Bryozoan faunas at the Tortonian-Messinian transition. A palaeoenvironmental case study from Crete Island, eastern Mediterranean

2021  43  26 geodiversitas Geodiversitas est une revue en flux continu publiée par les Publications scientifiques du Muséum, Paris Geodiversitas is a fast track journal published by the Museum Science Press, Paris Les Publications scientifiques du Muséum publient aussi / The Museum Science Press also publish: Adansonia, Zoosystema, Anthropozoologica, European Journal of Taxonomy, Naturae, Cryptogamie sous-sections Algologie, Bryologie, Mycologie, Comptes Rendus Palevol Diffusion – Publications scientifiques Muséum national d’Histoire naturelle CP 41 – 57 rue Cuvier F-75231 Paris cedex 05 (France) Tél. : 33 (0)1 40 79 48 05 / Fax : 33 (0)1 40 79 38 40 [email protected] / http://sciencepress.mnhn.fr © Publications scientifiques du Muséum national d’Histoire naturelle, Paris, 2021 ISSN (imprimé / print) : 1280-9659/ ISSN (électronique / electronic) : 1638-9395 Directeur De la publication / Publication director : Bruno David, Président du Muséum national d’Histoire naturelle réDacteur en chef / editor-in-chief : Didier Merle assistant De réDaction / assistant editor : Emmanuel Côtez ([email protected]) Mise en page / Page layout : Emmanuel Côtez coMité scientifique / scientific board : Christine Argot (Muséum national d’Histoire naturelle, Paris) Beatrix Azanza (Museo Nacional de Ciencias Naturales, Madrid) Raymond L. Bernor (Howard University, Washington DC) Alain Blieck (chercheur CNRS retraité, Haubourdin) Henning Blom (Uppsala University) Jean Broutin (Sorbonne Université, Paris, retraité) Gaël Clément (Muséum national d’Histoire naturelle, Paris) Ted Daeschler (Academy of Natural Sciences, Philadelphie) Bruno David (Muséum national d’Histoire naturelle, Paris) Gregory D. Edgecombe (The Natural History Museum, Londres) Ursula Göhlich (Natural History Museum Vienna) Jin Meng (American Museum of Natural History, New York) Brigitte Meyer-Berthaud (CIRAD, Montpellier) Zhu Min (Chinese Academy of Sciences, Pékin) Isabelle Rouget (Muséum national d’Histoire naturelle, Paris) Sevket Sen (Muséum national d’Histoire naturelle, Paris, retraité) Stanislav Štamberg (Museum of Eastern Bohemia, Hradec Králové) Paul Taylor (The Natural History Museum, Londres, retraité) couverture / cover : Réalisée à partir des Figures de l’article/Made from the Figures of the article. Geodiversitas est indexé dans / Geodiversitas is indexed in: – Science Citation Index Expanded (SciSearch®) – ISI Alerting Services® – Current Contents® / Physical, Chemical, and Earth Sciences® – Scopus® Geodiversitas est distribué en version électronique par / Geodiversitas is distributed electronically by: – BioOne® (http://www.bioone.org) Les articles ainsi que les nouveautés nomenclaturales publiés dans Geodiversitas sont référencés par / Articles and nomenclatural novelties published in Geodiversitas are referenced by: – ZooBank® (http://zoobank.org) 1365 GEODIVERSITAS • 2021 • 43 (26) © Publications scientifiques du Muséum national d’Histoire naturelle, Paris. www.geodiversitas.com Pierre MOISSETTE National & Kapodistrian University of Athens, Faculty of Geology and Geoenvironment, Department of Historical Geology and Paleontology 15784, Athens (Greece) and CR2P (CNRS, MNHN, UPMC, Sorbonne Université), Département Origines et Évolution, UMR7207, Muséum national d’Histoire naturelle, case postale 38, 57 rue Cuvier, F-75231 Paris cedex 05 (France) [email protected] Assimina ANTONARAKOU George KONTAKIOTIS National & Kapodistrian University of Athens, Faculty of Geology and Geoenvironment Department of Historical Geology & Paleontology, 15784, Athens (Greece) [email protected] [email protected] Jean-Jacques CORNÉE Géosciences Montpellier, Université de Montpellier Université des Antilles, CNRS, Pointe à Pitre, Guadeloupe, FWI (France) [email protected] Vasileios KARAKITSIOS National & Kapodistrian University of Athens, Faculty of Geology and Geoenvironment Department of Historical Geology & Paleontology, 15784, Athens (Greece) [email protected] Submitted on 5 February 2020 | accepted on 6 June 2020 | published on 16 December 2021 Bryozoan faunas at the Tortonian-Messinian transition. A palaeoenvironmental case study from Crete Island, eastern Mediterranean KEY WORDS Bryozoa, palaeobathymetry, palaeoenvironment, palaeogeography, Late Miocene, Greece, Crete, Aegean Sea. urn:lsid:zoobank.org:pub:14A6956D-54AD-48D2-9C5E-BA380EDACAA4 Moissette P., Antonarakou A., Kontakiotis G., Cornée J.-J. & Karakitsios V. 2021. — Bryozoan faunas at the TortonianMessinian transition. A palaeoenvironmental case study from Crete Island, eastern Mediterranean. Geodiversitas 43 (26): 1365-1400. https://doi.org/10.5252/geodiversitas2021v43a26. http://geodiversitas.com/43/26 ABSTRACT Four sedimentary sections were logged and sampled from upper Tortonian-lower Messinian outcrops on the island of Crete (Greece). The collected material yielded about 60 bryozoan species belonging to nine different colonial morphotypes. A few species are stenobathic, indicating either shallowor deep-water environments, but most of them are eurybathic (with bathymetric ranges extending in some instances from the shelf down to several hundreds of metres). Bryozoan communities point to sea-level variations modulated by local tectonics. Deep circalittoral environments were recognized at the base of three sections (whereas the fourth section starts with infralittoral/shallow circalittoral environments passing to deep circalittoral. Shallow bathyal habitats follow upwards, succeeded in turn by assemblages indicative of deep circalittoral and subsequently shallow circalittoral/infralittoral depths. Inferred bathymetric fluctuations are diachronous throughout the island. Shallow-water species found associated, sometimes abundantly, with rarer deep-water faunas in a few levels, are interpreted as transported by currents. Several dysoxic episodes have been also detected: likely the result of water 1366 GEODIVERSITAS • 2021 • 43 (26) Moissette P. et al. INTRODUCTION At the end of the Miocene, during the Tortonian-Messinian Transition (TMT), sea water exchanges between the Mediterranean and the Atlantic Ocean began to be restricted (e.g., Kontakiotis etal. 2019; Corbí etal. 2020). This restriction culminated by the end of the Messinian when the Betic and Rifian corridors were closed (Martín etal. 2001; Capella etal. 2017; Krijgsman etal. 2018). This led to the well-known Messinian Salinity Crisis (MSC) characterised by the widespread deposition of thick evaporites in deep and/or marginal sub-basins of the entire Mediterranean Sea (Hsü etal. 1973; Ryan etal. 1973; Aksu etal. 2018; Haq etal. 2020; Manzi etal. 2020). An abundant literature has been devoted to the timing and the causes of this event, and a consensus progressively emerged (CIESM 2008; Manzi etal. 2013; Roveri etal. 2014, 2018, 2020). To explain the large volume of evaporites, connections between the Atlantic Ocean and the Mediterranean Sea are considered necessary. Atlantic waters entered the Mediterranean either through the Betic and Rifian corridors or even the Messinian Gibraltar Corridor (Achalhi etal. 2016; Krijgsman etal. 2018). The first step of the MSC should have occurred in the 5.97-5.60 Ma interval (Gautier etal. 1994; Krijgsman etal. 1999; Manzi etal. 2013). During this interval, the Primary Lower Gypsum unit (PLG), the Terminal Carbonate Complex (TCC; Esteban 1979), and