GEOLOGICA CARPATHICA, 49, 6, BRATISLAVA, DECEMBER 1998
PIENINIA OBLONGA SKELETAL PARTS OR ENDOPARASITES
OF KERATOSA SPONGES ?
Department of Geology and Paleontology, Faculty of Science, Comenius University, Mlynská dolina,
842 15 Bratislava, Slovak Republic
(Manuscript received June 10, 1998; accepted in revised form September 1, 1998)
Abstract: The problematic bodies Pieninia oblonga Borza & Miík common in Barremian to Upper Eocene shallow-
water sediments were considered as algae, rudist fragments, echinoderm remains, or sclerites of Gorgonaceae. They
were recently found inside the skeleton of sponges, probably Keratosa. The growth stages of these bodies are described.
The question remains whether they are unusual sclerites or parasites. The way of their loosening from the skeletons
was not observed.
Key words: Western Carpathians, Paleocene, biohermal limestones, Pieninia problematic bodies, Octocorallia,
Problematic bodies named Pieninia oblonga were described
by Borza & Miík (1976) and considered to be of algal
origin. Later they were found in many countries and various
opinions concerning their systematic position were advanced
(see in the following chapters). Surprising occurrences of
Pieninia oblonga in skeletal parts of presumed sponges or
hydrozoans (I received from E. Köhler the first thin sections
of them) initiated this new study.
Pieninia oblonga Borza & Miík 1976
+ 1976 Pieninia oblonga n.sp. Borza & Miík: p. 6577, Pl. 14
1976 Forma indeterminata, Luperto Sinni: Pl. 58, Figs. 45.
1978 Pieninia oblonga, Knauer & Gellai: Pl. 4, Figs. 12.
1979 Pieninia oblonga, Miík: p. 708, Pl. 8, Figs. 16.
1984 Pieninia oblonga, Bignot, Haas & Poignant: p. 437438, Pl. I,
1986 Pieninia oblonga, Granier: p. 103108, Pl. I, Figs. at.
1986 Pieninia oblonga, Császár: p. 165, Pl. XX, Fig. 1.
1987 Pieninia oblonga, Granier: p. 51, Pl. 43, Figs. do.
1988 Pieninia oblonga, Granier: p. 61, Pl. 12, Figs. do.
1988 Pieninia oblonga, Øehánek: p. 255, Fig. 4.
1991 Pieninia oblonga, Miík, Sýkora, Mock & Jablonský: p. 64, Pl.
XVIII, Fig. 3.
1991 Pieninia oblonga, Schlagintweit: p. 2829, Pl. 8, Figs. 15.
The bodies have the form of a biaxial ellipsoid with radial-
fibrous structure consisting of calcite fibres diverging from a
small central canal. The surface is sometimes covered with
small protuberances. According to the original description
the length ranges from 0.100 mm to 0.450 mm.
Pieninia oblonga was quoted from Slovakia, Italy, Hungary,
Croatia, Spain and Austria (that is from the Carpathian Mts.,
Dinaric Mts., Appenines, Betics and Eastern Alps). We also
found it in Urgonian facies of Algeria, Constantine (Tellian
The stratigraphical range known up to now is: Barremian
Upper Eocene (Priabonian).
In the first paper (Borza & Miík 1976) a Barremian
Thanetian range was supposed. Later, we found Pieninia ob-
longa in Ilerdian limestone containing Discocyclina seunessi
and Nummulites exilis, as well as in Lower Eocene limestone
with Cuvillierina vallensis (Miík et al. 1991, p. 50, 64). Re-
cently I observed it in Upper Eocene (Priabonian) limestone
with Chapmannina gassinesis, locality Podbánske (Pl. I:
Fig. D). Øehánek (1987) mentioned it without details from
the Upper Jurassic; verification is needed.
In the Western Carpathians, it is most common in
Barremian-Aptian, Senonian and Paleocene limestones.
