GEOLOGICA CARPATHICA, 49, 6, BRATISLAVA, DECEMBER 1998
RHAETIAN SCLERACTINIAN CORALS IN THE
and JOZEF MICHALÍK
Paleobiological Institute, Polish Academy of Sciences, Twarda 51/56 , 00-818 Warszawa, Poland; firstname.lastname@example.org
Geological Institute, Slovak Academy of Sciences, Dúbravská cesta 9, 842 26 Bratislava, Slovak Republic; email@example.com
(Manuscript received June 29, 1998; accepted in revised form November 3, 1998)
Abstract: The paper presents the distribution and taxonomy of the Rhaetian coral fauna in the Tatric, Fatric, Hronic
and Silicic paleogeographical zones of the Western Carpathians. The taxonomic spectrum of the corals from the Fatric
zone is the most typical of the Carpathians, and resembles that of the Alps in the abundance of reimaniphylliid corals
and frequent phaceloid growth forms, but it differs from the Alpine spectrum in its lower generic diversity. The
assemblage from the northernmost zone, formed by the Tatric Superunit, is closest to the Early Jurassic fauna from the
British Isles. Two new species are described: Zardinophyllum carpathicum sp.n. and Stylophyllopsis bobrovensis sp.n.
Key words: Western Carpathians, Rhaetian, coral build-ups, Scleractinia.
The aim of the present paper is to show the composition of
the coral fauna of the latest Triassic in the Western
Carpathians, and to revise some earlier taxonomic
determinations (Roniewicz 1974). Up to now, Late Triassic
scleractinians from this region have been described from the
High Tatra-, Western Tatra-, Low Tatra-, Malé Karpaty-,
Stratená- and Stráov Mts. (Goetel 1917; Kolosváry 1957,
1958a,b, 1963, 1966a,b, 1967; Roniewicz 1974; Roniewicz
& Michalík 1991a,b).
In the Carpathians, Triassic Scleractinia are present in
Anisian- and younger deposits. The earliest of these corals
were described from the Anisian limestones of Aggtelek,
Hungary (Scholz 1972), and constitute a sort of biostrome
composed mainly of phaceloid skeletons. Some Mid-Trias-
sic corals were described by Kolosváry from the Bükk- and
Mecsek Mts. (1958a,b, 1966a). However, in the light of the
new stratigraphic data (Riedel et al. 1988), the age of the
corals from the Bükk Mts. that were supposed to be Ladin-
ian (Kolosváry 1958a,b) must be changed to the Late Trias-
sic. The reported Mid-Triassic age of corals from the Mec-
sek Mts. (Kolosváry 1966a) has not been confirmed to date.
In addition, the age and taxonomy of the Slovak corals con-
sidered by Kolosváry to be of Mid-Triassic age (1958c,d,
1963, 1966b, 1967) still need revision. Recently, some Scler-
actinia from Slovakia have been collected from the Ladin-
ian/Carnian Wetterstein type limestones from the Malé Kar-
paty Mts. (Roniewicz & Buèek, in preparation). The most
interesting aspect of this discovery is that the corals repre-
sent solitary and phaceloid forms taxonomically close to the
Carnian fauna from the St. Cassian beds of the Dolomites. In
the Carpathians, the corals played a minor role only in orga-
nogenic constructions dominated by calcareous sponges and
hydrozoans (Michalík et al. 1993).
In other parts of the Carpathians, abundant Carnian coral
faunas were described by Papp (1911) from the Bakony Mts.
(Hungary), and from the Romanian Carpathians (Braºov) by
In Upper Triassic limestone facies of the Western Car-
pathians, corals show the whole morphological spectrum from
solitary and phaceloid to various massive forms (Kolosváry
1957, 1958c,d, 1963, 1966b, 1967; Roniewicz 1974). In this
region, Rhaetian Scleractinia constitute a widely distributed
and significant component of the fauna (Michalík 1973, 1974,
1977, 1978, 1979, 1980, 1982; Ga¿dzicki 1974; Roniewicz
1974): these corals will be considered in the present paper.
Corals of this age are to be found in the following tectonic su-
perunits: Tatric, Fatric, Hronic and Silicic nappe systems,
which crop out in diverse mountains, such as the High Tatra
Mts. (annotated simply as the Tatra Mts. in the Polish litera-
ture), Western Tatra-, Low Tatra-, Malá Fatra-, Ve¾ká Fatra-,
Povaský Inovec-, Tribeè-, Stráov-, Malé Karpaty- and
The bulk of the material considered here was collected by
one of us (J. Michalík, see Michalík 1979, 1982 for sample
locations) and is deposited in the Slovak National Museum,
Bratislava (abbreviated as SNM; collection of about 170
specimens). Some specimens belong to the collection of
S. Buèek (Geological Survey of the Slovak Republic, Brat-
islava). A large collection from the Tatra Mts. (above 100
specimens collected chiefly by A. Ga¿dzicki, A. Radwañski
& E. Roniewicz in the 1960s) is housed at the Institute of
Paleobiology, Polish Academy of Sciences, Warsaw
Endotheca vesiculous or tabuloid dissepiments infilling
the central and interseptal parts of the corallite.
