Bernecker, Michaela

Senowbari-Daryan, Baba

English

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An International Journal  of Palaeontology and GeobiologySeries A /Reihe AMitteilungen der Bayerischen Staatssammlung für Paläontologie und GeologieMünchen 2010ZittelianaZitteliana An International Journal of Palaeontology and Geobiology Series A/Reihe A 50Jubilee Volume50Zittelianacontents/InHALt BaBa SenowBari-Daryan & Michaela Bernecker Amblysiphonella agahensis nov. sp., and Musandamia omanica nov. gen., nov. sp. (Porifera) from the Upper Triassic of Oman 3alexanDer nützel A review of the Triassic gastropod genus Kittliconcha Bonarelli, 1927 – implications for the phylogeny of Caenogastropoda 9anDrzej kaiM & Maria aleSSanDra contiA problematic zygopleuroid gastropod Acanthostrophia revisited 21Gernot arpAmmonitenfauna und Stratigraphie des Grenzbereichs Jurensismergel/Opalinuston- Formation bei Neumarkt i.d. Opf. (oberstes Toarcium, Fränkische Alb) 25Volker Dietze Über Ammonites Humphriesianus umbilicus QuenSteDt, 1886 an seiner Typus-Lokalität (östliche Schwäbische Alb, Südwestdeutschland) 55Volker Dietze, Günter SchweiGert, GerD Dietl, wolfGanG auer, wolfGanG DanGelMaier,  roGer furze, Stefan GräBenStein, Michael kutz, elMar neiSSer, erich SchneiDer & DietMar  SchreiBer Rare Middle Jurassic ammonites of the families Erycitidae, Otoitidae and  Stephanoceratidae from southern Germany 71wolfGanG witt Late Miocene non-marine ostracods from the Lake Küçükçekmece region, Thrace (Turkey) 89 jérôMe prieto Note on the morphological variability of Keramidomys thaleri (Eomyidae, Mammalia) from Puttenhausen (North Alpine Foreland Basin, Germany) 103Martin pickforD Additions to the DehM collection of Siwalik hominoids, Pakistan: descriptions and interpretations 111Michael krinGS, nora Dotzler, thoMaS n. taylor & jean Galtier Microfungi from the upper Visean (Mississippian) of central France: Structure  and development of the sporocarp Mycocarpon cinctum nov. sp. 127Zlatko kVaČek & joSef BoGnerAracistrobus, an enigmatic non-araceous fossil from the Eurasian Oligocene and Miocene 137Instructions for authors/Hinweise für Autoren 143Zitteliana München, 30.06.2010 Issn 1612-412XA 50 143 seitenAn International Journal of Palaeontology and GeobiologySeries A/Reihe AMitteilungen der Bayerischen Staatssammlung für Paläontologie und Geologie50Editors-in-Chief/Herausgeber: Gert Wörheide, Michael KringsProduction and Layout/Bildbearbeitung und Layout: Martine Focke, Manuela SchellenbergerBayerische Staatssammlung für Paläontologie und GeologieBayerische Staatssammlung für Paläontologie und GeologieRichard-Wagner-Str. 10, D-80333 München, Deutschlandhttp://www.palmuc.deemail: zitteliana@lrz.uni-muenchen.deFür den Inhalt der Arbeiten sind die Autoren allein verantwortlich.Authors are solely responsible for the contents of their articles.Copyright © 2010 Bayerische Staassammlung für Paläontologie und Geologie, MünchenDie in der Zitteliana veröffentlichten Arbeiten sind urheberrechtlich geschützt. Nachdruck, Vervielfältigungen auf photomechanischem, elektronischem oder anderem Wege sowie die Anfertigung von Übersetzungen oder die Nutzung in Vorträgen, für Funk und Fernsehen  oder im Internet bleiben – auch auszugsweise – vorbehalten und bedürfen der schriftlichen Genehmigung  durch die Bayerische Staatssammlung für Paläontologie und Geologie, München.ISSN 1612-412XDruck: Gebr. Geiselberger GmbH, Altöttingeditorial BoardA. Altenbach, MünchenB.J. Axsmith, Mobile, ALF.T. Fürsich, ErlangenK. Heißig, MünchenH. Kerp, MünsterJ. Kriwet, StuttgartJ.H. Lipps, Berkeley, CAT. Litt, BonnA. Nützel, MünchenO.W.M. Rauhut, MünchenB. Reichenbacher, MünchenJ.W. Schopf, Los Angeles, CAG. Schweigert, StuttgartF. Steininger, Eggenburgcover illustrations: (from left to right) Shell of the gastropod Loxonema regium De koninck from the Carboniferous of Belgium (redrawn from De koninck 1881); Solitary coral Caninia sp. from the Carboniferous of England (redrawn from raMSBottoM in Mckerrow 1978); Tooth of the rare ruminant Orygotherium escheri Von Meyer from the Miocene of Germany (after röSSner & MörS 2001). Back cover: Atrium of the Munich Palaeontological Museum, view from the main entrance.Umschlagbilder: (von links nach rechts) Gehäuse der Schnecke Loxonema regium De koninck aus dem Karbon von Belgien (neu gezeichnet nach De koninck 1881); Solitärkoralle Caninia sp. aus dem Karbon von England (neu gezeichnet nach raMSBottoM in Mckerrow 1978); Zahn des seltenen Wiederkäuers Orygotherium escheri Von Meyer aus dem Miozän von Deutschland (nach röSSner & MörS 2001). Rückseite: Lichthof des Paläontologischen Museums München, Blick vom Haupteingang.3Abstract Two chambered sponges – A. agahensis nov. sp. and Musandamia omanica nov. gen., nov. sp. – from the Norian reefal limestones of Jebel Agah, southeast of the Musandam Mountains, Oman, are described. Both species are the first reported from the Triassic of the Arabian Peninsula. The fossil association is also mentioned. Key words: Porifera, “Sphinctozoa”, Amblysiphonella, Musandamia, Norian, Jebel Agah, Oman.KurzfassungZwei gekammerte Schwämme – A. agahensis nov. sp. und Musandamia omanica nov. gen., nov. sp. – werden aus dem norischen Riffkalken des Jebel Agah, südöstlich des Musandam Gebirges (Oman), beschrieben. Beide Arten stellen die ersten Nachweise für die Trias der Arabischen Halbinsel dar. Die Vergesellschaftung der Fossilien wird ebenfalls erwähnt.Schlüsselwörter: Porifera, “Sphinctozoa”, Amblysiph-onella, Musandamia, Nor, Jebel Agah, Oman.1. Introduction and geographic position of the locality Hypercalcified sponges, particularly the “sphinctozoans”, are the most abundant reef builders in Upper Triassic reefs and reefal deposits of Oman. Carnian sponges and other reef organisms from various localities in Oman were described by Senowbari-Daryan et al. (1999) and Senowbari-Daryan & bernecker (1997, 2009). Results from microfacies investiga-tions of Norian-Rhaetian reefs of Oman and descriptions of the reef organisms were published by bernecker (1996, 2005). Senowbari-Daryan & yancey (2007) and Senowbari-Daryan & Maurer (2008) described the Norian sponges of Musandam Mountains, NW of the Jebel Agah, where A. agahensis nov. sp. and Musandamia omanica nov. gen. nov. sp., were found. Jebel Agah is located between the carbonate succession of the Musandam shelf to the northwest and the Semail Ophiolite to the southeast (Textfig. 