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Rhabdomesid and cystoporid bryozoans from the Permian of Deh-e Mohammad, Shotori Mountains (northeastern Iran)
Geobios 42 (2009) 133–140
Original article
Rhabdomesid and cystoporid bryozoans from the Permian of Deh-e Mohammad, Shotori Mountains (northeastern Iran)§ Bryozoaires rhabdome´sides et cystoporides du Permien de Deh-e Mohammad, Montagnes Shotori (nord-est de l’Iran) Andrej Ernst a,*, Baba Senowbari-Daryan b, Koorosch Rashidi c a
Institut für Geowissenschaften der Christian-Albrechts, Universität zu Kiel, Ludewig-Meyn-Strasse 10, 24118 Kiel, Germany b Institut für Paläontologie, Universität Erlangen–Nürnberg, Loewenichstrasse 28, 91054 Erlangen, Germany c University of Payam-e Noor, Ardakan, Iran Received 19 January 2007; accepted 23 September 2008 Available online 11 November 2008
Abstract Seven bryozoan species belonging to the Order Rhabdomesida and Order Cystoporida are described from the Permian deposits exposed near the small town of Deh-e Mohammad, Shotori Mountains (northeastern Iran): Rhabdomeson cf. consimile Bassler, Pamirella nitida Gorjunova, Clausotrypa conferta Bassler, Streblotrypa (Streblotrypa) elegans Sakagami, Streblotrypa (Streblascopora) supernodata nov. sp., Cystodictya sp., and Filiramoporina cf. kretaphilia Fry and Cuffey. The described fauna identifies the age of the Jamal Formation at the locality near Deh-e Mohammad as Lower Permian. It displays palaeobiogeographic connections to the Lower Permian of Pamir (Tajikistan), Indonesia, Thailand and Kansas (North America). # 2008 Elsevier Masson SAS. All rights reserved. Résumé Sept espèces de bryozoaires qui appartiennent à l’Ordre Rhabdomesida et à l’Ordre Cystoporida sont décrites dans les dépôts permiens affleurant près de la petite ville de Deh-e Mohammad, Montagnes Shotori (nord-est de l’Iran) : Rhabdomeson cf. consimile Bassler, Pamirella nitida Gorjunova, Clausotrypa conferta Bassler, Streblotrypa (Streblotrypa) elegans Sakagami, Streblotrypa (Streblascopora) supernodata nov. sp., Cystodictya sp., et Filiramoporina cf. kretaphilia Fry et Cuffey. La faune décrite attribue un âge permien inférieur à la Formation Jamal, au niveau de la localité proche de Deh-e Mohammad. Elle affiche des relations paléobiogéographiques avec le Permien inférieur du Pamir (Tadjikistan), du Timor (Indonésie), de la Thaïlande et du Kansas (Amérique du Nord). # 2008 Elsevier Masson SAS. Tous droits réservés. Keywords: Bryozoa; Taxonomy; Lower Permian; Palaeobiogeography Mots clés : Bryozoa ; Taxonomie ; Permien inférieur ; Paléobiogéographie
1. Abbreviations The following abbreviations are used in measurements: Statistics:
N X SD CV MIN MAX
Measured characters (in mm):
§
Corresponding editor: Jean Vannier. * Corresponding author. E-mail address: [email protected] (A. Ernst).
number of measurements mean standard deviation coefficient of variation minimal value maximal value
AW
0016-6995/$ – see front matter # 2008 Elsevier Masson SAS. All rights reserved. doi:10.1016/j.geobios.2008.09.001
width of autozooecial aperture
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ADB WB AB ND AcD
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distance between aperture centres along branch branch width axial bundle diameter node diameter acanthostyle diameter
2. Introduction Bryozoans are abundant in Permian deposits in Iran, but they are poorly investigated. The earliest work, treating the Permian bryozoans of Iran, is published by Fantini Sestini (1965). This publication includes the description of three species from the Ruteh Limestone of Central Elborz (northern Iran). Later, Sakagami (1980) described 15 species from the Permian of central Iran (Abadeh region). This fauna displayed a close relation to the Dzhulfian (= Changhsingian) stage, as well as connections to the Permian of Pamir, Russian platform, Salt Range, Kashmir, South China, and Timor. Recently, Ernst et al. (2006a, 2006b) described two bryozoan faunal associations from the Jamal Formation, Iran. Thirty-one bryozoan species were identified from the Jamal Formation, exposed in Lakaftari area, south of the town of Bagher-Abad, northeast of Esfahan (central Iran). This association is regarded as Middle Permian (Murgabian) in age, and shows affinities to Australia, Indonesia, Thailand and Transcaucasia. Six bryozoan species were described from the Jamal Formation, exposed in Bagh-e Vang (Shotori Mountains, northeast Iran). This fauna indicates a Lower Permian age and shows palaeogeographic affinities to the bryozoan fauna known from Western Australia and Urals. The new bryozoan material, collected from the Jamal Formation exposed near the town of Deh-e Mohammad, Shotori Mountains (northeastern Iran), is described in the present publication. The bryozoans of this locality exhibit palaeogeographic connections to the Lower Permian of Pamir (Tajikistan), Indonesia, Thailand, and Kansas (North America). 3. Lithology and stratigraphy The material was collected from a locality situated northeast of the village Deh-e Mohammad in Shotori Mountains, northeastern Iran (Fig. 1). Permian rocks at this locality disconformably overlie the Upper Carboniferous Sardar Formation (Fig. 2). The Sardar Formation is composed of sandstones, sandy marls and light green shales. The Permian at this locality is represented by the Jamal Formation that contains at its base bioclastic limestones, succeeded by pure limestones, marly limestones with abundant Tubiphytes, algae, some sponges and echinoids, followed by conglomerates and breccias with rare sponges. The sphinctozoan sponge Amblysiphonella cf. vesiculosa vesiculosa (De Koninck) was described from this locality by Senowbari-Daryan et al. (2006). This part of the section is interpreted to be a reef mound. The middle part of the section consists of medium to thick bedded grey limestones with intercalated cherts and also Tubiphytes, small foraminifers, and a few bryozoans. The upper part of the section contains dolomites and cherts lacking fossils.
Fig. 1. Map showing the geographic position of the Permian locality near the town of Deh-e Mohammad.
4. Material and methods Taxonomic descriptions were based on thin sections. The rock materials, as well as the thin sections, are housed at the Institute of Paleontology, University of Erlangen–Nürnberg, Germany (Senowbari-Daryan, Permian, Deh-e Mohammad).
A. Ernst et al. / Geobios 42 (2009) 133–140
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Fig. 2. Lithostratigraphical column of the Permian deposits near the town of Deh-e Mohammad.
5. Systematic palaeontology Phylum BRYOZOA Ehrenberg, 1831 Class STENOLAEMATA Borg, 1926 Order RHABDOMESIDA Astrova and Morozova, 1956 Family RHABDOMESIDAE Vine, 1884 Genus Rhabdomeson Young and Young, 1874 [= Coeloconus Ulrich, 1889] Type species: Rhabdomeson progracile Wyse Jackson and Bancroft, 1995, by subsequent designation by Wyse Jackson
and Bancroft, 1995 (ICZN Opinion 1874); Lower Carboniferous, England. Rhabdomeson consimile Bassler, 1929 Fig. 3(A, B) 1929. Rhabdomeson consimile Bassler, p. 69, Pl. 13, Figs. 5–8. 1976. Rhabdomeson cf. consimile Bassler - Liu, p. 152, Pl. 75, Fig. 7. 1981. Rhabdomeson consimile Bassler - Yang, Lu and Xia, p. 93, Pl. 4, Fig. 7.