392 RONIEWICZ and MICHALÍK
Growth form shape of coral skeleton resulting from the
mode of life. The solitary mode produces various corallum
shapes: cylindrical, conical, etc. The phaceloid form, con-
sidered to be a result of a pseudocolonial mode of life
(Coates & Jackson 1987), shows skeletons built of subcy-
lindrical branches (=corallites) each branch ending with a
calice lacking connections with others with the exception
of the site of budding. The colonial mode produces colonies
of various structural types based on the different relation-
ships between neighbouring corallites: (1) cerioid-corallites
isolated by walls; (2) thamnasterioid-corallites lacking
walls and interconnected by septa; (3) meandroid-corallites
arranged into series; (4) plocoid-corallites divided from
each other by coenosteum, which is a common colonial
skeletal tissue. The shapes of colonies may be massive,
lamellate or branching.
Ornamentation microarchitectural structures on the sep-
tal flanks: granulations, pennules (Gill 1967: structures in the
form of balconies) or menianes (Gill 1967: structures in the
form of long or short ledges paralleling the distal septal margin).
Septal spines thick trabecula-like elements, differing
from trabeculae in lacking radial structure, and in being
formed from bundles of fibres; constitute septa in the sub-
order Stylophyllina (see also Roniewicz 1989).
Trabecula a structural element of septa having fibres ra-
dially arranged in relation to the central axis (centre of calci-
fication). Trabeculae may be differentiated into small trabec-
ulae (from ca. 20 to 50 micrometres in diameter) and thick
trabeculae (above 50 micrometres, usually 100200 micro-
metres; Roniewicz 1989; Morycowa & Roniewicz 1995).
Preservation of the material
The material considered consists of ca. 270 fragments of
corals from more than 100 localities distributed throughout
the above-mentioned mountain chains. The original aragonite
skeletons have been completely calcified, and, in the majority
of specimens, the internal structure has been destroyed. The
phaceloid corals especially show extensively changed
skeletons, their corallites being filled with blocky calcite or
deformed by pressure. The best preserved specimens were
selected for preparing thin sections, and the paper presents the
results based on investigation of ca. 270 thin sections. Al-
though poorly preserved, the skeletons display many structural
details enabling reconstruction of their morphology; taxonom-
ically significant features may be observed in fragments of
We consider the entire collection as a faunistic sample that
is representative of the region constituting the west-central
part of the Carpathians.
Relationships of the Rhaetian coral faunas of the
Western Carpathian paleogeographical zones
Tatric paleogeographical zone. In the limestones that form
part of the Tatric Superunit, corals have been found at a few
localities, in dark limestones cropping out in the Western Tatra
Mts. (Bobrovec Valley and the SW part of the Chocho³ow Val-
ley: Radwañski 1968; Roniewicz 1974: Fig. 1, Table 1).
Colonies of lamellate and phaceloid form constitute a small
patch at a single locality (Za Kýèerou), while isolated
branches of Stylophyllopsis media and lamellate colonies of
Rhaetiastraea tatrica may be found at a number of localities in
the Chocho³ow Valley (e.g. Skorusi leb stream section,
In contrast to the assemblages from the Fatric and Hronic
zones, the corals from the Tatra Superunit show, up to now,
only one species common to any of the other Carpathian
faunas, namely Phacelostylophyllum medium (now
Stylophyllopsis media) in the Fatra Formation (Michalík 1982,
Pl. 18: Fig. 6). The fauna contains Pamirophyllum, a genus
common to the Norian/Rhaetian fauna of the Pamirs in Cen-
tral Asia (Melnikova & Roniewicz 1990), and the ubiquitous
genus Stylophyllopsis, which is known from both Eurasia and
America from the uppermost Triassiclowermost Jurassic.
Another element, of the fauna is Heterastrea, a genus hitherto
not known earlier than the Lower Jurassic (cf. Taxonomy, p.
394 and Table 1). The genus Rhaetiastraea, described for the
first time from the Tatric, is known in the West Tethyan Lias-
sic deposits (see p. 398).