1). Geology and tectonic development of this area is summarized by robertSon et al. (1990) and recently by Maurer et al. (2008). Microcoprolites of this locality were described by Senowbari-Daryan & bernecker (2005). The studied material, including two rock pieces and one thin section, is stored in the Bayerische Staatssammlung für Paläontologie und Geologie, Munich, under accession numbers BSPG 2009 I 77 and 2009 I 78a & b.  2. Systematic palaeontologyClass  Demospongea SollaS, 1875Subclass  Ceractinomorpha levi, 1953Order Agelasida verrill, 1907Suborder Porata Seilacher, 1962Family Sebargasiidae De laubenfelS, 1955Subfamily Sebargasiinae Senowbari-Daryan, 1990Genus Amblysiphonella SteinMann, 1882Type species:  Amblysiphonella barroisi SteinMann, 1882.Additional species included: See Senowbari-Daryan (1990) and Senowbari-Daryan & Garcia-belliDo (2002). Amblysiphonella agahensis nov. sp.Pl. 1, Fig. 1Derivat io nominis :  Occurrence of the species in Jebel Agah.*Author for correspondence and reprint requests; E-mail: basendar@pal.uni-erlangen.deAmblysiphonella agahensis nov. sp., and Musandamia omanica nov. gen., nov. sp. (Porifera) from the Upper Triassic of OmanByBaba Senowbari-Daryan1* & Michaela Bernecker21Geozentrum Nordbayern, Department of Palaeontology, University of Erlangen-Nürnberg, Loewenichstrasse 28, 91054 Erlangen, Germany2German University of Technology, PO Box 1816, Athaibah PC 130 Muscat, Sultanate of OmanManuscript received November 11, 2009; revised manuscript accepted February 22, 2010Zitteliana 3 - 7 1 Pl., 1 Textfig. München, 30.06.2010 ISSN 1612 - 412XA504Holotype:  Pl. 1, Fig. 1.Locus typicus:  Reefal limestones of Jebel Agah, south of Musandam Mountains (see Fig. 1).Age:  Norian.Diagnosis :  Multi-branched species of the genus Amblysi-phonella with hemispherical to spherical chambers. Chamber walls pierced with single or branched pores. Wall of spongo-coel contains large pores, which continue as small tubes into chamber interiors. Individual chambers clearly visible from outside of sponge. Chamber interiors whole. Material :  One multi-branched specimen (BSPG 2009 I 77).  Description: This dichotomously multi-branched sponge is composed of numerous hemispherical to spherical chambers. After increasing the chamber diameter from the initial stage, at least three individual branches originate from a large chamber. This large chamber reaches a diameter of approximately 50 mm and a height of 13 mm. The diameter of the individual branches is ~30 mm, which corresponds to the diameter of the chambers. The chamber height varies between 8 and 12 mm. The chamber walls are about 1.4 mm thick, and are pierced by single or dichotomously branched pores of more or less equal distribution. They reach diameters of up to 0.6 mm. A spongocoel of 6-8 mm and assignable to the retrosiphonate type passes through the individual branches. The wall of the spongocoel has the same thickness as the chamber walls. The spongocoel wall is pierced by different large and rimmed pores up to 2 mm in diameter. Several large pores continue into the chamber interiors as small tubes. Chamber interiors are devoid of additional filling skeleton or vesiculae. Comparison:  Amblysiphonella agahensis nov. sp. repre-sents the first Triassic species of the genus from the Arabian Peninsula. A. agahensis differs from all other Triassic species of that genus listed in Senowbari-Daryan & Di Stefano (1988) and Senowbari-Daryan (1990) by the branching mode of growth and the size of the chambers.   Genus Musandamia nov. gen.Derivat io nominis :  Named after the occurrence of the sponge in Musandam Mountains. D i a g n o s i s :  Amblysiphonella-like perforate thalamid sponge with spongocoel of ambisiphonate type. Primary Textfigure 1: Geological map of the area with the location of Jebel Agah. Plate 1: Amblysiphonella agahensis nov. sp. and Musandamia omanica nov. gen., nov. sp. from the Norian of Jebel Agah, Oman. Scale bars in Figs 1–3 = 10 mm. (1) Ambysiphonella agahensis nov. sp., holotype; section through several branches showing the hemispherical to spherical chambers. Several large pores of the spongocoel continue into the chamber interiors as small tubes (BSPG 2009 I 77). (2) Musandamia omanica nov. gen., nov. sp., holotype, longitudinal section showing the sponge with moderately flattened chambers with relatively thick chamber walls. The wall of the spongocoel is thin and pierced by a single large pore. The sponge serves as a substrate for an inozoan sponge. Polished rock surface (BSPG 2009 I 78a). (3) Same as Pl. 1, Fig. 2., holotype specimen (thin section photograph in transmitted light) (BSPG 2009 I 78b). (4) Musandamia omanica nov. gen., nov. sp. SEM-microphotograph showing the spherulites of the rigid skeleton. (5) Similar to Pl. 1, Fig. 4.6lites are of different sizes, reaching diameters of up to 100 µm (Pl. 1, Figs 4–5). Spherulites indicate the primary aragonitic mineralogy of the skeleton. Spicules have not been observed.Fossi l  associat ion:  The following organisms associated with the sponge were identified in one thin section: Heterastri-dium sp., Microtubus communis flüGel, and chaetetid sponges gen. and sp. indet. The organisms are encrusted with microbial crusts (“Spongiostromata” sensu Pia in hirMer 1927). The index fossil Microtubus communis and the foraminifer Hirsuto-spirella pilosa, illustrated by Senowbari-Daryan & bernecker (2005), are suggestive of a Norian-Rhaetian (most probably Norian) age of the investigated material and described sponges. AcknowledgementsFinancial support was provided by the “Sonderfonds für wissenschaftliche Arbeiten an der Universität Erlangen-Nürnberg” to B. Senowbari-Daryan. We thank Novus West Asia for granting permission to publish the species, and C. tollanD of Oolithica Geoscience (Cheltenham, UK) for the sampling. Helpful comments from reviewer M. noSe (Munich) are gratefully acknowledged.    3. References bernecker, M. (1996): Upper Triassic reefs of the Oman Mountains: Data from the South Tethyan margin. – Facies 34: 41–76.bernecker, M. (2005): Late Triassic reefs from the Northwest and South Tethys: distribution, setting and biotic composition. – Facies, 51: 442–453.laubenfelS, M. W., de (1955): Porifera. – In: R. C. Moore (Ed.), Treatise on Invertebrate Palaeontology, Part E, Archaeocyatha and Porifera; New York, NY & Lawrence, KS (Geological Society of America and University of Kansas Press), 21-112.Maurer, f., rettori, r. & Martini, r. (2008): Triassic stratigraphy, facies and evolution of the Arabian shelf in the northern United Arab Emirates. – International Journal of Earth Sciences, 97: 765–784.lévi, C. (1957): Sur une nouvelle classification des Démosponges. – Comptes Rendus des séances de l’Académie des Sciences de Paris, Série IIa, 236: 853–855.Pia, J.