 
2) allbr FÜRSICH 6. WEHDT 1979 
N 
, 
C3 
C2 
Cl 
13 
• 
Stratigraphie scheme of the Car lIian of the Southem Alps with the proposed source for the "erratie 
boulders" with Cassian fauna. (eombined from FÜRSICH & WENDT 1977 and RUSSO et al. 
1991) 
Catbodc-ray lumincsccnce 
Fc ++-contcnt 
Mn ++-contcnt 
o.idi.la, Milieu reducfDI 
~ 
Burial diagenesis 
+ 
Time 
Fig. 3: 
~ 
Solu -
AI. oDite 
The interpretation scheme that was 
for the luminescence behaviour 
created by a cathode ray. The relative 
ranges of dissolution of cabonates and 
the amount of available Mn I I and 
Fe I I-ions during burial diagenesis are 
shown in dependance of Eh-evolution. 
The upper graph shows the evolution 
of the resulting luminescence 
intensity. 
epifluorescence behavior of mineralized skeletons. 
These two filters are the longewave high-per-
fOllnance narrow-band pass filter BP 546nm / 12 / 
LP 590nm (green, no. 487715) with a red 
ßuorescence and the high perfOlluance wide-
bandpass filter BP 450-490 nm / LP 520 nm (blue, 
no. 487709) with a yellow ßuorescence. The light 
source was a Hg-high-pressure vapour lamp 
attached to an Axiophot microscope (ZEISS). Pho-
tographs were taken with an Ektachrome Tungsten 
film 160 ASA. The intensity of emitted light can 
be to estimate the content of organic matter in 
the skeletal C3lbonate. 
The second energy source used is that of an 
electron beam produced in a cathode, the lumine-
scence resulting being called cathodoluminescence 
(CL). The investigations were made at the Institut 
für Interdisziplinäre Paläontologie at Erlangen, 
kindly supported by Dr. J. MEHL, using a Techno-
syn 8200 Mk 11 apparatus, equiped with a cold 
cathode. The polished and not covered tbin 
sections were bombed with electrons in an He-at-
mosphere at 0.17-0.20 torr pressure at a voltage of 
256 
• 
• • 
• 
• 
•• 
••• • • 
• • • • 
•• • • • 
Fig. 4: 
• 
I 
.' . 
• 
• • • 
· .. ' 
• • 
• • 
. , .. 
. .' 
• • • • • • 
• 
• • • 111 11 
Genetic classification of basal skeleton 
in the "Coralline Sponges" as shown 
in a trabecular sphinctozoan. I: 
primary basal skeleton with trabecules 
and cbamber-roof 11: secondary basal 
skeleton (vesicules) ill: tertiary basal 
skeleton (synvivo-diagenetically filled 
vesicular voids) 
12-15 kV and a current of 420-450 J.LA, thus 
getting stable conditions. Photographs were taken 
with an Agfachrome 1000 RS film (1000 ASA). In 
contrast to Hg-Iamp light the energy of a cathode-
ray is high enough to excite Mn I I -ions enclosed in 
the CaC03-lattice. This effect may be subdued, if 
Fe I I -ions are present in the lattice at the same 
time. Because the presence of Mn I I and Fe I I is 
depending on the diagenetic milieu, in which the 
mineral phase is fonned, the CL of carbonates 
shows a characteristic evolution with burial 
diagenesis (see fig. 3), making interpretations on 
original mineralogy of carbonate phases and a 
relative chronology of diagenetic events possible. 
One of the results from this study is, that the 
main advantage gained by the use of the two 
techniques for palaeobiological investigations is 
the much higher resolution of certain micro-
structural details in the skeleton. Additionally UV-
L and further short wave ligth can successfully be 
used to estimate the content of organic material in 
skeletal carbonates, giving some hints on their 
genesis. CL yields good results concerning the 
degree and relative chronology of diagenetic 
alterations of the skeleton, also leading to 
infounations on the original mineralogy. 
4. Te. minology 
For microstructural types the tellninology 
according to JONES (1979) was used, while for 
skeletal elements of sphinctozoans the terminology 
worked out by FINKS (1983) and SENOWBARI-
DARY AN (1990) was taken over. 
Nevertheless, recent developments in the 
of coralline sponges afford some 
modifications. It seems to hold valid for all 
coralline sponges, that skeletogenesis is ocwling in 
three steps: a first one, in which a rigid, coherent 
framework is built, in which the sponge houses 
(primary basal skeleton). In a second step basal 
parts of the skeleton are left by the soft tissue 
leaving behind thin, unpellneable membranes 
inserted into the primary basal skeleton, mostly 
called vesicula (secondary basal skeleton). The 
resulting voids in the primary skeleton are filled 
with e.g. mucopolysaccharids and other acidic 
macromolucules, which induce the third step of 
skeletogenesis. This step comprises passive 
calcification events, so that the voids can 
completely be filled up with calcium carbonate in 
the end (tertiary basal skeletons). By their relative 
position skeletal elements can be readily classified 
according to during which of the three steps they 
were built (see fig. 4). 
The described genetic classification is 
important in two respects: Firstly it allows 
interpretations on skeletogenesis, the position of 
the living sponge relative to the skeleton, and on 
the function of the skeleton in palaeobiological 
investigations. Secondly it forces to discriminate 
structures that seem similar, but are different in 
origin and function. An example for this are the 
numerous types of filling tissue found inside the 
chambers of sphinctozoans. While the reticular, 
trabecular and septal filling tissue belong to the 
primary basal skeleton, vesicular, "spore like" and 
"pisolithic" belong to the secondary. Tubular filling 
tissues can be fonned by the secondary or primary 
basal skeleton. An identification of these two may 
lead to severe misinterpretations, because tubular 
structures in the primary skeleton may serve as an 
aquiferous system, while tubes originating from the 
secretion of the secondary basal skeleton can never 
have this function, due to the unpenlleability of 
these skeletal structures. 
5. Systematic descriptions 
Phylum Porifera GRANT 1872 
Classis Demospongiae SOLLAS 1875 
Subclassis Tetractinomorpha LEVI 1973 
Ordo Astrophorida SOLLAS 1887 
Familia Geoüdae GRAY 1867 
Genus Cassianothalamia REITNER 1987a 
Cassianothalamia zardinii REnNER 1987a 
(Text-fig. 5, pl. I, pl. IV/fig. 1-3) 
257 
1985 ?Stylothalamia n. sp.: REnNER & 
ENGESER, p.170, pl.4, fig.8-12 
1987a Cassianothalamia zardinii n.g. n.sp.: 
REnNER, p.571 ff., pIs 1-3 
1988 Cassianothalamia zardinii REnNER: 
ENGESER & APPOLD, p.73 ff. , pIs 1-2 
1989 Cassianothalamia zardinii REl'l'N1T.'ER: 
GAUIRET & CUIF', p.4 ff., pl.l, fig.l, pl.2, 
figs 1-2,6, pl.3, figs 2-3 
1989 Cassianothalamia zardinii REl'l'NER: 
, 
SENOWBARI-DARY AN, p.481, text-fig.3, 
pl.2, figs 9-10, pl.7, figs 1-6, pl.8, figs 9-10 
1990 Cassianothalamia zardinii REnNER: CUIF 
et al., p.24 ff., text-figs 2E, 3E, 4, pl.l , figs 
2,8-10, pl.2, fig.5 
1990 Cassianothalamia zardinii REnNER: 
SENOWBARI-DARY AN, p. 136, text-fig.8, 
pl.5, figs 1-6, pl.6, figs 9-10, pl.24, fig.8, 
pl.51, fig.9-1O, pl.52, fig.I-5 
1990 Cassianothalamia zardinii REnNER: 
REnNER, p.34, text-fig.l 
1991a Cassianothalamia zardinii REnNER: 
RElTNER, p.202 
1991 Cassianothalamia zardinii RElTNER: 
RUSSO et al., pl.52, figs 4-5, pl.53, figs 1-6 
1992 Cassianothalamia zardinii REITNER: 
REnNER, p. 140, text-fig.28, p1.15 
Material: 44 specimens in 14 thin sections 
were investigated, whereof 4 were stained with a 
solution of ali711rine-s and potassium-hexa-
cyanoferrate-ill. Autoepifluorescence was observed 
in 8, CL in 3 thin sections. 
Description: The primary basal skeleton of 
C. zardinii is constructed of 0.4 mm tlat chambers, 
whose roofs, pierced by exopores of 0.1-0.2 mm 
diameter, are sustained by 0.1-0.3 mm wide 
trabecules (for exact dimensions see table 1). The 
trabecules rest on the roof of the preceding 
chamber, sharply separated from the latter. While 
the chambers are widely overlapping, they form 
half-spherical aggregates up to 3 cm in diameter. 
This ideal fOlIll can be m.odified by the f011ll of 
small crypts, in which this sponge houses (see pl. 