This small fauna from the High-Tatra unit significantly dif-
fers at suprageneric level from the other Tethyan Triassic fau-
nas. Although three genera represent the order Stylophyllina
(Stylophyllopsis, Pamirophyllum and Heterastraea), charac-
terized by non-trabecular septa built of septal spines, and typi-
cal of the TriassicLiassic interval, only one genus, Rhaetias-
Fig. 1. Sketch drawings showing differences in arrangement of endothecal elements in individual species of Retiophyllia: 1. R. gracilis
Roniewicz, 2. R. gosaviensis Roniewicz,3. R. paraclathrata Roniewicz, 4. R. clathrata (Emmrich).
RHAETIAN SCLERACTINIAN CORALS IN THE WESTERN CARPATHIANS 393
Table 1: Rhaetian Scleractinia in the Western Carpathians. Distribution of considered taxa in the Tatric (T), Fatric (F), Hronic (H) and Silicic
(S) paleogeographical zones, and in the Alps; brackets indicate forms of morphology similar to the Alpine taxon (cf.). Measurements: d
diameter of the corallite measured from the middle of the wall; e.estimated; snumber of septa. The column entitled Septal apparatus
organization presents some examples of a variable arrangement of septa in Retiophyllia: S1, S2 septa of succeeding size orders (which
may or may not correspond to order of appearance).
nr of septa
R e m a r k s
M a t e r i a l
1. Zardinophyllum carpathicum
Tatra Mts: SNM 9,11; Low Tatra / Ve¾ká
Fatra Mts: SNM 10, 12, 13
2. Pinacophyllum lejowae
s ca. 35
Tatra Mts: SNM 1, 2; ZPAL HVI / 113
116, 120122; Stráovské Vrchy: SNM 3,
8; Malé Karpaty Mts: SNM 5
3. Stylophyllum robustum
Tatra Mts: ZPAL HVI / 41, 79, 86, 108,
129, 130, 145. Low Tatra / Ve¾ká Fatra
Mts: SNM 151
4. Stylophyllopsis media
s 45 (56)
removed from Stylophyllum to
Stylophyllopsis due to solid
Tatra Mts: ZPAL HVI / 17, 14, 73
5. Stylophyllopsis bobrovensis
s ca. 80
Tatra Mts: ZPAL HVI / 141144
6. Stylophyllopsis cf. polyactis
fragmentary corallite, septa
relatively solid, septal spines of
max. 0.6 mm
Malá Fatra Mts: SNM 16
7. Heterastrea gracilis
resembling H. stricklandi
Dunc. and H. tomesi Dunc.
from the Lias of England
(comp. Beauvais 1976)
Tatra Mts: ZPAL HVI / 137
8. Pamirophyllum tatricum Ron. In
Melnikova & Roniewicz1990
s 50ca. 80
Tatra Mts: ZPAL HVI / 138143
Roniewicz & Michalik 1991
Tatra Mts: SNM Z 20075
10. Retiophyllia fenestrata
s ca. 45
Tatra Mts: ZPAL HVI/46
11. Retiophyllia frechi
Malá Fatra Mts : SNM Z 54
12. Retiophyllia gosaviensis
Alpine form has up to 120
septa due to high number of
Stráovské Vrchy: SNM 94, 95. Ve¾ká
Fatra Mts: thin section 258/6. Tatra Mts:
ZPAL HVI / 15
13. Retiophyllia gracilis
Tatra Mts: ZPAL HVI/49
14. Retiophyllia clathrata
Tatra Mts: ZPAL HVI / 21, 28, 67, 72;
Ve¾ká Fatra Mts: SNM 113, 114, 117;
Malé Karpaty Mts: SNM, Buèek´s Coll.
15. Retiophyllia cf. oppeli
s < 40
Alpine form has more than 50
septa arranged into 78 systems
Tatra Mts: SNM 41,46,71; Stráovské
Vrchy: SNM 39; Malé Karpaty: SNM 44
16. Retiophyllia paraclathrata
Zigzag septa: Tatra Mts: ZPAL HVI/ 35
38, 66, 119, 123, 132;. Stráovské Vrchy
Mts: SNM 83; Malá Fatra Mts: SNM 93.
straight septa: Ve¾ká Fatra Mts: SNM 38.
Malé Karpaty Mts: Buèek Coll. Nos.13
17. Retiophyllia sp. 1
s > 80
Malá Fatra Mts: SNM 119
18. Distichophyllia sp.
d e. 20
Stratenská Hornatina: SNM 120
19. Chondrocoeania schafhaeutli
Tatra Mts: ZPAL HVI/ 45, 78, 8083, 109,
125; SNM 17, 19; Malá Fatra Mts: SNM
20. Pamiroseris rectilamellosa
d ca. 5
Tatra Mts: ZPAL HVI/ 47, 48, 50, 51, 61,
69, 7476, 126; SNM 31, 36; Malá Fatra
Mts: SNM 32
21. Rhaetiastraea tatrica
s < 45
Liassic Moroccan form with
4565 septa (compare Beauvais
Tatra Mts: ZPAL HVI / 8, 10, 12, 13,
22. Astraeomorpha confusa
s < 16
Tatra Mts: ZPAL HVI / 25, 26, 9498, 118,
133. SNM 46
394 RONIEWICZ and MICHALÍK
traea, belongs to the trabecular corals, the minitrabecular re-
imaniphylliids, a group typical of the Upper Triassic (Ro-
niewicz 1989), being absent.