(1927): Thallophyta. – In: M. hirMer: Handbuch der Paläobo-tanik, vol. 1; Munich & Berlin (R. Oldenbourg), 31-136. Pickett, J. (2002): Order Heteractinida Hinde, 1887. – In: J. N. A. hooPer & R. W. M. van SoeSt (Eds), System Porifera: A Guide to the Classification of Sponges, vol. 2; Dordrecht (Kluwer Aca-demic/Plenum Publishers), 1121–1139. Pickett, J. w. & Jell, P. A. (1983): Middle Cambrian Sphinctozoa (Porifera) from New South Wales. – Memoirs of the Association of Australasian Palaeontologists, 1: 85–92.robertSon, a. h. f., bloMe, c. D., cooPer, D. w. J., keMP, a. e. S. & Searle, M. P. (1990): Evolution of the Arabian continental margin in the Dibba Zone, Northern Oman Mountains. – In: a. h. f. robertSon, M. P. Searle & a. c. rieS (Eds), The Geology and Tectonics of the Oman Region; London (The Geological Society), 3–27. Seilacher, A. (1962): Die Sphinctozoa, eine Gruppe fossiler Kalk-schwämme. – Akademie der Wissenschaften und der Literatur, Abhandlungen der mathematisch-naturwissenschaftlichen Klasse, 1961/10: 720–790.Senowbari-Daryan, B. (1990): Die systematische Stellung der tha-lamiden Schwämme und ihre Bedeutung in der Erdgeschichte. – Münchner Geowissenschatliche Abhandlungen, 21: 1–325.aragonitic mineralogy of rigid skeleton with spherulitic mi-crostructure. Spiculae lacking.Discussion:  The general appearance of the genus Mu-sandamia with ring-chambers and perforated chamber walls is similar to the genus Amblysiphonella. Both genera also have a spherulitic microstructure. The spongocoel of Ambly-siphonella is of retrosiphonate type, but in Musandamia it is clearly ambisiphonate. Sphinctozoans with ambisiphonate spongocoel types are known from the Jurassic-Cretaceous calcisponges Barroisia SteinMann or Spharocoelia SteinMann, as well as from the Cambrian heteractinid sponge genus Nucha Picket & Jell (Picket & Jell 1983; Picket 2002; see Senowbari-Daryan 1991; Senowbari-Daryan & Garcia belliDo 2002). Perforated thalamid sponges characterized by ambisiphonate spongocoels are not known among the possibly demospongid thalamid sponges from the younger Paleozoic and Triassic time interval. The only sponge with a possibly ambisiphonate type of spongocoel is the Triassic genus Enoplocoelia, but this genus is defined as aporate, in contrast to the porate Musandamia.      Musandamia omanica nov. sp.Pl. 1, Figs 2–5Derivatio nominis:  Occurrence of the sponge in Oman.Holotype:  Pl. 1, Figs 2 & 3, showing the same specimen.Locus typicus:  See Textfigure 1.Age:  Norian.Diagnosis :  Chambered sponges with spongocoel of am-bisiphonate type and perforated chamber walls. Chambers flat-tened with thick chamber exo- and interwalls. Ring-chambers connected with sponcocoel by a large opening.  Material :  One specimen; BSPG 2009 I 77 and 2009 I 78a & b.Description:  The sole specimen of this sponge is cut in a longitudinal section. The sections reveal that the organism is composed of numerous ring-like and moderately flattened chambers of rectangular appearance. The internally chambered construction, reflecting the outer segmentation, is clearly re-cognizable also from the outside. The diameter of the sponge, corresponding to the diameter of the chambers, is approxima-tely 12 mm. The height of the individual chambers is usually ~1 mm, but may reach a maximum of 1.3 mm. Chamber walls are perforated by the evenly distributed pores, each of which is approximately 0.2 mm in diameter. A spongocoel of the ambisiphonate type (sensu Seilacher 1962) with a diameter of 4 mm passes through the whole sponge. The communication of the chamber interior with the spongocoel is established by a large opening 0.6–1.0 mm in diameter. The chamber interior is devoid of a secondary skeleton and vesiculae. After obtaining a thin section, which is illustrated in Pl. 1, Fig. 3, the remains of the sponge were analysed under SEM. The investigation shows the spherulitic microstructure of the rigid skeleton. Spheru-727: 17–21.Senowbari-Daryan, b. & Garcia-belliDo, D. C. (2002): “Sphincto-zoa”: Chambered Sponges (Polyphyletic). – In: J. n. a. hooPer & r. w. M. van SoeSt (Eds), System Porifera. A Guide to the Classification of Sponges; Dordrecht (Kluwer Academic/Plenum Publishers), 1511–1533. Senowbari-Daryan, b. & Maurer, F. (2008): Upper Triassic (Norian hypercalcified sponges from the Musandam Peninsula (United Arab Emirates and Oman). – Facies, 54: 433–460. Senowbari-Daryan, b. & yancey, T. (2007): A sphinctozoan sponge from the Late Triassic (Norian) of the United Arab Emirates. –  Journal of Paleontology, 81: 1128–1131.SollaS, W. J. (1875): Sponges. – In: Encyclopedia Britanica, 9th edition; London, p. 451.SteinMann, G. (1882): Pharetronen-Studien. – Neues Jahrbuch für Mineralogie, Geologie und Paläontologie, 2: 139–191.verrill, A. E. (1907): Porifera of the Bermuda Islands. – New Haven, Connecticut, Transactions of the Academy of Arts and Sciences, 12: 330–344. Senowbari-Daryan, B. (1991): “Sphinctozoa”: An Overview. – In: J. reitner & h. keuPP (Eds), Fossil and Recent Sponges; Berlin (Springer), 224–241. Senowbari-Daryan, b. & bernecker, M. (1997): Barbafera carnica Senowbari-Daryan: Ein ungewöhnlicher Riffbildner aus dem Karn von Oman. – Geologische Blätter für Nordost-Bayern und angrenzende Gebiete, 47: 431–446.Senowbari-Daryan, b. & bernecker M. (2005): Crustacean coprolites of the Late Triassic Tethys: Biogeography from the Mediterranian to the Middle East. – Journal of Alpine Geology, 47: 137–144.Senowbari-Daryan, b. & bernecker M. (2009): Carnian (Ujpper Triassic) reef biota from limestone blocks of different tectonic settings in the Oman Mountains. – Beringeria, 40: 3–53.Senowbari-Daryan, bernecker, M., kryStyn, l. & Siblik, M. (1999): Carnian reef biota from a megabreccia of the Hawasina Complex (Al Aqida), Oman. – Rivista Italiana di Paleontologia e Stratigrafia, 105: 327-342.Senowbari-Daryan, b. & Di Stefano, P. (1988): Amblysiphonella maxima n. sp., a new sphinctozoan sponge from Upper Triassic reefs in Sicily. – Bollettino della Società Paleontologica Italiana, 

