www.geologicacarpathica.sk
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA
, OCTOBER 2011, 62, 5, 397—412 doi: 10.2478/v10096-011-0029-4
Introduction
The number and abundance of hypercalcified sponge taxa –
particularly those of chambered construction, the “sphincto-
zoa”, described in this paper – occurring in Permian reefs
and reefal deposits decreased considerably after the dramatic
mass extinction crises at the end of the Paleozoic. Almost all
species of “sphinctozoans” disappeared and holdovers of this
group are not known from the Early Triassic (Scythian) and
from the Middle Triassic pioneer reefs. The absence of Permian
“sphinctozoan” genera in the Middle Triassic also exhibits
the disappearance of this group at the generic level. Morpho-
logically identical or similar “sphinctozoans” appear first in
Norian reefs.
The pioneer Triassic reefs, and consequently the
“sphinctozoans”, appeared first in the Middle Anisian and
became more abundant during the Ladinian to Late Triassic.
The majority of Middle Anisian “sphinctozoans” are abso-
lutely different genera, not known from the Permian reefs.
Anisian reefs and reef builders are known from several local-
ities in the world (Flügel & Senowbari-Daryan 2001), but
the most information about the paleontological contents of
Anisian reefs comes from the Dolomites of northern Italy
(Gaetani et al. 1981; Fois & Gaetani 1984; Scheuber 1990;
Senowbari-Daryan et al. 1993), Austria (Schafhauser 1997),
China (Enos et al. 1997; Lehrmann et al. 1998) and Hungary
(Scholz 1972; Senowbari-Daryan & Velledits 2007a,b). The
known “sphinctozoan” taxa of the Middle Triassic and those
described in this paper are morphologically small-sized,
Sponges from the Middle Triassic reef limestone of the
Aggtelek Karst (NE Hungary)
BABA SENOWBARI-DARYAN
1
, †SÁNDOR KOVÁCS and FELICITÁSZ VELLEDITS
2
This work is dedicated to our late colleague Dr. Sándor Kovács, who unfortunately died before the work was finished. He collected the ma-
jority of the sponge samples described in this paper and the main result of the stratigraphy of the reef carbonates is his contribution.
1
Geozentrum Nordbayern, Department of Paleontology, University of Erlangen-Nürnberg, Loewenichstrasse 28, 91054 Erlangen,
Germany; basendar@pal.uni-erlangen.de
2
Limestone Bt., Andrássy Gy. u. 6., 2120 Dunakeszi, Hungary; fvelledits@freemail.hu
(Manuscript received January 19, 2011; accepted in revised form March 17, 2011)
Abstract: The hypercalcified sponge fauna of the Middle Triassic (Anisian—Ladinian) reef limestone exposed between
Aggtelek-Jósvafő-Égerszög (northern Hungary) is described. Almost all the identified species are chambered sponges
(“sphinctozoa”). Only two fragments of a not determinable species of non-chambered species (“inozoa”) were identified.
Hexactinellid sponges are not found. The majority of the Middle Anisian “sphinctozoans” are absolutely different genera,
not known from the Permian reefs. The sponge fauna of the Triassic pioneer reefs in the Aggtelek Karst are distinctly
small-scaled occurring in “Tubiphytes”-dominated carbonates. The following taxa are described: Amblysiphonella sp.,
Celyphia zoldana Ott, Pisa & Farabegoli, Colospongia catenulata catenulata Ott, C. catenulata macrocatenulata Scholz,
Follicatena cautica Ott, Kovacsia baloghi (Kovács), Solenolmia manon manon (Münster), S. radiata Senowbari-Daryan
& Riedel, Olangocoelia otti Bechstädt & Brandner, Thaumastocoelia dolomitica Senowbari-Daryan, Zühlke, Bechstädt &
Flügel, Thaumastocoelia cf., Th. cassiana Steinmann.
Key words: Anisian—Ladinian, NE Hungary, Aggtelek, reef, sponges, “sphinctozoans”.
while those of the Late Triassic (especially of the Norian—
Rhaetian) are large-scale representatives. “Sphinctozoans”
appear first in the Middle Anisian and become more abun-
dant during the Ladinian to Late Triassic.
It was Scholz (1972) who first gave detailed information
on the fauna and flora of the Aggtelek reef, but his age deter-
mination (Pelsonian—middle Illyrian) was not correct, be-
cause he considered the reef limestone heteropic with the
Steinalm Limestone. His dating was based on dasycladales,
and foraminifera originated from the Pelsonian Steinalm
Limestone. The latest detailed (1 : 10,000) geological map-
ping (Velledits et al. 2011) has revealed, that the Steinalm
Limestone was not heteropic with the Aggtelek reef. The
Steinalm Limestone is overlain by a deep-water limestone
(Jenei Limestone, see Figs. 1a,b, 2 and 3).
This deep-water intercalation constitutes the underlying
unit of the Aggtelek reef. Scholz’s inaccurate age determina-
tion was cited in the literature many times e.g. Kovács
(1978a), Flügel (1982, 2002), Senowbari-Daryan et al. (1993),
Payne et al. (2006).
Sponges of the Hungarian Ladinian—Carnian Wetterstein
Limestone were described by Balogh & Kovács (1976),
Kovács (1978a,b), and Flügel et al. (1991/1992).
The studied “sphinctozoans” originate from the Aggtelek
reef. It is exposed on the Aggtelek Karst, between Aggtelek-
Jósvafő-Égerszög (Fig. 1a) and in the Baradla Cave between
Aggtelek and Jósvafő (Fig. 1b). It forms a belt, which ex-
tends 7 km long in a NW—SE direction and its width varies
between 1—3 km (Fig. 1a).
398
SENOWBARI-DARYAN, KOVÁCS and VELLEDITS
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA, 2011, 62, 5, 397—412
Fig. 1. a – Geological map of the Aggtelek-Jósvafő area (Velledits et al. 2011) and the sampling points (asterisks). Dashed line depicts the
vertical projection of the Baradla Cave. Asterisks – locations of the samples from surface exposures. b – Middle Triassic formations re-
vealed by the Baradla Cave with the location of the sampling points.
399
SPONGES FROM THE MIDDLE TRIASSIC REEF LIMESTONE (AGGTELEK, HUNGARY)
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA, 2011, 62, 5, 397—412
Geologic-stratigraphic setting
The Triassic formations building up the Aggtelek Karst
belong to the Silica Nappe which is the uppermost nappe of
the Inner Western Carpathians (Kozur & Mock 1973; Mello
et al. 1997). According to its lithostratigraphic succession
the Silica Nappe corresponds to the “Juvavicum” of the
Northern Calcareous Alps (NCA) (Lein 1987; Tollmann
1987; Kozur 1991), recently considered to be the “Hallstatt
mélange” (Gawlick et al. 1999). In the Middle—Late Triassic
both the Western Carpathians and the Northern Calcareous
Alps were situated on the northern shelf of the north-western
end of the opening Neotethys (Tollmann 1987; Kovács
1997; Haas et al. 2001; Schmid et al. 2008). The Triassic rift-
ing had a profound influence on the evolution of the Silica
Nappe and thus also on the evolution of the area under study.
According to Kovács (1989) three units (Aggtelek,
Szőlősardó and Bódva) of the Silica Nappe formed a fairly
uniform ramp from the Early Triassic until the late Pelso-
nian. The ramp was dissected due to rifting (Fig. 2). From
that time onward the evolution of the three units was differ-
ent. The major part of the Aggtelek Unit survived as a plat-
form up to the Tuvalian, while the Szőlősardó Unit
represented the shelf-slope and the periplatform environment
and the Bódva Unit corresponded to the pelagic basin bor-
dering the opening ocean.
In the Aggtelek Unit in the late Pelsonian (Binodosus Sub-
zone, Velledits et al. 2011) the uniform Steinalm Platform
was drowned and dissected. Basins and highs were formed.
In the NW part of the studied area the lower—middle? Illyrian
basinal carbonates were overlain by a platform margin reef
(reef stage 1) developed on a morphological high. According
to our present knowledge this is the oldest known Triassic
platform margin reef within the Alpine-Carpathian region.
The reef association is dominated by sphinctozoans and
microproblematicas. The fossils are characteristic of the
Wetterstein-type reef communities (Velledits et al. 2011).