coeval basinal sediments were deposited. The PLG is topped by a major erosional surface which has been identified both onshore and offshore (e.g., Lofi etal. 2011). The second step happened in the 5.60-5.54 Ma interval; it is characterised by the Resedimented Lower Gypsum unit (RLG) and halite deposition. The third step of the MSC occurred in the 5.545.33 Ma interval. It is typified by the deposition of the Upper Gypsum unit and sediments displaying freshwater influences (Lago Mare facies), prior to a marine reflooding at ca 5.33 Ma (base of the Pliocene). The island of Crete (Aegean Sea, Greece) offers excellently exposed Miocene to Pliocene marine deposits that yielded abundant, diverse, and generally well-preserved fossil organisms. Apart from foraminifera, previous studies dealing with fossil communities are relatively few (Agiadi etal. 2017). Among invertebrates, bryozoans are rather common, but have not been studied in detail, even in the late Miocene (Moissette etal. 1993, 2018; Drinia etal. 2009). Bryozoans are benthic sessile invertebrates that can confidently be used for palaeoenvironmental reconstructions (e.g., Schopf 1969; Harmelin 1988; Moissette 2000; Amini etal. 2004; Taylor 2005). Studies on present-day Mediterranean deep-water bryozoans are scarce, but fossil communities have been the subject of much less attention (Moissette& Spjeldnaes 1995; Rosso 2005; Di Geronimo etal. 2005; Moissette etal. 2017). The purposes of this paper are thus: 1) to contribute to a better knowledge of late Miocene bryozoans and 2) use them for palaeoenvironmental reconstructions. Four sections (Potamida, Keramoutsi, Kapariana, and Faneromeni) have been chosen (Moissette etal. 2018) in three different sedimentary basins (Chania, Heraklion, and Sitia) in a west-east transect along the Island of Crete, in order to column stratification and enhanced productivity. These processes were probably boosted by a combination of changes in oceanic circulation, climate, global sea-level, and the local/regional tectonics (in Crete and/or the marine gateways between the Mediterranean and the Atlantic). RÉSUMÉ Faunes de bryozoaires à la limite Tortonien-Messinien. Étude de cas des paléoenvironnements de l’île de Crète, Méditerranée orientale. Quatre coupes lithostratigraphiques ont été levées et échantillonnées dans les affleurements du Tortonien supérieur-Messinien inférieur de l’île de Crète (Grèce). Le matériel prélevé a livré environ 60 espèces de bryozoaires appartenant à neuf morphotypes coloniaux différents. Quelques espèces sont sténobathes, indiquant des environnements littoraux ou profonds, mais la plupart sont eurybathes, avec des intervalles bathymétriques s’étendant parfois depuis le plateau continental jusqu’à plusieurs centaines de mètres de profondeur. Les communautés de bryozoaires révèlent des variations du niveau marin modulées par la tectonique locale. Des environnements du circalittoral profond sont reconnus à la base de trois des coupes (mais la quatrième coupe commence par l’infralittoral/circalittoral côtier, passant ensuite au circalittoral profond). Des habitats du bathyal supérieur apparaissent ensuite, suivis d’assemblages indiquant le circalittoral profond, puis le circalittoral côtier/infralittoral. Ces fluctuations bathymétriques sont fortement diachrones à travers l’île. Des espèces littorales sont par ailleurs associées, parfois en grand nombre, avec de plus rares faunes profondes dans quelques niveaux où elles ont été transportées par des courants. Plusieurs épisodes dysoxiques ont été également identifiés, résultant probablement d’une stratification croissante de la colonne d’eau et d’une augmentation de la productivité organique. Ces processus ont été vraisemblablement favorisés par une combinaison de changements dans la circulation océanique, le climat, le niveau marin global et la tectonique locale/régionale (en Crète et/ou au niveau des corridors marins entre la Méditerranée et l’Atlantique). MOTS CLÉS Bryozoaires, paléobathymétrie, paléogéographie, paléoenvironnement, Miocène supérieur, Grèce, Crète, Mer Égée. 1367 Bryozoan faunas at the Tortonian-Messinian transition GEODIVERSITAS • 2021 • 43 (26) analyse the changes that occurred in bryozoan faunas before the Messinian Salinity Crisis (MSC). Our sampling starts approximately at 7.58 Ma, c.220 ka after the Tortonian Salinity Crisis (TSC at 7.8 Ma; Krijgsman etal. 2000), and terminates at 6.72 Ma, c.750 ka before the onset of the MSC (5.97 Ma; Manzi etal. 2013, 2018). Therefore, the studied sediments document the TMT, a crucial time interval for the palaeoceanographic history of the Mediterranean Sea. GEOLOGICAL BACKGROUND Around the middle/late Miocene boundary, the area now occupied by the island of Crete (Fig. 1) became divided into a series of basins bounded by faults and filled mostly by marine sediments (van Hinsbergen& Meulenkamp 2006; Zachariasse etal. 2008, 2011). During the late Miocene, marls, clayey limestones and sapropels deposited, later replaced by carbonates, diatomites and evaporites (Drinia etal. 2004, 2007b; Karakitsios etal. 2017a, b; Moissette etal. 2018; Antonarakou etal. 2019; Kontakiotis etal. 2019, 2020). MATERIAL AND METHODS The four sections studied in this paper have been logged and sampled during several field work campaigns (Figs 2-5) and some data have been already published in Moissette etal. (2018). Depending mostly on outcropping conditions, an average sampling interval of approximately 1m has been used (range TUNISIA ITALY SICILY TURKEY LIBYA GREECE 20 km Pre-NeogeneNeogene Heraklion B Sitia Chania Potamida Faneromeni Kapariana Gavdos A Rethymnon Keramoutsi 200 km CRETE 35°30' 35° 35°30’ 35° 26° 26°25°30’ 25°30’25° 25°24°30’ 24°30’ 24° 24° 23°30’ 23°30’ fig. 1. — A, Situation map of Crete within the eastern Mediterranean; B, geological sketch map of the island of Crete (after Krijgsman et al. 1994), with location of the studied sections. 1368 GEODIVERSITAS • 2021 • 43 (26) Moissette P. et al. between about 1.6m in Keramoutsi and 0.6m in Potamida section). An overall number of 232 samples were collected, among which 171 yielded at least one bryozoan specimen. For each sample, 500 g of dry sediment were soaked for several hours in diluted hydrogen peroxide and later washed under running water through five mesh sieves (2mm, 1mm, 0.5mm, 0.25mm, and 0.125mm). The residues were oven-dried and all identifiable fossils were picked and counted under a stereomicroscope. The time framework and the stratigraphic correlation of the studied sections was based on the qualitative and semi-quantitative analysis of the planktonic foraminifera identified in the samples (Moissette etal. 2018; Kontakiotis etal. 2019; this study). The chronology of the studied sections is based on the assemblage-based concept of the marker species, which was initially developed by Zachariasse (1975) and elaborated by several authors (e.g., Hilgen etal. 1995; Antonarakou etal. 2007; Karakitsios etal. 2017a Lozar etal. 2018; Kontakiotis etal. 2019; Vasiliev etal. 2019; Zachariasse etal. 2021) in Mediterranean Neogene sections. A total of c.300 specimens from the >0.125mm size fraction of the planktonic foraminiferal assemblage were collected from the washed residue to obtain information about the abundance of marker species as a percentage of the total planktonic foraminiferal fauna. The sections were dated by using presence/absence patterns and coiling of planktonic foraminiferal marker species based on