1), by differential growth, when the basal 
skeleton is overgrown by ' encrusters, and by 
budding. If C. zardinii reaches a certain size, a 
central spongocoel is fonned, which contains 
endopores of slightly larger diameter than that of 
the exopores. The material of the primary basal 
skeleton is adense, irregular micrite (RElTNER 
1987a observed crystal sizes of 1 JUD). Epi-
tluorescence is very bright which shows that 
organics, responsible for the secretion of the 
skeleton, are still contained. CL shows only low 
luminescence, hinting at a little diagenetic 
alteration. Both observations point at an original 
mineralogy of high-Mg calcite, which already was 
I 
1 
-
-
• 
• • '. . . 
, 
• 
258 
• 
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.' ... 
. '. . 
, ' 
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. '. . 
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• . . , , 
, ' 
, 
. , 
• • • 
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--:, 
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, 
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.... . . 
, " , 
.. .. . 
. . .... ' . 
, 
, 
, 
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, ' 
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, 
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. ,.. . . , 
... " - ..... 
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. . . \ 
• • • 
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•• • 
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• • 
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.. . .. . 
, 
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=.,.,... . 
, , 
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.. , 
.. . . .. 
• • • • • 
• • • • 
• • 
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• • 
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. , 
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. 
.. . 
• 
. . . .' 
.. , . 
Fig. 5: A: Growth strategy of the basal skeleton of Cassianothalamia zardinii B: Biomineralization 
process of Cassianothalamia zardinii (For explanation see text) 
postulated by REITNER (1987a) and ENGESER & 
APPOLD (1988), who speak of 15-25 Mol-% 
MgC03. In some cases the trabecules show areas 
of recrystallisation at their flanks and in the top, 
where they broaden to fOlln the segment roof. In 
these areas there is no epifluorescence and CL, 
probably due to late diagenetic Fe I I -uptake 
(earlier cements show brightly luminescing 
rims).The trabecules always show a thin, denser 
envelope. In very thin thin sections one can 
observe, that thread-like structures of reddish-
brown colour nm through the central part of the 
trabecules and spread into the segment roofs, 
surrounding the exopores. 
The secondary and tertiary basal skeleton is 
secreted to a very variable extent. While almost 
completely missing in some specimens, it fills up 
nearly all of the skeleton's inner voids in others. 
The secondary skeleton comprises vesicula, that 
are inserted into the primary skeleton, dearly 
attached to the primary elements and arranged in 
"fronts". The latter show no dear orientation, but 
may nm obliquely through the skeleton. Voids, 
isolated from the environment by primary and 
secondary skeletal elements, are usually sediment-
free or may be filled with skeletal carbonate of the 
tertiary basal skeleton. If they remain hollow, they 
often show a different succession of early dia-
genetic cements, than voids in contact with the 
surrounding medium. 
Both secondary and tertiary basal skeleton 
show the same mineralogy, luminescence characte-
ristics and microstructure as the primary, except 
for the tertiary basal skeleton being distinctly 
laminated under normal and epifluorescence light. 
The primary as weil as the secondary and 
tertiary skeleton are able to entrap encmsters 
(mostly GirvaneIJa-like, thin micritic crusts), sedi-
ment particles and in rare cases scleres. The latter 
always are slightly sigmoid monaxones and totally 
replaced by late diagenetic, non-Iuminescent Fe-
calcite, pointing at an original siliceous minera-
logy. Secretion of secondary and tertiary basal ske-
leton, disturbance of the architecture of the primary 
basal skeleton and encrustation are clearly 
correlated. 
• • 
• • • • • 
• 
. '. . 
. .' ., 
• • 
• '. .,.. - - :......110 • • 
•• ' •• If'-.":*~\ \ • 
.' . " ... . 
---_ • • • ",.-', .,', I •• 
· \ . '. . 
- -• r··· ..... \" • 
.' . .  . \ . 
. .' . . .. \ . . 
• • • • • • 
• • 
• • 
• 
• 
. . . . . . , 
• 
. ... -- ... _-
• 
• 
I,. ~ , 
• • 
• • • • • 
• 
• • • 
• • • • 
• 
• • • • • 
• 
• 
1 
• • 
. .' 
. . , . 
• 
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• • 
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• 
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• 
259 
• 
• 
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• 
• 
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J) 
3 
Fig. 6: A: Growth strategy of the basal skeleton of Cryptocoelia zitteli B: Biomineralization process of 
Cryptocoelia zitteli (for explanation see text) 
Remarks: REITNER (1992) assigned C. 
zardinii to the Geoüdae, due to the finding of 
sterraster microscleres in some specimens. The 
species has until now been found in the lower 
Camian Cassian beds, the Upper Call1ian 
Lechkogel beds (ENGESER & APPOLD 1988) and 
in Norian Cipit boulders in Turkey 
(SENOWBARI-DARY AN 1990). 
Demospongiae inc. sed. 
Genus Cryptocoelia STE 1882 
Cryptocoelia zitteli STE 1882 
(Text fig. 6, plate Wfig. 1 & 2, pI. IV/fig. 4 & 5) 
zitteli n. gen n. sp.: 
,p.176, pL7, fig.5, pL5, fig.4 
1967a Cryptocoelia zitteli STE : OTT, 
p.42, pL9, fig.5-7 
1968 Cryptocoelia zitteli STE : DIECI 
et al., p.149, pl.33, fig.2 
1971 Cryptocoelia zitteli STE : JAB-
LONSKY, p.342, text-figs 8-9 
1973 Cryptocoelia zitteli STE : JAB-
LONSKY, p.185, pU, figs 1-2, pL2, figs 1-
2 
1973 Cryptocoelia zitteli STE :WOLFF, 
text-fig.4/3 
1974 Cryptocoelia zitteli STE : ASSE-
RETO & MONOD, text-fig.14/A 
non 1978 Cryptocoelia zitteli STE : 
SENOWBARI-DARY AN (in: FLÜGEL et 
al.), p.I71, pL24, fig. 4, pL26, fig.l, pL28, 
fig.3-4 
non 1980 Cryptocoelia zitteli STE 
SENOWBARI-DARY AN, pl.3, fig.3 
• 
• 
1980 Cryptocoelia zUteli STE : DULLO 
(in: DULLO & LEIN), pLI, fig.8, pl.3, fig.3 
1981 Cryptocoelia zUteli STE : SE-
NOWBARI-DARYAN, pLI, fig.I-2 
non 1981 Cryptocoelia zitteli STE 
NOWBARI-DARY A,N, pL2, fig.2 
: SE-
1981 Cryptocoelia zUteU STE 
SEK et al., pI. 10, fig.2,4 
1983 Cryptocoelia zitteli 
NOWBARI-DARY AN 
p.183, pI. 6, fig.3 
: TURN-
1983 Cryptocoelia zUteli STE 
RICH, pL5, fig.l 
1987 Cryptocoelia zUteli S 
et al., pl.3, fig.9/A 
& 
. SE-
•• 
SCHAFER, 
: BEN-
:DULLO 
1990 Cryptocoelia zUteli STE : SE-
NOWBARI-DARY AN, pL29, figs 3-4, 
pl.34, fig.l 
non 1990 Cryptocoelia zitteli STE 
SENOWBARI-DARY AN, pL29, fig.lA 
• 
• 
1991 Cryptocoelia zitteli STE : BOlKO 
et al., p. 140, pl.41, pI. 42, fig 1 
• 
Material: 10 specimens in 12 thin sections 
were investigated, whereof 6 were stained with a 
solution of alizarine-s and potassium-hexacyano-
fenate-III. Autoepifluorescence was observed in 6, 
CL in 3 thin sections. 
Description: The basal skeleton of C. zitteli 
shows the same overall appearance as that of C. 
zardinii, i.e. it is composed of ca. 1 mm flat 
chambers overlapping each other and sustained by 
somewhat irregular 0.1-0.2 mm wide trabecules 
and pierced by small round exopores of 0.1-0.2 mm 
diameter. Through the skeleton runs a retro-
siphonate, narrow (0.8 mm in diameter) spongo-
coel with endopores. In contrast to C. zardinii these 
chambers fOIm up to I cm wide cylinders wich 
may be branched. The bases of the trabecules are 
sha1ply separated from the roof of the preceding 
chamber by thin micritic lines (probably of micro-
bial origin, because they sometimes contain fila-
mentous structures), while their tops broaden to 
form the segment roof. 
The secondary and tertiary basal skeleton 
contains the same elements like C. zardinii, i.e. 
vesicula and laminated deposits of skeletal car-
bonate inside the voids of the primary and 
secondary basal skeleton. Vesicula are often seen to 
close off ontogentic older parts of the skeleton. 
Here too, there is a correlation between disturbance 
of skeletal architecture, encrustation (c. zitteli is 
very often incrusted by small Tubiphytes-skeletons) 
and secretion of secondary and tertiary skeletal 
elements. In one specimen it was possible to ob-
serve a monaxone sclere, built into a trabecule of 
the basal skeleton. It is assumed to belong to C. 
zitte/i, because other foreign particles were never 
seen being incorporated into the skeleton. It shows 
the same diagentic alteration as those in C. 
zardinii, thus probably being of originally silicious 
composition. 
Despite the great similarities in general 
appearance, C. zitteli can clearly be distinguished 
from C. zardinii, due to its peculiar microstructure. 