A sphinctozoan sponge from the Late Triassic (Norian) of the United Arab Emirates.

Amblysiphonella maxima n. sp., a new sphinctozoan sponge from Upper Triassic reefs in Sicily.

Barbafera carnica Senowbari-Daryan:

Carnian (Ujpper Triassic) reef biota from limestone blocks of different tectonic settings in the Oman Mountains.

Carnian reef biota from a megabreccia of the Hawasina Complex (Al Aqida),

Crustacean coprolites of the Late Triassic Tethys: Biogeography from the Mediterranian to the Middle East.

Die Sphinctozoa, eine Gruppe fossiler Kalkschwämme.

Die systematische Stellung der thalamiden Schwämme und ihre Bedeutung in der Erdgeschichte.

Evolution of the Arabian continental margin in the Dibba Zone,

Late Triassic reefs from the Northwest and South Tethys: distribution, setting and biotic composition.

Middle Cambrian Sphinctozoa (Porifera) from New South Wales.

One specimen; BSPG

Order Heteractinida Hinde,

Porifera of the Bermuda Islands.

Sphinctozoa”: An Overview. – In: J. reitner & h. keuPP (Eds), Fossil and Recent Sponges;

Sphinctozoa”: Chambered Sponges (Polyphyletic). – In:

Sponges. – In: Encyclopedia Britanica, 9th edition; London,

Sur une nouvelle classification des Démosponges.

System Porifera: A Guide to the Classification of Sponges,

Treatise on Invertebrate Palaeontology, Part E, Archaeocyatha and Porifera;

Triassic stratigraphy, facies and evolution of the Arabian shelf in the northern United Arab Emirates.