the planktonic foraminiferal biostratigraphy of Krijgsman etal. (1994, 1995). Ages for the planktonic foraminiferal bioevents recognized (Table 1) are derived from the astrochronology based on magnetostratigraphy and astronomical tuning of sedimentary cycle patterns (Lourens etal. 2004). Each bryozoan specimen, fragment or whole colony, was counted as one unit in the >0.25mm size fraction and a semi-quantitative representation (very rare, rare, frequent, common, abundant) was then produced (Figs 2-5). The palaeoenvironmental analysis was based on the known ecological requirements of living representatives: colonial growth forms (zoarial forms) and species (Hageman etal. 1998; Moissette 2000; Amini etal. 2004; Taylor 2005). Selected well-preserved specimens were ultrasonically cleaned and then observed and photographed with a scanning electron microscope (JEOL JSM-6360) at the National and Kapodistrian University of Athens (Department of Historical Geology-Paleontology). The bathymetric (bionomical) zonation used in this paper follows that of Pérès& Picard (1964). The depth limits are estimates related to light and temperature levels along latitudinal and longitudinal gradients, also depending on bottom substrate characteristics: infralittoral (0-40m), shallow circalittoral (40-80m), deep circalittoral (80-200m), and upper bathyal (200-500m). The approximate equivalent terms used by van Morkhoven etal. (1986) are respectively: inner shelf (inner neritic), mid shelf (mid neritic), outer shelf (outer neritic), and upper slope (upper bathyal). RESULTS AND INTERPRETATIONS Biostratigraphy Most results related to the Potamida, Kapariana and Faneromeni sections have already been published by Moissette etal. (2018). New data for the 84-m thick Keramoutsi section (Heraklion basin; 35°16’34.77”N, 25°01’38.88”E) are given below and illustrated in Figure 3. The planktonic foraminiferal analysis of the study sections revealed the succession of 10 planktonic bioevents (Table 1) and the Tortonian-Messinian boundary (7.24 Ma). Overall, all study sections cover the TMT spanning the time interval between 7.58 and 6.72 Ma. The Faneromeni section is slightly older than the other three sections, with the dextral to sinistral (d/s) coiling change of the Globorotalia scitula group recorded at the base of this section (Moissette etal. 2018). The biostratigraphic evidence from the top of the sections further points to an age younger than, at least, the Last Occurrence (LO) of G.nicolae at 6.72 Ma, for Faneromeni and Kapariana compared to Keramoutsi and Potamida sections respectively (Figs 2-5). table 1. — Planktonic foraminiferal bioevents identified in the four studied sections (Moissette et al. 2018; Kontakiotis et al. 2019). Abbreviations: LO, Last Occurrence; LCO, Last Common Occurrence; FO, First Occurrence; FCO, First Common Occurrence. Stage Planktonic Foraminiferal Bioevents Stratigraphic level (m) Chronostratigraphy Potamida Keramoutsi Kapariana Faneromeni Age (Ma) Messinian (10) LO G. nicolae – – 37.5 47.7 6.72 (9) FO G. nicolae 34.4 65.5 33.0 41.0 6.83 (8) LCO G. scitula group (sin) – – 31.0 32.4 7.08 (7) FCO G. miotumida group 28.8 44.5 – 24.6 7.24 Tortonian (6) Influx G. menardii 4 within the range of G. menardii 5 26.8 – 15.1 22.2 7.28 (5) Paracme end G. scitula group (dex) 26.8 – – 22.2 7.28 (4) FO G. menardii 5 (dex) 7.5 – 13.2 19.4 7.36 (3) LO C. parvulus 4.3 28.0 7.0 12.8 7.45 (2) LCO G. menardii 4 (sin) 0.5 21.0 – 8.2 7.51 (1) G. scitula group coiling change D/S – – – 3.5 7.58 1369 Bryozoan faunas at the Tortonian-Messinian transition GEODIVERSITAS • 2021 • 43 (26) Description of the sections anD of their Bryozoan communities The studied sediments are mostly composed of marls and clayey limestones. Additionally, sandstones, sandy marls, bioclastic limestones, ferruginous concretions, and bivalve shell beds are common (Moissette etal. 2018). The associated skeletal organisms comprise abundant foraminifera (benthic and planktonic), common to rare bivalves, rare to very rare ostracods, echinoids, pteropods, and very rare gastropods, scaphopods, decapods, brachiopods and fishes. Bryozoan remains occur in almost 3/4 of the collected samples, but they are generally minor components with only a few fragments and species in each sample. The main results (including distribution of the species and semi-quantitative abundances) are presented in Figures 2-5 . Marl Clayey limestone Sandstone Halimeda limeston e Ferruginous bed Bivalve shells Crisia denticulata Crisia aculeata Entalophoroecia sp. Exidmonea atlantica Scrupocellaria sp. Crisia sp. Ybselosoecia typica Nellia tenella Reteporella sp. Smittina cervicornis Turbicellepora sp. Scrupocellaria scrupea Chaperia annulus Cheiloporina campanulata Disporella hispida Margaretta cereoides Smittina canavarii Steginoporella montenati Annectocyma major Canda rectangulata Celleporina canariensis Cellaria salicornioides Bufonellaria muriella Calpensia nobilis Biflustra savartii Celleporina cf. siphuncula Escharina cf. dutertrei Escharoides coccinea Lagenipora lepralioides Mecynoecia delicatula Myriapora truncata Onychocella angulosa Puellina sp. Schizotheca fissa Tubulipora sp. MESSINIAN –2 10 10 20 –1 20 0 0 39 30 m 10 30 20 40 m 1 30 TORTONIAN SECT. POTAMIDA A SECT. POTAMIDA B Dysoxia Transported elements DysoxiaDysoxia frequent (11-20) rare (6-10) very rare (1-5) common (21-50) abundant (>51) 7.28 Ma 7.24 Ma 6.83 Ma 7.24 Ma deep circalittoral upper bathyal deep circalittoral shallow circ.- infralittoral samples Batopora rosula Anguisia verrucosa Orbitulipora excentrica Bryocryptella torquata 7.51 Ma 7.45 Ma 7.36 Ma fig. 2 . — Schematic sedimentary log of Potamida composite section with sample location and semi-quantitative abundances of bryozoan species. 1370 GEODIVERSITAS • 2021 • 43 (26) Moissette P. et al. Dysoxia 10 20 1 30 9' 40 50 Transported elements Annectocyma major Crisia fistulosa Crisia denticulata Entalophoroecia sp. Exidmonea atlantica Crisia sp. Ybselosoecia typica Canda rectangulata Celleporina canariensis Chlidonia pyriformis Nellia tenella Reteporella sp. Smittina cervicornis Tubulipora sp. Cellaria salicornioides Buffonellaria muriella Calpensia nobilis Celleporina cf. siphuncula Cheiloporina campanulata Margaretta cereoides frequent (11-20) rare (6-10) very rare (1-5) common (21-50) abundant (>51) Marl Clayey limestone Laminated marl Sandstone Bivalve shells Ferruginous bed Sandy marl Adeonella polystomella Biflustra savartii Cellaria fistulosa Cupuladria cf. canariensis Discoporella reussiana Escharoides coccinea Hincksina sp. Hippoporina sp. Mecynoecia delicatula Scrupocellaria sp. Onychocella angulosa Turbicellepora sp. Steginoporella montenati Schizotheca fissa Patinella radiata Scrupocellaria scrupea 80 m 70 60 50 40 30 20 10 0 MESSINIANTORTONIAN Crisia aculeata 7.24 Ma upper bathyal deep circ. infralittoral shallow circalittoral-infralittoral deep circalittoral lagoonal shallow circ.- samples Dysoxia 7.51 Ma 7.45 Ma 7.24 Ma 6.83 Ma Gemellipora eburnea Batopora rosula Tervia irregularis Anguisia verrucosa Orbitulipora excentrica fig. 3 . — Schematic sedimentary log of Keramoutsi section with sample location and semi-quantitative abundances of bryozoan species. 