Differently from this in the SE part of the studied region
a basin existed from the late Pelsonian until the end of the
Illyrian (Fig. 1a). During the late Illyrian—Early Ladinian the
reef prograded to the SE, where reef stage 2 was established.
Meanwhile, on the NW part of the platform a lagoon was
formed behind reef stage 2.
In the evolution of the Aggtelek reef two different stages
can be distinguished. (1) Reef stage 1 of Illyrian age. It crops
out only in the NW part of the studied area. Reef stage 1 can
Fig. 2. Middle Triassic deposi-
tional model and Triassic
lithostratigraphy of the Ag-
gtelek and neighbouring units.
(Modified after Kovács 1997.)
Light grey shading indicates
the studied area. R1 – Wet-
terstein Formation reef facies,
stage 1. R2 – Wetterstein
Formation reef facies, stage 2.
be followed on the edge
of the platform in the
neighbourhood of a basin
over a distance of 3.5 km
(Fig. 1a). Consequently we
can conclude that it was a
platform margin reef.
In the Early Ladinian
(Curionii Zone or some-
what
later)
the
reef
(stage 1) prograded into the
SE situated basin, where
(2) reef stage 2 came into
existence. Its age is late
Illyrian—Early Ladinian.
Because reef-building or-
ganisms do not allow exact
biostratigraphic age deter-
mination, the age of reef
stage 1 was determined by
the age determination of the
underlying and intercalat-
ing deep-water limestone,
and with the overlying la-
goonal unit. The conodonts
400
SENOWBARI-DARYAN, KOVÁCS and VELLEDITS
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA, 2011, 62, 5, 397—412
(Gondolella liebermani, G. szaboi, Gladigondolella budurovi
det. by S. Kovács) and radiolaria (Pararuesticyrtium (?) cf.
illyricum, Eptingium cf. ramovsi, Pseudostylosphaera japonica
det. P. Dumitrica) from the underlying basinal carbonates of
reef stage 1 refer to middle Illyrian (Trinodosus Zone up to the
greater part of the Reitzi Zone). The deep-water intercalation
in the upper part of reef stage 1 contains conodonts (Gondolella
fueloepi, G. trammeri det. S. Kovács) referring to Avisianum
Subzone or younger age (Fig. 3). In the overlying lagoonal
limestone Diplopora annulata appears in large quantity, indi-
cating a late Illyrian—Fassanian age.
The age of reef stage 2 was determined with the help of
conodonts from the underlying strata. G. trammeri, G. tran-
sita, G. transita/pseudolonga and much younger forms Gon-
dolella gr. “Bakalovi” (sensu Kovács 1994) indicating late
Illyrian to Fassanian (upper Secedensis to Curionii Zones)
age. The cessation time of reef stage 2 is not known because
the overlying strata is tectonically sheared.
Fig. 3. Stratigraphical positions of the samples.
Fig. 4. “Sphinctozoans” from the Anisian-Ladinian Aggtelek reef limestones, NE part of Hungary. Scale in A—F and K = 2 mm, in G—J = 5 mm.
A – Follicatena cautica Ott. Longitudinal section through a specimen composed of several spherical to barrel-shaped chambers. The cham-
bers communicate with others and with the outside by single openings. Chamber interiors are filled with vesiculae. 79/1/4b. B – Colospongia
catenulata catenulata Ott. Longitudinal section through five chambers. Some vesiculae are secreted in one chamber interior. U9b. C – Colo-
spongia catenulata macrocatenulata Scholz. Magnification of two chambers from a specimen shows the pronounced and even perforation of
the chamber walls. 79/9d. D – 1 – Longitudinal section through two chambers of Solenolmia radiata Senowbari-Daryan & Riedel and 2 – Co-
lospongia catenulata catenulata Ott. Note the alternating growth stages of both sponges: the wall of the early three chambers of ...
Continues on the next page
401
SPONGES FROM THE MIDDLE TRIASSIC REEF LIMESTONE (AGGTELEK, HUNGARY)
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA, 2011, 62, 5, 397—412
Fig. 4. Text continues from the previous page. ... Colospongia is missing and the three chambers are lying on the first chamber wall of Sole-
nolmia. Despite this the wall of the fourth chamber of Colospongia is well developed and here Solenolmia has no wall. E – Colospongia
catenulata catenulata Ott. Longitudinal section through a branched specimen shows the uneven perforation of the chamber walls (Fig. 7). 79/4/1.
F – 1 – Chambered sponge? or foraminifer? gen. et sp. indet. and 2 – Colospongia catenulata catenulata Ott. 79/2a. G – Follicatena cau-
tica Ott. Sections through two specimens show similar characteristics as Fig. A. 79/3b. H – Follicatena cautica Ott. Longitudinal section sim-
ilar to Fig. A. 79/2/2a. I – Follicatena cautica Ott. Longitudinal section similar to Fig. A. 79/2/2a. J – Amblysiphonella sp. Longitudinal
section through numerous low rectangular to crescent-shaped chambers. In the upper part of the sponge the spongocoels are cut. 79/17/1b.
K – Olangocoelia otti Bechstädt & Brandner. Section through numerous small spherical chambers arranged in rows. Chambers are connected
either directly or by a short and narrow tube (arrows). 79/51.
402
SENOWBARI-DARYAN, KOVÁCS and VELLEDITS
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA, 2011, 62, 5, 397—412
Fig. 5. Ratio of chamber height/chamber width in Colospongia
catenulata catenulata Ott (dots, 19 measurements) and Colospongia
catenulata macrocatenulata Scholz (rectangle, 24 measurements).
Samples 79/1—18 originate from reef stage 1 (Illyrian),
whereas samples U1—23 come from reef stage 2 (upper Illyr-
ian—Lower Ladinian).
Systematic paleontology
R e m a r k s: The systematic classification of Fink & Rigby
(2004) with minor modifications is used to describe the
sponges in this paper.
Phyllum: Porifera Grant, 1836
Class: Demospongea Sollas, 1875
Order: Vaceletida Finks & Rigby, 2004
Family: Colospongiidae Senowbari-Daryan, 1990
Synonymy: Colospongiidae Boiko & Belyaeva (in Boiko et
al. 1991); Parauvanelliidae Wu, 1991; Imbricatocoeliidae
Wu, 1991 (see Finks & Rigby 2004: p. 697)
Subfamily: Colospongiinae Senowbari-Daryan, 1990
Genus: Colospongia Laube, 1864
T y p e s p e c i e s : Manon dubium Münster, 1841.
F u r t h e r s p e c i e s: All specimens of Colospongia with
stratigraphic range and geographic distribution are listed by
Senowbari-Daryan & Garcia Bellido (2002).
R e m a r k s: Two subspecies of Colospongia are known:
the type species C. catenulata catenulata from the Wetter-
stein Limestone (Ladinian—Carnian) described by Ott (1967)
and C. catenulata macrocatenulata described by Scholz
(1972) from the Illyrian part (reef stage 1) of the Aggtelek
reef, about 2 km northeast of Aggtelek. According to Scholz
this subspecies differs from C. catenulata catenulata Ott
(1967) by the large chamber diameters (in C. catenulata
catenulata: 1.3—6.0 mm; in C. catenulata macrocatenulata
up to 9 mm). In fact, we also found in the investigated mate-
rial two different-sized Colospongia. The first one (as illus-
trated in Fig. 4B,E) is small and reaches a maximum
chamber diameter of 2.3 mm with a maximum chamber
height of 2.5 mm (Fig. 5). The second species (as in Fig. 4C;
Fig. 6H—I) is a large one with a maximum diameter of
8.5 mm and height of 9 mm (Fig. 5). Data for the thickness
of chamber walls and pore diameters of both species are al-
most identical.
The dimensions of the first species of investigated material
are smaller than the average data reported of Colospongia
catenulata catenulata by Ott (1967) (see above). The dimen-
sions of the large species correspond to the data given for Colo-
spongia catenulata macrocatenulata by Scholz (1972). Based
on chamber dimensions we describe both subspecies separately.
