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Revista de la Sociedad Geológica de España 29 (1)

ISSN (versión impresa): 0214-2708 ISSN (Internet): 2255-1379

LATE APTIAN CORALS FROM THE SOUTH IBERIAN SUB-BASIN (CRETACEOUS; EASTERN SPAIN)

Corales del Aptiense superior de la subcuenca Suribérica (Cretácico; Este de España)

Hannes Löser1 and Patrick Zell2

1 Estación Regional del Noroeste, Instituto de Geología, Universidad Nacional Autónoma de México, Blvd. Luis Donaldo Col. Los Arcos, Colosio S/N y Madrid, 83250 Hermosillo, Sonora, Mexico; [email protected] 2Institut für Geowissenschaften, Universität Heidelberg, Im Neuenheimer Feld 234, 69120 Heidelberg, Germany; [email protected]

Abstract: A small coral fauna from the Aptian Calizas con Rudistas del Caroch Formation 18 km NNE of Requena (Valencia) is described. A total of 21 species in 11 genera of the suborders Ar- cheocaeniina, Caryophylliina, Faviina, Meandrinina, Rhipidogyrina, Microsolenina, and Styli- nina are reported. For the genus Angelismilia the range of its stratigraphic distribution is extended. For the formerly monospecific genus Nudacolumastrea another two species are descri- bed. The studied fauna is typical for the Early Cretaceous; almost all species were indicated in the early Aptian, but less in the Albian or Cenomanian. Only one genus has a range beyond the Cenomanian. The fauna is dominated by plocoid colonies (genera Cryptocoenia, Holocystis and Nudacolumastrea; 13 species) whereas cerioid, flabelloid, meandrinoid, phaceloid, and solitary forms are subordinated. Palaeobiogeographic relationship exists with the lower Albian of Mont- mell Formation in East Iberia and various Mexican faunas of Aptian to Albian age.

Key-words: Corals, Spain, Cretaceous, Taxonomy, Aptian, Calizas con Rudistas del Caroch For- mation.

Resumen: Se describe una pequeña fauna de corales del Aptiense superior de las Calizas con Rudis- tas de la Formación Caroch. El afloramiento se sitúa a 18 km al NNE de Requena (Valencia). En total, se encontraron 21 especies en 11 géneros de los subórdenes Archeocaeniina, Caryophylliina, Faviina, Meandrinina, Rhipidogyrina, Microsolenina, y Stylinina. Para el género Angelismilia se extendió su rango estratigráfico. Por otro lado, para el género Nudacolumastrea, que estaba definido como mo- noespecifico, se describen dos especies más. La fauna estudiada es típica del Cretácico Inferior; casi todas las especies fueron atribuidas al Aptiense inferior, aunque se encontraron unas pocas caracte- rísticas del Albiense o Cenomaniense. Sólo un género tiene una distribución que sobrepasa el Ceno- maniense. La fauna esta dominada por colonias plocoides (género Cryptocoenia, Holocystis y Nudacolumastrea; 13 especies) mientras que las colonias cerioides, flabeloides, meandrinoides, fa- celoides, y corales solitarios son menos abundantes. Existen relaciones paleobiogeográficas con la fauna del Albiense inferior de la Formación Montmell en el este de Iberia y con otras faunas de Mé- xico de edad Aptiense y Albiense.

Palabras claves: Corales, España, Cretácico, Taxonomía, Aptiense, Calizas con Rudistas de la Formación Caroch

Löser, H. y Zell, P. (2016): Late Aptian corals from the South Iberian Sub-Basin (Cretaceous; Eastern Spain). Revista de la Sociedad Geológica de España, 29(1): 3-20.