Opinions about taxonomical appurtenance
The affinity of Pieninia oblonga to algae (probably Codia-
cae) was suggested in the first paper (Borza & Miík 1976, p.
65). Mamet & Roux (1982, p. 962) stated that its kind of
growth is strongly reminiscent of the genus Nuia (alga of un-
certain taxonomic position). Bignot et al. (1984, p. 138) pre-
sumed that P. oblonga could correspond to cylindrical foliated
expansions of Mollusca, more precisely of rudists; they con-
sidered that our occurrences in Paleocene limestones were re-
deposited. Schlagintweit (1991, p. 28) suggested they could
pertain to echinoderms (Echinodermnatur). Radoièiæ (1983,
p. 80) interpreted them as sclerites derived from Octocorallia,
most probably Gorgonaceae. Granier (1986, p. 105) argued
that P. oblonga belonged to calcareous skeleton particles
(sclerites) of Alcyonaria, close to the recent genus Eunice
P. oblonga occurs mainly in bioherms, in reefal and pararee-
fal facies (Miík 1979). According to Schlagintweit (1991, p.
28), it is typical of the reef or platform margin (facial zone 5 of
Wilson). According to Granier (1986, p. 105), it is the most
characteristic for the outer part of carbonate platforms, from
the barrier to the talus where it can be redeposited into calci-
Frequency in the Western Carpathians
BarremianAptian. In limestones of Urgonian facies s.l.
(Pl. I: Figs. AC, I), usually only 12 specimens (maximum 6
specimens) in thin section (20
20 mm) can be found. The fre-
quency of the P. oblonga in sets of thin sections was as fol-
lows: (a) In the pebbles of Urgonian limestones from the Albi-
an conglomerates, Central Western Carpathians (Miík et al.
1981, p. 3233), it occurred in four thin sections from 53
(about 8 %). (b) In pebbles of Urgonian limestones from the
Peri-Klippen zone (Miík & Sýkora 1981, p. 3744), it was
found in 18 from 143 thin sections (about 13 %); it is most fre-
quently found in limestones with orbitolinas 12/83, with
fragments of rudists 5/13. It was totally absent in the microfa-
cies with miliolids 0/9 which indicates that Pieninia be-
longs to the bioconstructing organisms. (c) In the pebbles of
Urgonian limestones from the Strihovce conglomerates of
Eocene age (Miík et al. 1991, p. 3033), it was present in two
thin sections from 18 (about 10 %). (d) In the pebbles of Urgo-
nian limestones from Proè conglomerates of PaleoceneLower
Eocene age (Miík et al. 1991, p. 3435), P. oblonga occurred
in 5 thin sections from 42 (about 12 %). (e) From the outcrops
of Urgonian Niná limestone (Pieniny Klippen Belt, Miík
1990, p. 44), it was found in two from 12 thin sections (about
17 %). It can be summarized that P. oblonga occurs in approx-
imately 12 % of thin sections (with the surface about
) from the limestones with Urgonian facies.
Senonian. The frequency in sandy limestones from out-
crops in the Brezovské Karpaty Mts. was 6/23, or nearly 26 %.
P. oblonga was associated with fragments of bivalves (mostly
rudists), orbitoids, coralline algae, echinoderm articles and
bryozoans. In pebbles of Senonian limestones with rudists, its
frequency from Eocene Strihovce conglomerates was 3/9
(about 33 %), but it was absent in limestones with
Pseudosiderolites vidali 0/6 (Miík et al. 1991, p. 35, 37).
Paleocene. The frequency in the pebbles of Montian-Thane-
tian bioherm limestones (Kambühel Limestone) from Eocene
Strihovce conglomerates (l.c., p. 3040) was 1/13 (about 8 %);
in the limestone pebbles of the same age from Paleocene
Lower Eocene Proè conglomerates, it was 2/12 (about 17 %)
(Miík et al. 1991, p. 48).
Single occurrences in the Ilerdian, Lutetian, Priabonian
could not be quantified.