Colospongia catenulata catenulata Ott, 1967
(Fig. 4B,D/2—E; Fig. 7; Fig. 8B)
1967 Colospongia catenulata n. sp. – Ott; p. 32, pl. 7, fig. 3—4, pl. 8,
fig. 1—5
1972 Colospongia catenulata Ott – Ott; pl. 1, fig. 1
1974 Colospongia catenulata Ott – Jablonsky; p. 194, pl. 68, fig. 1
1976 Colospongia catenulata Ott – Balogh & Kovács; p. 300, pl. 1, fig. 5
1983 Colospongia catenulata catenulata Ott – Senowbari-Daryan &
Schäfer; p. 181, pl. 2, fig. 3, 6
1986 Colospongia catenulata catenulata Ott – Senowbari-Daryan &
Abate; p. 63, pl. 1, fig. 7, pl. 2, fig. 1—2
1989 Colospongia catenulata catenulata Ott – Mastandrea & Rettori;
p. 17, pl. 1, fig. a—c; pl. 3, fig. 1
1990 Colospongia catenulata – Ciarapica et al.; Fig. 2/A (left: the
same specimen illustrated in Mastandrea & Rettori 1989)
1997 Colospongia catenulata catenulata Ott – Rüffer & Zamparelli;
pl. 28, fig. 3
2005 Colospongia catenulata catenulata Ott – Emmerich, Zamparelli,
Bechstädt & Zühlke; fig. 11/7, 10
2005 ?Colospongia sp. – Emmerich, Zamparelli, Bechstädt & Zühlke;
fig. 10/7—8
2006 Colospongia catenulata Ott – Nittel; pl. 7, fig. 1
? 2006 Colospongia catenulata Ott – Nittel; pl. 7, fig. 2
M a t e r i a l: In numerous thin sections (for the illustrated
specimens see plate explanations).
D e s c r i p t i o n: Colospongia catenulata catenulata is one
of the most beautiful “sphinctozoan” sponges and easy for de-
termination. The straight or slightly curved sponge is com-
posed of numerous spherical to barrel-shaped chambers
arranged one above another like a chain. The chamber size is
almost constant in the whole sponge stem. Chamber walls are
pierced with evenly sized pores, but occasionally with an un-
even distribution (Fig. 7). The diameter of the pores increases
moderately to the outside of the chamber walls. Chamber inte-
riors are usually hollow, with vesiculae rarely observed in
some chambers. The ratios of chamber heights/chamber
widths of C. catenulata catenulata are shown in Fig. 5.
O c c u r r e n c e a n d s t r a t i g r a p h i c a l r a n g e: Colo-
spongia is a long lasting sponge genus occurring from the
Carboniferous to the Late Triassic. C. catenulata catenulata is
known from several Ladinian—Carnian reef localities in the
former Tethyan realm (Austria, Greece, Italy, Slovakia). The
species is described from the Anisian reef limestones of
Latemar (Dolomites, Italy) by Emmerich et al. (2005). C.
catenulata catenulata was described in Hungary for the first
time from the Ladinian—Carnian Wetterstein Limestone of the
Alsóhegy Karstplateau, northern Hungary by Balogh &
403
SPONGES FROM THE MIDDLE TRIASSIC REEF LIMESTONE (AGGTELEK, HUNGARY)
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA, 2011, 62, 5, 397—412
Fig. 6. “Sphinctozoans” from the Anisian—Ladian Aggtelek reef limestones, NE part of Hungary. Scale in all figs. 5 mm. A—F – Follicatena
cautica Ott. A – Several longitudinal, oblique and cross-sections showing the abundance of sponges in the investigated material. For magnifi-
cation of the specimen indicated with an arrow see Fig. C. 79/3c. B – Section through a branched specimen showing the single openings or
sieve-plates in the chamber walls and the vesiculae within the chamber interiors. U3a. C – The magnification from Fig. A (white arrow)
shows four chambers with thin walls, the perforation pattern of chamber walls and the abundance of concentrical vesiculae within the cham-
bers. 79/3c. D – Section through two chambers showing the vesiculae within the chamber interiors and the sieve-plates in the chamber walls.
U16. E – Similar to Fig. D. The sieve-plate with several openings is clearly visible. U8c. F – Cross-sections through three chambers showing
the abundant vesiculae within the chamber interiors. 79/3b. G – Amblysiphonella sp. The specimen seems to be branched in the middle part or
the ‘normal’ growth has been interrupted. 79/17/1a. H – Colospongia catenulata macrocatenulata Scholz. Section through two specimens (or
a branched specimen?) with numerous spherical chambers. Chamber walls are pierced by evenly distributed pores. Some vesiculae are within
the chamber interiors. 79/9a. I – Colospongia catenulata macrocatenulata Scholz. Section through a specimen showing similar characteristics
to the sponge in Fig. H. 79/9b.
404
SENOWBARI-DARYAN, KOVÁCS and VELLEDITS
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA, 2011, 62, 5, 397—412
Amblysiphonella sp.
(Fig. 4J; Fig. 8C)
M a t e r i a l: Two specimens.
D e s c r i p t i o n: The specimens of this sponge are poorly
preserved and therefore all characteristics are not recogniz-
able. The diameter of both specimens, illustrated in Fig. 4J
and Fig. 8C is about 8 mm and they are composed of low rect-
angular to crescent-shaped chambers with heights of about
3 mm. Some chambers are not well developed. The sponge
seems to have two or more axial spongocoels, which are not
clearly recognizable. Also due to the recrystallization of the
skeleton only indications of the wall perforation can be de-
tected. Rarely vesiculae seem to be present in some chambers.
Family: Solenolmiidae Engeser, 1986
Synonymy: Deningeriidae Boiko (in Boiko et al. 1991)
Subfamily: Solenolmiinae Senowbari-Daryan, 1990
Genus: Solenolmia Pomel, 1872
Synonymy: Dictyocoelia Ott, 1967
T y p e s p e c i e s: Scyphia? manon Münster, 1841.
F u r t h e r s p e c i e s: See Senowbari-Daryan & Garcia-
Bellido (2002: p. 1530).
Solenolmia manon manon (Münster), 1841
(Fig. 9A—D)
1841 Scyphia? manon n. sp. – Münster; p. 29, pl. 1, fig. 15
1987 Solenolmia manon manon (Münster) – Senowbari-Daryan &
Riedel; p. 7, pl. 1, fig. 1—2, pl. 2, fig. 1, 3—4, pl. 3, fig. 7—8
1989 Solenolmia manon manon (Münster) – Mastandrea & Rettori;
p. 22, pl. 3, fig. 2
1990 Solenolmia manon manon (Münster) – Senowbari-Daryan;
p. 89, pl. 29, fig. 1—2 (cum syn.)
1991/92 Solenolmia manon manon (Münster) – Flügel et al.; pl. 1, fig. 9
1997 Solenolmia manon manon (Münster) – Rüffer & Zamparelli;
pl. 27, fig. 4, pl. 28, 5
2005 Solenolmia manon manon (Münster) – Emmerich, Zamparelli,
Bechstädt & Zühlke; fig. 11/3—5, 11
2006 Solenolmia manon manon Nittel; pl. 5, fig. 2 (magnification in
pl. 7, fig. 6)
M a t e r i a l: Numerous specimens (for the investigated
material see the plate explanations).
D e s c r i p t i o n: Specimens of this sponge are composed of
several spherical, hemispherical or barrel-shaped chambers
in a moniliform arrangement one after the other. The cham-
ber height is variable (2.0—6.5 mm) but their diameter in the
whole stem is almost constant (4—5.5 mm). Chamber interi-
ors are filled with a fine reticulate skeleton. Chamber walls
are thin and coarsely perforated, appearing as dark lines in
transmitted light. Compared with the sponge diameter, a rel-
atively narrow spongocoel of about 0.5—1 mm in diameter
(with spongocoel wall up to 1.5 mm) passes internally
through the whole sponge. The spongocoel wall is thick and
exhibits a fine vertically oriented lamellar structure.
O c c u r r e n c e: Solenolmia manon manon is known from
numerous Anisian—Carnian localities in the west Tethyan
realm (see Senowbari-Daryan & Garcia-Bellido 2002: p. 1530;
Kovács (1976). It was also described from the Carnian reef of
the Bükk Mountains (NE Hungary) by Flügel et al. (1991/92)
and now in this paper; both from reef stage 1 and reef stage 2.
Colospongia catenulata macrocatenulata Scholz, 1972
(Fig. 4C; Fig. 6H—I)
1972 Colospongia catenulata Ott,1967 ssp. macrocatenulata nov. ssp.
– Scholz; p. 344, pl. 4, fig. 4, pl. 5, fig. 3—6
M a t e r i a l: At least 5 specimens.