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Fig. 3. A–B. Rhabdomeson consimile Bassler, 1929, D1-6. A. Oblique thin section of the branch. B. Tangential section of the branch. C–E. Pamirella nitida Gorjunova, 1975. C. Oblique section of the branch, D1. D. Longitudinal section of the branch, displaying acanthostyles, D1. E. Tangential section of the branch, displaying autozooecial apertures and acanthostyles, D4. F–H. Clausotrypa conferta Bassler, 1929. F. Cross section of the branch, D4. G. Oblique section of the branch, D11. H. Oblique tangential section of the branch displaying autozooecial apertures, D11. I, J. Streblotrypa (Streblotrypa) elegans Sakagami, 1970. I. Cross section of the branch, D1. J. Oblique section of the branch, D1-6. K–N. Streblotrypa (Streblascopora) supernodata nov. sp. K. Longitudinal section of the branch,
A. Ernst et al. / Geobios 42 (2009) 133–140
1982. Rhabdomeson consimile Bassler - Kiseleva, p. 87, Pl. 16, Fig. 3. 1983. Rhabdomeson consimile Bassler - Yang and Lu, pp. 285–286, Pl. 3, Figs. 1–3. 1988. Rhabdomeson cf. consimile Bassler - Sakagami in Yanagida, Pl. 19, Figs. 5, 6, Pl. 20, Figs. 4, 6, 7. 1999. Rhabdomeson consimile Bassler - Sakagami, p. 91, Pl. 22, Fig. 6, Pl. 23, Figs. 1, 2. Material: Single specimen D1–6. Description: Ramose colony with small axial cylinder and distinct exozone, 1.23 mm in diameter. Axial cylinder 0.29 mm in diameter. Autozooecia budding in spiral pattern around axial cylinder, rhomboidal in cross-section. Autozooecial apertures oval, 0.11–0.17 mm wide, arranged in regular diagonal rows, spaced 0.24–0.28 mm from centre to centre, four longitudinally and eight diagonally in 2 mm. Single macroacanthostyle situated in interspaces between apertures, 0.036–0.048 mm in diameter. Microacanthostyles 0.02–0.0.03 mm in diameter, arranged in one row between macroacanthostyles. Superior hemiseptum blunt, obscure, positioned distally; inferior hemiseptum absent. Autozooecial walls 0.024–0.036 mm thick in endozone, 0.11–0.12 mm thick in exozone. Remarks: Rhabdomeson consimile Bassler, 1929 is similar to R. mammillatum (Bretnall, 1926) from the Lower Permian of Australia, but differs from the latter in presence of microacanthostyles and hemisepta. It differs also from R. bispinosum Crockford, 1944, from the Lower Permian of Western Australia, in having hemisepta and single instead of paired acanthostyles between apertures. Occurrence: Upper Permian; Indonesia. Upper Permian; Khabarovsk region (Russia). Lower to Upper Permian (? Artinskian to Guadalupian); Thailand. Permian; Mongolia. Permian; China (Western Xinjiang). Jamal Formation, Lower Permian; Deh-e Mohammad, Shotori Mountains, northeastern Iran. Family RHOMBOPORIDAE Simpson, 1895 Genus Pamirella Gorjunova, 1975 Type species: P. nitida Gorjunova, 1975, by original designation; Lower Permian (Artinskian), SE Pamir (Tajikistan). Pamirella nitida Gorjunova, 1975 Fig. 3(C–E) and Table 1 1975. Pamirella nitida - Gorjunova, p. 63, Pl. 10, Fig. 1. 1983. Pamirella nitida Gorjunova - Blake, p. 580, Fig. 288 (1a–d). 1985. Pamirella nitida Gorjunova - Gorjunova, p. 119, Pl. 8, Fig. 4. 1986. Pamirella nitida Gorjunova - Xia, p. 208, Pl. 1, Figs. 1–6.
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Table 1 Measurements of Pamirella nitida Gorjunova, 1975 (see explanations of abbreviations in the text).
AW ADB AcD
N
X
SD
CV
MIN
MAX
6 4 12
0.14 0.64 0.07
0.016 0.038 0.008
11.058 5.921 11.832
0.13 0.60 0.06
0.17 0.69 0.08
Material: D1, D4. Description: Ramose cylindrical colonies, 1.26 mm in diameter. Autozooecia tubular, growing in spiral pattern. Autozooecial diaphragms absent. Apertures oval, arranged in diagonal rows on the colony surface. Metazooecia rare, 0.02 mm in diameter. Acanthostyles abundant, 7-8 surrounding each autozooecial aperture. Microacanthostyles abundant. Remarks: Pamirella nitida Gorjunova, 1975 differs from a similar species P. pulchra (Bassler, 1929) from the Lower Permian (Artinskian) of Indonesia in having thinner branches (1.26 mm versus 2.50–3.00 mm in P. pulchra) and smaller apertures (0.13–0.17 mm versus 0.35 mm in P. pulchra; data from Bassler, 1929). Pamirella nitida is also similar to P. nepalensis Sakagami and Sakai, 1991, known from the upper part of Sisne Formation of Nepal Lesser Himalayas, but differs from it in having smaller apertures (0.13–0.17 mm versus 0.30 mm in P. nepalensis). Occurrence: Lower Permian (Artinskian); SE Pamir, Tajikistan. Angjie Formation (Lower Permian); Xainza, Northern Xizang, China. Jamal Formation, Lower Permian; Deh-e Mohammad, Shotori Mountains, northeastern Iran. Family NEMATOTRYPIDAE Spjeldnaes, 1984 Genus Clausotrypa Bassler, 1929 Type species: C. separata Bassler, 1929, by original designation; Lower Permian, Timor, Indonesia. Clausotrypa conferta Bassler, 1929 Fig. 3(F–H) and Table 2 1929. Clausotrypa conferta Bassler, p. 72, Pl. 14, Figs. 1–3. 1988. Clausotrypa sp. indet. - Sakagami in Yanagida, Pl. 19, Fig. 4. 1999. Clausotrypa conferta Bassler - Sakagami, p. 92, Pl. 23, Fig. 3. 2000. Clausotrypa sp. cf. C. conferta Bassler - Sakagami, p. 161, Fig. 9-2. Material: D1, D11. Description: Cylindrical branches, 1.44–1.80 mm in diameter. Autozooecia long, cylindrical, growing parallel to the branch axis for a long distance, turning gently to the colony surface. Autozooecial apertures oval, spaced 5–6 longitudinally in 2 mm. Diaphragms not observed. Tectitozooecia numerous, restricted mostly to exozone, in 1–3 rows separating
holotype D1–2. L. Tangential section of the branch, displaying autozooecial apertures and nodes, paratype D11. M. Oblique section of the branch displaying axial bundle of zooecia, holotype D1–2. N. Cross section of the branch, holotype D1–2. O, P. Cystodictya sp. O. Tangential section of the branch, D36. P. Oblique section of the branch, D36. Q. Filiramoporina cf. kretaphilia Fry and Cuffey, 1976, cross section of the branch, D100.
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Table 2 Measurements of Clausotrypa conferta Bassler, 1929 (see explanations of abbreviations in the text).
AW ADB
N
X
SD
CV
MIN
MAX
10 6
0.13 0.36
0.007 0.023
5.128 6.393
0.12 0.34
0.14 0.40
autozooecial apertures. Acanthostyles 0.02–0.03 mm in diameter, abundant, originating in the late endozone, having clear cores. Remarks: Clausotrypa conferta Bassler, 1929 differs from C. monticola (Eichwald, 1860) by having smaller acanthostyles and larger apertures (acanthostyle diameter 0.02–0.03 mm versus 0.02–0.08 mm in C. monticola, and aperture width 0.12– 0.14 mm versus 0.10–0.12 mm in C. monticola, data from Morozova and Krutchinina, 1986). Occurrence: Lower–Middle Permian; Indonesia. Permian; Thailand. Jamal Formation, Lower Permian; Deh-e Mohammad, Shotori Mountains, northeastern Iran. Suborder HYPHASMOPORIDAE Vine, 1886 Family HYPHASMOPORIDAE Vine, 1886 Genus Streblotrypa Vine, 1885 Subgenus Streblotrypa (Streblotrypa) Vine, 1885 Type species: Streblotrypa nicklesi Vine, 1885, by original designation; Middle Carboniferous, England. Streblotrypa (Streblotrypa) elegans Sakagami, 1970 Fig. 3(I, J) 1970. Streblotrypa elegans - Sakagami, pp. 65–66, Pl. 13, Figs. 5–9. 1976. Streblotrypa elegans Sakagami - Sakagami, Pl. 27, Fig. 4. 2000. Streblotrypa elegans Sakagami - Sakagami, p. 153, Figs. 7, 6. Material: D1, D1–6. Description: Ramose branching colonies, 0.66–0.90 mm in diameter. Branch cross sections rounded-polygonal in shape. Autozooecia tubular, with long thin bases, forming a distinct axial bundle, rounded to slightly polygonal in cross section. Axial bundle 0.22–0.51 mm in diameter. Autozooecial apertures oval, arranged in regular diagonal rows, 0.066–0.090 mm in diameter (0.075 mm, average), spaced 5 in 2 mm longitudinally, having distinct peristomes. Peristomal wall 0.02 mm thick. Short superior hemiseptum present; inferior hemiseptum very long, positioned near the place of bending outwards of the autozooecia. Abundant metazooecia between apertures, arranged in 4-5 rows. Remarks: Streblotrypa (Streblotrypa) elegans Sakagami, 1970 is similar to S. (S.) parva Morozova, 1965, known from the Upper Permian of Caucasus. However, S. (S.) elegans differs from S. (S.) parva by having long hemisepta. Occurrence: Lower Permian (?Lower Artinskian); Thailand. Jamal Formation, Lower Permian; Deh-e Mohammad, Shotori Mountains, northeastern Iran.