Upper Triassic (Norian hypercalcified sponges from the Musandam Peninsula (United Arab Emirates and Oman).

Upper Triassic reefs of the Oman Mountains: Data from the South Tethyan margin.

Amblysiphonella agahensis nov. sp., and Musandamia omanica nov. gen., nov. sp. (Porifera) from the Upper Triassic of Oman

Universität München: Elektronischen Publikationen

An International Journal  
of Palaeontology and Geobiology
Series A /Reihe A
Mitteilungen der Bayerischen Staatssammlung 
für Paläontologie und Geologie
München 2010
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50
Jubilee Volume50
Zitteliana
contents/InHALt
 
BaBa SenowBari-Daryan & Michaela Bernecker 
Amblysiphonella agahensis nov. sp., and Musandamia omanica nov. gen., nov. sp. 
(Porifera) from the Upper Triassic of Oman 3
alexanDer nützel 
A review of the Triassic gastropod genus Kittliconcha Bonarelli, 1927 – 
implications for the phylogeny of Caenogastropoda 9
anDrzej kaiM & Maria aleSSanDra conti
A problematic zygopleuroid gastropod Acanthostrophia revisited 21
Gernot arp
Ammonitenfauna und Stratigraphie des Grenzbereichs Jurensismergel/Opalinuston- 
Formation bei Neumarkt i.d. Opf. (oberstes Toarcium, Fränkische Alb) 25
Volker Dietze 
Über Ammonites Humphriesianus umbilicus QuenSteDt, 1886 an seiner 
Typus-Lokalität (östliche Schwäbische Alb, Südwestdeutschland) 55
Volker Dietze, Günter SchweiGert, GerD Dietl, wolfGanG auer, wolfGanG DanGelMaier,  
roGer furze, Stefan GräBenStein, Michael kutz, elMar neiSSer, erich SchneiDer & DietMar  
SchreiBer 
Rare Middle Jurassic ammonites of the families Erycitidae, Otoitidae and  
Stephanoceratidae from southern Germany 71
wolfGanG witt 
Late Miocene non-marine ostracods from the Lake Küçükçekmece region, Thrace (Turkey) 89 
jérôMe prieto 
Note on the morphological variability of Keramidomys thaleri (Eomyidae, 
Mammalia) from Puttenhausen (North Alpine Foreland Basin, Germany) 103
Martin pickforD 
Additions to the DehM collection of Siwalik hominoids, Pakistan: descriptions and interpretations 111
Michael krinGS, nora Dotzler, thoMaS n. taylor & jean Galtier 
Microfungi from the upper Visean (Mississippian) of central France: Structure  
and development of the sporocarp Mycocarpon cinctum nov. sp. 127
Zlatko kVaČek & joSef BoGner
Aracistrobus, an enigmatic non-araceous fossil from the Eurasian Oligocene and Miocene 137
Instructions for authors/Hinweise für Autoren 143
Zitteliana München, 30.06.2010 Issn 1612-412XA 50 143 seiten
An International Journal of Palaeontology and Geobiology
Series A/Reihe A
Mitteilungen der Bayerischen Staatssammlung für Paläontologie und Geologie
50
Editors-in-Chief/Herausgeber: Gert Wörheide, Michael Krings
Production and Layout/Bildbearbeitung und Layout: Martine Focke, Manuela Schellenberger
Bayerische Staatssammlung für Paläontologie und Geologie
Bayerische Staatssammlung für Paläontologie und Geologie
Richard-Wagner-Str. 10, D-80333 München, Deutschland
http://www.palmuc.de
email: zitteliana@lrz.uni-muenchen.de
Für den Inhalt der Arbeiten sind die Autoren allein verantwortlich.
Authors are solely responsible for the contents of their articles.
Copyright © 2010 Bayerische Staassammlung für Paläontologie und Geologie, München
Die in der Zitteliana veröffentlichten Arbeiten sind urheberrechtlich geschützt. 
Nachdruck, Vervielfältigungen auf photomechanischem, elektronischem oder anderem Wege 
sowie die Anfertigung von Übersetzungen oder die Nutzung in Vorträgen, für Funk und Fernsehen  
oder im Internet bleiben – auch auszugsweise – vorbehalten und bedürfen der schriftlichen Genehmigung  
durch die Bayerische Staatssammlung für Paläontologie und Geologie, München.
ISSN 1612-412X
Druck: Gebr. Geiselberger GmbH, Altötting
editorial Board
A. Altenbach, München
B.J. Axsmith, Mobile, AL
F.T. Fürsich, Erlangen
K. Heißig, München
H. Kerp, Münster
J. Kriwet, Stuttgart
J.H. Lipps, Berkeley, CA
T. Litt, Bonn
A. Nützel, München
O.W.M. Rauhut, München
B. Reichenbacher, München
J.W. Schopf, Los Angeles, CA
G. Schweigert, Stuttgart
F. Steininger, Eggenburg
cover illustrations: (from left to right) Shell of the gastropod Loxonema regium De koninck from the Carboniferous of Belgium 
(redrawn from De koninck 1881); Solitary coral Caninia sp. from the Carboniferous of England (redrawn from raMSBottoM in 
Mckerrow 1978); Tooth of the rare ruminant Orygotherium escheri Von Meyer from the Miocene of Germany (after röSSner 
& MörS 2001). Back cover: Atrium of the Munich Palaeontological Museum, view from the main entrance.
Umschlagbilder: (von links nach rechts) Gehäuse der Schnecke Loxonema regium De koninck aus dem Karbon von Belgien (neu 
gezeichnet nach De koninck 1881); Solitärkoralle Caninia sp. aus dem Karbon von England (neu gezeichnet nach raMSBottoM 
in Mckerrow 1978); Zahn des seltenen Wiederkäuers Orygotherium escheri Von Meyer aus dem Miozän von Deutschland (nach 
röSSner & MörS 2001). Rückseite: Lichthof des Paläontologischen Museums München, Blick vom Haupteingang.
3
Abstract 
Two chambered sponges – A. agahensis nov. sp. and 
Musandamia omanica nov. gen., nov. sp. – from the Norian 
reefal limestones of Jebel Agah, southeast of the Musandam 
Mountains, Oman, are described. Both species are the first 
reported from the Triassic of the Arabian Peninsula. The fossil 
association is also mentioned. 
Key words: Porifera, “Sphinctozoa”, Amblysiphonella, 
Musandamia, Norian, Jebel Agah, Oman.
Kurzfassung
Zwei gekammerte Schwämme – A. agahensis nov. sp. 
und Musandamia omanica nov. gen., nov. sp. – werden aus 
dem norischen Riffkalken des Jebel Agah, südöstlich des 
Musandam Gebirges (Oman), beschrieben. Beide Arten 
stellen die ersten Nachweise für die Trias der Arabischen 
Halbinsel dar. Die Vergesellschaftung der Fossilien wird 
ebenfalls erwähnt.
Schlüsselwörter: Porifera, “Sphinctozoa”, Amblysiph-
onella, Musandamia, Nor, Jebel Agah, Oman.
1. Introduction and geographic 
position of the locality 
Hypercalcified sponges, particularly the “sphinctozoans”, 
are the most abundant reef builders in Upper Triassic reefs 
and reefal deposits of Oman. Carnian sponges and other reef 
organisms from various localities in Oman were described by 
Senowbari-Daryan et al. (1999) and Senowbari-Daryan & 
bernecker (1997, 2009). Results from microfacies investiga-
tions of Norian-Rhaetian reefs of Oman and descriptions of 
the reef organisms were published by bernecker (1996, 2005). 
Senowbari-Daryan & yancey (2007) and Senowbari-Daryan 
& Maurer (2008) described the Norian sponges of Musandam 
Mountains, NW of the Jebel Agah, where A. agahensis nov. 
sp. and Musandamia omanica nov. gen. nov. sp., were found. 
Jebel Agah is located between the carbonate succession 
of the Musandam shelf to the northwest and the Semail 
Ophiolite to the southeast (Textfig. 1). Geology and tectonic 
development of this area is summarized by robertSon et al. 
(1990) and recently by Maurer et al. (2008). Microcoprolites 
of this locality were described by Senowbari-Daryan & 
bernecker (2005). 
The studied material, including two rock pieces and one 
thin section, is stored in the Bayerische Staatssammlung für 
Paläontologie und Geologie, Munich, under accession numbers 
BSPG 2009 I 77 and 2009 I 78a & b.
  