1371 Bryozoan faunas at the Tortonian-Messinian transition GEODIVERSITAS • 2021 • 43 (26) Gemellipora eburnea Batopora rosula Tervia irregularis Anguisia verrucosa Dysoxia 26 PLIOCENE 0 10 20 30 40 50 m 01 010 020 030 036 1 10 12' 14' 20 MESSINIANTORTONIAN Transported element s Annectocyma major Crisia fistulosa Crisia denticulata Entalophoroecia sp. Exidmonea atlantica Scrupocellaria sp. Crisia sp. Ybselosoecia typica Canda rectangulata Celleporina cf. siphuncula Chlidonia pyriformis Reteporella sp. Smittina cervicornis Turbicellepora sp. Cellaria salicornioides Scrupocellaria scrupea Buffonellaria muriella Calpensia nobilis Cheiloporina campanulata Disporella hispida Halysisis cf. diaphana Margaretta cereoides Smittina canavarii Steginoporella montenati Marl Clayey limestone Laminated marl Sandstone Bioclastic limestone Bivalve shells Ravinement surface Dysoxia frequent (11-20) rare (6-10) very rare (1-5) common (21-50) abundant (>51) Crisia aculeata Ma deep circ. upper bathyal deep circalittoral shallow circalittoral-infralittoral SECT. KAPARIANA E (KAP) SECT. KAPARIANA W (CAP) samples 7.24 7.24 Ma 7.08 Ma 6.83 Ma 6.72 Ma 7.28 Ma 7.36 Ma 7.45 Ma fig. 4 . — Schematic sedimentary log of Kapariana composite section with sample location and semi-quantitative abundances of bryozoan species. 1378 GEODIVERSITAS • 2021 • 43 (26) Moissette P. et al. remarks The gonozooid, characteristic of the genus, consisting of a simple elongated chamber with a terminal tubular ooeciostome (J.-G. Harmelin, personal communication), was not observed. The encrusting basis is visible in some of the studied specimens (Fig. 6B). The species created by Neviani (1895) from the Pliocene/Pleistocene of northern Italy, A.jullieni most probably corresponds to the encrusting basis of A.verrucosa. However, the homonymous A.jullieni described by Ostrovsky (1998) from the present-day Antarctic is a different species. Peristome diameter and length are smaller in A.verrucosa and the prominent verrucae associated with the pseudopores are absent in A.jullieni. Family tuBuliporiDae Johnston, 1838 type genus.— Exidmonea David, Mongereau& Pouyet, 1972, by subsequent designation of Mongereau (1969: 212). Genus Exidmonea David, Mongereau& Pouyet, 1972 Exidmonea atlantica (Forbes in Johnston, 1847) (Fig. 7C, D) Idmonea atlantica Forbes in Johnston, 1847: 278, pl. 48, fig. 3.— Buge 1966: 5, pl. B, figs 1-2. Idmidronea atlantica – Harmelin 1976: 182, pl. 32, figs 1-11.— Vávra 1983: 73, pl. 1, figs 1-4.— Zabala 1986: 658, text-figs 235a-d; pl. 28, fig. E.— Zabala& Maluquer 1988: 174, text-figs 569-572; pl. 33, fig. A.— El Hajjaji 1992: 44, pl. 2, fig. 11.— Haddadi-Hamdane 1996: p. 45, pl. 1, fig. 8, pl. 2, fig. 4. Exidmonea atlantica – Mongereau 1970: 30, pl. 1, fig. 1; pl. 2, figs 1, 5.— David etal. 1972: 84.— Moissette 1988: 48, pl. 6, figs 6-7.— Pouyet& Moissette 1992: 24, pl. 1, figs 7-8.—Moissette etal. 1993: 84, figs 3i-j.— Moissette& Spjeldnaes 1995: 780, pl. 1, fig. 11. Exidmonea triforis – Hayward& McKinney 2002: 104, fig. 49A-H, 50. o ccurrence .— Eocene: Italy, Austria, Hungary, Romania (Zágoršek 2003). Oligocene: Germany, Italy (Braga 2008). Early Miocene: France, Italy, Egypt (El Safori& El-Sorogy 1999). Middle Miocene: Austria, Hungary, Poland, France, Libya (Vávra 1983). Late Miocene: Algeria (Moissette 1988), Morocco (El Hajjaji 1992), Tunisia (Moissette 1997), Crete. Pliocene: France, Sicily (Pouyet& Moissette 1992), Algeria (Haddadi-Hamdane 1996), Tunisia. Pleistocene: Sicily (Di Geronimo etal. 1997; Rosso 2005), Rhodes (Moissette& Spjeldnaes 1995). This list of fossil records concerns Idmonea-Idmidronea-Exidmonealike species, i.e. a group of species with the same growth form without consideration of the species-specific morphological criteria which cannot be, in most cases, preserved in fossils (J.-G. Harmelin, personal communication). Recent: Eastern Atlantic (North Sea to Angola) and Mediterranean, at depths ranging from 10 to 850m. In the Mediterranean, this species is particularly abundant between 40 and 100m (Harmelin 1976). Description Vinculariiform colony. Frontal surface showing zoecial tubes alternating in two series of 4-5 tubes. Dorsal flattened or slightly concave, ornamented by thin inverted U-shaped growth lines. Gonozooid (brood-chamber) elongate, situated along the axis of a branch, between peristomes. remarks The oeciostome was not observed. Family terviiDae Canu& Bassler, 1920 Genus Tervia Jullien, 1882 Tervia irregularis (Meneghini, 1844) Idmonea irregularis Meneghini, 1845: 128. Tervia irregularis – Malecki 1963: 71, text-fig. 30; pl. 3, fig. 11.— Vávra 1975: p. 523, pl. 2, figs 3-4.— Harmelin 1976: 163, pl. 26, figs 1-11.— Harmelin& d’Hondt 1982: 9, pl. 3, fig. 4.— Zabala 1986: 656, text-fig. 234.— Moissette 1988: 49, pl. 6, figs 10-12.— Zabala& Maluquer 1988: 174, text-figs 563-568.— Pouyet& Moissette 1992: 25, pl. 1, figs 11-12.— El Hajjaji 1992: p. 48, pl. 2, fig. 10.— Moissette etal. 1993: 85, fig. 3h, l.— Moissette& Spjeldnaes 1995: 782, pl. 1, figs 9, 12.— Haddadi-Hamdane 1996: 46, pl. 1, fig. 10, pl. 2, figs 3, 8. occurrence.— Eocene: Poland, Hungary, Spain (Malecki 1963). Oligocene: Germany (Malecki 1963). Early Miocene: France (Buge 1957). Middle Miocene: Austria, Hungary (Moissette etal. 2006), France. Late Miocene: Algeria (Moissette 1988), Morocco (El Hajjaji 1992), Crete (Moissette etal. 1993). Pliocene: Netherlands (Lagaaij 1952), Belgium, France, Algeria, Sicily (Pouyet& Moissette 1992). Pleistocene: Rhodes (Moissette& Spjeldnaes 1995), Karpathos (Moissette etal. 2017). Recent. Eastern Atlantic (down to a depth of 2650m in the Bay of Biscay), Mediterranean (60300m), Pacific, Indian Ocean. remarks Only a few fragments have been found in two samples. This easily recognizable species is not illustrated here because of the relatively poor preservation state of the rare recovered specimens. Family DiaperoeciiDae Canu, 1918 Genus Ybselosoecia Canu& Lecointre, 1933 Ybselosoecia typica (Manzoni, 1878) (Fig. 7E-G) Filisparsa typica Manzoni, 1878: 10, pl. 8, fig. 30.— Canu 1909: 115, pl. 14, figs 25-26. Ybselosoecia typica – Buge 1956: 13, pl. 1, figs 3-4; pl. 2, figs 3-4.— Malecki 1963: 76, fig. 33; pl. 5, fig. 1.— Mongereau 1965: 317, fig. 1.— Vávra 1974: 362, pl. 2, figs 9-10.— Vávra 1984: 226, pl. 1, figs 2-3.— Moissette 1988: 56, pl. 7, figs 9, 13; 1997: 188, pl. 1, fig. 6.— El Hajjaji 1992: 56, pl. 2, figs 2-3.— Pouyet& Moissette 1992: 28, pl. 2, fig. 4.— Moissette etal. 1993: 87.— Moissette& Spjeldnaes 1995: 784, pl. 2, fig.4.— Haddadi-Hamdane 1996: 51, pl. 1, figs 11-12; pl. 2, fig. 11.— Pouyet 1997: 26, pl. 1, figs 1-4.— Zágoršek 2003: 119, pl. 4, figs 5-6.— Zágoršek 2010a: 31, pl. 15, figs 1-5.— Zágoršek etal. 2017: 115, fig. 2J. 1379 Bryozoan faunas at the Tortonian-Messinian transition GEODIVERSITAS • 2021 • 43 (26) occurrence.