Table 3 Measurements of Streblotrypa (Streblascopora) supernodata nov. sp. (see explanations of abbreviations in the text).
WB AB AW ND
N
X
SD
CV
MIN
MAX
7 3 5 4
0.90 0.25 0.09 0.055
0.155 0.044 0.011 0.012
16.783 17.436 12.087 21.477
0.75 0.22 0.07 0.050
1.23 0.30 0.10 0.070
S. (Streblotrypa) elegans has been recently discovered in the Permian of Oman (Ernst et al., 2008: 692, Figs. 8.1–8.5). Subgenus Streblotrypa (Streblascopora) Bassler, 1929 Type species: Streblotrypa fasciculata Bassler, 1929, by original designation; Upper Permian, Indonesia. Streblotrypa (Streblascopora) supernodata nov. sp. Fig. 3(K–N) and Table 3 Etymology: The species name refers to large nodes between apertures. Holotype: D1–2 (Fig. 3K, M–N). Paratypes: D11 (Fig. 3L) and D116. Type locality: Deh-e Mohammad, Shotori Mountains (northeastern Iran [Fig. 1]). Type horizon: Jamal Formation, Lower Permian. Diagnosis: Ramose branched colonies; axial bundle with 11–13 axial zooecia arranged in 3-4 rows; inferior hemisepta long, superior hemisepta short; 3-4 large nodes surrounding each autozooecial aperture. Description: Ramose branched colonies, 0.75–1.23 mm in diameter. Axial bundle 0.22–0.30 mm in diameter, containing 11–12 axial zooecia arranged in 3–4 rows of zooecia. Branch cross sections rounded-polygonal in shape. Autozooecia tubular, growing from the axial bundle, Autozooecial apertures oval, arranged in regular diagonal rows. Inferior hemisepta long, superior hemisepta short. Diaphragms present, rare to common, thin, curved proximally. Metazooecia abundant, 5–7 arranged in 2–3 rows between apertures, 0.02–0.04 mm in diameter. Massive nodes between apertures; 3–4 surrounding each aperture. Remarks: Streblotrypa (Streblascopora) supernodata nov. sp. is similar to S. (S.) irianica Sakagami, 2000 from the Middle Permian of Irian Jaya, Indonesia, in having nodes, but differs from the latter in having thinner branches (0.75–1.23 mm versus 2.6–3.0 mm in S. (S.) irianica) and smaller autozooecial apertures (0.10–0.17 mm versus 0.15–0.19 mm in S. (S.) irianica). Order CYSTOPORIDA Astrova, 1964 Family CYSTODICTYONIDAE Ulrich, 1884 Genus Cystodictya Ulrich, 1882 Type species: C. ocellata Ulrich, 1882, by original designation; Lower Mississippian, Kentucky, USA. Cystodictya sp. Fig. 3(O, P) and Table 4
A. Ernst et al. / Geobios 42 (2009) 133–140 Table 4 Measurements of Cystodictya sp. (see explanations of abbreviations in the text).
AW ADB
N
X
SD
CV
MIN
MAX
5 6
0.15 0.62
0.012 0.059
8.333 9.515
0.13 0.57
0.16 0.72
Material: Two fragments D36. Description: Narrow bifoliate branches. Autozooecia short, growing from thin mesotheca in 2-3 rows on each side. Diaphragms not observed. Lunaria absent; distinct peristomes around autozooecial apertures. Few small vesicles. Hemisepta absent. Remarks: The present material is similar to Cystodictya absoluta Morozova and Lisitsyn, 2002 from the Upper Carboniferous (Gzhelian) of the Russian Plate. The latter species has four autozooecial rows instead of 2–3 in the present material, as well as wider and more widely spaced apertures (aperture width 0.16–0.18 mm versus 0.13–0.16 mm in present material; aperture spacing along branch 0.80–0.82 mm versus 0.57–0.72 mm in present material). Occurrence: Jamal Formation, Lower Permian; Deh-e Mohammad, Shotori Mountains, northeastern Iran. Genus Filiramoporina Fry and Cuffey, 1976 Type species: Filiramoporina kretaphilia Fry and Cuffey, 1976, by original designation; Lower Permian (Wreford Megacyclothem), Kansas, USA. Filiramoporina cf. kretaphilia Fry and Cuffey, 1976 Fig. 3(Q) Compare: 1976. Filiramoporina kretaphilia Fry and Cuffey, pp. 4–7, Pl. 1, Figs. 1–9, Pl. 2, Figs. 1–8. Material: Single specimen D100. Description: Bifoliate branch 2.42 mm wide and 1.65 mm thick, oval in cross section. Autozooecia arranged in eight rows on both sides of colony surface, growing from a straight mesotheca, trapezoid at their bases. Extrazooidal skeleton consisting of structureless material. Tangential and longitudinal sections are missing. Remarks: The Iranian species is similar to Filiramoporina kretaphilia Fry and Cuffey, 1976, the only known species of the genus, in branch size, autozooecial shape and the kind of extrazooidal skeleton (Roger Cuffey, pers. comm., 2006). However, the available material is insufficient to make a closer determination. Occurrence: Wreford Megacyclothem, Lower Permian; Kansas, USA. Jamal Formation, Lower Permian; Deh-e Mohammad, Shotori Mountains, northeastern Iran. 6. Discussion Bryozoa show a high potential for palaeobiogeographic purposes in the Palaeozoic. Permian bryozoan faunas were described from different regions and can be used for a palaeobiogeographic comparison. The bryozoan fauna from the
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Permian of Deh-e Mohammad locality described in the present paper displays connections predominantly to the Lower Permian of Thailand and Indonesia (Rhabdomeson consimile Bassler, 1929, Clausotrypa conferta Bassler, 1929, Streblotrypa (Streblotrypa) elegans Sakagami, 1970) and Pamir, Tajikistan (Pamirella nitida Gorjunova, 1975). Of note is the finding of Filiramoporina cf. kretaphilia Fry and Cuffey, 1976. The genus Filiramoporina Fry and Cuffey, 1976, with a single known species, was described from the Lower Permian of Kansas, USA. Acknowledgements The material for the present investigation was sampled by one of the authors (K.R.) during his Ph.D. thesis. The investigations were financially supported by the Deutsche Forschungsgemeinschaft (DFG) to B. Senowbari-Daryan (Project Se 416/10). K. Rashidi thanks the scientific ministry of Iran for financial support to stay 6 months in Erlangen. He thanks also the staff and members of the Institute of Paleontology (Erlangen) for their kind assistance. We also thank J.R.P. Ross, Bellingham for her kind review of the manuscript. References Astrova, G.G., 1964. Novyi otryad paleozoiskikh mshanok [New Order of Paleozoic Bryozoa]. Paleontologicheskij zhurnal 1, 22–31 (in Russian). Astrova, G.G., Morozova, I.P., 1956. K sistematike mshanok otryda Cryptostomata [About systematics of the Order Cryptostomata]. Doklady Akademii Nauk SSSR 110, 661–664 (in Russian). Bassler, R.S., 1929. The Permian Bryozoa of Timor. Paläontologie von Timor 16, 37–90. Blake, D.B., 1983. Systematic descriptions for the Suborder Rhabdomesina. In: Robison, R.A. (Ed.), Treatise on Invertebrate Paleontology, Part G (1): Bryozoa (revised). Geological Society of America and University of Kansas Press, Lawrence, pp. 551–592. Borg, F., 1926. Studies on recent cyclostomatous Bryozoa. Zoologiska Bidrag fran Uppsala 10, 181–507. Bretnall, R.W., 1926. Description of some western Australian fossil Polyzoa. Bulletin of Western Australia Geological Survey 88, 7–33. Crockford, J., 1944. Bryozoa from the Permian of Western Australia. Part I. Cyclostomata and Cryptostomata from the northwest basin and Kimberly district. Proceedings of the Linnean Society of New South Wales 69, 139– 175. Eichwald, E. von, 1860. Lethaea rossica, ou Paléontologie de la Russie. I, Seconde section de l’ancienne période. Bryozoaires, Schweizerbart, Stuttgart, pp. 355–494. Ehrenberg, C.G., 1831. Symbolae Physicae, seu Icones et descriptiones Corporum Naturalium novorum aut minus cognitorum, quae ex itineribus per Libyam, Aegiptum, Nubiam, Dongalaam, Syriam, Arabiam et Habessiniam, studia annis 1820–1825, redirent. Pars Zoologica, 4, Animalia Evertebrata exclusis Insectis. Berolini. Ernst, A., Senowbari-Daryan, B., Hamedani, A., 2006a. Middle Permian Bryozoa from the Lakaftari area, central Iran. Geodiversitas 28, 543– 590. Ernst, A., Senowbari-Daryan, B., Rashidi, K., 2006b. Lower Permian Bryozoa of the Jamal Formation from Bagh-e Vang (Shotori Mountains, northeast Iran). Facies 52, 627–635. Ernst, A., Weidlich, O., Schäfer, P., 2008. Stenolaemate Bryozoa from the Permian of Oman (Aseelah Unit, Batain coast). Journal of Paleontology 82, 676–716.