The composition of the fauna from the Tatric zone indi-
cates that the coralliferous series may be HettangianSine-
murian in age. In its being dominated by stylophylline corals,
it resembles the lowermost Jurassic coral fauna from Great
Britain (Beauvais 1976).
Fatric paleogeographical zone. The most common Rha-
etian corals originate from the Fatric paleogeographical zone
(formed by a homonymous superunit) from dark-coloured
limestones of the Fatra Formation.
The corals occur mainly in two coralliferous horizons of
the Fatra Formation, named the lower and upper biostromatic
members, respectively (Michalík et al. 1979). The limestones
of this formation constitute part of the Krína tectonic unit.
The scleractinians from the widely distributed Krína Unit
belong to the most typical of the West-Carpathian coral fau-
nas, and are also close to the fauna from other regions of the
Tethyan belt. These are corals that were relatively often de-
scribed in geological and paleontological papers (Goetel 1917;
Ga¿dzicki 1974; Roniewicz 1974; Roniewicz & Michalík
1991a,b). The coralliferous beds in the Fatra Formation are
widely distributed (Michalík 1973, 1974, 1977).
The most typical members of this fauna belong to the
specifically well diversified genus Retiophyllia. The
skeletons of these branching, phaceloid corals either form
biostromes or, together with lamellate corals (Astraeomopha,
Pamiroseris), contribute to the formation of small patches.
Determination of these corals at the specific level is usually
impossible due to recrystallization of the skeletons. Other
common species are Pamiroseris rectilamellosa and
Astraeomorpha confusa. A very interesting faunistic element
found in the lagoonal part of this zone, is the solitary,
dwarfish coral Zardinophyllum carpathicum sp.n., the only
known Rhaetian representative of the genus (Roniewicz &
Michalík 1991a). Another coral genus known up to now only
from this zone is Meandrostylophyllum, which belongs to the
here poorly represented stylophylline corals.
Hronic paleogeographical zone. Corals are known in
light coloured limestone facies of the Norovica Formation,
constituting the Mojtín Limestone Member, and in the Hybe
Beds (Ga¿dzicki & Michalík 1980). The limestones of this
zone form part of the Choè tectonic unit. The proportion of
corals in this zone is smaller than in the Fatric zone. Coral
communities cited in earlier literature (e.g. Zázvorka &
Prantl 1936), yielded a number of coral species (Table 1).
In general, the taxonomic composition of the coral fauna
from the Hronic zone is similar to that from the Fatric zone.
The poor state of preservation of the corals renders
impossible any quantitative comparison between the faunas
of the two zones.
Silicic paleogeographical zone. This zone yielded a small
number of coral finds in the Silicic tectonic unit (sensu lato).
The corals in the collection, determined only to generic level
due to their uncomplete preservation, represent reimaniphyl-
liid phaceloid Retiophyllia sp. and solitary Distichophyllia
sp. Due to the scarcity of material any detailed comparison
with the faunas of the two preceding zones is impossible.
General characteristics of the Rhaetian
Scleractinia from Western Carpathians
In general, the taxonomic spectrum of the Rhaetian corals
from the Fatric and Hronic zones corresponds to that of the
Alpine Rhaetian ones, although the numbers and proportions
of species are different in the Carpathians and in the Alps.
The corals from the Western Tatra Mts. either constitute a
rather isolated, endemic Rhaetian fauna, or they represent an
Early Jurassic assemblage.
The most striking feature of the Rhaetian corals in the
Carpathians is their morphology, or, more precisely, the
growth forms of their skeleton (Table 1). These are mainly
branching, phaceloid forms, which are abundant and dominate
the assemblages, 180 specimens being represented in the
collections considered here. The phaceloid corals are well
diversified taxonomically (e.g. Pinacophyllum, Stylophyllum,
and above all, the abundant and specifically diverse genus
Retiophyllia). Then, in descending order of numbers, there are
lamellate colonies of the thamnasterioid (ca. 40 specimens),
cerioid (10 specimens), plocoid (10 specimens) and meandroid
types (1 specimen). Each of five genera (Heterastrea,
Pamirophyllum, Pamiroseris, Rhaetiastraea, Astraeomorpha)
of the above colonial corals is represented by only one species.
Of the colonial corals, the thamnasterioid and plocoid types
are rather common, while the cerioid and meandroid corals are
known only from single localities.