— Eocene: France, Italy, Germany, Austria, Romania, Poland (Malecki 1963; Zágoršek 2003). Oligocene: France (Buge 1964). Oligo-Miocene: Iran (Zágoršek etal. 2017). Early Miocene: France (Mongereau 1965). Middle Miocene: France, Austria, Hungary, Romania, Czech Republic, Poland (Vávra 1984; Moissette etal. 2006). Late Miocene: Morocco, Algeria (Moissette 1988), Tunisia (Moissette 1997), Crete (Moissette etal. 1993). Pliocene: Netherlands, France, Sicily (Pouyet& Moissette 1992), Algeria (Haddadi-Hamdane 1996), Tunisia (Buge 1956). Pleistocene: Rhodes (Moissette& Spjeldnaes 1995). This fossil species has mostly been recorded from the Mediterranean and Paratethys basins. It has been recovered in a few instances together with deep-water bryozoan faunas: Messinian of western Algeria (Moissette 2000) and Pleistocene of Rhodes (Moissette& Spjeldnaes 1995). Description Vinculariiform colony. Dichotomous branches mostly flattened. Frontal smooth, zoecial tubes indistinct, quincuncially arranged and isolated, with well-developed sub-circular peristomes. Dorsal slightly convex, with thin transversal growth A DEF BC fig. 8. — A-C, Crisia aculeata Hassall, 1841; A, internode, frontal view, POTB13: AMPG(IV) 2828a; B, internode, dorsal view, POTB13 2828b: AMPG(IV); C, internode with gonozooid, KAP22: AMPG(IV) 3339. D-F, Crisia denticulata (Lamarck, 1816); D, internode, frontal view, POTB13: AMPG(IV) 2829a; E, internode dorsal view, POTB13: AMPG(IV) 2829b; F, gonozooid, POTB13: AMPG(IV) 2829c. Scale bars: A, 500 µm; B-F, 200 µm. 1380 GEODIVERSITAS • 2021 • 43 (26) Moissette P. et al. lines and perforated by small pseudopores. Gonozooid enlarged, also perforated by small pseudopores, occupying the whole width of a branch and encompassing up to ten zoecial tubes. Ooeciostome with flared, transverse and convoluted rim. Suborder ARTICULINA Busk, 1859 Family crisiiDae Johnston, 1838 Genus Crisia Lamouroux, 1812 Crisia aculeata Hassall, 1841 (Fig. 8A-C) Crisia aculeata Hassall, 1841: 170, pl. 7, figs 3-4.— Harmer 1891: 132, pl. 12, fig. 4.— Hayward& Ryland 1985: 50, fig. 14.— Harmelin1990: 1602, figs 2, 3-6. occurrence.— Recent: Eastern Atlantic (50-60m in the North Sea, much deeper in Norway, the Bay of Biscay and Morocco: 1351000m) and Mediterranean (115-480m). Description Cellariiform colony. Internodes short (8-12 zooids). The first ramification arises from the third zooid. Pseudopores elongate and relatively rare. Gonozooid pyriform with a discrete ooeciostome situated near the base of the following zooid. remarks This species is poorly known and has rarely been illustrated. It has often been considered as a variety of C.eburnea (Linnaeus) to which it resembles. Harmelin (1990) distinguished a northern (Atlantic) and a southern form (Atlantic coast of Morocco, Mediterranean). The main differences are a higher number of zooids (9-12) per internode and a mitre-shaped gonozooid in the southern form. Crisia denticulata (Lamarck, 1816) (Fig. 8D-F) Cellaria denticulata Lamarck, 1816: 137. Crisia denticulata – Vávra 1975: 523, pl. 1, fig. 1.— Hayward& Ryland 1985: 54, fig. 16.— Poluzzi etal. 1988: 64, figs 15b-d.— Poluzzi& Rosso 1988: 99, pl. 1, fig. 1.— Zabala& Maluquer 1988: 165, text-figs 480-482, pl. 29B.— Bobies 1958: 153, pl. 13, figs 5-7.— Marcopoulou-Diacantoni& Wuest 1999: 552, pl. 2, fig. 6. occurrence.— Middle Miocene: Austria, Hungary, Czech Republic (Bobies 1958; Vávra 1975). Late Miocene: Sardinia, Sicily (Moissette etal. 2002). Pliocene: UK, Netherlands (Lagaaij 1952), Belgium, Crete (Marcopoulou-Diacantoni& Wuest 1999). Pleistocene: Sicily (Rosso 1987). Recent: western and eastern Atlantic (Canada to Gulf of Mexico, Madeira, Azores, Norway to Ghana), Mediterranean. This species lives at depths between 0 and 100m, but it is associated with deep-water-corals in Norway (Sula Reef: 275-295m; Mortensen& Fosså 2006) and in the Bay of Biscay (Calvet 1896). remarks C.denticulata closely resembles the fossil species C.hoernesi Reuss, which however has a greater number of zooids per internode (14-16). Class GYMNOLAEMATA Allman, 1856 Order CHEILOSTOMATIDA Busk, 1852 Suborder FLUSTRINA Smitt, 1868 Superfamily calloporoiDea Norman, 1903 Family QuaDricellariiDae Gordon, 1984 Genus Nellia Busk, 1852 Nellia tenella (Lamarck, 1816) (Fig. 9E) Cellaria tenella Lamarck, 1816: 135. Nellia oculata – Busk 1852: 18, pl. 64, fig. 6; pl. 65 (bis), fig. 4.— Moissette etal. 1993: 98, fig. 5i.— Moissette 1997: 192, pl. 2, figs 5-6. Nellia tenella – Ziko etal. 2016: 18, pl. 2, fig. 6. occurrence.— Eocene: UK, France, Italy, Germany, Romania (Braga 2008). Oligocene: USA, Malta, Libya (El Safori& Muftah 2007), UAE (Braga& Bahr 2003). Early Miocene: USA (Di Martino etal. 2017), Dominican Republic, France, Portugal, Egypt (El Safori& El-Sorogy 1999). Middle Miocene: Jamaica, Hungary (Moissette etal. 2006), Austria, France, Portugal, Egypt (Ziko etal. 2016). Late Miocene: Morocco (El Hajjaji 1992), Algeria (Moissette 1988), Crete, Turkey, Tunisia (Moissette 1997). Pliocene: Costa Rica, Panama, Dominican Republic, Jamaica (Taylor& Foster 1998; Cheetham etal. 1999). Pleistocene: Rhodes (PM, pers. obs.). Recent: eastern and western Atlantic, Indian Ocean, Pacific. This species is widely distributed in tropical to subtropical waters around the world. It lives at depths between 0 and 250m (Fransen 1986). Description Colony cellariiform. Internodes straight with quadrangular section and elongate zooids separated by shallow grooves. Opesia oval, occupying the greatest part of the zooidal frontal wall. Gymnocyst and cryptocyst well-developed proximally. Two small avicularia occur on the proximal part of the gymnocyst. Ovicells not observed. remarks Nellia oculata Busk is now regarded as a junior synonym of N.tenella (Lamarck). However, their very broad ecological, geographical and stratigraphical records may indicate either a ’’living fossil’’ species or a “cryptic species complex” (Winston etal. 2014; Di Martino etal. 2017). Family cupulaDriiDae Lagaaij, 1952 Genus Cupuladria Canu& Bassler, 1919 Cupuladria cf. canariensis (Busk, 1859) (Fig. 9A, B) Cupularia canariensis Busk, 1859: 66, pl. 23, figs 6-9. Cupuladria canariensis – Manzoni 1869, p. 26, pl. 2, fig. 17; 1877: 72, pl. 17, figs 56a-c.— Cipolla 1921: 31, pl. 2, figs 22-24.— Lagaaij 1952: 33, pl. 2, figs 1a-b.— Buge 1957: 139, pl. 9, fig. 5.— Annoscia 1963: 225, pl. 9, fig. 1; pl. 10, fig. 1; pl. 11, figs 1a-b; pl. 12, figs 1a-b.— Cook 1965: 197, text-figs 1a-f, pl. 1, fig. 1; pl. 3, fig. 4.— Prenant& Bobin 1966: 307, figs 101-102.— Baluk& Radwansky 1984: 21, pl. 1, figs 1-4; pl. 8, figs 1-4.— Zabala& Maluquer 1988: 89, fig. 112- 1381 Bryozoan faunas at the Tortonian-Messinian transition GEODIVERSITAS • 2021 • 43 (26) A C EFGH D B fig. 9. — A, B, Cupuladria cf. canariensis (Busk, 1859); A, frontal view of a whole colony, KER39: AMPG(IV) 3078a; B, dorsal view of a whole colony, KER39: AMPG(IV) 3078b; C, D, Discoporella reussiana (Manzoni, 1869); C, frontal view of a whole colony, KER30: AMPG(IV) 3102; D, dorsal view of a whole colony, KER30: AMPG(IV) 3101; E, Nellia tenella (Lamarck, 1816), detail of an internode showing two zooids in frontal view, KER17: AMPG(IV) 3150a; F-H, Canda rectangulata Udin, 1964; F, dorsal view of an internode fragment, FAN35: AMPG(IV) 3506a; G, frontal view of an internode fragment, FAN35: AMPG(IV) 3506b; H, Detail of the same fragment, FAN35: AMPG(IV) 3506b. Scale bars: A-D, 1 mm; E, H, 100 µm; F-G, 200 µm. 1382 GEODIVERSITAS • 2021 • 43 (26) Moissette P. etal. 114.— Pouyet& Moissette 1992: 33, pl. 3, figs 4-5.— Moissette etal. 1993: 92, figs 4g-i.— Haddadi-Hamdane 1996: 65, pl. 5, figs 1, 4.— Marcopoulou-Diacantoni& Wuest 1999: 555, pl. 3, fig. 1. occurrence.