140
A. Ernst et al. / Geobios 42 (2009) 133–140
Fantini Sestini, F.N., 1965. The geology of the upper Djadjerud and Lar valleys (North Iran); II, Palaeontology; Bryozoans, brachiopods and molluscs from Ruteh limestone (Permian). Revista Italiana di Paleontologia 71, 13–108. Fry, H.C., Cuffey, R.J., 1976. Filiramoporina kretaphilia: a new genus and species of bifoliate tubulobryozoan (Ectoprocta) from the lower Permian Wreford Megacyclothem of Kansas. The University of Kansas Paleontological Contributions Paper 84, 1–12. Gorjunova, R.V., 1975. Permskie mshanki Pamira [Permian bryozoans of the Pamir]. Trudy paleontologicheskogo instituta Akademii Nauk SSSR 148, 1–127 (in Russian). Gorjunova, R.V., 1985. Morfologia, sistema I filogeniya mshanok (Otryad Rhabdomesida) [Morphology, system and phylogeny of Bryozoa (Order Rhabdomesida)]. Trudy paleontologicheskogo instituta Akademii Nauk SSSR 208, 1–152 (in Russian). Kiseleva, A.B., 1982. Pozdnepermskie mshanki Yuzhnogo Primorya [Late Permian bryozoans of the South Primorye]. Nauka 1–128 (in Russian). Liu, X.L., 1976. Phylum Bryozoa. In: Palaeontological Atlas of north China. Inner Mongolia], Vol. 1. Palaeozoic Volume. Geological Publishing House, Peking, pp. 131–155 (in Chinese). Morozova, I.P., 1965. Bryozoa. In: Ruzhentsev, V.E., Sarycheva, T.G. (Eds.), The evolution and changes in the marine organisms at the Paleozoic– Mesozoic boundary. Trudy paleontologicheskogo instituta Akademii Nauk SSSR, 108pp. 183–197 (in Russian). Morozova, I.P., Krutchinina, O.N., 1986. Permian Bryozoans of the Arctis (Western Sector). Nauka 1–141 (in Russian). Morozova, I.P., Lisitsyn, D.V., 2002. Revision of bryozoans from the stratotype sections of the Gzhelian Stage (Upper Carboniferous). Paleontologicheskij zhurnal 6, 63–72 (in Russian). Sakagami, S., 1970. Addition to the Permian Bryozoa from Ko Muk, Peninsular Thailand. Contributions to the Geology and Palaeontology of Southeast Asia. 53. Geology and Palaeontology of Southeast Asia. Contributions to the geology and palaeontology of Southeast Asia 8, 43–68. Sakagami, S., 1976. On the Permian Bryozoa from the northern part of Sainbeyli, Central Turkey. Transactions and Proceedings of the Palaeontological Society of Japan, New Series 103, 398–405. Sakagami, S., 1980. Permian Ectoprocta (Bryozoa) from the Abadeh Region, Central Iran. Transactions and Proceedings of the Palaeontological Society of Japan. New Series 118, 269–289. Sakagami, S., 1999. Permian bryozoans from some localities in the Khao Hin Kling Area near Phetchabun, north-central Thailand. Bulletin of the Kitakyushi Museum of Natural History 18, 77–103. Sakagami, S., 2000. Middle Permian Bryozoa from Irian Jaya, Indonesia. Bulletin of the National Science Museum, Series C. Geology and Paleontology 26, 139–168.
Sakagami, S., Sakai, H., 1991. Fossil bryozoans from the lower Gondwana Sisne Formation, Nepal Lesser Himalayas. Transactions and Proceedings of the Palaeontological Society of Japan, New Series 162, 761–766. Senowbari-Daryan, B., Rashidi, K., Hamedani, A., 2006. Two Permian sphinctozoan sponges from the Shotori Mountains, northeast Iran. Geologica Carpathica 57, 427–432. Simpson, G.B., 1895. A handbook of the genera of the North American Paleozoic Bryozoa. 14th Annual Report of the State geologist (of New York) for the year 1894, pp. 407–608. Spjeldnaes, N., 1984. Upper Ordovician bryozoans from Ojl Myr, Gotland, Sweden. Bulletin of the Geological Institutions of the University of Uppsala, New Series 10, 1–66. Ulrich, E.O., 1882. American Paleozoic Bryozoa. The Journal of the Cincinnati Society of Natural History 5, 121–175 pp. 233–257. Ulrich, E.O., 1884. American Palaeozoic Bryozoa. The Journal of the Cincinnati Society of Natural History 7, 24–51. Ulrich, E.O., 1889. Contributions to micropalaeontology of the Cambro– Silurian rocks of Canada. Part 2. The Geological and Natural History Survey of Canada. P. William Poster Brown and Co, Montreal, pp. 26–48. Vine, G.R., 1884. Fourth report of the committee, consisting of Dr. H.C. Sorby and Mr. G.R. Vine, appointed for the purpose of reporting on fossil Bryozoa. In: Report of the British Association for the Advancement of Science (Southport, 1883). John Murray, London, pp. 161–209. Vine, G.R., 1885. Further notes on new species and other Yorkshire Carboniferous fossil Polyzoa described by Prof. John Phillips. Proceedings of the Yorkshire Geological and Polytechnic Society, new series 8, 377–393. Vine, G.R., 1886. Notes on the Yoredale Polyzoa of north Lancashire. Proceedings of the Yorkshire Geological and Polytechnic Society, new series 9, 78–98. Wyse Jackson, P.N., Bancroft, A.J., 1995. Generic revision of the cryptostome bryozoan Rhabdomeson Young and Young, 1874, with descriptions of two species from the Carboniferous of the British Isles. Journal of Paleontology 69, 28–45. Xia, F., 1986. Carboniferous and Early Permian bryozoans from Xainza, northern Xizang. Bulletin of Nanjing Institute of Geology and Palaeontology, Academia Sinica 10, 201–254 (in Chinese). Yanagida, J., A Research Group, 1988. Biostratigraphic study of Paleozoic and Mesozoic Groups in Central and Northern Thailand. An Interim Report. Department of Geology, Faculty of Sciences, Kyushu University, pp. 1–46. Yang, J., Lu, L., 1983. Upper Carboniferous and Lower Permian bryozoans from Kalpin of western Xinjiang. Paleontologia Cathayana 1, 259–317 (in Chinese). Yang, J., Lu, L., Xia, F., 1981. Upper Paleozoic bryozoans from Xizang. Palaeontology of Xizang 81–100 (in Chinese). Young, J., Young, J., 1874. New Carboniferous Polyzoa. The Quarterly Journal of the Geological Society 681–683.