The solitary corals are rare and poorly diversified, in
comparison with the Alpine Rhaetian ones. Hitherto, only
one species has been described as locally abundant (Zardino-
phyllum carpathicum sp.n.), while two others (Disticho-
phyllia sp. and Stylophylliopsis cf. polyactis) are represented
by single specimens.
The most typical element of the Rhaetian fauna in the Car-
pathians consists of corals of the family Reimaniphylliidae
(genus Retiophyllia, ca. 150 specimens). This is the most ubiq-
uitous and opportunistic group, which is known, in both marly
and calcareous facies, from the Alps and from other regions of
the Tethyan belt (Central Asia: Melnikova 1975; Northern
Australia: Sarti et al. 1992). On the other hand, corals belong-
ing to the Stylophyllidae did not find in the Carpathians an en-
vironment as favourable for them as that of the Alpine Zlam-
bach Beds, from which over 10 species have been described
(Roniewicz 1989). The exception to this is the development of
the stylophylline corals in the Tatric zone in the Western Tatra
Mts., but this fauna may belong to a younger sedimentary cy-
cle than those from the remaining paleogeographical zones.
The diversity of the Carpathian coral fauna considered here
comprises 20 species belonging to 13 genera. The above
statistics show that, in comparison with the rich assemblage of
Rhaetian corals of the Alpine region comprising at least 64
species belonging to 30 genera (Roniewicz 1989), the
Carpathian corals are only moderately diversified.
The systematics follow Chevalier & Beauvais (1987).
Synonymy is reduced to recent bibliography. All measurable
RHAETIAN SCLERACTINIAN CORALS IN THE WESTERN CARPATHIANS 395
features of the taxa discussed as well as the material here ex-
amined have been shown in the Table 1.
Suborder Pachythecalina Eliáová 1978
Family Zardinophyllidae Montanaro-Gallitelli 1975
Genus Zardinophyllum Montanaro-Gallitelli 1975
Zardinophyllum carpathicum sp.n.
199b Zardinophyllum sp. A: Roniewicz & Michalík, p. 362, Pl. 1:
Figs. 16, and Pl. 2: Figs. 35.
Syntypes: specimen SNM-, illustrated in Roniewicz &
Michalík 1991b, Pl. 1, Figs. 14.
Type locality: Baranie at Svätý Jakub, Low Tatra Mts.
Type horizon: Rhaetian Fatra Formation limestones,
Krína Nappe (Fatric paleogeographical zone).
Diagnosis. Elongated vermicular corallites with diameters
12.5 mm; number of septa irregular, estimated at about 16.
Material. Several dozen specimens, Table 1.
Remarks. The species differs from Zardinophyllum zardi-
ni Montanaro-Gallitelli (type species; Montanaro-Gallitelli
1975) in far smaller corallite dimensions. The species
belongs to the pachythecalines of the smallest diametres
(Pachydendron microthallos Cuif shows corallites 2 mm in
diametres: Cuif 1975).
Suborder Stylophyllina Beauvais 1981
Family Stylophyllidae Frech 1890
The continuous series of transitions from solitary to
phaceloid coralla, which may be observed in Stylophyllopsis
and Stylophyllum (compare Frech 1890), weakens the
usefulness of the phaceloid growth form for the taxonomy of
this group of corals. For the above reason, the taxon established
for phaceloid stylophyllids, Phacelostylophyllum Melnikova
1972, has been rejected here (see below at the synonymies of
Stylophyllum and Stylophyllopsis species).
Genus Pinacophyllum Frech 1890
Pinacophyllum lejowae Roniewicz 1974
1974 ?Pinacophyllum lejowae Roniewicz: p. 101, Pl. 1: Figs. 1, 2;
Genus Stylophyllum Reuss 1854
Stylophyllum robustum (Roniewicz 1974)
1974 Phacelostylophyllum robustum Roniewicz: p. 105, Pl. 3:
1984 Stylophyllum robustum (Roniewicz): Fantini-Sestini & Mot-
ta, p. 358, Pl. 30, Fig. 4.
Genus Stylophyllopsis Frech 1890
Stylophyllopsis media (Roniewicz 1974)
1974 Phacelostylophyllum medium Roniewicz: p. 106, Pl. 3: Fig.
5; Figs. 5, 6.
1982 Phacelostylophyllum medium Roniewicz: Michalík, Pl. 18,
Stylophyllopsis bobrovensis sp.n.
1974 ?Phacelostylophyllum sp.: Roniewicz, p. 107, Pl. 3: Fig. 4.
Syntypes: ZPAL H.VI/141-144; figured as above.
Type locality: Za Kýèerou Site at Bobrovec Valley in the
Western Tatra Mts., Slovakia.