— Middle Miocene: France, Spain, Austria, Poland (Baluk& Radwansky 1984). Late Miocene: Germany, Italy, Crete (Moissette etal. 1993). Pliocene: UK, Netherlands, Sicily (Pouyet& Moissette 1992), Algeria (Haddadi-Hamdane 1996), Crete (Marcopoulou-Diacantoni& Wuest 1999). Pleistocene: Sicily (Rosso 1987), Rhodes (Moissette 2012). Recent: eastern Pacific (Ecuador to northern Mexico), western and eastern Atlantic (Brazil, Caribbean, Gabon, Azores, Madeira and Canary islands to southern Portugal), southern Mediterranean (Rosso& Di Martino 2016). This warm-water species lives on sandy, more or less muddy bottoms at depths between 50 and 300m (Prenant& Bobin 1966). But it has also been found in much shallower (5-50m) and much deeper waters (down to 860m in the Sargasso Sea; Lagaaij 1963). Following Cadée (1979, 1981), C.canariensis seems however mostly a western Atlantic species. Records with this name must consequently be carefully checked, eliminating possible misidentifications (A. Rosso, personal communication). remarks Due to possible confusion with two other species of the same genus (C.biporosa (Canu& Bassler, 1923) and C.vindobonensis Baluk& Radwanski, 1984) a systematic revision is needed. Cadée (1979) also created a new subspecies, Cupuladria canariensis cavernosa, for Mio-Pliocene European specimens with intermediate characteristics between C.biporosa and C.canariensis (Cadée 1979, 1981). Genus Discoporella d’Orbigny, 1852 Discoporella reussiana (Manzoni, 1869) (Fig. 9C, D) Cupularia reussiana Manzoni, 1869: 27, pl. 2, fig. 19. Cupuladria reussiana – Annoscia 1963: 226, pl. 9, fig. 2; pl. 10, fig. 2; pl. 13, fig. 1; pl. 14, fig. 1a-b.— Prenant& Bobin 1966: 316, fig. 104.— Reguant 1969: 38, figs 1-3. Reussirella reussiana – Baluk& Radwansky 1984: 27, pl. 11, figs 1-2. Discoporella reussiana – Cook 1965: 219, text-fig. 2f; pl. 3, fig. 1.— Poluzzi 1975: 52, pl. 18, figs 1a-m, 2, 3.— Pouyet& Moissette 1992: 44, pl. 5, fig. 4-6.— Moissette etal. 1993: 96, figs 5c-d.— Haddadi-Hamdane 1996: 66, pl. 5, figs 2, 3, 8. occurrence.— Middle Miocene: Austria (Baluk& Radwansky 1984). Late Miocene: Italy, Crete (Moissette etal. 1993). Pliocene: Algeria (Haddadi-Hamdane 1996), Sicily (Pouyet& Moissette 1992). Pleistocene: Spain (Reguant 1969), Sicily, Umbria, central Italy (Bizzarri etal. 2015). Recent: the present-day occurrence of this species is doubtful (Cook 1965; Prenant& Bobin 1966). The nearest living species, R.doma, lives in tropical to subtropical waters of the Atlantic and Mediterranean, on sandy to muddy bottoms at depths of 10 to 370m (Cook 1965; Prenant& Bobin 1966). Description Lunulitiform colony. Frontal surface with alternating radial series of rhomboidal zooids, each bearing a distal vibraculum. The six cryptocystal denticles do not fuse in the middle of the opesia. The apical zone of the colony shows zooids covered by a calcified lamina. Basal surface depressed in the centre, displaying marked radial grooves and strong tubercles. remarks D.reussiana resembles very much Reussirella doma (d’Orbigny, 1851) to which it has been confused (Prenant& Bobin 1966). According to Cook (1965), the unfused cryptocystal denticles are especially characteristic of the western African Reussirella owenii (Gray, 1828). Superfamily BuguloiDea Gray, 1848 Family canDiDae d’Orbigny, 1851 Genus Canda Lamouroux, 1816 Canda rectangulata Udin, 1964 (Fig. 9F-H) Canda rectangulata Udin, 1964: 393, pl. 1, fig. 3.— Vávra 1979: 599, pl. 1, figs a, d, g; 1980: 58, pl. 2, figs 5-6.— Schmid 1989: 25, pl. 6, figs 1-5, 8.— Moissette 1997: 193, pl. 2, figs 11-12. o ccurrence .— Middle Miocene: Austria (Schmid 1989), Hungary (Moissette etal. 2007). Late Miocene: Tunisia (Moissette 1997). Description Colony vinculariiform. Frontal showing rectangular zooids organized in two alternating series with prominent distal and lateral edges. Cryptocyst concave, proximally well-developed and thus resulting in an elongate V-shaped opesia. Dorsal displaying enlarged triangular vibracularia with long setal grooves and large radicular pore. Frontal avicularia and ovicells not observed. remarks The closeness with the Recent Canda retiformis Pourtalès has been emphasized since the creation of C.rectangulata. A revision of both fossil and Recent material of the genus Canda is desirable. Genus Scrupocellaria van Beneden, 1845 Scrupocellaria cf. elliptica (Reuss, 1847) (Fig. 10A-C) Bactridium ellipticum Reuss, 1847: 56, pl. 9, fig. 7-8. Scrupocellaria elliptica – Reuss 1874: 148, pl. 11, figs 1-9.— Moissette 1988: 106, pl. 16, figs 5, 8.— Schmid 1989: 23, pl. 5, figs 1-7.— Zágoršek 2010a: 46, pl. 52, figs 1-6. Scrupocellaria sp. – Berning 2006: 32, figs 27-29. occurrence.— Middle Miocene: Austria (Schmid 1989), Czech Republic (Zágoršek 2010a). Late Miocene: Algeria (Moissette 1988), Spain (Berning 2006). Description Cellariiform colony. Each segment consists of two alternating series of about five zooids. Gymnocyst smooth. Oval opesia 1383 Bryozoan faunas at the Tortonian-Messinian transition GEODIVERSITAS • 2021 • 43 (26) occupying approximately one half of the zooecial length. Two large septula are discernible in the distal part of the opesial margin. Four spine bases occur on the outer distal part of the opesia and three on the inner angle, accompanied by one slightly larger scutal spine (scutum rarely preserved, covering almost half the opesia). No frontal avicularium. Lateral avicularia triangular and well developed. Dorsal surface showing small triangular vibracularia and radicular pores. No observed ovicells. remarks As noted by Schmid (1989) and Berning (2006), many fossil Scrupocellaria specimens have been mistakenly assigned to S.elliptica. Partly due to a frequent poor state of preservation this has been accompanied by insufficient description and illustration, notably concerning the presence and number of spine bases. Spines are not alluded to in Reuss (1847), but mentioned and partly illustrated in Reuss (1874). Although not reported, spines are visible on some specimens of the late Miocene of Algeria (Moissette 1988: pl. 16, fig. 8) on. Scutum and spines (at least 5) are clearly observable on the middle Miocene material (Schmid 1989: pl. 5, fig. 2 and fig. 4, respectively). Six spines (+scutum) are reported and illustrated by Berning (2006). Zágoršek (2010b) describes “spines often arranged in 4-5 pairs”. A revision of Neogene Mediterranean/ Paratethys Scrupocellaria is thus strongly necessary, notably to confirm the attribution of specimens with seven oral spines to S.elliptica. A new approach to the taxonomy of the polyphyletic genus Scrupocellaria has been presented by Vieira etal. (2014). Scrupocellaria scrupea Busk, 1852 (Fig. 10D-E) Scrupocellaria scrupea Busk, 1852: 83, pl. 9, figs 11-12.— Hincks 1880: 50, pl. 7, figs 11-14.— Prenant& Bobin 1966: 432, text-fig. 143.— Ryland& Hayward 1977: 138, text-fig. 66.— Zabala& Maluquer 1988: 99, text-figs 167-168.— Moissette& Spjeldnaes 1995: 788, pl. 3, figs 2-4.— Hayward& McKinney 2002: 27, fig. 11A-E. occurrence.— Pliocene: Calabria (Neviani 1900). Pleistocene: Calabria (Di Geronimo etal. 1997), Rhodes (Moissette& Spjeldnaes 1995). Recent: eastern Atlantic and Mediterranean from the surface to 150m (and down to 500m in the Gulf of Gascony: Jullien& Calvet 1903). Description Cellariiform colony. Internodes comprising two alternating series of zooids. Gymnocyst smooth. Oval opesia occupying slightly more than half of the zooecial length with two large septula in the distal part of the opesial margin Five spine bases on the distal rim (three on the outer edge and two on the inner part), accompanied by one slightly larger scutal basis on the internal edge. The scutum itself, rarely preserved, is rather large and its proximal lobe is more developed. Prominent triangular lateral avicularia. When present, the frontal avicularia are small and always located near the internal distal part of the ovicells, which are rounded, smooth and exhibit a small proximal fenestra. Dorsal surface with small triangular vibracularia. remarks The number of spine bases is relatively constant, but one of them, on the distal rim, is occasionally lacking. Only two spines are observable on ovicellate zooids. Superfamily cellarioiDea Lamouroux, 1821 Family cellariiDae Hincks, 1880 Genus Cellaria Ellis& Solander, 1786 Cellaria salicornioides Lamouroux, 1816 (Fig. 10F-H) Cellaria salicornioides Lamouroux, 1816: 127.