Original article
Rhabdomesid and cystoporid bryozoans from the Permian of Deh-e Mohammad, Shotori Mountains (northeastern Iran)§ Bryozoaires rhabdome´sides et cystoporides du Permien de Deh-e Mohammad, Montagnes Shotori (nord-est de l’Iran) Andrej Ernst a,*, Baba Senowbari-Daryan b, Koorosch Rashidi c a
Institut für Geowissenschaften der Christian-Albrechts, Universität zu Kiel, Ludewig-Meyn-Strasse 10, 24118 Kiel, Germany b Institut für Paläontologie, Universität Erlangen–Nürnberg, Loewenichstrasse 28, 91054 Erlangen, Germany c University of Payam-e Noor, Ardakan, Iran Received 19 January 2007; accepted 23 September 2008 Available online 11 November 2008
Abstract Seven bryozoan species belonging to the Order Rhabdomesida and Order Cystoporida are described from the Permian deposits exposed near the small town of Deh-e Mohammad, Shotori Mountains (northeastern Iran): Rhabdomeson cf. consimile Bassler, Pamirella nitida Gorjunova, Clausotrypa conferta Bassler, Streblotrypa (Streblotrypa) elegans Sakagami, Streblotrypa (Streblascopora) supernodata nov. sp., Cystodictya sp., and Filiramoporina cf. kretaphilia Fry and Cuffey. The described fauna identifies the age of the Jamal Formation at the locality near Deh-e Mohammad as Lower Permian. It displays palaeobiogeographic connections to the Lower Permian of Pamir (Tajikistan), Indonesia, Thailand and Kansas (North America). # 2008 Elsevier Masson SAS. All rights reserved. Résumé Sept espèces de bryozoaires qui appartiennent à l’Ordre Rhabdomesida et à l’Ordre Cystoporida sont décrites dans les dépôts permiens affleurant près de la petite ville de Deh-e Mohammad, Montagnes Shotori (nord-est de l’Iran) : Rhabdomeson cf. consimile Bassler, Pamirella nitida Gorjunova, Clausotrypa conferta Bassler, Streblotrypa (Streblotrypa) elegans Sakagami, Streblotrypa (Streblascopora) supernodata nov. sp., Cystodictya sp., et Filiramoporina cf. kretaphilia Fry et Cuffey. La faune décrite attribue un âge permien inférieur à la Formation Jamal, au niveau de la localité proche de Deh-e Mohammad. Elle affiche des relations paléobiogéographiques avec le Permien inférieur du Pamir (Tadjikistan), du Timor (Indonésie), de la Thaïlande et du Kansas (Amérique du Nord). # 2008 Elsevier Masson SAS. Tous droits réservés. Keywords: Bryozoa; Taxonomy; Lower Permian; Palaeobiogeography Mots clés : Bryozoa ; Taxonomie ; Permien inférieur ; Paléobiogéographie
1. Abbreviations The following abbreviations are used in measurements: Statistics:
N X SD CV MIN MAX
Measured characters (in mm):
§
Corresponding editor: Jean Vannier. * Corresponding author. E-mail address: [email protected] (A. Ernst).
number of measurements mean standard deviation coefficient of variation minimal value maximal value
AW
0016-6995/$ – see front matter # 2008 Elsevier Masson SAS. All rights reserved. doi:10.1016/j.geobios.2008.09.001
width of autozooecial aperture
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ADB WB AB ND AcD
A. Ernst et al. / Geobios 42 (2009) 133–140
distance between aperture centres along branch branch width axial bundle diameter node diameter acanthostyle diameter
2. Introduction Bryozoans are abundant in Permian deposits in Iran, but they are poorly investigated. The earliest work, treating the Permian bryozoans of Iran, is published by Fantini Sestini (1965). This publication includes the description of three species from the Ruteh Limestone of Central Elborz (northern Iran). Later, Sakagami (1980) described 15 species from the Permian of central Iran (Abadeh region). This fauna displayed a close relation to the Dzhulfian (= Changhsingian) stage, as well as connections to the Permian of Pamir, Russian platform, Salt Range, Kashmir, South China, and Timor. Recently, Ernst et al. (2006a, 2006b) described two bryozoan faunal associations from the Jamal Formation, Iran. Thirty-one bryozoan species were identified from the Jamal Formation, exposed in Lakaftari area, south of the town of Bagher-Abad, northeast of Esfahan (central Iran). This association is regarded as Middle Permian (Murgabian) in age, and shows affinities to Australia, Indonesia, Thailand and Transcaucasia. Six bryozoan species were described from the Jamal Formation, exposed in Bagh-e Vang (Shotori Mountains, northeast Iran). This fauna indicates a Lower Permian age and shows palaeogeographic affinities to the bryozoan fauna known from Western Australia and Urals. The new bryozoan material, collected from the Jamal Formation exposed near the town of Deh-e Mohammad, Shotori Mountains (northeastern Iran), is described in the present publication. The bryozoans of this locality exhibit palaeogeographic connections to the Lower Permian of Pamir (Tajikistan), Indonesia, Thailand, and Kansas (North America). 3. Lithology and stratigraphy The material was collected from a locality situated northeast of the village Deh-e Mohammad in Shotori Mountains, northeastern Iran (Fig. 1). Permian rocks at this locality disconformably overlie the Upper Carboniferous Sardar Formation (Fig. 2). The Sardar Formation is composed of sandstones, sandy marls and light green shales. The Permian at this locality is represented by the Jamal Formation that contains at its base bioclastic limestones, succeeded by pure limestones, marly limestones with abundant Tubiphytes, algae, some sponges and echinoids, followed by conglomerates and breccias with rare sponges. The sphinctozoan sponge Amblysiphonella cf. vesiculosa vesiculosa (De Koninck) was described from this locality by Senowbari-Daryan et al. (2006). This part of the section is interpreted to be a reef mound. The middle part of the section consists of medium to thick bedded grey limestones with intercalated cherts and also Tubiphytes, small foraminifers, and a few bryozoans. The upper part of the section contains dolomites and cherts lacking fossils.
Fig. 1. Map showing the geographic position of the Permian locality near the town of Deh-e Mohammad.
4. Material and methods Taxonomic descriptions were based on thin sections. The rock materials, as well as the thin sections, are housed at the Institute of Paleontology, University of Erlangen–Nürnberg, Germany (Senowbari-Daryan, Permian, Deh-e Mohammad).
A. Ernst et al. / Geobios 42 (2009) 133–140
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Fig. 2. Lithostratigraphical column of the Permian deposits near the town of Deh-e Mohammad.
5. Systematic palaeontology Phylum BRYOZOA Ehrenberg, 1831 Class STENOLAEMATA Borg, 1926 Order RHABDOMESIDA Astrova and Morozova, 1956 Family RHABDOMESIDAE Vine, 1884 Genus Rhabdomeson Young and Young, 1874 [= Coeloconus Ulrich, 1889] Type species: Rhabdomeson progracile Wyse Jackson and Bancroft, 1995, by subsequent designation by Wyse Jackson
and Bancroft, 1995 (ICZN Opinion 1874); Lower Carboniferous, England. Rhabdomeson consimile Bassler, 1929 Fig. 3(A, B) 1929. Rhabdomeson consimile Bassler, p. 69, Pl. 13, Figs. 5–8. 1976. Rhabdomeson cf. consimile Bassler - Liu, p. 152, Pl. 75, Fig. 7. 1981. Rhabdomeson consimile Bassler - Yang, Lu and Xia, p. 93, Pl. 4, Fig. 7.