Type horizon: Rhaetian limestones, Tatric cover sequence
(Tatric paleogeographic zone).
Derivation of the name: After the type locality.
Diagnosis. Phaceloid skeletons built of few corallites; coral-
lite diameters ca. 25 mm; septa nearly solid, ca. 80 in number,
differentiated into 5 size orders. Endothecal elements numer-
ous, wide, slightly deepened at the corallite centre.
Material. The type material only. In the samples, the spe-
cies occurs together with Pamirophyllum tatricum.
Remarks. It differs from the similar S. zitteli Frech 1890 by hav-
ing larger corallite diameters and numerous densely arranged septa.
Stylophyllopsis cf. polyactis Frech 1890
Material consists of a fragmentary, solitary corallum
resembling Stylophyllopsis polyactis in the diameter of the
calice and in relatively solid septa S1 built of regular, rather
thin septal spines (ca. 600 micrometres in diameter).
Stylophyllopsis polyactis is a common Rhaetian taxon in the
Alps (Roniewicz 1989).
Genus Heterastrea Tomes 1888
Genus Heterastrea is known from the lowermost Jurassic
beds. For more on the genus see Beauvais (1976).
Heterastrea gracilis (Roniewicz 1974)
1974 Stylophyllum gracile Roniewicz: p. 104, Pl. 2: Fig. 2; Fig. 4.
Genus Pamirophyllum Melnikova & Roniewicz 1990
Pamirophyllum tatricum Roniewicz in
Melnikova & Roniewicz 1990
1974 ?Stylophyllum sp.: Roniewicz, p. 105, Pl. 2: Fig. 1.
1974 ?Elysastrea sp.: Roniewicz, p. 111, Pl. 8: Figs. 1, 2.
1990 Pamirophyllum tatricum Roniewicz: in Melnikova & Ro-
niewicz, p. 87, Pl. 22: Figs. 14.
Genus Meandrostylophyllum Roniewicz & Michalík 1991
Meandrostylophyllum vesiculare Roniewicz & Michalík 1991
1991a Meandrostylophyllum vesiculare: Roniewicz & Michalík,
p. 159, Figs. 35.
Suborder Caryophylliina Vaughan & Wells 1943
Family Reimaniphylliidae Melnikova 1974
Genus Retiophyllia Cuif 1966
Hitherto, more than 10 species of Rhaetian Retiophyllia have
been described from the western Tethys, differing primarily in
396 RONIEWICZ and MICHALÍK
their corallite diameters, structure of endotheca, and ornamenta-
tion of septal faces (Roniewicz 1989). Commonly, the corals
with diameters ca. 5 mm and a ring of large peripheral dissepi-
ments observed in cross section were determined as Retiophyl-
lia sellae (Stoppani 1865) (see also Frech 1890; Roniewicz
1974; Fantini-Sestini & Motta 1984). However, recent studies
(Roniewicz 1989, and present paper) show that the species of
similar corallite diameter and appearance of calicular cross-sec-
tion may differ in the ornamentation of septal faces and the
structure of endotheca. Unfortunately, the structure of the endot-
heca in the R. sellae type material has never been examined. To
avoid misinterpretations, corals having granular septal ornamen-
tation and dissepiments traversing the lumen have been included
either in R. fenestrata (Reuss 1854) (diameters about 56 mm)
or R. gracilis Roniewicz 1989 (diameters about 34 mm), while
the corals with comparable diameters, but having short me-
nianes and a lumen filled with vesiculous dissepiments were de-
termined either as R. multiramis Roniewicz 1989 or R. para-
clathrata Roniewicz 1974, depending on the arrangement and
size of dissepiments and the septal number.
In the Carpathian collections (High Tatra-, Low Tatra-,
Stráov-, Malá Fatra Mts.), the corals of the sellae-like
pattern are frequent. This group contains at least four species
(gracilis, fenestrata, paraclathrata, and oppeli) which may
be distinguished from each other only when their endothecal
features are investigated (Fig. 1).
Some of the taxa discussed here were originally described
from the High Tatra Mts. under the name of Parathecosmilia
Roniewicz 1974 which included forms with straight septa and
an apparently parathecal wall (i.e. dissepimental in origin).
Recent observations of the Alpine reimaniphylliid corals, per-
fectly preserved in aragonite, proved that the shape of septa in
this group has no taxonomical significance (Roniewicz 1989).
This allows us to synonymize Parathecosmilia wit h Retio-
phyllia. In addition, recent observations show that in all these
corals the wall is purely epithecal, and not parathecal (Ro-
niewicz 1989; Roniewicz & Stolarski 1999).