D e s c r i p t i o n: The characteristics of the sponge skeleton
of C. catenulata macrocatenulata correspond to C. catenula-
ta catenulata, described above. The only difference between
the subspecies is their chamber sizes. For comparison of the
chamber sizes of both species the ratios of chamber heights/
chamber widths are shown in Fig. 5.
Occurrence and stratigraphical range: C. catenu-
lata macrocatenulata is known from the Illyrian—Lower La-
dinian reef limestones between Aggtelek and Jósvafő; both
from reef stage 1 and reef stage 2 (see Fig. 1a—b). Scholz
(1972) misdetermined the age of the reef as Pelsonian (see
chapter: Introduction, above).
Family: Sebargasiidae de Laubenfels, 1955
Genus: Amblysiphonella Steinmann, 1882
Type species: Amblysiphonella barroisi Steinmann, 1882.
Fig. 7. A branched specimen of Colospongia catenulata catenulata
Ott shows the unevenly distributed pores of the chamber walls. T – A
specimen of a “Tubiphytes”-like organism embedded in the cham-
ber of the sponge (drawn from Fig. 4E).
405
SPONGES FROM THE MIDDLE TRIASSIC REEF LIMESTONE (AGGTELEK, HUNGARY)
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA, 2011, 62, 5, 397—412
Fig. 8. “Sphinctozoans” from the Anisian—Ladian Aggtelek reef limestones, NE part of Hungary. Scale in all figs. 5 mm. A – Celyphia? sp.
Longitudinal section through several irregularly arranged chambers with thin chamber walls. 79/51. B – Colospongia catenulata catenulata
Ott. Longitudinal section through numerous spherical chambers with evenly perforated chamber walls. 79/3b/2. C – Amblysiphonella sp.
Longitudinal section through several rectangular chambers. The section exhibits in the middle part the existence of one (or several?)
spongocoel(s). 79/17/1a. D – Celyphia zoldana Ott, Pisa & Farabegoli. Section through several circular to rectangular chambers. U8c.
E – Celyphia zoldana Ott, Pisa & Farabegoli. Similar to Fig. D. 79/17/2b. F – Solenolmia radiata Senowbari-Daryan & Riedel. Cross
(small arrows) and longitudinal sections (large arrows) through several specimens. U9c.
synonymy list). Like the preceding species S. manon manon
was described from Hungary for the first time by Balogh &
Kovács (1976).
Solenolmia radiata Senowbari-Daryan & Riedel, 1987
(Fig. 4D/1; Fig. 8F)
1987 Dictyocoelia cf. D. manon minor – Dullo et al.; p. 532, pl. 3, fig. 2—5
1987 Solenolmia radiata n. sp. – Senowbari-Daryan & Riedel; p. 12,
pl. 3, fig. 1—6, pl. 4, fig. 1—4
M a t e r i a l: Several specimens in thin section U9c.
D e s c r i p t i o n: The specimens of this species are composed
of several barrel-like chambers with a diameter of 2.5—3 mm.
Chamber heights (2.5—4 mm) are distinctly higher than
chamber diameters. An axial spongocoel of 0.3—0.8 mm
(measured with the spongocoel wall) or approximately
0.4 mm (without the spongocoel wall) passes internally
through the sponge. Chamber walls are thin and appear as
dark lines in transmitted light. Chamber interiors are filled
with a moderately coarse reticulate fibre skeleton, which
appears arranged radially in cross-sections.
O c c u r r e n c e a n d s t r a t i g r a p h i c a l r a n g e: Sole-
nolmia radiata is known from the Carnian Wetterstein Lime-
stone of Austria (Dullo et al. 1987; Senowbari-Daryan &
Riedel 1987) and from the upper Illyrian—Lower Ladinian
part of the reef (stage 2) east of Aggtelek (Fig. 1a: U1—23),
northern Hungary (this paper).
406
SENOWBARI-DARYAN, KOVÁCS and VELLEDITS
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA, 2011, 62, 5, 397—412
Fig. 9. “Sphinctozoans” from the Anisian—Ladian Aggtelek reef limestones, NE part of Hungary. Scale in A—D = 5 mm, in E—F and J = 2 mm,
in G—I = 1 mm. A—D – Solenolmia manon manon (Münster). A – Longitudinal and oblique sections through several specimens showing the
spherical to hemispherical chambers with thin chamber walls and reticulate filling skeleton within the chamber interiors. The axial spongocoel
is about 1/5 of the whole sponge diameter. The arrow indicates a specimen of Colospongia catenulata catenulata Ott. 79/14b. B – Oblique
section through three chambers clearly exhibiting the reticulate filling skeleton and the thin chamber walls. 79/14a. C – Marginally...
Continues on the next page
407
SPONGES FROM THE MIDDLE TRIASSIC REEF LIMESTONE (AGGTELEK, HUNGARY)
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA, 2011, 62, 5, 397—412
Thin section L NC CH CDB DC TW
U16
4
4
0.8–1.3
0.7
2–2.4
0.1
U21
5.5
5.5 1–1.5 0.7–1.0 1.7–1.9
0.1
U21
3.5 3
1.1–1.3
0.7
1.2–1.3
0.6
Table 1: Biometrical data of Thaumastocoelia dolomitica Senow-
bari-Daryan, Zühlke, Bechstädt & Flügel. L – length of sponge,
NC – number of chambers, CH – chamber height, CDB – cham-
ber diameter at the base, DC – chamber diameter at the top (col-
lar), TW – thickness of exowall. All dimensions in mm.
Family: Celyphiidae de Laubenfels, 1955
Genus: Celyphia Pomel, 1872
T y p e s p e c i e s: Manon submarginatum Münster, 1841.
F u r t h e r s p e c i e s: See Senowbari-Daryan & Garcia-
Bellido (2002: p. 1524).
Celyphia zoldana Ott, Pisa & Farabegoli, 1980
(Fig. 8D—E)
1980 Celyphia zoldana n. sp. – Ott, Pisa & Farabegoli; p. 833, pl. 62,
fig. 1—4
1993 Celyphia zoldana Ott, Pisa & Farabegoli – Senowbari-Daryan,
Zühlke, Bechstädt & Flügel; p. 208, pl. 43, fig. 1—7, pl. 45, fig. 5—6,
pl. 47, fig. 1—2, pl. 48, fig. 1, 3—5, pl. 49, fig. 4—5, 9 (cum syn.)
1997 Celyphia zoldana Ott, Pisa & Farabegoli – Schafhauser; p. 107—108,
pl. 12, fig. 5
2005 Celyphia zoldana Ott, Pisa & Farabegoli – Emmerich, Zamparelli,
Bechstädt & Zühlke; fig. 9/1—7
M a t e r i a l: At least two specimens. Both are illustrated in
Fig. 8D—E.
D e s c r i p t i o n: The small specimens of this sponge are
composed of circular to rectangular chambers with monili-
form or occasionally glomerate arrangements. The spherical
to barrel-shape (in thin section circular to rectangular) cham-
bers are small and usually less than 2 mm in diameter. The
imperforate chamber walls are thick (0.2 mm) and pierced
by one or two ostia in each chamber with upraised edges.
Chamber interiors are without vesiculae. Specimens are usu-
ally attached to the substratum.
O c c u r r e n c e a n d s t r a t i g r a p h i c r a n g e: Celyphia
zoldana is known from the Anisian of the Dolomites, Italy
(Ott et al. 1980; Senowbari-Daryan et al. 1993 and further
references; Emmerich et al. 2005), Karawanken, Austria
(Schafhauser 1997), and from the reef of Aggtelek; both
from reef stage 1 and reef stage 2 (this paper).
Celyphia? sp.
(Fig. 8A)
M a t e r i a l: One specimen only.
D e s c r i p t i o n: The only specimen of this sponge is about
17.5 mm long and is composed of several irregularly arranged
chambers of about 6 mm in diameter. The sponge seems to be
broken in a longitudinal direction, because the chamber walls
are straight on one side and cannot be seen on the other side.
The chamber walls of about 0.1 mm thick are imperforate and
pierced by rare ostia of 0.1—0.2 mm in diameter.
R e m a r k s: Because of the imperforate chamber walls
with some ostia the attribution of this sponge to the genus
Celyphia is possible, but not certain.
Family: Thaumastocoeliidae Ott, 1967
Subfamily: Thaumastocoeliinae Senowbari-Daryan, 1990
Genus: Thaumastocoelia Steinmann, 1882
T y p e s p e c i e s : Thaumastocoelia cassiana Steinmann,
1882.