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Fig. 3. A–B. Rhabdomeson consimile Bassler, 1929, D1-6. A. Oblique thin section of the branch. B. Tangential section of the branch. C–E. Pamirella nitida Gorjunova, 1975. C. Oblique section of the branch, D1. D. Longitudinal section of the branch, displaying acanthostyles, D1. E. Tangential section of the branch, displaying autozooecial apertures and acanthostyles, D4. F–H. Clausotrypa conferta Bassler, 1929. F. Cross section of the branch, D4. G. Oblique section of the branch, D11. H. Oblique tangential section of the branch displaying autozooecial apertures, D11. I, J. Streblotrypa (Streblotrypa) elegans Sakagami, 1970. I. Cross section of the branch, D1. J. Oblique section of the branch, D1-6. K–N. Streblotrypa (Streblascopora) supernodata nov. sp. K. Longitudinal section of the branch,
A. Ernst et al. / Geobios 42 (2009) 133–140
1982. Rhabdomeson consimile Bassler - Kiseleva, p. 87, Pl. 16, Fig. 3. 1983. Rhabdomeson consimile Bassler - Yang and Lu, pp. 285–286, Pl. 3, Figs. 1–3. 1988. Rhabdomeson cf. consimile Bassler - Sakagami in Yanagida, Pl. 19, Figs. 5, 6, Pl. 20, Figs. 4, 6, 7. 1999. Rhabdomeson consimile Bassler - Sakagami, p. 91, Pl. 22, Fig. 6, Pl. 23, Figs. 1, 2. Material: Single specimen D1–6. Description: Ramose colony with small axial cylinder and distinct exozone, 1.23 mm in diameter. Axial cylinder 0.29 mm in diameter. Autozooecia budding in spiral pattern around axial cylinder, rhomboidal in cross-section. Autozooecial apertures oval, 0.11–0.17 mm wide, arranged in regular diagonal rows, spaced 0.24–0.28 mm from centre to centre, four longitudinally and eight diagonally in 2 mm. Single macroacanthostyle situated in interspaces between apertures, 0.036–0.048 mm in diameter. Microacanthostyles 0.02–0.0.03 mm in diameter, arranged in one row between macroacanthostyles. Superior hemiseptum blunt, obscure, positioned distally; inferior hemiseptum absent. Autozooecial walls 0.024–0.036 mm thick in endozone, 0.11–0.12 mm thick in exozone. Remarks: Rhabdomeson consimile Bassler, 1929 is similar to R. mammillatum (Bretnall, 1926) from the Lower Permian of Australia, but differs from the latter in presence of microacanthostyles and hemisepta. It differs also from R. bispinosum Crockford, 1944, from the Lower Permian of Western Australia, in having hemisepta and single instead of paired acanthostyles between apertures. Occurrence: Upper Permian; Indonesia. Upper Permian; Khabarovsk region (Russia). Lower to Upper Permian (? Artinskian to Guadalupian); Thailand. Permian; Mongolia. Permian; China (Western Xinjiang). Jamal Formation, Lower Permian; Deh-e Mohammad, Shotori Mountains, northeastern Iran. Family RHOMBOPORIDAE Simpson, 1895 Genus Pamirella Gorjunova, 1975 Type species: P. nitida Gorjunova, 1975, by original designation; Lower Permian (Artinskian), SE Pamir (Tajikistan). Pamirella nitida Gorjunova, 1975 Fig. 3(C–E) and Table 1 1975. Pamirella nitida - Gorjunova, p. 63, Pl. 10, Fig. 1. 1983. Pamirella nitida Gorjunova - Blake, p. 580, Fig. 288 (1a–d). 1985. Pamirella nitida Gorjunova - Gorjunova, p. 119, Pl. 8, Fig. 4. 1986. Pamirella nitida Gorjunova - Xia, p. 208, Pl. 1, Figs. 1–6.
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Table 1 Measurements of Pamirella nitida Gorjunova, 1975 (see explanations of abbreviations in the text).
AW ADB AcD
N
X
SD
CV
MIN
MAX
6 4 12
0.14 0.64 0.07
0.016 0.038 0.008
11.058 5.921 11.832
0.13 0.60 0.06
0.17 0.69 0.08
Material: D1, D4. Description: Ramose cylindrical colonies, 1.26 mm in diameter. Autozooecia tubular, growing in spiral pattern. Autozooecial diaphragms absent. Apertures oval, arranged in diagonal rows on the colony surface. Metazooecia rare, 0.02 mm in diameter. Acanthostyles abundant, 7-8 surrounding each autozooecial aperture. Microacanthostyles abundant. Remarks: Pamirella nitida Gorjunova, 1975 differs from a similar species P. pulchra (Bassler, 1929) from the Lower Permian (Artinskian) of Indonesia in having thinner branches (1.26 mm versus 2.50–3.00 mm in P. pulchra) and smaller apertures (0.13–0.17 mm versus 0.35 mm in P. pulchra; data from Bassler, 1929). Pamirella nitida is also similar to P. nepalensis Sakagami and Sakai, 1991, known from the upper part of Sisne Formation of Nepal Lesser Himalayas, but differs from it in having smaller apertures (0.13–0.17 mm versus 0.30 mm in P. nepalensis). Occurrence: Lower Permian (Artinskian); SE Pamir, Tajikistan. Angjie Formation (Lower Permian); Xainza, Northern Xizang, China. Jamal Formation, Lower Permian; Deh-e Mohammad, Shotori Mountains, northeastern Iran. Family NEMATOTRYPIDAE Spjeldnaes, 1984 Genus Clausotrypa Bassler, 1929 Type species: C. separata Bassler, 1929, by original designation; Lower Permian, Timor, Indonesia. Clausotrypa conferta Bassler, 1929 Fig. 3(F–H) and Table 2 1929. Clausotrypa conferta Bassler, p. 72, Pl. 14, Figs. 1–3. 1988. Clausotrypa sp. indet. - Sakagami in Yanagida, Pl. 19, Fig. 4. 1999. Clausotrypa conferta Bassler - Sakagami, p. 92, Pl. 23, Fig. 3. 2000. Clausotrypa sp. cf. C. conferta Bassler - Sakagami, p. 161, Fig. 9-2. Material: D1, D11. Description: Cylindrical branches, 1.44–1.80 mm in diameter. Autozooecia long, cylindrical, growing parallel to the branch axis for a long distance, turning gently to the colony surface. Autozooecial apertures oval, spaced 5–6 longitudinally in 2 mm. Diaphragms not observed. Tectitozooecia numerous, restricted mostly to exozone, in 1–3 rows separating
holotype D1–2. L. Tangential section of the branch, displaying autozooecial apertures and nodes, paratype D11. M. Oblique section of the branch displaying axial bundle of zooecia, holotype D1–2. N. Cross section of the branch, holotype D1–2. O, P. Cystodictya sp. O. Tangential section of the branch, D36. P. Oblique section of the branch, D36. Q. Filiramoporina cf. kretaphilia Fry and Cuffey, 1976, cross section of the branch, D100.
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Table 2 Measurements of Clausotrypa conferta Bassler, 1929 (see explanations of abbreviations in the text).
AW ADB
N
X
SD
CV
MIN
MAX
10 6
0.13 0.36
0.007 0.023
5.128 6.393
0.12 0.34
0.14 0.40
autozooecial apertures. Acanthostyles 0.02–0.03 mm in diameter, abundant, originating in the late endozone, having clear cores. Remarks: Clausotrypa conferta Bassler, 1929 differs from C. monticola (Eichwald, 1860) by having smaller acanthostyles and larger apertures (acanthostyle diameter 0.02–0.03 mm versus 0.02–0.08 mm in C. monticola, and aperture width 0.12– 0.14 mm versus 0.10–0.12 mm in C. monticola, data from Morozova and Krutchinina, 1986). Occurrence: Lower–Middle Permian; Indonesia. Permian; Thailand. Jamal Formation, Lower Permian; Deh-e Mohammad, Shotori Mountains, northeastern Iran. Suborder HYPHASMOPORIDAE Vine, 1886 Family HYPHASMOPORIDAE Vine, 1886 Genus Streblotrypa Vine, 1885 Subgenus Streblotrypa (Streblotrypa) Vine, 1885 Type species: Streblotrypa nicklesi Vine, 1885, by original designation; Middle Carboniferous, England. Streblotrypa (Streblotrypa) elegans Sakagami, 1970 Fig. 3(I, J) 1970. Streblotrypa elegans - Sakagami, pp. 65–66, Pl. 13, Figs. 5–9. 1976. Streblotrypa elegans Sakagami - Sakagami, Pl. 27, Fig. 4. 2000. Streblotrypa elegans Sakagami - Sakagami, p. 153, Figs. 7, 6. Material: D1, D1–6. Description: Ramose branching colonies, 0.66–0.90 mm in diameter. Branch cross sections rounded-polygonal in shape. Autozooecia tubular, with long thin bases, forming a distinct axial bundle, rounded to slightly polygonal in cross section. Axial bundle 0.22–0.51 mm in diameter. Autozooecial apertures oval, arranged in regular diagonal rows, 0.066–0.090 mm in diameter (0.075 mm, average), spaced 5 in 2 mm longitudinally, having distinct peristomes. Peristomal wall 0.02 mm thick. Short superior hemiseptum present; inferior hemiseptum very long, positioned near the place of bending outwards of the autozooecia. Abundant metazooecia between apertures, arranged in 4-5 rows. Remarks: Streblotrypa (Streblotrypa) elegans Sakagami, 1970 is similar to S. (S.) parva Morozova, 1965, known from the Upper Permian of Caucasus. However, S. (S.) elegans differs from S. (S.) parva by having long hemisepta. Occurrence: Lower Permian (?Lower Artinskian); Thailand. Jamal Formation, Lower Permian; Deh-e Mohammad, Shotori Mountains, northeastern Iran.