Paleocene limestones with Pieninia oblonga inside
the bioclasts of probable Keratosa sponges
Formerly only loose specimens of Pieninia oblonga were
known. We illustrate some of them in the Pl. I: Figs. AI. In the
1992 we succeeded in finding P. oblonga inside fragments of
supposed Coelenterata or Porifera (Pl. I: Figs. J, K; Pl. II: Figs.
AD; Pl. III: Figs. AE) and asked several experts about their
opinion. Helena Eliáová (State Geological Institute, Prague)
supposed from the microphotographs that the fragments could
Plate I: Pieninia oblonga Borza & Miík, occurrences of the
isolated bodies in limestones. Fig. A In the Urgonian facies
(Upper BarremianLower Aptian), associated with corals and algae.
Pebble from the Cenomanian conglomerates of the Manín Unit,
Ovèiarsko near ilina. Thin section No. 5150,
55. Fig. B The
same in cross-polarized light. Fig. C The same locality, thin
section No. 6149,
43. Fig. D In the Upper Eocene (Priabonian)
biohermal limestone. Podbánske. Thin section No. 14970,
E In the Paleocene Kambühel Limestone, Ve¾ký Lipník.
Fig. F In the Aptian limestone. Pebble from the Paleocene Proè
Conglomerate of the Pieniny Klippen Belt, Proè-d. Thin section No.
136. Fig. G In the Orbitolina-bearing limestone (Upper
BarremianLower Aptian), cobble from the Cenomanian
conglomerates, Manín Unit, Malé Hradisko pri iline. Thin section
43. Fig. H The same locality. Thin section No. 6154,
43. Fig. I Partly silicified specimen in the Urgonian facies with
Palorbitolina lenticularis (Upper BarremianLower Aptian).
Pebble from the Cenomanian conglomerates, Manín Unit, Hradná-
JRD-d. Thin section No. 7116,
43. Fig. J Pieninia oblonga
within the skeleton of Keratosa sponge or of Coelenterata.
Paleocene biohermal Kambühel Limestone. Ve¾ký Lipník,
K Juvenile stage of Pieninia oblonga, not well differentiated
from the skeletal mass. The same locality,
Fig. 1. Localities with Pieninia oblonga in sponge- or coelenter-
PIENINIA OBLONGA SKELETAL PARTS OR ENDOPARASITES OF KERATOSA SPONGES ? 403
Plate II: Pieninia oblonga Borza & Miík in the skeleton of the sponge Keratosa or of a Coelenterata fragment. Paleocene
Kambühel biohermal limestone, Peri-Klippen zone. Fig. A Settlement Dúbrava near Stará Turá. Thin section No. 22706,
40. Fig. B
The same. Fig. C Ve¾ký Lipník,
40. Fig. D The same.
PIENINIA OBLONGA SKELETAL PARTS OR ENDOPARASITES OF KERATOSA SPONGES ? 405
Plate III: Pieninia oblonga Borza & Miík in the skeleton of the sponge Keratosa of the Coelenterata bioclast. Paleocene Kambühel
biohermal limestone, Peri-Klippen zone. Fig. A Motenec,
27. Fig. B Ve¾ký Lipník,
30. Fig. C Motenec,
86. Fig. D The same.
belong to the corals Poritidae Gray or Actinacidae Vaughan et
Wells, but their hydrozoan nature could not be excluded (letter
1992). Dragica Turnek (Centre of Scientific Research of the
Slovenian Academy of Sciences and Arts, Ljubljana) in the let-
ter from 31. 3. 1992 deduced from the microphotographs that
the fragments with P. oblonga could belong to some sponges;
but parasites or algae are not excluded. Elzbieta Morycowa
(Jagellonian University, Cracow) also did not exclude their par-
asitical nature (letter 1992). Joachim Reitner (Geol. Paleont.
Institute, University Göttingen) considered the bioclasts with
Pieninia as calcitized Keratosa which seems to be most likely.