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The late Aptian is a critical time span for coral evolution. Pacific (Japan; Löser et al., 2002) and the Iberian Peninsula. Coral diversity increased gradually from the Hauterivian on to Apart from these areas, isolated faunas are known from nort- reach a diversity peak during the early Aptian (Löser, 2013b; hern Africa (Algeria; Masse and Chiki-Aouimeur, 1982), the Löser et al., 2013a), but the oceanic Anoxic Event 1a during Tatra Mts (Poland, Morycowa and Lefeld, 1966; for stratigra- the early Aptian and subsequent demise of the carbonate plat- phy see Masse and Uchman, 1997), and Greece (Löser, 2013a). forms limited coral growth and diversity substantially (Skelton Late Aptian coral fauna from the Iberian Peninsula are repor- and Gili, 2012). ted mainly from North and East Iberia by various authors (Al- Whereas early Aptian coral faunas were distributed world- loiteau, 1947; Schöllhorn, 1998; Tomás et al., 2008; wide, late Aptian coral faunas are limited to three larger areas: Bover-Arnal et al., 2012). Compared to the early Aptian, the the Western Hemisphere (Texas, USA; Puebla and Sonora, Me- total number of faunas is low. Here we present another late Ap- xico; Löser and Minor, 2007; Löser et al., 2013b), the Western tian coral fauna from the southern part of the Iberian Range.

Fig. 1.- Map of the Iberian Peninsula with inset of the Embalse del Buseo region, at Chera village, Valencia province (a; simplified after Instituto Geográfico Nacional de España, 2014) and Aptian palaeogeographic map of Tethyan/eastern Atlantic regions (b; map based on Masse et al., 2000 and Skelton and Gili, 2012). The studied section is located on the eastern flank of the Pico Ropé peak at the Camino de Chelva (39°37'14.6"N 0°58'23.69"W).

Revista de la Sociedad Geológica de España, 29(1), 2016 Hannes Löser and Patrick Zell 5

Study area

The South Iberian sub-basin was formed during Late Juras- sic-Early Cretaceous times as a consequence of increased tec- tonic activity and block tectonics affecting the Iberian Basin situated in Central Spain (Van Wees and Stephenson, 1995). As a result of abrupt relative sea-level changes, the upper Barremian to upper Albian succession of the Iberian Basin reflects a large va- riety of depositional environments including fluvial, deltaic and transitional facies with intercalated marine wedges (e.g. Van Wees et al., 1998). The study area lies about 18 km NNE of Requena (Valencia), north of the village Chera, on the north-western margin of the Parque Natural Geológico de Chera-Sot de Chera (Fig. 1a). The studied section (Fig. 2) is situated on the eastern flank of the Pico Ropé peak at the Camino de Chelva. The Pico Ropé peak pro- vides an almost horizontal series of Barremian to Turonian well- exposed succession of about 300 m in thickness. In the Chera region, including the Pico Ropé locality, the Ap- tian interval is represented by deltaic and shallow marine sedi- ments of the Calizas con Rudistas del Caroch Formation, deposited on the western margin of the South Iberian sub-basin (García, 1977; Arias et al., 1979; Mas, 1981; Fig. 1b). The up- permost member of the formation, the Calizas del Buseo Mbr, is sandwiched between siliciclastic-dominated strata of the under- lying Arenas y Arcillas del Burgal Mbr (deltaic, Gargasian- Clansayesian; e.g., Vilas et al., 1982) and the overlying Calizas, Margas y Areniscas de Sacaras Fm (lagoonal, Albian; e.g., Meléndez and López-Gómez, 2003). Shallow marine fossils of the Calizas del Buseo Mbr, mainly rudistid bivalves of the genus Toucasia, oysters and benthic foraminifers, are abundant in several horizons of the succession at the Pico Ropé peak. Lithologies are dominated by organic- rich, thin- to medium bedded wacke- to grainstones. Three in- tercalated marly intervals rich in charcoal are present between the base and the middle part of the succession. The corals described here derive from a fossil-rich, 0.3 m- thick limestone with a bioclastic grainstone matrix of the Calizas del Buseo Mbr, immediately above 2 m-thick marls (Fig. 2). The diverse fauna represents an in situ record of a rudist, coral and brachiopod dominated biostrome; gastropods and oysters are less represented. High Corg. and siliciclastic contents as well as char- coal layers present throughout the entire succession indicate that the shallow marine environment under consideration was affected by phases of lowered oxygenation at the sea-floor and by a pro- nounced deltaic complex. The preservation is good; fossils are preserved three-dimensionally without evidence of deformation and erosion due to transportation. Gastropod shells are preserved; bivalves exhibit both their valves. Former aragonitic shell was re- crystallized to calcite. Assemblaged rudists (Polyconites verneuili Bayle in De Verneuil et al., 1860; Horiopleura lambertiDouvillé, 1889; Toucasia sp.) and microfossils in underlying (Salpingopo- rella muehlbergi Lorenz, 1902; Charentia cuvillieri Neumann, 1965) and overlying (Cuneolina parva Henson, 1948) beds indi- cate a late Aptian age for the horizon under consideration.