— Audouin 1826: 236; Savigny 1817: pl. 6, fig. 7.— Prenant& Bobin 1966: 382, text-fig. 124.— Ryland& Hayward 1977: 124, text-figs 56A, 59.— Poluzzi& Padovani 1984: 109, fig. 5d.— Zabala 1986: 307, text-fig. 84:3a-e; pl. 3, figs E-F.— Moissette 1988: 104, pl. 17, figs 1-2.— Zabala& Maluquer 1988: 94, text-figs 137-140; pl. 2, fig. H.— Schmid 1989: 18, pl. 4, figs 3-6.— El Hajjaji 1992: 129, pl. 6, fig. 7.— Moissette& Spjeldnaes 1995: 788, pl. 3, fig. 1.— Haddadi-Hamdane 1996: 71, pl. 6, figs 1213.— López de la Cuadra& García-Gómez 1996: 158, figs 1C, 3A-F, 4. occurrence.— Early Miocene: France (Pouyet 1991). Middle Miocene: Austria (Schmid 1989), Hungary (Moissette etal. 2006), Czech Republic, Poland (Pouyet 1997). Late Miocene: Morocco (El Hajjaji 1992), Algeria (Moissette 1988). Pliocene: Algeria (Haddadi-Hamdane 1996), Crete. Pleistocene: Sicily (Poluzzi& Padovani 1984), Rhodes (Moissette& Spjeldnaes 1995). Recent: eastern Atlantic (Morocco, Madeira, up to the Shetland Islands), Mediterranean, Red Sea. The species has been recorded from the Atlantic at depths of 0-360m (and even down to 636m in Morocco). In the Mediterranean, it occurs from the shallow infralittoral down to 280m. remarks A number of specimens, especially those from the Faneromeni section, resemble C.salicornioides var. normani, a variety created by Hastings (1946) and considered as a distinct species, C.normani, by Prenant& Bobin (1966). However, López de la Cuadra& García-Gómez (1996) considered that the slight morphological differences (more slender internodes and larger avicularia in C.normani) do not justify the erection of a distinct taxon. Suborder ASCOPHORA Levinsen, 1909 Infraorder HIPPOTHOOMORPHA Gordon, 1989 Superfamily hippothooiDea Busk, 1859 Family pasytheiDae Davis, 1934 Genus Gemellipora Smitt, 1873 Gemellipora eburnea Smitt, 1873 (Fig. 10I) Gemellipora eburnea Smitt, 1873: 35, pl. 7, figs 152-156.— Harmer 1957: 994, pl. 69, figs 28-29.— Moissette 1988: 190, pl. 30, figs 6, 9, 12. Gemellaria punctata – Seguenza 1880: 127, pl. 12, figs 14-14a.— Neviani 1900: 148, pl. 16, figs 89. 1384 GEODIVERSITAS • 2021 • 43 (26) Moissette P. et al. Pasythea eburnea – Busk 1884: 5, pl. 34, fig. 1a-f.— Canu& Bassler 1928: 151, pl. 8, figs 11-12. occurrence.— Early Miocene: Indonesia (Di Martino& Taylor 2014). Middle Miocene: Hungary (Moissette etal. 2006). Late Miocene: Dominican Republic (Cheetham etal. 1999), Algeria (Moissette 1988), Calabria. Pliocene: Sicily (Rosso 2002). Pleistocene: Calabria, Sicily (Rosso& Di Geronimo 1998; Rosso 2005), Rhodes (Moissette& Spjeldnaes 1995), Karpathos (Moissette etal. 2017). Holocene: southern Italy (Di Geronimo etal. 2001). Recent: western and eastern Atlantic (Brazil, Caribbean, Azores, Madeira, Bay of Biscay, Gulf of Cadiz), Indian Ocean (Indonesia), Pacific (Hawaii, New Zealand). This is a deep-water species found in the Atlantic at depths between 60m and 3300m (Harmer 1957; Harmelin 1977). Description Cellariiform colony. Internodes made of one to four pairs of zooids. In each pair, the zooids are separated by a thin groove, placed back to back, slightly twisted from one another, and oriented at about 90 degrees of the previous/ A E FG HI B CD fig. 10. — A-C, Scrupocellaria cf. elliptica (Reuss, 1847); A, view of some zooids showing opesia with 4 spine bases on the outer distal part and 3 spine bases + 1 scutal spine on the inner distal rim, FAN36: AMPG(IV) 3587; B, zooids with 7 spine bases and the scutum, FAN35: AMPG(IV) 3586a; C, dorsal surface of a colony fragment with small vibracularia and radicular pores, FAN35: AMPG(IV) 3586b; D, E, Scrupocellaria scrupea Busk, 1852; D, ovicellate colony fragment, FAN35: AMPG(IV) 3590a; E, detail frontal view showing opesia with five spine bases, FAN35: AMPG(IV) 3590b; F-H, Cellaria salicornioides Lamouroux, 1816; F, part of a slender internode, FAN18: AMPG(IV) 3513a; G, detail view showing denticles and endotoichal ovicells, POTB13: AMPG(IV) 2821a; H, avicularia in frontal and lateral view (arrow), POTB13: AMPG(IV) 2821b; I, Gemellipora eburnea Smitt, 1873, broken internode showing two zooids with two oval scars separated by a thin interzooidal groove, CAP032: AMPG(IV) 3426. Scale bars: A-E, G, I, 100 µm; F, H, 200 µm. 1385 Bryozoan faunas at the Tortonian-Messinian transition GEODIVERSITAS • 2021 • 43 (26) following pair. Zooids elongate, displaying a smooth frontal surface with scarce inconspicuous pores. A few zooids bear in their central part a small oval scar (separated by the thin groove dividing two zooids) corresponding to the start of a lateral branch. Apertures almost circular with two very small indentations on the proximal corners. Neither avicularia nor ovicells. remarks Smitt (1873) describes an encrusting base (Smitt 1873: pl. 7, fig. 152) from which the erect, typical form arises. Another creeping colony (Smitt 1873: pl. 9, fig. 178) is erroneously attributed by the same author to G.eburnea. The confusion was evidenced by Canu& Bassler (1928). Infraorder UMBONULOMORPHA Gordon, 1989 Superfamily lepralielloiDea Vigneaux, 1949 Family BryocryptelliDae Vigneaux, 1949 Genus Bryocryptella Cossman, 1906 Bryocryptella torquata (Jullien, 1903) (Fig. 11A-C) Cryptella torquata Jullien in Jullien& Calvet, 1903: 77, pl. 7, fig. 5a-c. Porella torquata – Norman 1909: 300, pl. 39, figs 5-8.— Barroso 1912: 33, figs 6, 6a-e. Bryocryptella torquata – Álvarez 1991: 93, fig. 5; pl. 10, figs A-E. occurrence.— Recent: eastern Atlantic (Gulf of Gascony, Madeira). Deep-water species recorded at depths from 80m to 300m (Álvarez 1991). Description Colony vinculariiform. Narrow subcylindrical branches formed by 3 - 4 alternating longitudinal rows of zooids opening on the same frontal side. Zooids hexagonal separated by deep furrows. Frontal ventricose with about 20 large pores. Aperture subcircular; a small avicularium is sometimes present on its proximal edge. Peristome more or less developed, devoid of pores. Dorsal slightly convex showing marked grooves corresponding to zooecial limits and a few pores. Ovicells not observed. remarks This species was erroneously identified as Characodoma sp. in Moissette etal. (2018). Family tessaraDomiDae Jullien& Calvet, 1903 Genus Tessaradoma Norman, 1869 Tessaradoma boreale (Busk, 1860) (Fig. 11D) Onchopora borealis Busk, 1860: 213, pl. 28, figs 6-7. Porina borealis – Hincks 1880: 229, pl. 31, figs 4-6.— Neviani 1891: 120, pl. 4, figs 4-5. Tessaradoma boreale – Gautier1962: 222.— Hayward& Ryland 1979: 242, text-fig. 104.— Moissette 1988: 123, pl. 20, fig. 5.— Zabala& Maluquer 1988: 142, text-fig. 343.— El Hajjaji 1992: 225, pl. 13, fig. 2.— Pouyet& Moissette 1992: 53, pl. 7, fig. 3.— Moissette& Spjeldnaes 1995: 790, pl. 3, fig. 5. o ccurrence .