The primary skeletal elements are quite distinctly 
laminated, i.e. they are composed of discrete 
elements, having a dark envelope and containing 
coarser crystalline, thus lighter, material. In speci-
mens of good preservation, Autofluorescence is 
very bright in the coarser crystalline inner part of 
these elements. In specimens with stronger dia-
genetic alteration, CL shows bright colours in these 
areas, of the same colour and intensity as early 
diagenetic cements, pointing at relatively early 
solution events. 
Some specimens are totally recrystallized by 
late diagenetic non-Iuminescent Fe-calcite. In these 
ske1etal elements are only distinguishable by 
pyrite secreted in small crystals around them, in 
some instances even tracing the lamination. The 
strong recrystallization of some specimens and 
early diagenetic recrystallization events point, 
260 
when contrasted with the preservation of Mg 
calcitic C. zardinii, at an originally arago'ntic 
mineralogy of the basal skeleton. 
Remarks: With the discovery of a siliceous spicule 
in C. zitle/i, a demosponge affinity is highly 
probable (hexactinellids are not known to form 
calcareous basal skeletons). By its peculiar 
rnicrostructure it is clearly set off from other 
sphinctozoan groups. To avoid confusion with 
other species (e.g. Solenolmia manon, which is 
sirnilar in general appearance), only species 
described in literature, which show this lamination, 
were taken up into the synonmy list. Wether all 
species contained in the family Cryptocoelidae 
erected by SENOWBARI-DARYAN (1990) 
possess this distinctive feature, which rnight be an 
acceptable argument for monophyletic origin, 
seems doubtful. 
C. zit/eli is a frequent part of biohermal 
limestones from the Ladinian-Carnian western 
Tethys (DULLO et al. 1987, TURNSEK 1984) and 
the upper Triassic of the southwestern Parnir 
(BOIKO et al. 1991). Other species of this genus 
are known from the Norian and Rhaetian of Sicily 
(SENOWBARI-DARY AN 1980, 1990). 
Genus Jab/onskya SENOWBARI-DARYAN 1990 
Jablonskya andrusovi (JABLONSKy) 1975 
(Text-fig. 7, pI. II1fig. 3-5, pI. IIIIfig. 2) 
1975 C%spongia andrusovi n. sp.: JAB-
LONSKY, p.267 ff., pis 1-3 
1978 Follicatena cautica OTT: SENOWBARI-
DARYAN (in: FLÜGEL ET AL.), p.167, 
p1.28, fig.2 
1981 C%spongia andrusovi JABLONSKY: 
SENOWBARI-DARY AN, pl.5, fig.2 
1983 Colospongia andrusovi JABLONSKY: 
SENOWBARI-DARY AN & sCHÄFER, 
p.181, pl.2, figs 2,7 
1987 Colospongia andrusovi JABLONSKY: 
DULLO et al., p.532, p1.4, fig.2 
1989 "Colospongia" andrusovi JABLONSKY: 
SENOWBARI-DARY AN, p.475, pl.2, fig.8, 
pl.l1, figs 7-9 
1990 Jab/onskya andrusovi (JABLONSKY): 
SENOWBARI-DARY AN, p.140, text-fig. 
15, p1.9, figs 7-9, p1.49, figs 1,3-4, p1.51, 
fig.8 
1991 C%spongia andrusovi: RUSSO et al., 
pI. 50, fig.2, pI. 51, fig.4 
•• partim 1992 Ce/yphia submarginata STER): 
REITNER, p. 131 ff., text-fig. 24/e-f, text-
fig. 25/a, pI. 10, fig.l; non: text-fig. 24/a-<1, 
text-fig. b, pUO, figs 2-5, pis 11-12 
Material: 11 specimens (partly fragmented) 
in 5 thin sections were investigated, whereof 4 
were stained with a solution of alizarine-s and 
t4..4-.r:):" I '~ ""-'.*::..: ", ,. 
.. ~' /<.r;~ . 
. ~ ') "'\ ', '. 
. 1 ·' • • I •• 1\ ' 
· ~ .. , . ' 1 " ~""!!, fI" 
· · ·. i'~ . . ' " , ' ~'. · _· .. ... :'>l 
-' I ' . ,I - . I , ' ..•. • 'j. ' "\!'·'l 
. . I ' . ,.', 1 ' \ " .."} 
. .... I I' I . . . .. : . • , . ~ 
• ', • ' • • ' JIo .\1- . . .,,:,. ' rA ., 
• f , ' I I . ' ''' "' 
· . . , .~. ~
· . . .1 . .. .' ; - 1.,.. ;..... 
. - . " \ . . - /7,' '" I " . . . /1' .. I_ J ,)".', ., ... . I ".' '. V (I., 
', 1" .. -1 ... / • , '1 
.. . .. ~ 
- , ', ' ,' ," ', 11 ' ,. ', 
1.- •• ',' , 1. ·0: 
..... . . .. '. , I. 
- ' .. . 
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• • • 
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1 
3 
261 
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, I 
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. . J l\ . . • . • 
• " ~ ' ••.. , J ~ . ~-
.. \, . / . . , 
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· . , I · " -' . IJ . 
· . " . 
· ... ". ', /AI ·\I . . '.' , :.. ~. 
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,/ .' . ." 
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• • • • 
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. . .. . ·.· 1'- ... ", ., ,=. 'J 
· . , \ "" ... ' "1 -" ....:(.·V_ 
"; . " .' . . ' .' '( , / -.,' " . ~ ",. .... ...,. . 
"... ) . ". --: I . . " l .....  I ' 
-" '.' \ 1 - ... " . 
- - . . 
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. .." . ." '" ", . ' \ '/'" . ~. .. " .. ". 
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. ' . .' . . ' " /' - . . 
· , , " ~." 
• 
• • 
• 
" . . " 
• 
• • • • 
• 
• • 
4 
Fig. 7: Growth strategy ofthe basal skeleton of Jablonskya andrusovi (For explanation see text) 
potassium-hexacyanoferrate-lli. Intensive autoepi-
fluorescence was observed in 4, CL in 2 thin 
sections. 
Description: J. andrusovi possesses a primary 
basal skeleton, that is composed of several 
spherical or barrel-folilled segments of up to 1 cm 
diameter, ananged in a catenulate manner. Tbe 
walls have a thickness of ca. 0.4 mm and are 
pierced by a lot of regularily arranged exopores of 
0.1 mm diameter. In the top of the segment or 
losely spaced on the outer walls appear larger, up 
to 1 mm wide openings (oscula), surrounded by a 
collar-like extension of the wall. Tbe walls contain 
irregularily branching 0.05 wide "channels", which 
never lead outside the wall. It was shown by 
• 
SENOWBARI-DARYAN (1989), that these are 
not spicules, due to their ultrastructure, so that he 
tel med them as "pseudospicules" . 
Tbe microstructure of the primary basal skeleton is 
irregular micritic, being of an original high-Mg 
calcite composition (11 Mol% MgC03 after 
RUSSO et al 1991), like in C zardinii. Auto-
epifluorescence shows high intensity, especially in 
the described "channel-like" structures, pointing at 
contained organies, while CL is only weak. a hint 
at little diagenetic alteration. This primary basal 
skeleton is not secreted, where it rests on a 
substrate, be it a preceding segment or an allen 
substrate. Succeeding chambers are often separated 
by thio layers of micrit coDtaioiog 
particles. 
The segments of the primary skeleton are 
mostly filled with vesicular filling tissue -
only one small specimen lacking these structures -
comprising the secondary basal skeleton. These 
vesicula are ca 0.2 mm, at the maximum 0.4 mm 
thick, adding an inner layer of up to 0.7 mm 
thickness to the segmental walls of the primary 
basal skeleton and thus dosing the exopores. It is 
also fot med at the base of segments, such that 
intersegmental walls gain the three-Iayered 
appearance. Vesicules can also be contained inside 
the exopores. 
The vesicules show an arrangement with the 
concave sides pointing out of the segment. In some 
cases one or two exopores are left open, with a 
succession of vesicules pointing towards them with 
their concave side. The last voids, being in contact 
with the environment by these open exopores are 
often filled with sediment. They sometimes have a 
tubular form, leading to the osculum at the top of 
the preceding osculllm. Similar tubular structures 
may run through the center of segments, 
interconnecting them. Because they are fonned by 
impenneable vesicules, they can not be inter-
pretated asretrosiphonate spongocoels as has been 
done by SENOWBARI-DARY AN & SCHÄFER 
(1983). At the osculllm they are frequently 
sustained by bundles of monaxone scleres, pointing 
back into the chamber.They were probably 
primarily siliceous, as shown by their replacement 
by highly Illminescent (CL) early diagenetic 
cements and non-lllminescent late diagenetic Fe-
calcite. 
Another frequent and peculiar structure are 
small, roundish objects of up to 0.4 mm diameter 
and with a larninated appearance. They are in dose 
contact to the vesicular filling tissue, actllally being 
fonned by it and may contain smaIl sedimentary 
partides as cores. They thus resemble the elements 
of the pisolithic filling tissue of Pisothalamia 
described by SENOWBARI-DARY AN & RIGBY 
(1988). 