Fig. 2.- Sedimentary column of the Lower Cretaceous at Pico Ropé peak. The coral fauna described here was collected from a fossili- ferous horizon from the upper part of the member (arrow). 1: conglomerate; 2: crossbedded conglomerate; 3: bedded sandstone; 4: crossbedded sandstone; 5: clay; 6: sandy limestone; 7: silty limestone; 8: crossbedded sandy limestone; 9: marl; 10: limestone.

Revista de la Sociedad Geológica de España, 29(1), 2016 6 LATE APTIAN CoRALS

Methods clmax, large lumen; clmin, small lumen; Coral specimens were cut and polished. Thin sections in crd, distance of calicular series; both transversal and longitudinal orientation were prepared crw, width of calicular series; where possible. Thin sections were scanned by passing light s, number of radial elements in adult calices; through them and using a flat bed scanner with an optical res- sd, density of radial elements. olution of 6,400 dpi. Scanned images were then transferred to Measurements in millimetres. The number of the measured grey scale bit maps. Their quality was amended by histogram specimen is indicated above the values. contrast manipulation (contrast stretching) where possible. To gain more insight into the intraspecific variation of fos- Suborder Archeocaeniina Alloiteau, 1952 sil corals and to obtain a better strategy for comparing species, calicular dimensions were systematically measured. To Family Actinastraeidae Alloiteau, 1952 achieve statistical significance, the largest number of possible Actinastraeopsis Sikharulidze, 1977 measurements was taken. This number was mainly deter- Actinastraeopsis birleyoe (Gregory, 1899) mined by the size and quality of the thin section and the size of the single calices in relation to the size of the thin sections. Material For each type of measurement (calicular diameter and BSPG 2014 XVIII 13; 2 thin sections. distance, width and distance of calicular row) in one thin Synonymy section the following values were obtained: *v 1899 Cladophyllia Birleyoe Gregory, p. 457, text-fig. 1 n number of measurements min-max lowest and v 2013c Actinastraeopsis birleyoe – Löser, fig. 1.1-1.2 highest measured value [here more detailed synonymy] µ arithmetic mean (average) σ standard deviation Dimensions v coefficient of variation according to K Pearson (13) µ±σ first interval c min 4.7 Thin sections were measured and values were calculated c max 5.8 using the Palaeontological Database System PaleoTax, module cl min 5.8 PaleoTax/Measure (http://www.paleotax.de/measure). Charac- cl max 4.8 ters visible on the fossils were compared against those on spe- s 34 cimens in world-wide fossil coral collections and an associated image database (ca. 23,000 specimens, ca. 11,200 illustrated, Remarks located in the Estación Regional de Noroeste (ERNo), Sonora, The species was only found as isolated fragments. Mexico). Data storage and processing were carried out using occurrence the PaleoTax database program. To compare the studied fauna Late Barremian to early Aptian (Gerhardtia sartousiana with other coral faunas outside the study area, a computer data- to Deshayesites forbesi zones) of Germany (Bayern) All- base of about 2,700 world-wide coral localities with coral indi- gäuer Helvetikum, Mitteleck. Early Aptian (Prode- cations was used. For details see also Löser (2013b) and Löser shayesites fissicostatus and Deshayesites forbesi zones) of and Minor (2007). The herein studied material is stored at the UK (Isle of Wight) Atherfield. Early Aptian of Venezuela Bavarian State Collection of Palaeontology and Geology (Mu- (Sucre) Cumaná, Las Cinco Ceibas. Earliest Albian nich, Germany) under the signature 2014 XVIII. (Leymeriella tardefurcata Zone) of Spain (Catalunya, Ta- rragona) Comarca del Baix Penedés, Municipi del Mont- Taxonomy mell, Marmellà, Can Xuec. Middle Albian of Mexico (Sonora) Municipio San Pedro de la Cueva, Tepache, Lam- Collection abbreviations are as follows: pazos area, Espinazo de Diablo (ERNo L-134201). BSPG, Bayerische Staatssammlung für Paläontologie und Geologie, München, Germany; Suborder Caryophylliina Vaughan and Wells, 1943 ERNo, Universidad Nacional Autónoma de México, Insti- tuto de Geología, Estación Regional de Noroeste, Hermosillo, Family Caryophylliidae Dana, 1846 Mexico; MGSB, Museo Geológico del Seminario de Barcelona, Spain; Angelismilia Reig, 1988 USNM, United States National Museum, Washington, Remarks D.C., USA. Angelismilia is clearly a caryophylliid genus with compact septa. Septal perforations do not exist, but numerous pali. The following abbreviations are used describing the di- The genus is revised (Löser, 2015a). mensions of the corals: c, outer calicular diameter; Angelismilia portisi (d'Angelis d'ossat, 1905) c max, larger outer calicular diameter; Fig. 3.1 c min, smaller outer calicular diameter; Material ccd, distance between calicular centres; BSPG 2014 XVIII 25; 2 thin sections.