Thaumastocoelia dolomitica Senowbari-Daryan, Zühlke,
Bechstädt & Flügel, 1993
(Fig. 9G—I)
1993 Thaumastocoelia dolomitica n. sp. – Senowbari-Daryan, Zühlke,
Bechstädt, Flügel; p. 210, pl. 54, fig. 1—4
2005 Thaumastocoelia dolomitica Senowbari-Daryan, Zühlke, Bech-
städt and Flügel – Emmerich, Zamparelli, Bechstädt & Zühlke;
fig. 11/6
M a t e r i a l: Three specimens.
D e s c r i p t i o n: The tiny and moniliform stems of this
sponge are composed of cup-like or funnel-shaped chambers
with a distinctly collar-like edge on the upper part of the
chambers. The diameter of the upper part of the chambers
(collar) varies between 1.2 mm and 2.24 mm in different
specimens. There is no spongocoel, but the chamber roofs
are pierced by several small pores. The chamber walls are
thin, like the roofs. Some ostia seem to be located in the ex-
owall (Fig. 10). The biometrical data of Th. dolomitica are
summarized in Table 1 and a reconstruction is given by
Senowbari-Daryan et al. (1993).
Fig. 9. Text continues from the previous page. ... longitudinal section through numerous chambers showing similar characteristics of the
sponge to Fig. A. 79/12b. D – Section through four chambers showing similar characteristics to Fig. A. 79/14/d. E – Kovacsia baloghi
(Kovács). Marginally longitudinal section through four chambers exhibiting the imperforate chamber walls with only two ostia (arrows). The
thin, dark line appearing within the chamber interiors indicates the spongocoel wall, which is filled with micritic sediment or cement in
part. The white arrow indicates another specimen cut in cross-section. 79/17/1e. F – Thaumastocoelia cf. Th. cassiana Steinmann. Longitu-
dinal section through several chambers filled with spary calcite cement. Chamber exowalls are pierced with ostia. The arrow indicates the dou-
ble-layered interwall with an osculum, cut marginally. The ostia of the exowalls show internally a dark line, which is interpreted as vesiculae,
closing the ostium. U13. G—I – Thaumastocoelia dolomitica Senowbari-Daryan, Zühlke, Bechstädt & Flügel. G – Section through a poorly
preserved specimen with six cup-like or funnel-shaped chambers. The youngest chamber is incompletely formed. U21. H – Similar section to
Fig. G. U21. I – Section through five funnel-shaped chambers. The first chamber is cut only on the collar and the youngest chamber is incom-
plete. The upraised collar-like edge of the chambers is clearly visible. U16. J – Olangocoelia otti Bechstädt & Brandner. Section through nu-
merous small spherical chambers arranged in rows, partly in two rows. Some chambers are incompletely formed (see arrow). 79/26.
408
SENOWBARI-DARYAN, KOVÁCS and VELLEDITS
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA, 2011, 62, 5, 397—412
R e m a r k s: Thamastocoelia dolomitica is one of the
smallest “sphinctozoan” sponges. The dimensions of
sponge, chambers and chamber elements of the Hungarian
specimens correspond to the dimensions of the type material
from the Anisian of the Italian Dolomites. The only differ-
ences between the specimens of the type material and speci-
mens from Hungary are the distinctly developed collars at
the upper edge of the chambers in the Hungarian specimens.
The specimen illustrated from the Anisian reef limestones of
Latemar (Dolomites, Italy) by Emmerich et al. (2005) also
shows well developed collars like the Hungarian specimens.
O c c u r r e n c e a n d s t r a t i g r a p h i c r a n g e: Thaumas-
tocoelia dolomitica is known from the Anisian of the type lo-
cality (Reflugio Berti/Comelico, Dolomites, Italy) and from
the Anisian reef limestones of Latemar (Dolomites, Italy). The
third locality where Th. dolomitica occurs is the upper part of
the Aggtelek reef (upper Illyrian—Lower Ladinian: reef
stage 2). In all three localities it is a rare sponge species.
Thaumastocoelia cf. Th. cassiana Steinmann, 1882
(Fig. 9F)
M a t e r i a l: One specimen only.
D e s c r i p t i o n: The one specimen of this sponge reaches a
length of 25 mm and is composed of six hemispherical cham-
bers in which the oldest and youngest chambers are broken.
The chamber diameter is more or less constant in the whole
sponge, reaching about 6 mm. Each chamber roof is double-
layered and pierced by one opening (or perhaps occasionally
by more, as they are not situated in the middle of the roofs).
Chamber exowalls are pierced by ostia of up to 1 mm in dia-
meter. Thickness of the chamber walls is about 0.4—0.5 mm.
The chamber interiors have no filling skeleton or vesiculae.
R e m a r k s: The asiphonate construction of the sponge,
the double-layered chamber roofs and the ostia in the exo-
wall are similar to Thaumatocoelia cassiana. The chamber
roofs of Th. cassiana are pierced by several openings which
could not be proven for certain in this sponge. In addition
Th. cassiana is known only from the Carnian time interval;
the present species is from the upper Illyrian—Lower Ladin-
ian part (reef stage 2) of the reef.
Genus: Follicatena Ott, 1967
T y p e s p e c i e s: Follicatena cautica Ott, 1967.
F u r t h e r s p e c i e s: See Senowbari-Daryan & Garcia-
Bellido (2002).
Follicatena cautica Ott, 1967
(Fig. 4A,G—I; Fig. 6A—F)
1967 Follicatena cautica n. sp. – Ott; p. 22, pl. 1, fig. 1—7
1978b Follicatena cautica Ott – Kovács; p. 301, pl. 4, fig. 5
1978b Follicatena cf. cautica Ott – Kovács; p. 302, pl. 4, fig. 2
1989 Follicatena cautica Ott – Mastandrea & Rettori; p. 20, pl. 2,
fig. a—c
1990 Follicatena cautica – Ciarapica et al.; Fig. 2/A (right)
1990 Follicatena cautica Ott – Senowbari-Daryan; p. 116, pl. 41,
fig. 4 (cum syn.)
1997 Follicatena cautica Ott – Rüffer & Zamparelli; pl. 28, fig. 3
2005 Follicatena cautica Ott – Emmerich, Zamparelli, Bechstädt &
Zühlke; fig. 11/7, 12—13
M a t e r i a l: Numerous specimens.
D e s c r i p t i o n: Follicatena cautica is – along with Sole-
nolmia manon manon and Colospongia catenulata – the
most abundant sponge species in the investigated reef lime-
stones. To show the variable morphology of the sponge, sev-
eral specimens are illustrated. The chamber arrangement of
the single or branched sponge is either moniliform
(Fig. 4A,G—I) or occasionally glomerate (Fig. 6A). Diameter
of the chambers varies between 3 mm and 8 mm. The cham-
bers communicate with each other and with the outside either
by a single opening (Fig. 4A,G,I) or by sieve-plates (Fig. 6E).
Chamber walls are thin, but they can appear thicker, if the
concentrically multilayered vesiculae are secreted within the
chamber interior. Vesiculae are also abundant within the
chamber interiors (Fig. 6A—F). The wall between two cham-
bers may be single or double-layered.
O c c u r r e n c e a n d s t r a t i g r a p h i c a l r a n g e: Follica-
tena cautica is known from the Ladinian—Carnian of several
localities in the Alpine—Mediterranean region (see Senowbari-
Daryan & Garcia-Bellido 2002 and synonymy list). Boiko
(1986) described it from the Carnian of the Pamir Moun-
tains. It occurred in the Aggtelek reef both in reef stage 1 and
in reef stage 2.
Genus: Kovacsia nov. gen.
D e r i v a t i o n o m i n i s: This genus name is dedicated to
our late colleague Sándor Kovács, who described the sponge
in question as Sollasia for the first time.
D e s c r i p t i o n: An imperforate chambered sponge with
spherical to barrel-shaped chambers arranged one above the
Fig. 10. Thaumastocoelia dolomitica Senowbari-Daryan, Zühlke,
Bechstädt & Flügel. The section shows the upraised edges of the
funnel-shaped chambers and some openings in the chamber exo-
walls and interwalls (drawn from Fig. 9I).