Remarks: The Mg calcitic mineralogy 
distinguishes J.andrusovi dearly from the Genus 
C%spongia, with which it shares a lot of other 
characteristics. The dosure of the exopores by the 
secondary basal skeleton and their rarer 
appearances in vertical sections often provoked 
confusions with other apo rate sphinctozoans such 
as Follicatena (SENOWBARI-DARY AN 1978) 
and Ce/yphia submarginata (REnNER 1992). 
Up til now J. andrusovi has only been found 
in Call1ian reef limestones of the Western Tethys 
(Dolomites, Northern Calcareous Alps, Carpathi-
ans, former Yugoslavia and Turkey). 
262 
Porifera inc. sed. 
Genus Amb/ysiphone//a STE 
Amb/ysiphone//a strobiliformis DIE CI et aI. 1968 
(text-fig. 8, pI. IWfig. 1, 3 & 4, pI. IV/fig. 6) 
1968 Amb/ysiphone//a strobilijormis n. sp. : 
DIE CI et aI., p.142, text-fig.9, pI.29, figs 1-
3, pl.33, fig.3 
Material: 5 specimens in 3 thin sectioos 
were investigated, all being stained with a solution 
of alizarine-s and potassillm-hexacyonoferrate-lII. 
Auto-epifluorescence was observed in 2 thin 
sections, CL in 1. 
Description: A. strobiliformis has a 
catenulate primary basal skeleton, whose flat 
chambers are of trapezoidal fonn and of up to 2.8 
cm width and up to 4 mm height. The segments get 
broader with ontogeny while staying constant in 
height. Each segment, sometimes separated by a 
layer of micritic crusts, rests on the preceding one 
with the oarrower end, such that the whole 
skeleton gets fllnnelshaped with a dear outer 
segmentation. If the chambers reach a certain 
width (about 1 cm) a narrow secondarily 
retrosiphonate spongocoel is instalied, which later 
(at a segment width of ca 2 cm) wideos to fonn a 
funnel in the segment roof (primarily 
retrosiphonate). The segment walls are about 0.6 
mm thick, getting much thicker at the upper edges 
of the segments. In their outer portion they contain 
irregular branching pores of 0.05 mm diameter, 
while the segment roof is pierced by openings of up 
to 0.23 mm diameter. The spongocoel wall is quite 
thin and penetrated by irregular pores. The 
skeleton is rooted by skeletal material intruding 
into pores of the substrate. 
The microstructure of the primary basal 
skeleton is spherulitic with spherulits of 0.1 mm 
diameter. Due to intensive diagenetic alteration by 
replacement in fOlIl1 of Fe-calcite, auto-
epifluorescence as weIl as CL show weak 
luminescence colours, a hint to originally 
aragonitic composition. In some instances, 
spherulits contain small cores of intense 
epifluorescence, caused by remaining organics. 
The secondary and tertiary basal skeleton 
are weakly developed in A. strobiliformis. Only 
some of the charnbers contain thick vesicula, which 
dose off large, basal parts in the respective 
chamber. Inside these spaces, the inner side of the 
segmental wall may be covered with another layer 
of skeletal carbonate with an orthogonal 
microstructure. It is interpreted as the tertiary basal 
skeleton, being produced by epitactic growth of the 
spherulits in voids left by the living tissue of the 
sponge. The same process is responsible for the 
fact, that a lot of pores of the outer walls are dosed 
secondarily. 
f T 
-
r t t 
r..--I . . . .. " 
1 
263 
2 3 
, 
1. .. , ,- - - Il-. . ... . ... 
. .. . o. . . . \. • .. 
• • • ",' ',, ' 1 : • • • 
. . 
r . . '. .. . . . . \., .; " . '" . ,' .. ": . '/ ' ,"" 
• • 
..... • : . •..... -.!. :~ ... 
' .. C-" -"" - , -~ • • • • . • 
- ... 
• • • . . 
,.
. . .. . " 
• • • 
- .e 
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• • • 
• 
• 
• 
• • 
• 
• 
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• 
• • 
• 
• 
• • • • 
• 
• . . 
• • • • 
• • 
• • • 
______ ~ -_.- 1 
4 
~- 3 ~-- 2 
Fig. 8 : A: Growth strategies of the basal skeleton of A mblysiphone/Ja strobiliformis B: Biomineralization 
process of Amblysiphone/Ja strobiliformis (for explanation see text) 
6. Palaeobiological reconstructions 
6.1. Palaeobiological interpretation of general 
features of sphinctozoan coralline sponges 
Sphinctozoans, with their chambered basal 
skeleton, have long been interpreted as sponges, 
though their eloser affinities remained doubtful. 10 
some cases there were proposals to ascribe them to 
hydrozoans (c. zit/eli; STE 1882) or 
even to algae (Celyphia submarginata; RAUFF 
1914). Arguments for their poriferan origin where 
spicules (Barroisia; RAUFF 1914) or tubular 
systems inside the cbambered skeleton combined 
with a system of exo- and endopores (SEILACHER 
1962, OIT 1967a). The latter can be used as an 
argument, because it implies a filtering life habit 
(compare experimental results of BALSAM & 
VOGEL, 1973, referring to archaeocyathids). 
Further evidence for the above assumption came 
from the (re-)discovery and elose examination of 
recent poriferans with calcareous basal skeletons 
and especia1ly of the sphinctozoan Vaceletia 
crypta. All species described in this study show 
either of these features and microstructural 
sirnilarities to recent cpralline sponges, so that 
their poriferan nature is confinned. 
As has been noted earlier, this study tries to 
ouiline palaeobiological r models for different 
sphinctozoan species separately, so that details will 
follow in the succeding sections. But two general 
features are common to all sphinctozoans, such 
that their interpretation is best dicussed in this 
section. These are the possession of achambered 
basal skeleton and the fonIIation ofvesicula. 
The fonnation of massive, calcareous, basal 
skeletons is known from a variety of sponges today. 
Basically the secretion of such a skeleton should 
follow the general pattern, proposedly valid for all 
metazoans and given by the following model 
(following WHEELER & SIKES 1989): Ca' '-ions 
attach to free ca1boxyl-groups of the so-called 
soluble matrix (i.e. soluble in EDT A-solution) 
fOlilled by polypeptids in an ordered manner. The 
structure and composition of these polypeptids (ß-
sheets) determines the nucleation rate and 
mineralogy (aragonite or calcite) of the resulting 
precipitate. To develop the ability of the soluble 
matrix to capture Ca' '-ions, a second set of 
organic macromolecules, the insoluble matrix, is 
needed, on which the soluble matrix can be 
attached. Details for this process are still unknown 
for sponges, but it seerns probable that exo-
pinacodenn cells produce the soluble matrix, while 
specialized cells of lophocyte or collencyte origin 
("Iarge cells with granules" in Acanthochaetetes, 
REITNER 1991b) provide the insoluble matrix. 
During the described process, the participating 
organic substances are enriched in the precipitate, 
and can be made visible by using epifluorescence 
• 
nucroscopy. 
As this luminescence can be observed in all 
species under examination here, it is highly 
probable, that a similar process is responsible for 
their skeletogenesis. It was especially RAUFF who 
argued in a very polemic manner for a diagenetic 
origin of calcareous basal skeletons, especially in 
his very detailed work on Barroisia (RAUFF 
1914). That the basal skeletons of sphinctozoans 
were produced by the organisrns themselves can be 
derived from two other facts too. These are the 
frequent overgrowth by epizoans (SElLACHER 
1962, ZIEGLER 1964a) and by borings 
(STE 1882, OTI 1967a). In this study a 
boring occured only once in C. zardinii, but 
epizoans are very frequent (mainly Inicritic crnsts 
of microbial origin, Tubiphytes and coralline 
sponges in some cases). As a matter of fact, these 
two arguments do not show, that the skeleton was 
produced during lifetime, but that calcification had 
to occur shortly after or during death at the latest. 
Another i mporta nt interpretation, that can be 
derived from encrustations and borings of each 
individual chamber is, that each chamber of a 
sphinctozoan is the result of a discrete calcification 
step. This lead HERAK (1944) to regard the single 
chamber as representing the individual, while the 
whole skeleton represented a colony of first order 
(called "Person"), which in turn can form 
branching colonies of second order. 
The basal skeleton in coralline sponges 
serves three main functions: 
1. It enhances the resistability against 
mechanical stress. SC CHER & PLEWKA 
(1981) computed a far higher resistability against 
mechanical stress of the basal skeleton in 
Ceratoporella nichoJsoni, a chaetetid coralline 
sponge, than in scleractinians. 
264 
2. It provides the sponge with a permanent 
substrate, which can be resettled after ecological 
crisis or scavenging. Some coralline sponges are 
known to store special cells for reproduction and 
nutrition ("thesocytes", known from Acantho-
chaetetes weIlsi. MerJia normani and Petrobiona 
massiliana, VACELET 1991, REITNER 1992). 