Revista de la Sociedad Geológica de España, 29(1), 2016 Hannes Löser and Patrick Zell 7

Fig. 3.- S1, Angelismilia portisi (d'Angelis d'ossat, 1905), BSPG 2014 XVIII 25, transversal thin section. 2, Plesiosmilia renevieri (Koby, 1896), BSPG 2014 XVIII 43, transversal thin section. 3-5, Nudacolumastrea montsiani (Bataller, 1936), BSPG 2014 XVIII 31; 3, transversal thin section, 4, transversal thin section, detail 5, longitudinal thin section. 6-8, Nudacolumastrea stefani Löser and Zell, 2015, Holotype of Nudacolumastrea stefani, BSPG 2014 XVIII 19; 6, transversal thin section, 7, transversal thin section, detail, 8, lon- gitudinal thin section. 9-10, Nudacolumastrea sp., BSPG 2014 XVIII 24; 9, transversal thin section, 10, transversal thin section, detail, 11, Aulastraeopora harrisi (Wells, 1932), BSPG 2014 XVIII 4, transversal thin section. Scale bar 1mm.

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Fig. 4.- 1-3, Diplogyra arasensis (Alloiteau, 1946-47), BSPG 2014 XVIII 21; 1, transversal thin section, 2, transversal thin section, de- tail, 3, longitudinal thin section. 4-6, Diplocteniopsis sp., BSPG 2014 XVIII 14; 4, transversal thin section, 5, transversal thin section, detail, 6, longitudinal thin section. 7-9, Montlivaltoides ngariensis He and Xiao, 1990, BSPG 2014 XVIII 17; 7, transversal thin section, 8, transversal thin section, detail, 9, longitudinal thin section. 10-12, ?Thalamocaeniopsis sp., BSPG 2014 XVIII 27; 10, transversal thin section, 11, transversal thin section, detail, 12, longitudinal thin section. Scale bar 1mm.