409
SPONGES FROM THE MIDDLE TRIASSIC REEF LIMESTONE (AGGTELEK, HUNGARY)
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA, 2011, 62, 5, 397—412
Fig. 11. Kovacsia baloghi (Kovács). A – The holotype of the spe-
cies drawn from Kovács (1978a: fig. 6C) and B – from Fig. 9E
(this paper). Both species show the thin spongocoel wall followed
the chamber exowalls. The holotype shows two openings in one
chamber roof and only one opening in three chamber roofs. Rarely
ostia are cut in the chamber exowalls of both specimens.
other. Chamber size is more or less constant in the whole
stem. Ostia are seldom found in the chamber exowalls. There
is an axial spongocoel following the exowalls with an ex-
tremely thin wall and not everywhere the same diameter.
T y p e s p e c i e s: Sollasia? baloghi Kovács, 1978a.
Kovacsia baloghi (Kovács, 1978a) nov. comb.
(Fig. 9E, Fig. 11)
1978a ?Sollasia baloghi n. sp. – Kovács; p. 686; fig. 5B, 6C
M a t e r i a l: One specimen only.
D e s c r i p t i o n: This sponge species is documented by three
specimens only. The holotype, illustrated in Kovács (1978a:
fig. 6C; redrawn in Fig. 11) is a curved specimen and is com-
posed of eleven spherical chambers. Kovács gives the diameter
of the chambers as 2.5—2.7 mm and their height 2.0—2.6 mm.
The specimen in our collection is composed of four cham-
bers with a maximum diameter of 4 mm and is moderately
larger than the holotype described originally by Kovács
(1978a). The axial canal of approximately 1 mm in diameter
is cut in the two middle chambers. The holotype described
by Kovács exhibits only one in three chamber roofs but there
are two small openings in one chamber roof (see Fig. 11A).
The chamber walls are imperforate, but in two chambers
large ostia are cut marginally (Fig. 11B). The most charac-
teristic feature of the sponge is the thin spongocoel wall,
which is widened in the middle of the chambers (clearly seen
in the holotype, see Fig. 11A), more or less following the
chamber exowalls. As in the holotype the chamber interiors
of our specimen are filled with calcite cement, but the interior
of the spongocoel has a different filling. Such a widening
spongocoel in the middle of the chambers is not known from
the other “sphinctozoan” genera, either from the Paleozoic or
from the Triassic. For this character the new genus Kovacsia
is introduced (see also remarks).
The white arrow in Fig. 9E points to the second specimen
cut in a transversal section. This specimen exhibits the spon-
gocoel appearing as a circle within the chamber.
An aporate chambered sponge with an axial cylindrical
spongocoel is characteristic for e.g. the Permian genera Gir-
tyocoelia Cossmann (1909), or Shotorithalamia Senowbari-
Daryan et al. (2006) but the annulated spongocoel wall
following the chamber exowalls is known only in the new
genus Kovacsia.
R e m a r k s: The chamber walls of Kovacsia baloghi were
characterized by “two layers” according to Kovács (1978a).
In his description he wrote (p. 687): “The imperforate wall
consists of an inner, dark, micritic layer 0.06—0.18 mm in
thickness and an outer, light, recrystallized layer, which is of
0.35 mm maximum thickness and is missing on several
parts”. The “outer, light and recrystallized” layer, mentioned
by Kovács is an incrustation and is not secreted by the
sponge. Consequently the chamber wall consists only of a
single micritic layer appearing as a dark micritic layer in
transmitted light. This is clearly recognizable in our speci-
men illustrated in Fig. 9E.
O c c u r r e n c e a n d s t r a t i g r a p h i c a l r a n ge: Kovacsia
baloghi is known only from the Anisian (Illyrian; reef
stage 1) part of the reef limestone of the Aggtelek reefs,
northeast Hungary (Kovács 1978a; and this paper).
Order uncertain
Family: Olangocoeliidae Bechstädt & Brandner, 1970
Genus: Olangocoelia Bechstädt & Brandner, 1970
T y p e s p e c i e s: Olangocoelia otti Bechstädt & Brandner,
1970.
Olangocoelia otti Bechstädt & Brandner, 1970
(Fig. 4K; Fig. 9J)
1970 Olangocoelia otti n. sp. – Bechstädt & Brandner; p. 68, pl. 14,
fig. 1—2, pl. 15, fig. 1—4
1990 Olangocoelia otti Bechstädt & Brandner – Senowbari-Daryan;
pl. 45, fig. 9—10; pl. 46, fig. 1—2 (cum syn.)
1990 Olangocoelia otti Bechstädt & Brandner – Scheuber; p. 72,
pl. 11, fig. 6
1993 Olangocoelia otti Bechstädt & Brandner – Senowbari-Daryan,
Zühlke, Bechstädt & Flügel; p. 218, pl. 40, fig. 1—5, pl. 41,
fig. 1—4; pl. 42, Fig. 1—2, pl. 45, fig. 7, text- fig. 10
1997 Olangocoelia otti Bechstädt & Brandner – Schafhauser; p. 109,
pl. 12, fig. 7
2004 Olangocoelia otti Bechstädt & Brandner – Finks & Rigby;
p. 681, fig. 5a—b
2005 Olangocoelia otti Bechstädt & Brandner – Emmerich, Zamparelli,
Bechstädt & Zühlke; Fig. 11/1—2
M a t e r i a l: Several “colonies”.
D e s c r i p t i o n: The “colonies” of this sponge are composed
of numerous circular to oval chambers arranged in irregularly
410
SENOWBARI-DARYAN, KOVÁCS and VELLEDITS
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA, 2011, 62, 5, 397—412
curved lines. The chambers are arranged on one layered lines,
but occasionally also on two-layered lines. The chains of
Olangocoelia surround the small irregular cavities, which are
usually filled with cement or, rarely, with micritic sediment.
The interior of the chambers was hollow and later they became
filled with calcite cement. Ostia are rare within the chamber
walls. The connection of individual chambers is established ei-
ther directly or by a small tube. Vesiculae do not occur, either in
our material or in the material described by Senowbari-Daryan
et al. (1993), but according to Bechstädt & Brandner (1970:
p. 69) they occur within some chambers. Senowbari-Daryan et
al. (1993: p. 220) observed “dichotomously branched tubes
which stretch out from the chambers into the sediments, but
never into the cavity surrounded by the chambers” (see Senow-
bari-Daryan 1993: text-fig. 10). The detailed description of such
tubes and also Olangocoelia otti is undertaken by Bechstädt &
Brandner (1970) and by Senowbari-Daryan et al. (1993).
The systematic position of Olangocoelia as a “sphinctozo-
an” sponge or foraminifer is uncertain (Senowbari-Daryan et
al. 1993).
O c c u r r e n c e: Olangocoelia otti is known from several
Anisian—Ladinian localities in the Dolomites, northern Italy
(Bechstädt & Brandner 1970; Zorn 1971, 1972; Blendinger
1983; Fois & Gaetani 1984; Senowbari-Daryan et al. 1993;
Emmerich et al. 2005), Karawanken (Schafhauser 1997), Hun-
gary (this paper) and Carnic Alps (Pfeiffer 1988). It is known
both from reef stage 1 and reef stage 2 of the Aggtelek reef.
Discussion
The “sphinctozoan” fauna of the Illyrian—Lower Ladinian
Aggtelek reef was previously described by Scholz (1972)
and the present paper. The “sphinctozoans” of the Ladinian—
Carnian Wetterstein Limestone of Alsóhegy Karstplateau, in
northern Hungary (20 km north-east of the Aggtelek reef)
were described by Balogh & Kovács (1976) and Kovács
(1978a,b). Several taxa, including the genera Vesicocaulis,
Uvanella, Cryptocoelia, Stylothalamia, Paravesicocaulis
and “Verticillites” described by these authors were not found
in the investigated material of the Aggtelek reef limestones.
These genera are also not known from other Anisian locali-
ties in the world and are “typically” sponges of Wetterstein
reef limestone (Ladinian—Carnian).
Some general notes are thought necessary concerning
some sponge taxa of the Ladinian—Carnian Wetterstein lime-
stones described by the above mentioned authors from the
Alsóhegy Karstplateau: the Species, described as ?Cysto-
thalamia sp. and illustrated in pl. 3, fig. 1 by Balogh &
Kovács (1976) is most probably “Cystothalamia” bavarica,
attributed now to the genus Alpinothalamia Senowbari-
Daryan (1990). “Verticillites” triassicus Kovács (1978a) is
attributed as type species to the genus Senowbaridaryana by
Engeser & Neumann (1986). All these genera are typically
“sphinctozoans” of the Carnian, but may also occur rarely in
the Ladinian time interval.