The microfacies of these samples are as follow:
Boundstones to floatstones (wackestones) of cream colour;
Paleocene age. Localities:
1. Ve¾ký Lipník (Pl. II: Figs. C, D; Pl. III: Fig. B). Algae are
dominating: Lithophyllum sp., Lithothamnium sp., Amphiroa
propria (Lemoine), Elianella elegans (Pfender & Basse) =
Parachaetetes asvapatii Pia, Ethelia alba (Pfender) =
Polystrata, rare Dasycladales including Acicularia sp.
Abundant sessile foraminifers: nubecularids, Haddonia
heissigi Hagn, Miniacina multicamerata Scheibner, Bullopora
sp., Thurammina sp., also rotalids, miliolids and Anomalina
sp. Rarely echinoid spines, fragments of bivalves with
prismatic structure, gastropods, serpulids, corals; some
fragment of Porifera (?) containing Pieninia oblonga Borza &
Miík; also loose specimens of P. oblonga are present. Sandy
admixture of scattered quartz, single grains of plagioclase and
2. Motenec (Pl. III: Figs. A, C, D). Algae especially
Corallinaceae are dominant: Lithothamnium sp., Lithophyllum
sp., Amphiroa propria (Lemoine), Distichoplax biserialis
(Dietrich), Elianella elegans (Pfender & Basse), Ethelia alba
(Pfender), Acicularia sp. Among the foraminifers the
encrusting forms prevail: nubecularids, Bullopora cf.
tuberculata Sollas, Planorbulina cretae (Marsson),
Miniacina ? multiformis Scheibner, also miliolids, Alveolina
(Glomalveolina) primaeva Reichel, Anomalina sp. Rare
fragments of bryozoans, echinoderm spines and plates, tubes
of serpulid worms, ostracods, fragments of corals. Pieninia
oblonga Borza & Miík is present inside the fragments of
Porifera (?) and loose specimens are also present. Rare
admixture of quartz grains (max. 0.55 mm). Mud matrix is
dominant, sometimes with small agglutinated pellets. Rare
voids are rimmed with pigmented radiaxial cement, while a
clear isometric mosaic occupies the central parts.
3. Stará Turá-Drahý vrch-II. Fragments of Hydrozoa, cor-
als, Bryozoa and bivalves are dominant. Algae are rare in
comparison with the previous localities: Lithophyllum sp.,
Mesophyllum sp., Elianella elegans (Pfender & Basse), Aci-
culella sp. Foraminifers are surprisingly rare: Haddonia heis-
sigi Hagn, Miniacina sp.: further some serpulids, echinoid
spines, ostracods and gastropods. One fragment of Porifera
(?) with immature Pieninia oblonga; loose specimens are
absent. The limestone contains a considerable amount of
clastic quartz (silt), single grains of chromspinelids and tour-
maline, tiny fragments of dolomites and volcanites. Small
pellets are frequent.
4. Dúbrava near Stará Turá (Pl. II: Figs. A, B; Pl. III: Fig. E).
Coralline algae overgrown with encrusting foraminifers: Had-
donia heissigi Hagn, Thurammina sp. and nubecularids; abun-
dant Elianella elegans Pfender & Basse, rare ostracods and cor-
als. Sponge fragment probably of Keratosa filled by small yel-
lowish and larger white Pieninia oblonga.
The distance between the first and the last localities is about
250 km (Fig. 1). Localities 1 and 2 were found by E. Köhler,
loc. 3 by S. Buèek, and loc. 4 by myself (M. Miík).
Paleocene biohermal Kambühel limestones occur only as
blocks and pebbles derived from a ridge which bordered the
Pieniny Klippen Belt on its inner side. A map of all known
localities has been published by Miík (1996, Fig. 1).
The bodies of Pieninia oblonga are irregularly distributed
within the branches of bioclasts belonging to Porifera or Co-
elenterata. The process of their growth can be described as
follows. The bodies are at first very small and not well differ-
entiated from the surrounding framework (Pl. I: Fig. K).