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Synonymy Remarks *v 1905 Trochosmilia Portisi Angelis d'ossat, p. 223, pl. The morphology of the material corresponds completely 15, figs. 2 a-f to the type species but differs by smaller calicular dimension. v 2015a Angelismilia portisi (d'Angelis d'ossat, 1905) – The assignation of this species (that was originally described Löser, p. 29, figs. 1, 2 from the early Aptian of Catalunya) to Nudacolumastrea, and the finding of the same species in the early Albian of Sonora Dimensions (Mexico) allows a range extension of the genus. (25) occurrence c 18.2-24 Early Aptian of Spain (Catalunya, Tarragona) Comarca del s 110 Montsià, Municipi d‘Ulldecona. Early Albian of Mexico (Sonora) Municipio Ures, Cerro de oro (ERNo L-4840). Description Solitary turbinate coral. Calicular outline elliptical. Septa com- Nudacolumastrea sp. pact, in cross section thick close to the wall, thinner toward the Figs. 3.9-10 centre. Symmetry of septa radial and regularly hexameral. Cy- Material cles of septa subregular. Septal cycles differ in length and thick- BSPG 2014 XVIII 24; 2 thin sections. ness. First three septal cycles extended to the calicular centre, later cycles are shorter. Septa rarely connected to each other. Septal la- Dimensions teral face with thorns, inner margin slightly swollen in places. Pali (24) n min-max µ σ cv µ±σ present, at least on four septal cycles in up to three crowns. Most cl min 35 3.03-4.45 3.74 0.34 9.2 3.40-4.09 septa of the first two cycles are connected to the columella. Costae cl max 35 3.55-5.58 4.44 0.52 11.9 3.91-4.97 hardly present. Synapticulae absent. Columella composed of va- ccd 40 3.37-5.18 4.22 0.41 9.9 3.80-4.63 rious lamellae. Endotheca consists of numerous small dissepi- s 24 ments. Wall compact, septothecal. Epitheca present. Remarks Remarks This is the first indication of the genus outside of its type area, The morphology of the material corresponds com- the Montmell Formation (lowermost Albian) of East Iberia. Un- pletely to the type species but differs by larger calicular di- fortunately the only specimen is fragmentarily preserved. mension. occurrence Earliest Albian (Leymeriella tardefurcata Zone) of Spain Nudacolumastrea stefani Löser and Zell, 2015 (Catalunya, Barcelona) Comarca del Alt Penedés, Municipi del Figs. 3.6-8 Castellví de la Marca, Can Pascual; Comarca del Baix Penedès, Material Municipi Masllorenç, Masarbones, field W (ERNo L-7247). BSPG 2014 XVIII 6, 7, 16, 18, 19, 34, 37; 15 thin sections. Synonymy Suborder Faviina Vaughan and Wells, 1943 *v 2015 Nudacolumastrea stefani Löser and Zell, p. 162, figs. 6.1-3 Family Columastraeidae Alloiteau, 1952 Dimensions σ σ Nudacolumastrea Löser and Zell, 2015 (19) n min-max µ cv µ± Remarks cl min 20 2.71-3.78 3.24 0.28 8.7 2.95-3.52 Critical review of type material helped to improve the knowl- cl max 20 3.02-4.04 3.50 0.27 7.9 3.22-3.78 edge on the newly introduced genus. The genus is not monospe- ccd 20 3.46-4.46 3.93 0.26 6.6 3.67-4.19 cific as originally assumed. There exists named and undescribed s 24 material with a range from the late Aptian to the early Albian. Remarks Nudacolumastrea montsiani (Bataller, 1936) The species was discussed in detail in Löser and Zell Figs. 3.3-5 (2015). Material BSPG 2014 XVIII 31; 2 thin sections. Family Eugyridae Felix, 1903 Synonymy *v 1935 Phyllocoenia Montsiani Bataller, p. 196 Diplogyra Eguchi, 1936 Diplogyra arasensis (Alloiteau, 1947) Dimensions (31) n min-max µ σ cv µ±σ Figs. 4.1-3 cl min 7 1.83-2.90 2.48 0.41 16.6 2.06-2.89 Material cl max 6 2.61-3.92 3.13 0.50 16.2 2.62-3.64 BSPG 2014 XVIII 21; 2 thin sections. ccd 11 2.33-3.54 2.99 0.43 14.6 2.55-3.43 Synonymy s 24 *v 1947 Eugyra arasensis Alloiteau, p. 197, pl. 2, fig. 4, text-fig. 2