Acknowledgments: The investigations were carried out by
Felicitasz Velledits during a short stay in Germany supported
by the “Deutscher Akademischer Austauschdienst”. The au-
thors thank the journal referees for their valuable comments.
References
B
alogh K. & Kovács S. 1976: Sphinctozoa from the reef facies of
the Wetterstein Limestone of Alsóhegy—Mount (South Gemer-
icum, West Carpathian, Northern Hungary). Acta Mineral.
Petrogr. Szeged 22, 2, 297—310.
Bechstädt T. & Brandner R. 1970: Das Anis zwischen St. Vigil und
dem Höhlensteintal (Pragser und Olanger Dolomiten,
Südtirol). Festband Geol. Inst., 300-Jahr-Feier, Univ. Inns-
bruck, 9—103.
Blendinger W. 1983: Anisian sedimentation and tectonics of the M.
Pore—M. Cernera area (Dolomites). Riv. Ital. Paleont. Stratigr.
69, 2, 175—208.
Boiko E.V. 1986: ‘Sphinctozoans’ from the Late Triassic Reefs
from the SE Pamir. In: Sokolov B.S. (Ed.): Phanerozoic reefs
and corals of the USSR. Acad. Nauk USSR, Moscow, 3—11 (in
Russian).
Boiko E.V., Belyaeva G.V. & Zhuravleva I.T. 1991: Sfinktozoa
fanerozoya territorii SSSR. [Phanerozoic ‘sphinctozoans’ from
the Territory of USSR.] Nauka, Moscow, 1—23 (in Russian).
Ciarapica G., Cirilli S., Martini R., Rettori R., Zaninetti L. & Salvini-
Bonnard G. 1990: Carbonate buildups and associated facies in
the Monte Facito Formation (Southern Appenines). Boll. Soc.
Geol. Ital. 109/1, 151—164.
Cossmann M. 1909: Rectification de nomenclature. Rev. Critique
Paléontologie 13, 67.
de Laubenfels M.W. 1955: Porifera. In: Moore R.C. (Ed.): Treatise
on invertebrate palaeontology. Part E: Archaeocyatha and
Porifera. Geol. Soc. Amer., Univ. Kansas Press, New York &
Lawrence, 21—112.
Dullo W.-Ch., Flügel E., Lein R., Riedel P. & Senowbari-Daryan B.
1987: Algen, Kalkschwämme und Mikroproblematika aus un-
terkarnischen Riffkalken des Bosruck—Gipfels (Nördliche
Kalkalpen, Österreich). Jb. Geol. B.—A 129 (3 + 4), 525—543.
Emmerich A., Zamparelli V., Bechstädt T. & Zühlke R. 2005: The
reefal margin and slope of a Middle Triassic carbonate plat-
form: The Latemar (Dolomites, Italy). Facies 50, 573—614.
Engeser T. 1986: Nomenklatorische Notiz zur Gattung Dictyocoelia
Ott 1967 (“Sphinctozoa”, Porifera). Neu. Jb. Geol. Paläont.,
Mh. 1986 (10), 587—590.
Engeser T. & Neumann H.H. 1986: Ein neuer verticillidider
“Sphinctozoe” (Demospongiae, Porifera) aus dem Campan der
Krappfeld—Gosau (Kärnten, Österreich). Mitt. Geol. Paläont.
Inst. Univ. Hamburg 61, 149—159.
Enos P., Jiayoung W. & Yangii Y. 1997: Facies distribution and re-
treat of Middle Triassic platform margin, Guizhou Province,
south China. Sedimentology 44, 563—584.
Finks R.M. & Rigby J.K. 2004: Hypercalcified sponges. In: Kaesler
R.L. (Ed.): Treatise on invertebrate paleontology. Part E:
Porifera (revised). Vol. 3. Geol. Soc. Amer., Univ. Kansas,
Boulder, Kansas, 585—764.
Flügel E. 1982: Evolution of Triassic reefs: Current concepts and
problems. Entwicklung der trias—Riffe: Kenntnisstand und
Probleme. Facies, 297—327.
Flügel E. 2002: Triassic Reef patterns. In: Kiessling W., Flügel E.
& Golonka J. (Eds.): Phanerozoic reef patterns. SEPM Spec.
Publ., Tulsa 72, 391—463.
Flügel E. & Senowbari-Daryan B. 2001: Triassic reefs of the
Tethys. In: Stanley G.D. (Ed.): The history and sedimentology
of ancient reef systems. Pleneum Press, New York, 217—249.
Flügel E., Velledits F., Senowbari-Daryan B. & Riedel P. 1991/1992:
411
SPONGES FROM THE MIDDLE TRIASSIC REEF LIMESTONE (AGGTELEK, HUNGARY)
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA, 2011, 62, 5, 397—412
Rifforganismen aus “Wettersteinkalken” (Karn?) des Bükk-Ge-
birges, Ungarn. Geol. Paläont. Mitt. Innsbruck 18, 35—62.
Fois E. & Gaetani M. 1984: The recovery of reef-building commu-
nities and the role of cnidarians in carbonate sequences of the
Middle Triassic (Anisian) in the Italian Dolomites. Palaeonto-
graph. Amer. 54, 191—200.
Gaetani M., Fois E., Jadoul F. & Nicora A. 1981: Nature and evolu-
tion of Middle Triassic build-ups in the Dolomites (Italy).
Mar. Geol. 44, 25—57.
Gawlick H.J., Frisch W., Vecsei A., Steiger T. & Böhm F. 1999:
The change from the rifting to thrusting in the Northern Cal-
careous Alps as recorded in Jurassic sediments. Geol. Rundsch.
87, 644—657.
Grant R.E. 1836: Animal kingdom. In: Todd R.B. (Ed.): The Cyclo-
paedia of anatomy and physiology. Vol. 1. Sherwood, Gilbert
& Piper, London, 107—118.
Haas J., Hámor G., Jámbor Á., Kovács S., Nagymarosy A. &
Szederkényi T. 2001: Geology of Hungary. Eötvös University
Press, Budapest, 1—317.
Jablonský E. 1974: Segmentierte Kalkschwämme (Sphinctozoa) aus
Wettersteinkalken einiger Gebirge der Westkarpaten. Acta
Geol. Geogr. Univ. Comenianae, Geol. 26, 189—202.
Kovács S. 1978a: New “sphinctozoan” sponges from the North
Hungarian Triassic. Neu. Jb. Geol. Paläont., Mh. 1978 (11),
685—697.
Kovács S. 1978b: Newer Calcareous sponges from the Wetterstein
Reef limestone of Alsóhegy Karstplateau (Silica Nappe, West-
ern Carpathians, North Hungary). Acta Mineral. Petrogr.
Szeged 23, 2, 299—317.
Kovács S. 1989: Geology of North Hungary: Paleozoic and Meso-
zoic terranes. In: Kecskeméti T. (Ed.): 21st Europ. Micropale-
ont. Colloq. Guidebook. Hung. Geol. Soc., Budapest, 15—36.
Kovács S. 1997: Middle Triassic rifting and facies differentiation in
Northeast Hungary. In: Sinha A.K. (Ed.): Geodynamic do-
mains in the Alpine-Himalayan Tethys. IBH Publishing Co.
Pvt. Ltd., Oxford, New Delhi, Calcutta, 375—397.
Kozur H. 1991: The evolution of the Meliata-Hallstatt ocean and its
significance for the early evolution of the Eastern Alps and
Western Carpathians. Palaeogeogr. Palaeoclimatol. Palaeoecol.
87, 109—135.
Kozur H. & Mock R. 1973: Die Bedeutung der Trias—Conodonten für
die Stratigraphie und Tektonik der Trias in den Westkarpaten.
Geol. Paläont. Mitt. Innsbruck 3, 2, 1—14.
Laube G.C. 1864: Bemerkungen über die Münster’schen Arten von
St. Cassian in der Münchner Paläontologischen Sammlung. Jb.
Geol. Reichsanst. 14, 402—412.
Lehrmann D.J., Wei J. & Enos P. 1998: Controls on facies architec-
ture of a large Triassic carbonate platform: the great bank of
Guizhou, Nanpanjiang Basin, South China. J. Sed. Res. B68,
311—326.