2. Systematic palaeontology
Class  Demospongea SollaS, 1875
Subclass  Ceractinomorpha levi, 1953
Order Agelasida verrill, 1907
Suborder Porata Seilacher, 1962
Family Sebargasiidae De laubenfelS, 1955
Subfamily Sebargasiinae Senowbari-Daryan, 1990
Genus Amblysiphonella SteinMann, 1882
Type species:  Amblysiphonella barroisi SteinMann, 1882.
Additional species included: See Senowbari-Daryan 
(1990) and Senowbari-Daryan & Garcia-belliDo (2002). 
Amblysiphonella agahensis nov. sp.
Pl. 1, Fig. 1
Derivat io nominis :  Occurrence of the species in Jebel 
Agah.
*Author for correspondence and reprint requests; E-mail: basendar@pal.uni-erlangen.de
Amblysiphonella agahensis nov. sp., and Musandamia omanica nov. gen., 
nov. sp. (Porifera) from the Upper Triassic of Oman
By
Baba Senowbari-Daryan1* & Michaela Bernecker2
1Geozentrum Nordbayern, Department of Palaeontology, University of Erlangen-Nürnberg, 
Loewenichstrasse 28, 91054 Erlangen, Germany
2German University of Technology, PO Box 1816, Athaibah PC 130 Muscat, Sultanate of Oman
Manuscript received November 11, 2009; revised manuscript accepted February 22, 2010
Zitteliana 3 - 7 1 Pl., 1 Textfig. München, 30.06.2010 ISSN 1612 - 412XA50
4
Holotype:  Pl. 1, Fig. 1.
Locus typicus:  Reefal limestones of Jebel Agah, south 
of Musandam Mountains (see Fig. 1).
Age:  Norian.
Diagnosis :  Multi-branched species of the genus Amblysi-
phonella with hemispherical to spherical chambers. Chamber 
walls pierced with single or branched pores. Wall of spongo-
coel contains large pores, which continue as small tubes into 
chamber interiors. Individual chambers clearly visible from 
outside of sponge. Chamber interiors whole. 
Material :  One multi-branched specimen (BSPG 2009 I 
77). 
 