— Middle Miocene: France. Late Miocene: Morocco (El Hajjaji 1992), Algeria (Moissette 1988), Calabria. Pliocene: northern Italy, Sicily (Pouyet& Moissette 1992). Pleistocene: Sicily, Calabria (Rosso 2005), Rhodes (Moissette& Spjeldnaes 1995), Karpathos (Moissette etal. 2017). Recent: Atlantic (western, but mostly eastern), Arctic, Mediterranean. T.boreale is predominantly a deep-water (Cheetham 1972), cryophilic (max. 13° C) species (60-3500m in the Atlantic, 50-1300m in the Mediterranean). Description Vinculariiform colony. Slender cylindrical branches. Oval, elongate zooids arranged in alternating series. Frontal smooth and finely striated, with a row of marginal pores. A spiramen is situated proximally to the semicircular aperture, which bears a short tubular peristome. Avicularia (1 - 3) and ovicell not observed. remarks T.gracile (Sars) is generally considered as a synonym of T.boreale. Family ascosiiDae Jullien, 1883 Genus Kionidella Koschinsky, 1885 Kionidella excelsa Koschinsky, 1885 (Fig. 11E-F) Kionidella excelsa Koschinsky, 1885: 68, pl. 7, figs 5-12.— Malecki 1963: 133, fig. 58, pl. 15, fig. 1.— Braga 1975, p. 147, pl. 3, fig. 67.— Moissette 1988: 192, pl. 31, figs 4-6.— Zágoršek 2001: 554, pl. 19, fig. 4.— Zágoršek 2003: 176, pl. 28, fig. 6. Fedora excelsa – Waters 1891: 29, pl. 4, fig. 6. occurrence.— Eocene: Germany, Austria, Italy, Slovakia, Hungary, Romania, Poland (Zágoršek 2003). Late Miocene: Algeria (Moissette 1988). Description Small conical, hollow and rather elongated conescharelliniform colonies. Hexagonal zooids arranged in alternating radial series. Smooth convex frontal. Pear-shaped aperture with a large anter separated by two strong cardelles from a smaller poster. One or two well-developed oval avicularia are generally present on the lateral sides of the zooid. Ovicell not observed. remarks As noted by several authors (e.g., Zágoršek 2003), the avicularia are more or less developed and may either occur in pairs, or alone, or lack completely. The ovicells have rarely been observed. 1386 GEODIVERSITAS • 2021 • 43 (26) Moissette P. et al. A D E F BC fig. 11. — A-C, Bryocryptella torquata (Jullien, 1903); A, colony fragment, frontal view, POTB13: AMPG(IV) 2815a; B, colony fragment, dorsal view, POTB13: AMPG(IV) 2815b; C, detail view of zooids, FAN31: AMPG(IV) 3500a; D, Tessaradoma boreale (Busk, 1860), colony fragment, FAN28: AMPG(IV) 3613. E-F, Kionidella excelsa Koschinsky, 1885; E, colony fragment, basal view, FAN7: AMPG(IV)3570a; F, frontal view of a few zooids, FAN16: AMPG(IV) 3572. Scale bars: A, 500 µm; B, E, 200 µm; C, F, 100 µm; D, 1 mm. 1387 Bryozoan faunas at the Tortonian-Messinian transition GEODIVERSITAS • 2021 • 43 (26) Superfamily conescharellinoiDea Levinsen, 1909 Family BatoporiDae Neviani, 1900 Genus Batopora Reuss, 1867 Batopora rosula (Reuss, 1847) (Fig. 12A-G) Cellepora rosula Reuss, 1847: 78, pl. 9, fig. 17; 1867: pl. 1, fig. 7ac; pl. 2, fig. 1a-c. Batopora rosula – Manzoni 1877: 54, pl. 2, fig. 6.— Malecki 1963: 135, pl. 15, fig. 4.— Braga 1975: 147, pl. 3, figs 10-11.— Cook& Lagaaij 1976: 351, pl. 3, figs 2-3; pl. 4, figs 1-2.— Moissette etal. 1993: 113, figs 7g-i.— Haddadi-Hamdane 1996: 121, pl. 2, fig. 4.— Moissette 1996: 193, figs 1A-C.— Pizzaferri& Braga 2000: 61, fig. 2.— Zágoršek 2010b: 168, pl. 83, fig. 4. Lacrimula sp. – Moissette 1988: 193, pl. 31, figs 7-9. occurrence.— Eocene: Italy, Romania (Malecki 1963; Braga 1975). Early Miocene: Spain, Corsica, Italy, Malta (Moissette 1996). Middle Miocene: Austria, Hungary (Moissette etal. 2006), Czech Republic (Zágoršek 2010b), northern Italy, Calabria, Malta (Moissette 1996). Late Miocene: Algeria (Moissette 1988), Sardinia, Sicily, Calabria (Pizzaferri& Braga 2000), Malta (PM, pers. obs.), Crete (Moissette etal. 1993). Pliocene: Spain, Algeria (Haddadi-Hamdane 1996), northern Italy, Sicily, Crete (PM, pers. obs.), Karpathos (PM, pers. obs.). Pleistocene: Rhodes (PM, pers. obs.). This fossil species has exclusively been recorded from the Mediterranean-Paratethys realm (Moissette 1996). Three present-day Batopora species are known from the Indo-Pacific at depths between 285 and 880m (Cook& Lagaaij 1976; Hayward& Cook 1979). Description Small conical conescharelliniform colonies with a flattened base an apical tube comprised of kenozooids and terminated by a small pit. Hexagonal zooids arranged in concentric alternating series. Frontal convex with fairly large pores. Large circular aperture located in the distal part of each zooid. No avicularia. Rare small broken hyperstomial ovicells are visible (Fig. 12A). remarks A few juvenile colonies are present in a fair number of samples. This was also observed by several authors (Cook& Lagaaij 1976; Pizzaferri& Braga 2000). Family orBituliporiDae Canu& Bassler, 1923 Genus Orbitulipora Stolickza, 1862 Orbitulipora excentrica Seguenza, 1880 (Fig. 12H-I) Orbitulipora excentrica Seguenza, 1880: 130, pl. 12, figs 22-22a.— Neviani 1900: 188, pl. 17, figs 15-16.— Waters 1919: 90, text-fig. 2a-c.— Rosso& Sanfilippo 1991: 202, pl. 1, figs 1-5; pl. 2, figs 1-8.— Moissette etal. 1993: 113, figs a-c. occurrence.— Late Eocene: Italy (Waters 1919). Early Miocene: Sardinia (Rosso& Sanfilippo 1991). Late Miocene: Calabria (Rosso& Sanfilippo 1991), Crete (Moissette etal. 1993). Orbitulipora and the orbituliporiform morphotype (fossil and Recent) are considered as deep-water markers and living representatives as typical of muddy bottoms (Cook 1981; Rosso& Sanfilippo 1991). Description Discoidal bilaminar morphology (orbituliporiform). A short kenozooidal tube occurs at the apical part of each colony. Subcircular to subhexagonal zooids arranged in irregular concentric series and progressively increasing in size from the apex to the base. Frontal convex with relatively large pores. Very large circular aperture located in the centre of each zooid. No avicularia. A few ovicells (or their scars) are visible at the growing edge of some colonies. They are hyperstomial, spherical, as large as a zooid and perforated by pores similar to those of the zooecial frontal. DISCUSSION taphonomy Most bryozoans (and other skeletal organisms) are well preserved, except a number of yellowish, abraded fragments of littoral species (e.g., Adeonella polystomella, Margaretta cereoides, Reteporella sp., Smittina cervicornis, and Steginoporella montenati) indicating transport from shallow to deep waters. Examples of such transported material are found in three sections but only in a few beds (among which Potamida B13; Kapariana CAP18; Keramoutsi KER22, KER28), where numerous shallow-water bryozoan remains are found together with fewer deeper water elements. palaeoenvironmental significance of the late miocene Bryozoans of crete Depth A general trend is observed in all studied sections (Figs 2-5). Starting in the late Tortonian, a relatively deep basal assemblage (deep circalittoral) is later replaced by an upper bathyal one. A shallowing-upward sequence follows, with deeper circalittoral conditions succeeded by shallow circalittoral to infralittoral ones. A slight dissimilarity is observed in the Keramoutsi section with very shallow settings (infralittoral-lagoonal) at the base, overlain by shallow circalittoral environments and later the same sequence as in the other three sections. This pattern (Fig. 6), within different sub-basins, is interpreted as resulting mostly from the stepped closure of the Betic-Rifian corridors. It is validated by the analysis of the associated benthic faunas of foraminifera, bivalves, and ostracods (Moissette etal. 2018). However, slight differences are noticed between the two results with the recognition in the present paper of shallower environments (deep circalittoral) at the base of all sections. Correlations between the provided planktonic foraminiferal biostratigraphic zonation and the palaeoenvironmental scheme based on bryozoan (and other) faunas are difficult. 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