Experimental results on this topic where gained by 
JACKSON & PALUMBI (1979). 
3. The basal skeleton can be seen as a 
container for Ca' , -waste deposits, which occur 
during metabolic Ca' , -detoxification by the 
sponge. This is especially true for the tertiary basal 
skeleton (for discussion see REnNER 1992). 
The mechanical strength of the skeleton is 
considerably diIninuished by the segmentation in 
sphinctozoans. The advantage of this strategy of 
skeletogenesis is, that the organisms gain a much 
higher speed in vertical growth. This is the reason 
why sphinctozoans are generally more readily 
found in bafilestones than in true framestones. At 
least this is true for the material underlying this 
study, with the exception of C. zardinii, which 
houses in smal I crypts in coralline sponge-
hydrozoan framestones (compare FAGERS'IR0M 
1984). 
Except for possessing a basal skeleton, there 
is hardly any other feature common to all coralline 
sponges. The only exception is the possession of 
vesicula, which is also known from skeletons of 
archaeocyaths, scleractinans, brachiopods, 
bryozoans and rndists (ZIEGLER & R lETSCHEL 
1970). Their distinct nature, which lead us to 
designate them as secondary basal skeleton, has 
been recognized early. SElLACHER (1962) and 
OTI (1967a) interpreted them as residues of an 
retreating epithelium, in analogy to the scleracti-
nians. In VaceJetia crypta they are known to be 
formed by thin organic lamella, which are 
produced by the basal endopinacoderm, and 
separate the upper part of the skeleton inhabitated 
by the sponge tissue from the basal part, which has 
been left by the sponge during upward retreat and 
is filled with an organic mucus (VACELET 1979, 
REnNER 1992; compare also SENOWBARl-
DARY AN 1990). They are impermeable, which 
also can be seen in fossil species, as voids 
deliInited by vesicula are nearly always sediment-
free in contrast to other voids. Due to their 
fonnation at the base of the living tissue, they are 
of great value in palaeobiological reconstruction, 
because they enable to detect strategies of retreat in 
the basal skeletons. 
The function of such lamellas is primarily to 
separate parts of the basal skeleton, which can not 
be supplied with food, from the remaining living 
tissue. This is shown by specimens in our material, 
where vesicula are fOIllled in the vicinity of 
encrnstations covering the pores, and of contact 
zones with other skeletalized organisms. Food 
• 
crises may also force the organism to retreat from 
the basal parts of skeletons, and last but not least 
the sponge may reach its maximum size of the soft 
body, which leads to the same result. The 
remaining voids below the vesicula are filled with 
an organic mucus, which may calcify in a later 
stage (tertiary basal skeleton). Because the 
secretion of the tertiary basal skeleton may be 
interpreted as a Ca I l-detoxification (see above), 
the vesicula may also have an osmoregulatory 
function. 
The interpretation of vesicula is complicated 
by the fact, that there are sirnilar structures called 
tabulae in chaetetid Acanthochaetetes weIlsi, 
which are fOfllled by the basal exopinacoderm, and 
enclose thesocytes (see above). The importance of 
this strategy will be discussed in the section on J. 
andrusovi. 
6.2. Palaeobiology of Cassianothalamia zardinii 
(text -fig. 5) 
The basal skeleton of C. zardinii shows very close 
sirnilarities to the recent sphinctozoan species 
Vaceletia crypta in all respects, except for its 
mineralogy being Mg calcitic. This already lead 
REnNER (1987) and ENGESER & APPOLD 
(1988) to compare these two fOHIIS. Nevertheless 
the basal skeleton has to be seen as a convergence, 
due to their known systematic positions, which 
shows the high systematic value of skeleton 
mineralogy. 
Like Vaceletia crypta, C. zardinii erects its 
flat cbambers successively, as is shown by the clear 
separation line between the base of trabecules and 
the roof of the preceding chamber as weil as by 
• 
enclosed encrusters. The creation of a new cbam-
ber may have occured according to the following 
scheme, in analogy to Vaceletia crypta and as 
shown by fine structures: Part of the soft tissue 
emigrates from the already existing cbamber and 
erects an organic matrix, preforming the later 
skeleton. This organic matrix is composed of an 
organic envelope (sometimes seen as a thin 
redbrown line bordering the primary basal 
skeleton) containing a central thread forwing a 
network of organic fibres extending into the roof 
and the tlanks of the trabecule (in some cases seen 
in fossil material). The whole envelope is filled 
with organic mucus, which induces the calci-
fication progressing from the center outwards. 
Remaining organics can be seen by auto-
epifluorescence in the resulting precipitate. The 
recrystallization areas in the trabecules may be due 
to an incomplete calcification. The intel nal 
structure of theresulting skeleton is very regular, 
but gets irregular, if the calcification process is 
disturbed (encmstation, rapid vertical growth). 
The overall form of the skeleton is balf-
spherical, but may be cbanged by the form of the 
265 
crypt inhabitated or by encrustations. C. zardini 
shows a high potential of regeneration by asexual 
budding. That this is provided by reproductives 
enclosed by the vesicules is not plausible, 
the latter are not permeable and fragmented 
specimens are very rare. The resettlement was 
rather due to remaining soft tissue in the surficial 
regions of the skeleton. The spongocoel, formed in 
late ontogenetic stages, served the reduction of the 
way the water current had to pass through the 
living tissue. In analogy to Vaceletia crypta the 
exopores can be interpieted as prosopores and the 
endopores as apopores. In earlier ontogenetic 
stages the function of apopores may have been 
taken over by apical pores. The diameter of these 
pores (140~) does surely not represent that ofthe 
ostia, because the latter havediameters of only 50 
~ (BAR 1982). It is much more plau-
sible that the skeleton was enclosed by a thin exo-
pinacodermallayer, narrowing the pores. 
The vesicula were most probably produced 
as described for Vaceletia crypta above. Their 
position shows, that the living tissue of C. zardinii 
could inhabit several cbambers. Position and extent 
of the living portion of the skeleton flucluated 
highly, showing distinct phases of retreat, probably 
due to food crises or similar factors. The cor-
relation of encrustation or other factors of growth 
disturbance and the secretion of a secondary and 
tertiary basal skeleton proves, that the latter play a 
role in the physiological reaction potential of the 
organism, when exposed to ecologic stress. In some 
cases it was observed, that the secondary and 
tertiary basal skeleton enclose alien particles and 
scleres, functionally reminding of pearls. The 
encapture of spicules shows no systematic orien-
tation, so they play no integral part in skeleton 
formation. 
6.3. Palaeobiology of Cryptocoelia zitteli (text-fig. 
6) 
In spite of the overall sirnilarity in skeletal 
architecture with C. zardinii, C. zitteli shows a 
, 
very different strategy of skeletogenesis. While the 
skeleton is also fOIllled by flat, clearly distin-
guished chambers, supported by trabecules, the 
elements of the prirnary skeleton are laminated. 
These structures point to the fact, that the skeleton 
of each segment was raised successively by joining 
discrete elements, and not in a rash, progressing 
manner as in C. zardinii. In specimens of good 
preservation, these elements are shown by auto-
epifluorescence to resembIe small cushions, pro-
vided with an envelope and filled with ca.bonate 
material rich in organics. The assertion of OTI 
(1967a), that the trabecules grow downwards is not 
plausible, and surely due to section effects. Rather 
the segment was built upwards by piling up the 
elements. 
A recent pendant is not known to the 
authors. Only the secretion of calcillmcarbonate in 
an extrapinacodellllal layer of mucus, as observed 
in Acanthochaetetes, may serve as an inter-
pretation. This would mean, that the elements, 
• 
constituting the trabecules, where fonned in a 
small void between the basal pinacodenn and the 
already finished part of the basal skeleton. An 
alternative could be, that small containers with an 
envelope of organic material were produced by the 
organism, and then stacked to fOlm the skeletal 
elements. The organic mucus inside these 
containers then induced calcification. Here, as in 
C. zardinii, the calcification of the secondary and 
tertiary skeleton followed the same basic process as 
that of the primary basal skeleton, as is shown by 
the occurrence of the same rnicrostructural and 
lurninescence features. 
The appearance of vesicules, a tertiary basal 
skeleton, and scleres have to be interpreted as in C. 
zardinii. That vesicula close off greater parts of the 
skeleton from the surrounding is rare. This may be 
due to the cylindrical, branchy habit, which 
reduces the distance between exo- and endopores. 
Additionally to this the narrow, retrosiphonate 
spongocoel served the same function. This growth 
fonn is also responsible for the fact, that C. zille/i 
is able to in habit a milieu of higher sedimentary 
input. A rapid vertical growth is also seen in the 
much greater segment height, which frequently has 
the consequence, that the trabecules get highly 
irregular. Tubiphytes, which is very often seen 
enclosed in the skeleton, was most certainly a 
commensal, and points to quite long growth 
intelluptions between the erection of segments. 