When they acquire larger dimensions, their radial-fibrous
structure becomes distinct but the colour in thin sections still
remains yellowish. During the further growth they accomo-
dated to the form of the branches: some of them are strongly
elongated or bent (Pl. II: Figs. AC), while others are almost
isometric (Pl. II: Fig. D). Large individuals from the final
stage are already well differentiated from the skeletal frame-
work and acquire a white colour (e.g. Pl. I: Fig. 1; Pl. II:
Fig. C; Pl. III: Fig. D). The hypothesis about the calcite
composition of Pieninia and aragonite of Pieninia-bearing
skeleton was not checked; the probability that aragonite
could be preserved in these Paleocene limestones is almost
The attribution of the Pieninia-bearing bioclasts to Gorgo-
naceae or to Keratosa sponges bears some difficulties. Recent
Gorgonaceae possess a horny organic skeleton and spicules of
high Mg-calcite, recent Keratosidae display a skeleton consist-
ing of spongine. The structure of the Pieninia-bearing bio-
clasts is well preserved and excludes a postmortal calcitiza-
tion. It may be only accepted that the ancestors of these groups
possessed a calcitic skeleton.
The loose bodies of Pieninia oblonga occur with Pieninia-
bearing bioclasts in the same thin sections (Pl. I: Fig. E; Pl.
II: Fig. F). We do not know the process involved in the loos-
ening of the bodies of Pieninia from the skeleton. If the skel-
etons were from other material than low Mg-calcite, their
postmortal dissolution could have taken place. No sign of
such a dissolution (corrosion) on the skeletons was observed.
If the Pieninia was an endoparasite, the loosening could be
mediated by its dissolution activity. No such dissolution from
inside was observed. There is hope that new, more favorable
occurrences from other localities will solve the question.
The Pieninia-bearing skeletal fragments were found only
in the Paleocene limestones. In the Barremian-Aptian lime-
stones (Urgonian facies) where Pieninia is much more fre-
quent and fragments of Coelenterata currently occur in the
same thin sections, no analogic Pieninia-bearing skeletons
were found. Perhaps Pieninia oblonga represents objects of
different origin but we consider this probability to be very low.
PIENINIA OBLONGA SKELETAL PARTS OR ENDOPARASITES OF KERATOSA SPONGES ? 407
The problematic bodies Pieninia oblonga Borza & Miík
(1976) occur in shallow-water limestones ranging from
Barremian to Late Eocene. They were found up to now in
Slovakia, Italy, Hungary, Austria, Croatia, Spain and Algeria.
They were considered to be algae, expansions of rudists,
echinoderm fragments and mostly sclerites of Gorgoniaceae.
Pieninia occurs most frequently in Barremian-Aptian lime-
stones of the Western Carpathians from 268 thin sections
31 contained Pieninia (frequency about 12 %). In 1992,
Pieninia was found in the fragments of Coelenterata or sponge
skeletons in Paleocene biohermal limestones. Skeletal
fragments were not definitely determined. After J. Reitners
opinion they belong to Keratosida sponges. Recent Keratosa
as well as Gorgoniaceae possess a skeleton consisting of
organic matter and spicules of Gorgoniaceae of unstable high-
Mg calcite. Perhaps their ancestors might had a low-Mg
calcite skeleton. The possibility Pieninia is an endoparasite
was discussed. The process by which Pieninia was loosened
from the skeleton is not clear. We call attention to this
interesting phenomenon with the hope that some better
findings will resolve the problem.
Acknowledgements: The author would like to express his
gratitude to RNDr. E. Köhler, DrSc. and RNDr. S. Buèek,
CSc. for the granting of samples, RNDr. H. Eliáová, CSc.
(Prague), Dr. D. Turnek (Ljubljana), Prof. E. Morycowa
(Cracow) and Prof. J. Reitner (Göttingen) for their kind
consultations of microphotographs, Dr. B. Granier (France)
and RNDr. J. Michalík, DrSc. (SAV Bratislava) for valuable
comments on the manuscript.
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