The species presented below are ordered in the 1
representing corals with septa ornamented by granulations,
and with large, tabuloid dissepiments traversing the central
part of the corallite and the 2
group comprising the corals
with septa ornamented by short menianes, and lacking
tabuloid dissepiments in the central part of the corallite. In
corals of both groups, synapticular projections may develop
at the corallite periphery.
Retiophyllia fenestrata (Reuss 1854)
pars 1974 Parathecosmilia sellae (Reuss): Roniewicz, p. 110, Pl.
6: Figs. 13, Pl. 7: Figs 1, 2; Figs. 9, 10.
1989 Retiophyllia fenestrata (Reuss): Roniewicz, p. 59, Pl. 9: Fig.
6, Pl. 10: Fig. 1, Pl. 13: Fig. 1.
Retiophyllia frechi Roniewicz 1989
Pl. I: Fig. 1
1989 Retiophyllia frechi Roniewicz: p. 48, Pl. 7: Figs. 18, Pl. 9:
Fig. 5, Pl. 13: Fig. 2.
Retiophyllia gosaviensis Roniewicz 1989
Pl. I: Fig. 2; Fig. 1.2
?1974 Parathecosmilia sp.: Roniewicz, p. 111, Pl. 6: Fig. 4.
1989 Retiophyllia gosaviensis Roniewicz: p. 55, Pl. 9: Fig.1, Pl.
11: Figs. 4, 5.
Retiophyllia gracilis Roniewicz 1989
1989 Retiophyllia gracilis Roniewicz: p. 58, Pl. 9: Fig. 9, Pl. 10:
Fig. 2, Pl. 13: Fig. 3.
Retiophyllia clathrata (Emmrich 1853)
Pl. I: Figs. 3, 4ac; Fig. 1.4
1890 Thecosmilia clathrata Emmrich: Frech, p. 15, Pl. 4: Figs. 1,
1974 Retiophyllia clathrata (Emmrich): Roniewicz, p. 108, Pl. 4:
Fig. 1, Pl. 5: Fig. 3; Fig. 7.
Nomenclatural note: In the Emmrich collection housed at the
Geiseltal Museum, Halle, three specimens determined as The-
cosmilia clathrata Emmrich represent the large-corallite form
(Frech 1890, Pl. 4: Figs. 1, 5, 7: d 7.09.5 mm, locality Wunder-
graben in vicinity of Ruhpolding and Schwarzachen near Ber-
Plate I: Fig. 1. Retiophyllia frechi Roniewicz 1989: phaceloid cor-
allites in cross-section showing rare, thick septa. Specimen No.
SNM-, Istebné Valley. Fig. 2. Retiophyllia gosaviensis
Roniewicz 1989: phaceloid corallites in cross-section. Septa numer-
ous and subequal in thickness, large dissepiments crossing the axis
(at upper right corner), epitheca preserved, apophyses present (ar-
row). Synapticulae are lacking. Tectonically deformed corallite is
visible at the upper left side. Specimen SNM-, Domaniské
Stráne Hill. Figs. 3, 4a,b,c. Retiophyllia clathrata (Emmrich 1854):
3 phaceloid corallites in cross-section. Septa thin and numerous,
synapticulae numerous, axial space filled with blocky calcite; speci-
men SNM-, Rohoník Valley. 4a cross-section of corallites
showing well marked internal synapticular wall; 4b cross-sec-
tion, internal portion of septum showing obliquely arranged me-
nianes (arrow); 4c a fragment showing numerous synapticulae
constituting a sort of incomplete synapticular wall, traces of a very
thin epithecal wall covering the septal ends are to be noticed at right
lower corner; specimen from the Buèek collection No. 535/86. Malé
Karpaty Mts. Fig. 5a,b. Retiophyllia cf. oppeli (Reuss 1865): 5a
a calice in cross-section showing septa peripherally enlarged and
provided with thick synapticular projections (arrow), pellicular epi-
theca has been destroyed; 5b phaceloid corallites in cross-sec-
tion; some corallites with peripherally thickened septa and other
corallites with thin septa; pellicular epitheca partially preserved;
specimen No. SNM-, Bobrovèek, Smoliacke Hrádky Hill. Fig.
6. Chondrocoenia schafhaeutli (Winkler 1861). A plocoid colony in
cross-section. Intercalicular, perithecal space is very narrow as the
calices are densely crowded; thick columella and characteristic
anastomosis of septa (at left) are to be noticed. Specimen No. SNM-
, Trnovec Valley. All the specimens were derived from the Rha-
etian Fatra Formation, Western Carpathians. Scale bars in mm.
RHAETIAN SCLERACTINIAN CORALS IN THE WESTERN CARPATHIANS 397
398 RONIEWICZ and MICHALÍK
gen) and one belongs to the small-corallite form (unfigured, d
5.56.0 mm; Schwarzachen near Bergen). The series of speci-
mens of larger diameters, well illustrated by Frech, is here es-
tablished as the type series of R. clathrata (Emmrich).