Lein R. 1987: Evolution of the Northern Calcareous Alps during
Triassic times. In: Flügel H.W. & Faupl P. (Eds.): Geodynamics
of the Eastern Alps. Deutick, Wien, 85—102.
Mastandrea A. & Rettori R. 1989:
On the occurrence of a sphincto-
zoan (Porifera) assemblage in the carbonatic bodies of Monte
Facito Formation (Southern Apennines). Atti Soc. Nat. Modena
120 (1989), 15—26 (in Italian).
Mello J., Elečko M., Pristaš J., Reichwalder P., Snopko L., Vass D.,
Vozárová A., Gaál ., Hanzel V., Hók J., Kováč P., Slavkay
M. & Steiner A. 1997: Explanations to the geological map of
the Slovak Karst, 1 : 50,000. Vyd. D. Štúra, Bratislava, 1—255
(in Slovak).
Münster G.F. 1841: Beiträge zur Geognosie und Petrefakten-Kunde
des südöstlichen Tirols, vorzüglich der Schichten von St. Cas-
sian. Bayreuth, 1—152.
Nittel P. 2006: Beiträge zur Stratigraphie und Mikropaläontologie
der Mitteltrias der Innsbrucker Nordkette (Nördlicher Kalkal-
pen, Austria). Geo. Alp. 3, 93—145.
Ott E. 1967: Segmentierte Kalkschwämme (Sphinctozoa) aus der al-
pinen Mitteltrias und ihre Bedeutung als Riffbildner im Wetter-
steinkalk. Bayer. Akad. Wiss. Math.-Naturwiss. Kl., Abh., N.F.
131, 1—96.
Ott E. 1972: Mitteltriadische Riffe der Nördlichen Kalkalpen und
altergleiche Bildungen aus Karaburun und Chios (Ägäis). Mitt.
Gesell. Geol. Bergbaustud. 21, 251—276.
Ott E., Pisa G. & Farabegoli E. 1980: Celyphia zoldana sp. n., a reef
building “sphinctozoan” sponge in Anisian limestones of the
Southern Dolomites. Riv. Ital. Paleont. 85, 829—842.
Payne J.L., Lehrmann D.J., Christensen S., Wei J. & Knoll A. 2006:
Environmental and biological controls on the initiation and
growth of a Middle Triassic (Anisian) reef complex on the Great
Bank of Guizhou, Guizhou province, China. Palaios 21, 325—343.
Pfeiffer J. 1988: Paleontology and microfacies of platform margin in
the Carnic Alps (Austria, Middle Triassic). Facies 19, 33—60.
Pomel A. 1872: Paléontologie ou description des animaux fossils de
la province d’Oran. 5. Spongiaaires, Oran, 1—256.
Rüffer T. & Zamparelli V. 1997: Facies and biota of Anisian to Car-
nian Carbonate platforms in the Northern Calcareous Alps
(Tyrol and Bavaria). Facies 37, 115—136.
Schafhauser M. 1997: Stratigraphie und Fazies in der Mitteltrias der
Südkarawanken (Kärnten/Österreich) im Vergleich zur litho-
stratigraphischen Entwicklung des angrenzenden Südalpins. Un-
publ. Ph.D. Thesis, Technische Universität Berlin, Berlin, 1—161.
Scheuber M. 1990: Der Spitzkalk von Recoaro (Vicentinische Al-
pen, Norditalien): Sedimentologie, Paläontologie und Paläo-
geographie eines mitteltriassischen Sedimentationsraumes.
Facies 23, 57—96.
Schmid S.M., Bernoulli D., Fügenschuh B., Matenco L., Schefer S.,
Schuster R., Tischler M. & Ustaszewski K. 2008: The Alpine-
Carpathian-Dinaridic orogenic system: correlation and evolu-
tion of tectonic units. Swiss. J. Geosci. 101, 139—183.
Doi: 10.1007/s00015—008—1247—3
Scholz G. 1972: An Anisian Wetterstein Limestone Reef in North
Hungary. Acta Mineral. Petrogr. Szeged 20, 2, 337—362.
Senowbari-Daryan B. 1990: Die systematische Stellung der thala-
miden Schwämme und ihre Bedeutung in der Erdgeschichte.
Münchner Geowiss. Abh., Reihe A (Geologie und Palaeont.)
21, 1—325.
Senowbari-Daryan B. & Abate B. 1986: Zur Paläontologie, Fazies
und Stratigraphie der Karbonate innerhalb der “Formazione
Mufara” (Obertrias, Sizilien). Natur. Sicil., Ser. IV 10 (1—4),
50—104.
Senowbari-Daryan B. & Garcia-Bellido D.C. 2002: “Sphinctozoa”:
Chambered sponges (Polyphyletic). In: Hooper J.N.A. & Van
Soest R.W.M. (Eds.): System Porifera. A guide to the classifi-
cation of sponges. Kluwer Acad./Plenum Publ., New York,
1511—1533.
Senowbari-Daryan B. & Riedel P. 1987: Revision der triadischen
Arten von Solenolmia Pomel 1872 (Dictyocoelia Ott 1967)
(“Sphinctozoa”, Porifera) aus den alpin-mediterranen Raum.
Mitt. Bayer. Staatssamml. Paläont. Hist. Geol. 27, 5—20.
Senowbari-Daryan B. & Schäfer P. 1983: Zur Sphinctozoen-Fauna
der obertriadischen Riffkalke (“Pantokratorkalke”) von Hydra,
Griechenland. Geologica et Palaeont. 17, 179—205.
Senowbari-Daryan B. & Velledits F. 2007a: Axopora aggtelekensis
Scholz 1972, originally described as Hydrozoa, is attributed to
the new genus Anisophytes n. gen. (Cyanophyta). Riv. Paleont.
Stratigr. Ital. 113, 3, 357—368.
Senowbari-Daryan B. & Velledits F. 2007b: Aggtecella, a new genus
of Corallinales (Rhodophyta) from the Anisian of the Aggtelek-
Rudabánya Mountains, NE Hungary. Facies 53, 401—407.
Senowbari-Daryan B., Zühlke R., Bechstädt Th. & Flügel E. 1993:
412
SENOWBARI-DARYAN, KOVÁCS and VELLEDITS
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA, 2011, 62, 5, 397—412
Anisian (Middle Triassic) Buildups of the Northern Dolomites
(Italy): The recovery of reef communities after the Permian/
Triassic Crisis. Facies 28, 181—256.
Senowbari-Daryan B., Rashidi K. & Hamedani A. 2006: Two Per-
mian “sphinctozoan” sponges from the Shotori Mountains
(eastern Iran). Geol. Carpathica 57, 6, 427—432.
Sollas W.J. 1875: Sponges. In: Encyclopedia Britannica. 9
th
edition.
Encyclopedia Britannica, Inc., London, 1—451.
Steinmann G. 1882: Pharetronen-Studien. Neu. Jb. Miner., Geol.
Paläont. 2, 139—191.
Tollmann A. 1987: Neue Wege in der Ostalpengeologie und die
Beziehungen zum Ostmediterran. (New directions on the Geol-
ogy of the Eastern Alps and their connection to the Eastern
Mediterranean). Mitt. Österr. Geol. Gesell. 80, 47—113.
Velledits F., Péró Cs., Blau J., Senowbari-Daryan B., †Kovács S.,
Piros O., Pocsai T., Szügyi-Simon H., Dumitrică P. & Pálfy J.
2011: The oldest Triassic platform margin reef from the Al-
pine-Carpathian Region (Aggtelek, NE Hungary): platform
evolution, reefal biota and biostratigraphic framework. Riv.
Paleont. Stratigr. Ital. 117, 221—268.
Wu Y.-Sh. 1991: Organisms and communities of Permian Reef of
Xiangbo, China. Int. Acad. Publ., Beijing, 1—192.
Zorn H. 1971: Paläontologische, stratigraphische und sedimentolo-
gische Untersuchungen des Salvatoredolomits (Mitteltrias) der
Tessiner Kalkalpen. Schweiz. Paläont. Abh. 91, 1—90.
Zorn H. 1972: Mikrofazielle Analyse eines mitteltriadischen
Riffkomplexes in den Tessiner Kalkalpen. Mitt. Gesell. Geol.
Bergbaustud. 21, 123—142.