6.4. Palaeobiology of Jablonskya andrusovi (text-
fig.7) 
After the two sphinctozoan species described in the 
previous sections, which have a skeleton composed 
of tlat segments sustained by trabecules, we now 
turn to a "typical" sphinctozoan with a primary 
basal skeleton built up by spherical, hollow 
chambers. J. andrusovi is very similar to C. 
zardinii in skeletal mineralogy (Mg calcite), 
microstructure (irregular micritic with some 
Jaminations), diagenesis, and organic content (both 
skeletons with intense auto-tluorescence and 
remains of organic matrix in foun of red-brown 
fibres). The only difference in these respects seeIOS 
to be the occurence of "pseudospiculae" in the 
walls of J. andrusovi , which are interpreted as an 
additional element of the organic matrix respon-
sible for calcification. Although these facts suggest, 
that skeletogenesis occured in the same way in both 
mentioned species, it is unplausible, that J. 
andrusovi should construct its segmental walls by 
an organic matrix, raised by a not yet differentiated 
cell assemblage emigrated from the preceding 
266 
chamber, because such a structure would be 
extremly instable. It is more plausible, that the 
calcification occured in the cortex of an individual 
already established with an active choanosome on 
the preceding chamber. This special way of basal 
skeleton formation will be discussed in more detail 
in the next section. 
The exopores, piercing the walls of J. 
andrusovi, do not show the dimensions of ostia 
(max. 50 ~). Thus it is problematic to decide, 
wether they were inhalant or exhalant openings. In 
the text fig.7 illustrating . the palaeobiology of J. 
andrusovi, the first alternative was favoured, 
although it is as likely, that the real inhalant ostia 
lay in the cortex around the exopores, having been 
closed during calcification. That they are 
represented by the channel-like struktures in the 
chamber wall is unprobable, because these never 
lead outside the wall. The larger openings in the 
apex of the charnbers and on their side were surely 
exhalant openings or oscula. 
The most striking feature of J. andrusovi is 
its secondary basal skeleton, developed as adense 
vesicular filling tissue. The following observations, 
partly already made by SENOWBARI-DARY AN 
(1990) and FINKS (1990) are important for the 
interpretation: 
l. If one aSSIImes, that vesicula represent 
remains of a retreating soft tissue with the concave 
side showing into the direction of retreat, the soft 
tissue was withdrawn from the chamber in phases, 
beginning at the base of the chamber and 
progressing upwards and outwards. 
2. By this process the exopores are closed by 
a vesicular layer coveriog the inside of the chamber 
walls. Three cases can be distinguished: The pore 
is closed inwards from the outside, the pore is 
closed outwards from the inside and the pore 
• 
remarns open. 
3. The vesicular filling tisue fonns tubular 
syteIOS, which interconnect segments or lead 
outside and are attached to oscula. 
From these observations the following 
picture about the skeletogenesis of J. andrusovi 
arises: In the beginning stands a fully differentiated 
individual, provided with a SUbdeIßlaJ cortex and 
resting on the precee.ding chamber. At a certain 
point, not exactly fixable, the cortex calcifies with 
the exception of the contact area to the substrate 
(where probably no cortex was fonned). Later, the 
soft tissue withdraws from the newly built 
chamber, leaving behind vesicula. Partly soft 
tissue, resting on the chamber walls (probably the 
exopinacodenn) is drawn into the chamber by this 
process. The soft tissue leaves the chamber through 
some exopores and the oscula. At the oscula, the 
exhalant channels leading towards them are 
pictured by vesicula, including scleres stabilizing 
them. In the end, a new individual is established on 
the newly left charnber. 
This bebaviour can be interpreted as a 
special of a well known phenomenon in 
sponges, the tissue regression. During this process 
the soft tissue is reduced, thereby losing its 
choanosomal organization and changing a lot of 
cell types into arcbaeocytes. Such tissue reductions 
are called forth by ecological crises 
and/or reproductive phases (for more detailed 
information see SIMPSON 1984). A special case of 
tissue reduction is the production of reductiae in 
Ephydatia jIuviatilis, where large voids appear in 
the sponge tissue during the metamorphosis, lined 
by endopinacodenn (as seen from illustrations in 
WEISSENFELS 1989). This endopinacodenn rnay 
well bave been responsible in J andrusovi for 
secretion of vesicula. The seasonal occurence of 
such processes explains segmentation in this case 
in an elegant way. 
The tubular systems enclosed in the 
vesicular filling tissue show that the basal skeleton 
probably still bad some storage function, either for 
nutritive and/or reproductive cells. Such storage 
function of the basal skeleton are known from a 
number of recent coralline sponges (Acantho-
chaetetes. Mer/ia. and Petrobiona, see V ACELET 
1990, REITNER 1989, 1991a) and gemmulae, 
special reproductive cells also known from marine 
species in the recent, were found in the fossil 
sphinctozoan Ce/yphia submarginata by RElTNER 
(1992). 1f this were the case in J andrusovi. it is 
possible, that the vesicula were secreted by the 
exopinacoderm, in analogy to the tabulae of 
Acanthochaetetes, which seclude the stored theso-
cytes from the rest of the soft tissue. The frequent 
occurrence of fragmented specimens may bave 
been a way of propagation for the reproductives 
(propagation by fragmentation is also known in 
recent sponges, see BATIERSH I I .L & BERG-
QUIST 1990). 
Another function of the dense vesicular 
filling tissue was of course to stabilize the hollow 
chambers. It can additionally be observed, that ve-
sicula are secreted around intruded foreign 
particles to foun pisoid-like structures. They would 
thus bave a function as pearls. 
6.5. Palaeobiology of Amblysiphone//a strobili-
jormis (text-fig. 8) 
Like J andrusovi, A. strobiliformis has a primary 
basal skeleton composed of large, hollow chambers 
arranged in a catenulate manner. Nevertheless it is 
different in not possessing such adense vesicular 
filling tissue. 1f we assume, that the secondary 
skeleton is always secreted, when the soft tissue 
retreats, several chambers of A. strobiliformis must 
bave been inhabited at the same time. This is 
additionally suggested by the spongocoel 
penetrating most of the segments. The exopores 
point at the following course of the water current: 
267 
the narrow channels on the side of the chamber 
were inhalant pores, while the wider exopores in 
the segment roof and the endopores in the 
spongocoel functioned as exhalant pores. This 
interpretation is possible, in contrast to J. 
andrusovi, the exopores show a clear 
differentiation in size. 
The difficulty in expJaining the secretion of 
large, hollow chambers was already hinted at in the 
preceding section. To solve this question, we first 
turn to the spherulitic microstructure of A. 
strobiliformis. Such a microstructure is known 
from the recent coralline sponge Astrosclera 
willeyana, where the spherulits, composing the 
basal skeleton, are secreted around seed nuclei 
containing high amounts of organics (GAU'IRET 
1986). Their existence in A. strobiliformis is 
proved by the appearance of centers with intense 
auto-fluorescence inside the spherulits. These seed 
nuclei are produced intracellularily (according to 
REnNER (1991b) in vacuols of special cells 
derived from the pinacodelIn, according to WOOD 
(1991) in surficial arcbaeocytes) and then 
transported to their destination, where they grow 
on epitactically until they merge. This would mean, 
that A. strobiliformis possessed a specialized 
subdermal zone as the zone of destination for the 
seed nuclei. Such subpinacodermal zones are well 
known from a number of sponge taxa and called 
cortex (for details see SIMPSON 1984). According 
to VACELET (1971) a cortex has the function to 
protect and stabilize the sponge. 
Thus the growth of A . strobiliformis can be 
described in the following way: In a fully 
differentiated sponge provided with a cortex, 
calcification starts with the intracellular production 
of seed nuclei by specialized cells. The seed nuclei 
are transported into the cortex and start to grow 
epitactically. Because it is known, that the 
possession of a cortex may inhibit growth, it is 
probable, that calcification marks a point where the 
maximum of growth is reached. This is also 
suggested by the very constant height of the 
segments. After calcification the soft tissue extends 
, 
out ofthe newly formed chamber. 
Except for chamber height, A. strobiliformis 
shows a clear ontogeny in the succeei!ing 
chambers: the breadth of each segment is 
continually increased. 1f a certain breadth is 
reached, a secondary retrosiphonate spongocoel is 
instalied, which later widens to fOlm a primary 
retrosiphonate spongocoel. This clearly shows, that 
the installation of a spongocoel is a function of 
chamber volume, in the way, that it reduces 
transport ways of the filtered water in the chamber. 
The occasional appearance of vesicules, shutting 
off basal parts of large chambers, rnay bave served 
the same function. 
The capability to grow epitactically is 
retained by the spherulits. This is shown by the 
fact. that the oarrow iobalant cbaonels are closed 
by epitactic growth, and by the forlllation of an 
additional inner layer of the walls of an orthogonal 
microstructure (as part of the tertiary basal 
skeleton). In both processes remains of organic 
substaoces in dead portions of the basal skeleton 
may have been of great irnportance. The pisoidal 
structures, fixed to the vesicular filling tissue, have 
to be interpreted as pearls, like in J. andrusovi. 