Revista de la Sociedad Geológica de España, 29(1), 2016 10 LATE APTIAN CoRALS v 2013b Diplogyra arasensis (Alloiteau, 1946-47) – Löser, ral face with thorns, inner margin T-shaped or swollen. Pali p. 11, figs. 4g-i [here detailed synonymy] or paliform lobes absent. Some septa may be attached to the columella. Costae present but short. Synapticulae ab- Dimensions sent. Columella lamellar, discontinuous. Endotheca con- (21) n min-max µ σ cv µ±σ sists of tabulae. Wall subcompact, parathecal. Coenosteum crw 15 1.73-2.66 2.21 0.29 13.1 1.92-2.50 absent. crd 15 3.30-4.86 3.95 0.42 10.7 3.52-4.38 Remarks sd 6/5mm The present material differs from the type species of the genus by distinct calices. All other formally described Description species comes from one outcrop and are very probably sy- Meandroid colony. Calicular rows long and straight. nonymous with the type species. Corallites indistinct. Septa compact, in cross section thick close to the wall, thinner toward the centre. No septal sym- Plesiosmiliids (informal group) metry. Septa not connected to each other. Septal lateral face Remarks smooth (probably due to preservation), inner margin Intention and constitution of the formal group is dis- swollen in places. Pali and paliform lobes absent. Costae cussed in Löser (2013b). present, non-confluent. Synapticulae and columella absent. Endotheca consists of numerous tabulae. Wall compact, Montlivaltoides He and Xiao, 1990 parathecal. Coenosteum broad, consists of costae and exothecal dissepiments. Budding intracalicinal, polysto- Remarks modeal. Montlivaltoides is a poorly defined genus. Although occurrence Xinyi He provided photographs of the type material of See Löser (2013b). the type species (M. typicus), the morphology did not become clear because the material is very poorly pre- Suborder Meandrinina Alloiteau, 1952 served. The type material is strongly recrystallized. Therefore the microstructure and the calicular centre Family Phyllosmiliidae Felix, 1903 could not be clearly deciphered. It is possible that the genus represents a phaceloid coral that has a structure Diplocteniopsis Zlatarski, 1968 comparable to Plesiosmilia, lacking a columella. The Remarks genus ranges from the Aptian to Cenomanian. Apart The type material of Diplocteniopsis curvicalix from Montlivaltoides there exists a yet unnamed genus Zlatarski, 1968, type species of Diplocteniopsis was avai- that is almost identical but has a lamelar columella as in lable for study and also topotypical material from the type Plesiosmilia. locality. The genus is one of the earliest member of the Phyllosmiliidae family. It differs from the similar Phrag- Montlivaltoides ngariensis He and Xiao, 1990 mosgyra Reig oriol, 1994 by a columella that is vertically Figs. 4.7-9 not continuously developed. The genus was originally as- Material signed to the family Diplocteniopsidae Zlatarski, 1968 that BSPG 2014 XVIII 17; 2 thin sections. becomes a junior synonym of the Phyllosmiliidae. Synonymy * 1990 Montlivaltoides ngariensis He and Xiao, p. 155, Diplocteniopsis sp. 249, pl. 18, fig. 8, pl. 17, fig. 7 Figs. 4.4-6 Material Dimensions BSPG 2014 XVIII 14; 2 thin sections. (17) c 12.4-12.8 Dimensions s 48 (14) c 10-12 Description s 11-22 Phaceloid colony. Calicular outline irregular circular. sd 7/5mm Septa compact. Microstructure of small-sized trabeculae, septa with a median dark line. Septa in cross section Description slightly thicker close to the wall, becoming slightly thinner Solitary flabelloid coral with distinct calices. Septa toward the centre. Symmetry of septa irregular, but three compact, in cross section slightly thicker close to the wall, size orders can be distinguished. Septal generations differ becoming slightly thinner toward the centre. Symmetry of in length and thickness. Septa not connected to each other. septa irregular, but two to three size orders can be distin- Septal lateral face smooth, inner margin smooth. Pali and guished. Cycles of septa subregular. Septal generations dif- paliform lobes absent. Costae present. Synapticulae, colu- fer in length and thickness. Septa of the first cycle rarely mella, and wall absent. Endotheca consists of numerous connected to each other in the calicular centre. Septal late- tabulae.