Description: This dichotomously multi-branched sponge 
is composed of numerous hemispherical to spherical chambers. 
After increasing the chamber diameter from the initial stage, at 
least three individual branches originate from a large chamber. 
This large chamber reaches a diameter of approximately 50 
mm and a height of 13 mm. The diameter of the individual 
branches is ~30 mm, which corresponds to the diameter of the 
chambers. The chamber height varies between 8 and 12 mm. 
The chamber walls are about 1.4 mm thick, and are pierced 
by single or dichotomously branched pores of more or less 
equal distribution. They reach diameters of up to 0.6 mm. A 
spongocoel of 6-8 mm and assignable to the retrosiphonate 
type passes through the individual branches. The wall of the 
spongocoel has the same thickness as the chamber walls. The 
spongocoel wall is pierced by different large and rimmed pores 
up to 2 mm in diameter. Several large pores continue into the 
chamber interiors as small tubes. Chamber interiors are devoid 
of additional filling skeleton or vesiculae. 
Comparison:  Amblysiphonella agahensis nov. sp. repre-
sents the first Triassic species of the genus from the Arabian 
Peninsula. A. agahensis differs from all other Triassic species of 
that genus listed in Senowbari-Daryan & Di Stefano (1988) 
and Senowbari-Daryan (1990) by the branching mode of 
growth and the size of the chambers.   
Genus Musandamia nov. gen.
Derivat io nominis :  Named after the occurrence of the 
sponge in Musandam Mountains. 
D i a g n o s i s :  Amblysiphonella-like perforate thalamid 
sponge with spongocoel of ambisiphonate type. Primary 
Textfigure 1: Geological map of the area with the location of Jebel Agah. 
Plate 1: Amblysiphonella agahensis nov. sp. and Musandamia omanica nov. gen., nov. sp. from the Norian of Jebel Agah, Oman. Scale bars in 
Figs 1–3 = 10 mm. 
(1) Ambysiphonella agahensis nov. sp., holotype; section through several branches showing the hemispherical to spherical chambers. Several 
large pores of the spongocoel continue into the chamber interiors as small tubes (BSPG 2009 I 77). (2) Musandamia omanica nov. gen., nov. 
sp., holotype, longitudinal section showing the sponge with moderately flattened chambers with relatively thick chamber walls. The wall of the 
spongocoel is thin and pierced by a single large pore. The sponge serves as a substrate for an inozoan sponge. Polished rock surface (BSPG 2009 
I 78a). (3) Same as Pl. 1, Fig. 2., holotype specimen (thin section photograph in transmitted light) (BSPG 2009 I 78b). (4) Musandamia omanica 
nov. gen., nov. sp. SEM-microphotograph showing the spherulites of the rigid skeleton. (5) Similar to Pl. 1, Fig. 4.