6.6. Comparison with other coralline sponges 
Because comparisons always are in need of a 
thorough classification the following one of 
architectural types in fossil and recent coralline 
sponges is proposed, mainly resting on the works 
ofRElTNER & KEUPP (1989), REITNER (1992) 
and WOOO (1991) 
l. the crnstal type: The basal skeleton is 
produced by biomineralization occuring at the base 
of the soft tissue. Two subtypes can be distin-
guished: 
a) simple crnstal type: simple, solid crnsts 
are secreted (known from Hispidopetra); 
Ceratoporella mediates to the chaetetid crnstal 
type by inserting tubes into its basal crnst. 
b) chaetetid cmstal type: The basal crnst 
contains tubes subdivided by horizontal tabulae; 
surficial astrorhizae may be developed (known 
from Acanthochaetetes. Merlia and fossil 
chaetetids) . 
2. the matrix type: biomineralization occurs 
in an organo-spicular matrix, penetrating the 
sponge tissue. The following subtypes exist: 
a) stromatoporoid matrix type: the 
encrnsting basal skeleton is composed of vertical 
and horizontal elements forming a more or less 
regular network. Threedimensional astrorhizae 
systems or mamelones maY occur (known from 
Minchinellidae, Astrosclera, Calcijibrospongia 
and fossil stromatoporoids). 
b) inozoid matrix type: branching basal 
skeletons with an intelllai structure like that of 
stromatoporoids. Prominent chanoel systems may 
penetrate the whole skeleton (early ontogenetic 
Astrosclera, fossil inozoaos and some fossil 
stromatoporoids) . 
c) sphinctozoid matrix type: Horizontal and 
vertical elements of the basal skeleton are arranged 
to fOI 111 achambered skeleton, sometimes pene-
trated by spongocoels (knmm from recent 
Vaceletia crypta and some fossil sphinctozoaos). 
The archaeocyaths (if sponges at all), 
established as a separate type by WOOO (1991), 
can be identified as representatives of the matrix 
type. 
The first two studied species, C.zardinii and 
C. zitteli, with their trabecular skeletons, are easily 
fitted into this scheme as representatives of the 
sphinctozoid matrix type. This is also true for the 
268 
fossil taxa Verticillitidae and Vaceletiidae, 
established by REITNER & ENGESER (1985), as 
is shown by comparisons with Vaceletia crypta. 
From these two taxa C. zardinii is only different in 
spiculation and mineralogy of the basal skeleton. 
Other fossil taxa of the same architectural type are 
Solenolmia ("Dictyocoelia'~ manon and the genus 
Zardinia, which even shares the same mineralogy 
of the basal skeleton with C. zardinii. Interesting 
comparisons can also be made with archaeocyaths 
(see DEBRENNE & VACELET 1984. ZHU-
RA VLEV 1989). . 
C. zitteli with its very special mode of 
biomineralization can only be compared with a few 
fossil representatives in this respect. There are only 
some members of the family Cryptocoeliidae and 
some Palaeozoic stromatoporoids with a similar 
laminated microstructure. 
In contrast the two other studied species, J. 
andrusovi and A. strobiliformis, represent a third 
basic architectural type within the coralline 
sponges, which we call the cortex type. It is 
characterized by the fact, that biomineralization of 
a basal skeleton occurs in a specialized, subdermal 
zone of the more or less spherical sponge 
individual. Thus the classical sphinctozoans are 
distributed in this classification among two groups, 
notwithstanding possible transitional forms. The 
cortex type is not known from the recent. 
The growth strategy of J. andrusovi with its 
retreat from the chambers, is weIl known from 
other fossil taxa. As examples may serve: the genus 
Colospongia (see OTT 1967b) from the Triassic 
and the genus Salzburgia (see SENOWBARl-
OARYAN & DI STEFANO 1988). There are also 
great similarities with the glomerate sphinctozoan 
Alpinothalamia (see SENOWBARl-OARY AN 
1990). 
A . strobiliformis, last but not least, re-
presents the most widespread type of sphinctozoan, 
being composed of more or less hollow chambers, 
pierced by numerous pores. With its aragonitic 
basal skeleton of a spherulitic microstructure it is 
part of the family Sebargasiidae, responsible for the 
, 
sphinctozoan radiation in the Upper Carboniferous 
and the PeIlllian (FINKS 1990). If they compose a 
monophylum is doubtable though, after all we 
know about the strong tendency of sponges, to 
develop secondary basal skeletons in independent 
evolutionary lineages. Of interest is the inter-
pretation of pisoid-like structures inside the 
skeleton of A. strobiliformis as pearls, which liken 
structures in Amblysiphonella? bullifera (SE-
NOWBARl-DARY AN & RIGBY 1988) and the 
"spore-like" filling tissue of Intrasporeocoelia 
described by RIGBY et al. (1988). If the 
interpretation is true, these structures should not 
have a high systematic value. The same is true for 
the pisoid-like structures in J. andrusovi, which are 
also known from the Pennian Pisothalamia 
(SENOWBARI-DARY AN & RIGBY 1988). 
The model given for J. andrusovi may also 
serve to get a better understanding of the 
sphinctozoaos, which possess an aporate skeleton. 
A lot of them have a prominent vesicular filling 
tissue, so that it is very plausible, that a 
calcitication of the cortex without keeping pores 
open forced the soft tissue to retreat from the 
skeleton through the larger opeoings (probably 
oscula) always present in these taxa (compare 
Follicatena cautica in OTI 1967a, Gyrtiocoelia 
beedi in SENOWBARI-DARY AN & DI 
STEF ANO 1988, and Gyrtiocoelia typica in 
FINKS 1990). In some aporate taxa with a 
spherulitic microstructure (Thanmastocoelidae, 
OTI 1967a) the loss of pores may be due to the 
epitactic growth of the spherulits. 
The possession of pores may serve as a 
feature to subdivide the sphinctozoaos of cortex 
type into further groupings: 
3) the cortex type: biomineraliUltion of the 
basal skeleton occurs in the cortex of more or less 
spherical individuals. The following subtypes cao 
be distinguished: 
a) primarily aporate cortex type: The 
calcified cortex contaios no pores, but only 
occasional openings (probably oscula). 
b) secondarily aporate cortex type: pores in 
the calcified cortex are closed during retreat of the 
soft tissue from the chamber by vesicula. 
c) porate cortex type: pores are left open 
during the calcification of the cortex and the 
vesicula. 
7. Conclusions 
1. The erratic boulders with Cassian fauna from 
the lower Carnian of the Dolomites contain a 
diverse fanna of sphinctozoans, which are 
suitable for palaeobiological reconstructions 
due to their excellent preservation. 
2. Luminescence techniques are successfully be 
nsed for investigations in biomineralization, 
because of their capability to resolve certaio 
structures better than by nOlmal light. 
Additionally they render the possibility to 
estimate the role of orgaoics in 
biomineralization and the course of diagenesis. 
3. The comparison of sphinctozoans with recent 
"CoralIine Sponges" is very valuable for the 
reconstruction of their palaeobiology. The 
iosight in genetic relationships forces to 
modify the traditional classification of basal 
skeletons. One of the consequences is the 
introduction of a new tenuioology for the 
elements of basal skeletons. Three different 
components cao be distinguished: the primary 
basal skeleton (fOlming a rigid network), the 
269 
secondary basal skeleton (vesicules, spread up 
in the former) and the tertiary basal skeleton 
(passively secreted ca.bonates in parts of the 
basal skeleton, which have been left by the 
living tissue). 
4. Cassianothalamia zardinii RElTNER is a 
sphinctozoan with a trabecular architecture. Its 
basal skeleton is analogous to that of recent 
Vaceletia crypta (V ACELET). The Mg 
calcitic, irregular-micritic basal skeleton is 
buHt by the calcification of an organie matrix, 
which penetrates the Iiving tissue. 
5. Cryptocoelia zitteli STE erects its 
aragonitic, trabecular basal skeleton by a 
successive stacking of discrete elements, which 
results in a lamellar microstructure. 
6. Jablonskya andrusovi (JABLONSKY) erects 
its Mg calcitic, irregular-micritic chambers by 
the calcification of a cortex. Later the tissue 
emigrates from the chamber, in connection 
with a tissue regression (triggered by ecologic 
or reproduktive cycles). During this the hollow 
chamber is filled up with vesicules, which 
close of the exopores. The basal skeleton 
possibly posessed a container function. 
7. Amblysiphonella strobiliformis DIE CI et al. 
constructs its chambers by the transport of 
intracellularily secreted, aragonitic seed nuclei 
into a cortex. Inside the cortex the seed nuclei 
grow epitactially to form spherolites. A greater 
part of the basal skeleton is constantly 
inhabitated. 
8. The analysed taxa can be seen as models for 
the greater part of sphinctozoans. As a 
consequence the morphological subdivision of 
the "CoralIine Sponges" into organizational 
types has to be modified: while the 
sphinctozoaos with a trabecular architecture 
alongside the stromatoporans represent those 
coralline sponges, which secrete their skeleton 
in an organic matrix, penetrating the living 
tissue (matrix type), others consist of large 
hollow chambers, thus representing another 
type. Here the calcification happens in a 
, 
subdermal cortex (cortex type). Inside this 
group of coralIine sponges three subtypes can 
be distioguished: the porate, the primary 
aporate and the secondary aporate. Taxa with 
a prominant intelllaI, primary skeleton as weIl 
as uvifOIm and glomerate fOlIns mediate to the 
matrix type. In the third type of coralline 
sponges, the cmstal type, taxa of sphinctozoid 
architecture are not known. 