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Fig. 5.- 1-3, Cryptocoenia aguilerai (Reyeros Navarro, 1963), BSPG 2014 XVIII 26; 1, transversal thin section, 2, transversal thin section, detail, 3, longitudinal thin section.4-6, Cryptocoenia annae (Volz, 1903), BSPG 2014 XVIII 20, 4, transversal thin section, 5, transversal thin section, detail, 6, longitudinal thin section. 7-9, Cryptocoenia atempa (Felix, 1891), BSPG 2014 XVIII 29, 7, transver- sal thin section, 8, transversal thin section, detail, 9, longitudinal thin section. 10-12, Cryptocoenia steinmanni (Fritzsche, 1924), BSPG 2014 XVIII 8, 10, transversal thin section, 11, transversal thin section, detail, 12, longitudinal thin section. Scale bar 1mm.

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Fig. 6.- 1, Holocystis cieszynica (Kolodziej and Gedl, 2000), BSPG 2014 XVIII 23, 1, transversal thin section, 2, transversal thin section, detail, 3, longitudinal thin section. 4-6, Holocystis dupini (d'orbigny, 1850), BSPG 2014 XVIII 30, 4, transversal thin section, 5, trans- versal thin section, detail, 6, longitudinal thin section. 7-9, Holocystis magnicanaliculata (Prever, 1909), BSPG 2014 XVIII 2, 7, trans- versal thin section, 8, transversal thin section, detail, 9, longitudinal thin section. 10-12, Holocystis nomikosi Löser, 2006, BSPG 2014 XVIII 38, 10, transversal thin section, 11, transversal thin section, detail, 12, longitudinal thin section. Scale bar 1mm.