6
lites are of different sizes, reaching diameters of up to 100 µm 
(Pl. 1, Figs 4–5). Spherulites indicate the primary aragonitic 
mineralogy of the skeleton. Spicules have not been observed.
Fossi l  associat ion:  The following organisms associated 
with the sponge were identified in one thin section: Heterastri-
dium sp., Microtubus communis flüGel, and chaetetid sponges 
gen. and sp. indet. The organisms are encrusted with microbial 
crusts (“Spongiostromata” sensu Pia in hirMer 1927). The 
index fossil Microtubus communis and the foraminifer Hirsuto-
spirella pilosa, illustrated by Senowbari-Daryan & bernecker 
(2005), are suggestive of a Norian-Rhaetian (most probably 
Norian) age of the investigated material and described sponges. 
Acknowledgements
Financial support was provided by the “Sonderfonds 
für wissenschaftliche Arbeiten an der Universität Erlangen-
Nürnberg” to B. Senowbari-Daryan. We thank Novus West 
Asia for granting permission to publish the species, and C. 
tollanD of Oolithica Geoscience (Cheltenham, UK) for the 
sampling. Helpful comments from reviewer M. noSe (Munich) 
are gratefully acknowledged.    
3. References 
bernecker, M. (1996): Upper Triassic reefs of the Oman Mountains: 
Data from the South Tethyan margin. – Facies 34: 41–76.
bernecker, M. (2005): Late Triassic reefs from the Northwest and 
South Tethys: distribution, setting and biotic composition. – 
Facies, 51: 442–453.
laubenfelS, M. W., de (1955): Porifera. – In: R. C. Moore (Ed.), 
Treatise on Invertebrate Palaeontology, Part E, Archaeocyatha and 
Porifera; New York, NY & Lawrence, KS (Geological Society of 
America and University of Kansas Press), 21-112.
Maurer, f., rettori, r. & Martini, r. (2008): Triassic stratigraphy, 
facies and evolution of the Arabian shelf in the northern United 
Arab Emirates. – International Journal of Earth Sciences, 97: 
765–784.
lévi, C. (1957): Sur une nouvelle classification des Démosponges. – 
Comptes Rendus des séances de l’Académie des Sciences de Paris, 
Série IIa, 236: 853–855.
Pia, J.(1927): Thallophyta. – In: M. hirMer: Handbuch der Paläobo-
tanik, vol. 1; Munich & Berlin (R. Oldenbourg), 31-136. 
Pickett, J. (2002): Order Heteractinida Hinde, 1887. – In: J. N. A. 
hooPer & R. W. M. van SoeSt (Eds), System Porifera: A Guide 
to the Classification of Sponges, vol. 2; Dordrecht (Kluwer Aca-
demic/Plenum Publishers), 1121–1139. 
Pickett, J. w. & Jell, P. A. (1983): Middle Cambrian Sphinctozoa 
(Porifera) from New South Wales. – Memoirs of the Association 
of Australasian Palaeontologists, 1: 85–92.
robertSon, a. h. f., bloMe, c. D., cooPer, D. w. J., keMP, a. e. 
S. & Searle, M. P. (1990): Evolution of the Arabian continental 
margin in the Dibba Zone, Northern Oman Mountains. – In: a. 
h. f. robertSon, M. P. Searle & a. c. rieS (Eds), The Geology 
and Tectonics of the Oman Region; London (The Geological 
Society), 3–27. 
Seilacher, A. (1962): Die Sphinctozoa, eine Gruppe fossiler Kalk-
schwämme. – Akademie der Wissenschaften und der Literatur, 
Abhandlungen der mathematisch-naturwissenschaftlichen Klasse, 
1961/10: 720–790.
Senowbari-Daryan, B. (1990): Die systematische Stellung der tha-
lamiden Schwämme und ihre Bedeutung in der Erdgeschichte. 
– Münchner Geowissenschatliche Abhandlungen, 21: 1–325.
aragonitic mineralogy of rigid skeleton with spherulitic mi-
crostructure. Spiculae lacking.
Discussion:  The general appearance of the genus Mu-
sandamia with ring-chambers and perforated chamber walls 
is similar to the genus Amblysiphonella. Both genera also 
have a spherulitic microstructure. The spongocoel of Ambly-
siphonella is of retrosiphonate type, but in Musandamia it is 
clearly ambisiphonate. Sphinctozoans with ambisiphonate 
spongocoel types are known from the Jurassic-Cretaceous 
calcisponges Barroisia SteinMann or Spharocoelia SteinMann, 
as well as from the Cambrian heteractinid sponge genus 
Nucha Picket & Jell (Picket & Jell 1983; Picket 2002; see 
Senowbari-Daryan 1991; Senowbari-Daryan & Garcia 
belliDo 2002). Perforated thalamid sponges characterized 
by ambisiphonate spongocoels are not known among the 
possibly demospongid thalamid sponges from the younger 
Paleozoic and Triassic time interval. The only sponge with 
a possibly ambisiphonate type of spongocoel is the Triassic 
genus Enoplocoelia, but this genus is defined as aporate, in 
contrast to the porate Musandamia.      
Musandamia omanica nov. sp.
Pl. 1, Figs 2–5
Derivatio nominis:  Occurrence of the sponge in Oman.
Holotype:  Pl. 1, Figs 2 & 3, showing the same specimen.
Locus typicus:  See Textfigure 1.
Age:  Norian.
Diagnosis :  Chambered sponges with spongocoel of am-
bisiphonate type and perforated chamber walls. Chambers flat-
tened with thick chamber exo- and interwalls. Ring-chambers 
connected with sponcocoel by a large opening.  
Material :  One specimen; BSPG 2009 I 77 and 2009 I 78a 
& b.
Description:  The sole specimen of this sponge is cut in 
a longitudinal section. The sections reveal that the organism 
is composed of numerous ring-like and moderately flattened 
chambers of rectangular appearance. The internally chambered 
construction, reflecting the outer segmentation, is clearly re-
cognizable also from the outside. The diameter of the sponge, 
corresponding to the diameter of the chambers, is approxima-
tely 12 mm. The height of the individual chambers is usually 
~1 mm, but may reach a maximum of 1.3 mm. Chamber walls 
are perforated by the evenly distributed pores, each of which 
is approximately 0.2 mm in diameter. A spongocoel of the 
ambisiphonate type (sensu Seilacher 1962) with a diameter of 
4 mm passes through the whole sponge. The communication 
of the chamber interior with the spongocoel is established by a 
large opening 0.6–1.0 mm in diameter. The chamber interior is 
devoid of a secondary skeleton and vesiculae. After obtaining a 
thin section, which is illustrated in Pl. 1, Fig. 3, the remains of 
the sponge were analysed under SEM. The investigation shows 
the spherulitic microstructure of the rigid skeleton. Spheru-
7
27: 17–21.
Senowbari-Daryan, b. & Garcia-belliDo, D. C. (2002): “Sphincto-
zoa”: Chambered Sponges (Polyphyletic). – In: J. n. a. hooPer 
& r. w. M. van SoeSt (Eds), System Porifera. A Guide to the 
Classification of Sponges; Dordrecht (Kluwer Academic/Plenum 
Publishers), 1511–1533. 
Senowbari-Daryan, b. & Maurer, F. (2008): Upper Triassic (Norian 
hypercalcified sponges from the Musandam Peninsula (United 
Arab Emirates and Oman). – Facies, 54: 433–460. 
Senowbari-Daryan, b. & yancey, T. (2007): A sphinctozoan sponge 
from the Late Triassic (Norian) of the United Arab Emirates. 
–  Journal of Paleontology, 81: 1128–1131.
SollaS, W. J. (1875): Sponges. – In: Encyclopedia Britanica, 9th edition; 
London, p. 451.
SteinMann, G. (1882): Pharetronen-Studien. – Neues Jahrbuch für 
Mineralogie, Geologie und Paläontologie, 2: 139–191.
verrill, A. E. (1907): Porifera of the Bermuda Islands. – New Haven, 
Connecticut, Transactions of the Academy of Arts and Sciences, 
12: 330–344. 
Senowbari-Daryan, B. (1991): “Sphinctozoa”: An Overview. – In: 
J. reitner & h. keuPP (Eds), Fossil and Recent Sponges; Berlin 
(Springer), 224–241. 
Senowbari-Daryan, b. & bernecker, M. (1997): Barbafera carnica 
Senowbari-Daryan: Ein ungewöhnlicher Riffbildner aus dem 
Karn von Oman. – Geologische Blätter für Nordost-Bayern und 
angrenzende Gebiete, 47: 431–446.
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of the Late Triassic Tethys: Biogeography from the Mediterranian 
to the Middle East. – Journal of Alpine Geology, 47: 137–144.
Senowbari-Daryan, b. & bernecker M. (2009): Carnian (Ujpper 
Triassic) reef biota from limestone blocks of different tectonic 
settings in the Oman Mountains. – Beringeria, 40: 3–53.
Senowbari-Daryan, bernecker, M., kryStyn, l. & Siblik, M. (1999): 
Carnian reef biota from a megabreccia of the Hawasina Complex 
(Al Aqida), Oman. – Rivista Italiana di Paleontologia e Stratigrafia, 
105: 327-342.
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maxima n. sp., a new sphinctozoan sponge from Upper Triassic 
reefs in Sicily. – Bollettino della Società Paleontologica Italiana, 


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Amblysiphonella agahensis nov. sp., and Musandamia omanica nov. gen., nov. sp. (Porifera) from the Upper Triassic of Oman

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