8. Acknowledgements 
The Deutsche Forschungsgemeinschaft (DFG) is 
gratefully acknowledged for finacial support (Re 
665/1 & 4) 
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271 
RAUFF,H. (1914): Barroisia und die Phare-
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, 
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274 
Plate I 
Fig. 1: CassianolhaJamia zardini REnNER 1987 
Two specimens growing together and being separated from each other by vesicula. They show several growth 
phases due to resettlement of the basal skeleton. The specimen to the left is restricted in growth by 
encrnstations, such that only a small rest is allowed to grow on in a columnar manner (arrow). The volume of 
this part of the basal skeleton is reduced by vesiculae. Scale bar is 3 mm. 
Fig. 2: Cassianolhalamia zardini REITNER 1987 
Specimen with a regular primary basal skeleton structure. The secondary and tertiary skeleton fill up nearly all 
of the skeleton's voids. The whole specimen is surrounded by rnicritic crnsts. Scale bar is 2 mm . 
• 
Fig. 3: Cassianolhalamia zardini REnNER 1987 
Part of a GirvaneLJa-like crnst growing on a segment roof. It is enclosed by the base of a trabecule of the 
following segment. Scale bar is 250 ~. 
Fig. 4: CassianolhaJamia zardini REnNER 1987 
A monaxone sclere [1], a sedimentary particle [2] , and GirvaneLJa-like crnsts [3] are enclosed by elements of 
the secondary and tertiary basal skeleton. The skeletal architecture is disturbed. Scale bar is 250 ~. 
Fig. 5: CassianolhaJamia zardini REITNER 1987 
Two specimens, one dwelling in a smaIl gap between a strornatoporoid and a chaetetid coralline sponge. The 
overall f01l1l of the basal skeleton matches the form of the gap. Qnly the ontogenetically youngest cbambers 
were inhabitated as shown by vesicula closing off the older chambers from intruding sediment (arrow). Scale 
bar is 2 mm. 
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Plate n 
Fig 1: Cryptocoe/ia zitte/i STE 1882 
Oblique section through a specimen. The lamellar microstructure of the basal skeleton is clearly recognizable 
[1] , as well as the narrow, retrosiphonate spongocoel [arrow]. The ontogenetic older parts ofthe skeleton on the 
rightside are cut horizontally and show the irregular, somewhat septal arrangement of the trabecules [2] and a 
dense secondary basal skeleton (vesicula). The segment roofs are covered by thin, micritic crusts. Scale bar is 2 
mm. 
Fig 2: Cryptocoe/ia zitte/i STE 1882 
A sclere integrated into the primary basal skeleton [arrows]. It protrudes from the se<4ment and is covered by 
the micritic crust also covering the segment roof. The originally siliciouis spicule is replaced by Fe-calcite. The 
basal skeleton contains vesicula [1] and deposits ofthe tertiary basal skeleton [2]. Scale bar is 1 mm. 
Fig. 3: JabJonskya andrusovi (JABLONSKY) 1973) 
Specimen with four chambers. The chamber walls [1] consist of micritic, irregular Mg calcite and contain a 
dense vesicular filling tissue constituting the secondary basal skeleton [2]. This fonus a second inner layer of 
the chambers, such that intersegmental walls gain a three-Iayered appearance. The chamber walls are pierced 
by Ollmerous pores (see tangential sectrion of a wall above specimen) which are closed by the secondary basal 
skeleton. The vesicular filling tissue contains pseudopisoids [3] and tubular systems [arrows]. Scale bar is 5 
mm. 
Fig. 4: JabJonskya andrusovi (JABLONSKY) 1973) 
Dense vesicular filling tissue; the thicker vesicula show larnination [arrow]. The pores below on the right side 
are closed by the vesicular inner layer of the chamber wall [2] . The vesicula point out ofthe segment with their 
concave sides. The vesicular void at the far right communicates with the surrounding environment by a pore 
left open [1]. Scale bar is 1 mm. 
Fig. 5: Jablonskya andrusovi (JABLONSKY) 1973) 
Pseudopisoid and vesicula inside a chamber. The pseudo-pisod shows a circumlamellar structure and is in 
direct contact with the vesicula. It contains a sedimentary particle as a core [1]. Scale bar is 250 J.UIl. 
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278 
Plateill 
Fig. I : Amblysiphone/la strobilijormis DIECI et al. 1968 . 
Whole specimen with a spongocoel [1], which develops from a primary retrosiphonate to an secondarily 
retrosiphonate. The chambers are trapezoidal, broadening upwards while keeping a constant height. The upper 
edges of the chambers are thickened. The outer walls contain labyrinthous pores, while the segment roofs 
contain wider, straight pores. Inside the chambers are some vesicules and pseudo-pisoids [arrow]. The first 
segment roots in a bryozoan [2]. Growth is renewed on the spongocoel of the youngest chamber. Scale bar is 5 
mm. 
Fig. 2: Jab/onskya andrusovi (JABLONSKY) 1973) . 
Part of a vesicular tube directly beneath an osculum. The tube is f01ll1ed by vesicula and sustained by collar-like 
arranged scleres [1] . The osculum possesses an elevated rim. The chamber walls contain irregular, chartnel-like 
voids [2] . Scale bar is 500 J.Ul1. 
Fig 3: Amb/ysiphone/la strobilijormis DIECI et al. 1968 
Outer part of a chamber, which has been secluded by a vesiculum [1]. The vesiclular void contains pseudo-
pisoids of spherulitic rnicrostructure [2] and and an inner walilayer of orthogonal to clinogonal rnicrostructure 
[4] added to the spherulitic segment wall. Scale bar is 500 J.Ul1. 
Fig. 4: The outer edge of a chamber. The wall contains labyrinthous, sometimes branching pores. The 
spherulitic rnicrostructure of the aragonitic basal skeleton is discernable. Scale bar is 500 J.Ul1 . 
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Plate IV 
Fig. 1: CassianolhaJamia zardini REnNER 1987 
A monaxone sclere entrapped in the primary basal skeleton. The sclere is passing through a void between two 
trabecula. The originally silicious sclere appears non-Iuminescent under epifluorescence (yellow fluorescence, 
high performance wide-band pass filter BP 450-490 nm, LP 520 nm) due to replacement by Fe-calcite. Scale 
bar is 125 ~ (auto-epifluorescence photograph). 
Fig. 2: Cassianotha/amia zardini RElTNER 1987 
Part of the periphery of the specimen in pI. I, fig. 2 under epifluorescence. Voids in the primary basal skeleton 
are filled with highly Illminescent material from the secondary and tertiary basal skeletpn. The tertiary skeleton 
shows a stepwise decrease in luminescence intensity towards the center of voids. Voids filled with diagenetic 
cements show no luminescence. Scale bar is 125 ~. 
Fig 3: CassianolhaJamia zardini RElTNER 1987 
Part of the primary basal skeleton with vesicula (secondary basal skeleton). The elements of the primary basal 
skeleton show a somewhat stronger luminescent rim. Voids of the skeleton are filled with early diagenetic 
cements in two phases: 1. a palisade cement with low lllminescence [1], 2. a non luminescent bladed calcite 
with a brightly orange luminescing rim [2] . This sequence is developed with much higher thicknesses inside 
the voids closed off by skeletal elements. In the last cementation phase a non-Iuminescent Fe-calcite is 
deposited [4]. Scale bar is 250 ~ (combined noc lila I light and cathode-ray luminescence photograph). 
Fig. 4: Cryptocoe/ia zilteli STE 1882 
The lamella of the primary basal skeleton show a bright luminescence under the cathode-ray [2] , enveloping 
weakly luminescent zones. Colour and intensity of the luminescence corresponds to that of early diagenetic 
cements [arrow). The envelopes of the trabecules and the vesicula themselves show hardly no luminescence. 
Scale bar is 500 ~ (cathode-ray luminescence). 
Fig 5: Cryptocoelia zilleli STE 1882 
Two trabecules with distinct lamination. Auto-epifluorescence light one recognizes, that the trabecules are 
constructed by distinct "cushions", which possess a weak luminescent outer envelope [2] and a brightly yellow 
luminescing interioc [1] . Between the trabecules vesicula [4] and deposits of the tertiary basal skeleton (3] have 
been deposited. Scale bar is 125 ~. 
Fig. 6 : Amb/ysiphoneJla strobiliformis DIECI et al. 1968 
Outer wall under epifluorescence (BP 450-490; LP 520).showing labyrinthous pores filled with brightly 
luminescent grurneleuse micrite [1] . The primary basal skeleton is strongly recrystallized and thus of 10w 
luminescence, except for some brightly luminescent centers of spherulits [arrow]. Scale bar is 125 ~. 
281