Table 3 Measurements of Streblotrypa (Streblascopora) supernodata nov. sp. (see explanations of abbreviations in the text).
WB AB AW ND
N
X
SD
CV
MIN
MAX
7 3 5 4
0.90 0.25 0.09 0.055
0.155 0.044 0.011 0.012
16.783 17.436 12.087 21.477
0.75 0.22 0.07 0.050
1.23 0.30 0.10 0.070
S. (Streblotrypa) elegans has been recently discovered in the Permian of Oman (Ernst et al., 2008: 692, Figs. 8.1–8.5). Subgenus Streblotrypa (Streblascopora) Bassler, 1929 Type species: Streblotrypa fasciculata Bassler, 1929, by original designation; Upper Permian, Indonesia. Streblotrypa (Streblascopora) supernodata nov. sp. Fig. 3(K–N) and Table 3 Etymology: The species name refers to large nodes between apertures. Holotype: D1–2 (Fig. 3K, M–N). Paratypes: D11 (Fig. 3L) and D116. Type locality: Deh-e Mohammad, Shotori Mountains (northeastern Iran [Fig. 1]). Type horizon: Jamal Formation, Lower Permian. Diagnosis: Ramose branched colonies; axial bundle with 11–13 axial zooecia arranged in 3-4 rows; inferior hemisepta long, superior hemisepta short; 3-4 large nodes surrounding each autozooecial aperture. Description: Ramose branched colonies, 0.75–1.23 mm in diameter. Axial bundle 0.22–0.30 mm in diameter, containing 11–12 axial zooecia arranged in 3–4 rows of zooecia. Branch cross sections rounded-polygonal in shape. Autozooecia tubular, growing from the axial bundle, Autozooecial apertures oval, arranged in regular diagonal rows. Inferior hemisepta long, superior hemisepta short. Diaphragms present, rare to common, thin, curved proximally. Metazooecia abundant, 5–7 arranged in 2–3 rows between apertures, 0.02–0.04 mm in diameter. Massive nodes between apertures; 3–4 surrounding each aperture. Remarks: Streblotrypa (Streblascopora) supernodata nov. sp. is similar to S. (S.) irianica Sakagami, 2000 from the Middle Permian of Irian Jaya, Indonesia, in having nodes, but differs from the latter in having thinner branches (0.75–1.23 mm versus 2.6–3.0 mm in S. (S.) irianica) and smaller autozooecial apertures (0.10–0.17 mm versus 0.15–0.19 mm in S. (S.) irianica). Order CYSTOPORIDA Astrova, 1964 Family CYSTODICTYONIDAE Ulrich, 1884 Genus Cystodictya Ulrich, 1882 Type species: C. ocellata Ulrich, 1882, by original designation; Lower Mississippian, Kentucky, USA. Cystodictya sp. Fig. 3(O, P) and Table 4
A. Ernst et al. / Geobios 42 (2009) 133–140 Table 4 Measurements of Cystodictya sp. (see explanations of abbreviations in the text).
AW ADB
N
X
SD
CV
MIN
MAX
5 6
0.15 0.62
0.012 0.059
8.333 9.515
0.13 0.57
0.16 0.72
Material: Two fragments D36. Description: Narrow bifoliate branches. Autozooecia short, growing from thin mesotheca in 2-3 rows on each side. Diaphragms not observed. Lunaria absent; distinct peristomes around autozooecial apertures. Few small vesicles. Hemisepta absent. Remarks: The present material is similar to Cystodictya absoluta Morozova and Lisitsyn, 2002 from the Upper Carboniferous (Gzhelian) of the Russian Plate. The latter species has four autozooecial rows instead of 2–3 in the present material, as well as wider and more widely spaced apertures (aperture width 0.16–0.18 mm versus 0.13–0.16 mm in present material; aperture spacing along branch 0.80–0.82 mm versus 0.57–0.72 mm in present material). Occurrence: Jamal Formation, Lower Permian; Deh-e Mohammad, Shotori Mountains, northeastern Iran. Genus Filiramoporina Fry and Cuffey, 1976 Type species: Filiramoporina kretaphilia Fry and Cuffey, 1976, by original designation; Lower Permian (Wreford Megacyclothem), Kansas, USA. Filiramoporina cf. kretaphilia Fry and Cuffey, 1976 Fig. 3(Q) Compare: 1976. Filiramoporina kretaphilia Fry and Cuffey, pp. 4–7, Pl. 1, Figs. 1–9, Pl. 2, Figs. 1–8. Material: Single specimen D100. Description: Bifoliate branch 2.42 mm wide and 1.65 mm thick, oval in cross section. Autozooecia arranged in eight rows on both sides of colony surface, growing from a straight mesotheca, trapezoid at their bases. Extrazooidal skeleton consisting of structureless material. Tangential and longitudinal sections are missing. Remarks: The Iranian species is similar to Filiramoporina kretaphilia Fry and Cuffey, 1976, the only known species of the genus, in branch size, autozooecial shape and the kind of extrazooidal skeleton (Roger Cuffey, pers. comm., 2006). However, the available material is insufficient to make a closer determination. Occurrence: Wreford Megacyclothem, Lower Permian; Kansas, USA. Jamal Formation, Lower Permian; Deh-e Mohammad, Shotori Mountains, northeastern Iran. 6. Discussion Bryozoa show a high potential for palaeobiogeographic purposes in the Palaeozoic. Permian bryozoan faunas were described from different regions and can be used for a palaeobiogeographic comparison. The bryozoan fauna from the
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Permian of Deh-e Mohammad locality described in the present paper displays connections predominantly to the Lower Permian of Thailand and Indonesia (Rhabdomeson consimile Bassler, 1929, Clausotrypa conferta Bassler, 1929, Streblotrypa (Streblotrypa) elegans Sakagami, 1970) and Pamir, Tajikistan (Pamirella nitida Gorjunova, 1975). Of note is the finding of Filiramoporina cf. kretaphilia Fry and Cuffey, 1976. The genus Filiramoporina Fry and Cuffey, 1976, with a single known species, was described from the Lower Permian of Kansas, USA. Acknowledgements The material for the present investigation was sampled by one of the authors (K.R.) during his Ph.D. thesis. The investigations were financially supported by the Deutsche Forschungsgemeinschaft (DFG) to B. Senowbari-Daryan (Project Se 416/10). K. Rashidi thanks the scientific ministry of Iran for financial support to stay 6 months in Erlangen. He thanks also the staff and members of the Institute of Paleontology (Erlangen) for their kind assistance. We also thank J.R.P. Ross, Bellingham for her kind review of the manuscript. References Astrova, G.G., 1964. Novyi otryad paleozoiskikh mshanok [New Order of Paleozoic Bryozoa]. Paleontologicheskij zhurnal 1, 22–31 (in Russian). Astrova, G.G., Morozova, I.P., 1956. K sistematike mshanok otryda Cryptostomata [About systematics of the Order Cryptostomata]. Doklady Akademii Nauk SSSR 110, 661–664 (in Russian). Bassler, R.S., 1929. The Permian Bryozoa of Timor. Paläontologie von Timor 16, 37–90. Blake, D.B., 1983. Systematic descriptions for the Suborder Rhabdomesina. In: Robison, R.A. (Ed.), Treatise on Invertebrate Paleontology, Part G (1): Bryozoa (revised). Geological Society of America and University of Kansas Press, Lawrence, pp. 551–592. Borg, F., 1926. Studies on recent cyclostomatous Bryozoa. Zoologiska Bidrag fran Uppsala 10, 181–507. Bretnall, R.W., 1926. Description of some western Australian fossil Polyzoa. Bulletin of Western Australia Geological Survey 88, 7–33. Crockford, J., 1944. Bryozoa from the Permian of Western Australia. Part I. Cyclostomata and Cryptostomata from the northwest basin and Kimberly district. Proceedings of the Linnean Society of New South Wales 69, 139– 175. Eichwald, E. von, 1860. Lethaea rossica, ou Paléontologie de la Russie. I, Seconde section de l’ancienne période. Bryozoaires, Schweizerbart, Stuttgart, pp. 355–494. Ehrenberg, C.G., 1831. Symbolae Physicae, seu Icones et descriptiones Corporum Naturalium novorum aut minus cognitorum, quae ex itineribus per Libyam, Aegiptum, Nubiam, Dongalaam, Syriam, Arabiam et Habessiniam, studia annis 1820–1825, redirent. Pars Zoologica, 4, Animalia Evertebrata exclusis Insectis. Berolini. Ernst, A., Senowbari-Daryan, B., Hamedani, A., 2006a. Middle Permian Bryozoa from the Lakaftari area, central Iran. Geodiversitas 28, 543– 590. Ernst, A., Senowbari-Daryan, B., Rashidi, K., 2006b. Lower Permian Bryozoa of the Jamal Formation from Bagh-e Vang (Shotori Mountains, northeast Iran). Facies 52, 627–635. Ernst, A., Weidlich, O., Schäfer, P., 2008. Stenolaemate Bryozoa from the Permian of Oman (Aseelah Unit, Batain coast). Journal of Paleontology 82, 676–716.