Description. Thin radial elements with sharp costal edges
equal in thickness. Septa S1 subequal to S2, approaching to
the axis, S3 longer than half the length of S1, S4 usually
longer than 1/3 of the radius, but in some systems rudimenta-
ry or absent, S5 sporadical. Internal septal portion thin, zig-
zag in shape, with prominent menianes. Peripheral parts with
numerous lateral projections which may link the septa to
form the synapticulae. Endotheca formed by thin and rather
uniform, not extended dissepiments, which slope axialward
from the periphery.
Retiophyllia cf. oppeli (Reuss 1865)
Pl. I: Fig. 5a,b
The form presented here resembles one of the Alpine
Rhaetian Thecosmilia oppeli Reuss forms described by Frech
from Grosser Zlambachgraben (Frech 1890, Pl. 3: Fig. 4E),
and R. oppeli (Reuss) described by Roniewicz (1989, Pl. 10:
Fig. 3) from Fischerwiese.
Retiophyllia paraclathrata Roniewicz 1974
1974 Retiophyllia paraclathrata Roniewicz: p. 108, Pl. 4: Fig. 2,
Pl. 6: Figs. 1, 2; Fig. 8.
In the collection, the specimens differ in the shape of septa
which may be either zigzag, or straight. Corals with zigzag
septa and a prominent lateral ornamentation resemble the ho-
lotype. The corals with straight septa have sellae-like corallite
pattern in cross section.
Retiophyllia sp. 1
A fragmentary coral similar to R. defilippi (Stoppani).
Genus Distichophyllia Cuif 1975
Large Distichophyllia-like coral, associated with a retio-
phyllian coral of the diameter of 56 mm.
Suborder Archaeocoeniina Alloiteau 1952
Family Actinastreidae Alloiteau 1952
Genus Chondrocoenia Roniewicz 1989
Chondrocoenia schafhaeutli (Winkler 1861)
Pl. I: Fig. 6
1974 Cyathocoenia alpina (Guembel): Roniewicz, p. 103, Pl. 1:
Figs. 3, 4; Fig. 3.
1989 Chondrocoenia schafhaeutli (Winkler): Roniewicz, p. 104, Pl.
33: Figs. 13, 5 (here additional synonymy included).
Suborder Faviina Vaughan & Wells 1943
Family Pamiroseriidae Melnikova 1984
Genus Pamiroseris Melnikova 1971
Pamiroseris rectilamellosa (Winkler 1861)
1974 Pamiroseris rectilamellosa (Winkler): Roniewicz, p. 114, Pl.
8: Fig. 3, Pl. 10: Figs. 13.
1989 Pamiroseris rectilamellosa (Winkler): Roniewicz, p. 111, Pl.
34: Figs. 35 (here additional synonymy included).
Genus Rhaetiastrea Roniewicz 1974
The genus, described originally from the Tatric Superunit, in
addition to the type species, contains the Liassic (Dommerian)
species from Morocco described under the name of
Trigerastrea minima Beauvais (Beauvais 1986, p. 47, Pl. 12:
Fig. 1). The Moroccan species resembles Rhaetiastraea
tatrica morphologically (cerioid colony and high density of
septal apparatus) and microstructurally (thin septa formed by
trabeculae provided with granulations). It differs from the
Tatric species by having a higher number of septa (4565 in
the Moroccan species, while 3445 in the Tatric species).
Rhaetiastrea tatrica Roniewicz 1974
1974 Rhaetiastrea tatrica Roniewicz: p. 113, Pl. 9: Figs. 13; Fig. 12.
Suborder Fungiina Verrill 1865, faute de mieux
Family Astraeomorphidae Frech 1890
Genus Astraeomorpha Reuss 1854
Astraeomorpha confusa (Winkler 1861)
1974 Astraeomorpha crassisepta Reuss: Roniewicz, p. 113, Pl. 8:
Figs. 4, 5.
1989 Astraeomorpha confusa (Winkler): Roniewicz, p. 96, Pl. 28:
Figs. 1, 2, Pl. 29: Figs. 13 (additional synonymy included).
Acknowledgments: The authors are grateful to Dr. E. Mory-
cowa and Dr. H. Eliáová for critical reading of the manuscript
and for inspiring comments. Dr. S. Buèek is acknowledged for
kindly offering his samples for examination. The photographs
and the majority of the thin sections were prepared in the Insti-
tute of Paleobiology, Polish Academy of Sciences. Financial
supports were provided by KBN Grant No. 6 PO4D 037 14
and VEGA Grant 4076. This contribution has been made in
the context of collaboration between the Polish and Slovak
Academies of Sciences.
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