140
A. Ernst et al. / Geobios 42 (2009) 133–140
Fantini Sestini, F.N., 1965. The geology of the upper Djadjerud and Lar valleys (North Iran); II, Palaeontology; Bryozoans, brachiopods and molluscs from Ruteh limestone (Permian). Revista Italiana di Paleontologia 71, 13–108. Fry, H.C., Cuffey, R.J., 1976. Filiramoporina kretaphilia: a new genus and species of bifoliate tubulobryozoan (Ectoprocta) from the lower Permian Wreford Megacyclothem of Kansas. The University of Kansas Paleontological Contributions Paper 84, 1–12. Gorjunova, R.V., 1975. Permskie mshanki Pamira [Permian bryozoans of the Pamir]. Trudy paleontologicheskogo instituta Akademii Nauk SSSR 148, 1–127 (in Russian). Gorjunova, R.V., 1985. Morfologia, sistema I filogeniya mshanok (Otryad Rhabdomesida) [Morphology, system and phylogeny of Bryozoa (Order Rhabdomesida)]. Trudy paleontologicheskogo instituta Akademii Nauk SSSR 208, 1–152 (in Russian). Kiseleva, A.B., 1982. Pozdnepermskie mshanki Yuzhnogo Primorya [Late Permian bryozoans of the South Primorye]. Nauka 1–128 (in Russian). Liu, X.L., 1976. Phylum Bryozoa. In: Palaeontological Atlas of north China. Inner Mongolia], Vol. 1. Palaeozoic Volume. Geological Publishing House, Peking, pp. 131–155 (in Chinese). Morozova, I.P., 1965. Bryozoa. In: Ruzhentsev, V.E., Sarycheva, T.G. (Eds.), The evolution and changes in the marine organisms at the Paleozoic– Mesozoic boundary. Trudy paleontologicheskogo instituta Akademii Nauk SSSR, 108pp. 183–197 (in Russian). Morozova, I.P., Krutchinina, O.N., 1986. Permian Bryozoans of the Arctis (Western Sector). Nauka 1–141 (in Russian). Morozova, I.P., Lisitsyn, D.V., 2002. Revision of bryozoans from the stratotype sections of the Gzhelian Stage (Upper Carboniferous). Paleontologicheskij zhurnal 6, 63–72 (in Russian). Sakagami, S., 1970. Addition to the Permian Bryozoa from Ko Muk, Peninsular Thailand. Contributions to the Geology and Palaeontology of Southeast Asia. 53. Geology and Palaeontology of Southeast Asia. Contributions to the geology and palaeontology of Southeast Asia 8, 43–68. Sakagami, S., 1976. On the Permian Bryozoa from the northern part of Sainbeyli, Central Turkey. Transactions and Proceedings of the Palaeontological Society of Japan, New Series 103, 398–405. Sakagami, S., 1980. Permian Ectoprocta (Bryozoa) from the Abadeh Region, Central Iran. Transactions and Proceedings of the Palaeontological Society of Japan. New Series 118, 269–289. Sakagami, S., 1999. Permian bryozoans from some localities in the Khao Hin Kling Area near Phetchabun, north-central Thailand. Bulletin of the Kitakyushi Museum of Natural History 18, 77–103. Sakagami, S., 2000. Middle Permian Bryozoa from Irian Jaya, Indonesia. Bulletin of the National Science Museum, Series C. Geology and Paleontology 26, 139–168.
Sakagami, S., Sakai, H., 1991. Fossil bryozoans from the lower Gondwana Sisne Formation, Nepal Lesser Himalayas. Transactions and Proceedings of the Palaeontological Society of Japan, New Series 162, 761–766. Senowbari-Daryan, B., Rashidi, K., Hamedani, A., 2006. Two Permian sphinctozoan sponges from the Shotori Mountains, northeast Iran. Geologica Carpathica 57, 427–432. Simpson, G.B., 1895. A handbook of the genera of the North American Paleozoic Bryozoa. 14th Annual Report of the State geologist (of New York) for the year 1894, pp. 407–608. Spjeldnaes, N., 1984. Upper Ordovician bryozoans from Ojl Myr, Gotland, Sweden. Bulletin of the Geological Institutions of the University of Uppsala, New Series 10, 1–66. Ulrich, E.O., 1882. American Paleozoic Bryozoa. The Journal of the Cincinnati Society of Natural History 5, 121–175 pp. 233–257. Ulrich, E.O., 1884. American Palaeozoic Bryozoa. The Journal of the Cincinnati Society of Natural History 7, 24–51. Ulrich, E.O., 1889. Contributions to micropalaeontology of the Cambro– Silurian rocks of Canada. Part 2. The Geological and Natural History Survey of Canada. P. William Poster Brown and Co, Montreal, pp. 26–48. Vine, G.R., 1884. Fourth report of the committee, consisting of Dr. H.C. Sorby and Mr. G.R. Vine, appointed for the purpose of reporting on fossil Bryozoa. In: Report of the British Association for the Advancement of Science (Southport, 1883). John Murray, London, pp. 161–209. Vine, G.R., 1885. Further notes on new species and other Yorkshire Carboniferous fossil Polyzoa described by Prof. John Phillips. Proceedings of the Yorkshire Geological and Polytechnic Society, new series 8, 377–393. Vine, G.R., 1886. Notes on the Yoredale Polyzoa of north Lancashire. Proceedings of the Yorkshire Geological and Polytechnic Society, new series 9, 78–98. Wyse Jackson, P.N., Bancroft, A.J., 1995. Generic revision of the cryptostome bryozoan Rhabdomeson Young and Young, 1874, with descriptions of two species from the Carboniferous of the British Isles. Journal of Paleontology 69, 28–45. Xia, F., 1986. Carboniferous and Early Permian bryozoans from Xainza, northern Xizang. Bulletin of Nanjing Institute of Geology and Palaeontology, Academia Sinica 10, 201–254 (in Chinese). Yanagida, J., A Research Group, 1988. Biostratigraphic study of Paleozoic and Mesozoic Groups in Central and Northern Thailand. An Interim Report. Department of Geology, Faculty of Sciences, Kyushu University, pp. 1–46. Yang, J., Lu, L., 1983. Upper Carboniferous and Lower Permian bryozoans from Kalpin of western Xinjiang. Paleontologia Cathayana 1, 259–317 (in Chinese). Yang, J., Lu, L., Xia, F., 1981. Upper Paleozoic bryozoans from Xizang. Palaeontology of Xizang 81–100 (in Chinese). Young, J., Young, J., 1874. New Carboniferous Polyzoa. The Quarterly Journal of the Geological Society 681–683.