Revista de la Sociedad Geológica de España, 29(1), 2016 Hannes Löser and Patrick Zell 13 occurrence tre. Symmetry of septa irregular. Cycles of septa irregular, but Aptian of China (Xizang [= Tibet] Autonomous Re- size orders can be distinguished. Septal generations differ in gion) Rutog county, Risum district, Jaggang, Qiekan. Late length and thickness. Septa of younger generations often con- early Albian (Douvilleiceras mammillatum Zone) of Spain nected to septa of preceding ones. Septal lateral face with pen- (Cantabria, Santander) Cala de Islares (ERNo L-133150). nulae and thorns, inner margin smooth. Pali or paliform lobes absent. Costae absent. Synapticulae occasional, mainly in the Plesiosmilia Milaschewitsch, 1876 space between corallites. Columella small, substyliform. En- Plesiosmilia renevieri (Koby, 1896) dotheca absent. Wall subcompact, synapticulothecal. Coenos- Fig. 3.2 teum absent. Budding extracalicinal. Material Remarks BSPG 2014 XVIII 43; 1 thin section. The material differs from the genus by much more per- Synonymy forated septa. *v 1896 Pleurosmilia Renevieri Koby, p. 18, pl. 2, fig. 5 Suborder Rhipidogyrina Roniewicz, 1976 Dimensions (43) Family Aulastraeoporidae Alloiteau, 1957 c 7.1-7.2 s 30 Aulastraeopora Prever, 1909 Aulastraeopora harrisi (Wells, 1932) Description Fig. 3.11 Solitary turbinate coral. Calicular outline circular. Septa Material compact, in cross section slightly thicker close to the wall, BSPG 2014 XVIII 4; 1 thin section. becoming slightly thinner toward the centre. Symmetry of Synonymy septa radial and regularly hexameral. Cycles of septa sub- *v 1932 Blothrocyathus harrisi Wells, p. 242, pl. 30, figs. 6, 6 regular. Septal cycles differ in length and thickness. Septa a, 7, pl. 31, figs. 3, 4 rarely connected to each other. Septal lateral face smooth v 2013b Aulastraeopora harrisi (Wells, 1932) – Löser, p. (probably due to preservation), inner margin smooth. Pali 31, figs. 10jk [here more detailed synonymy] or paliform lobes absent. Two opposite septa of the first cycle are attached to the columella. Costae and synapticu- Dimensions lae absent. Columella lamellar. Wall not compact, proba- (4) bly epithecal. c 12.7-13.3 occurrence s 24 Late Barremian to early Aptian of France (Haute-Savoie) Bonneville, Reignier. Early Aptian of Venezuela (Sucre) Description Cumaná, Las Cinco Ceibas (USNM I-312010). Cenomanian Solitary cylindric coral. Calicular outline circular. Septa (Neocardioceras juddii Zone) of France (Aude) Les Cor- compact, in cross section slightly thicker close to the wall, bières, Sougraigne, Prat-Périé (BSPG 2011 XXVI 120). becoming slightly thinner toward the centre. Symmetry of septa radial and regularly hexameral. Cycles of septa regu- Suborder Microsolenina Morycowa and Roniewicz, 1995 lar. Septal cycles differ in length and thickness. Septa not connected to each other. Septal lateral face with thorns, Family Leptophylliidae Vaughan, 1905 inner margin with auriculae in places. Pali or paliform lobes absent. Lonsdaleoid septa present. Costae, synapticulae, co- Thalamocaeniopsis Alloiteau, 1954 lumella absent. Wall compact, has the same structure as ?Thalamocaeniopsis sp. septa, is therefore septothecal. Figs. 4.10-12 occurrence Material Compare to Löser (2013b) for details. BSPG 2014 XVIII 19#2, 2014 XVIII 27; 4 thin sections. Suborder Stylinina Alloiteau, 1952 Dimensions (27) n min-max µ σ cv µ±σ Family Solenocoeniidae Roniewicz, 2008 cl min 10 4.77-8.09 6.15 0.91 14.8 5.24-7.06 Remarks cl max 10 7.34-11.23 9.26 1.41 15.2 7.85-10.68 The family Cyathophoridae should no longer be used, ccd 12 4.65-9.23 7.00 1.43 20.5 5.57-8.44 because the type material of the type species of the name s 8 51-76 61.50 7.98 12.9 54-69 giving genus Cyathophora does not present the charac- teristics ascribed to it. There is no other family name Description available for the moment except the Solenocoeniidae (see Cerioid colony. Calicular outline polygonal. Septa regu- also Löser 2015b). The study published by Morycowa larly perforated. Microstructure of large trabeculae. Septa in and Roniewicz (2016) is mainly based on material from cross section thick close to the wall, thinner toward the cen- Poland and Switzerland, and almost not on the poorly

Revista de la Sociedad Geológica de España, 29(1), 2016 14 LATE APTIAN CoRALS

Fig. 7.- 1-2, Cryptocoenia miyakoensis (Eguchi, 1936), BSPG 2014 XVIII 12, 1, transversal thin section, 2, transver- sal thin section, detail. 3-4, Holocystis elegans (Lonsdale, 1849), BSPG 2014 XVIII 33, 3, transversal thin section, 4, longitudinal thin section. Scale bar 1mm.

conserved type material of Cyathophora richardi *v 963 Procyathophora aguilerai Reyeros Navarro, p. 8, Michelin, 1843, type species of Cyathophora. It is ob- pl. 3, figs. 3, 5 vious that Cyathophora sensu stricto has long septa that v 2013b Cryptocoenia aguilerai (Reyeros Navarro, 1963) are connected to each other, whereas the material in- – Löser, p. 31, figs. 11a-c [here detailed synonymy] vestigated by Morycowa and Roniewicz (2016) has very short septa that are not connected to each other. The Dimensions conclusions made for the family taxon Cyathophoridae (26) n min-max µ σ cv µ±σ are meaningless. cl min 10 2.27-3.32 2.81 0.33 11.9 2.47-3.14 cl max 10 2.56-3.79 3.06 0.39 12.9 2.66-3.46 Cryptocoenia orbigny, 1849 ccd 10 3.18-4.39 3.83 0.39 10.3 3.43-4.22 Description s 24 Plocoid colony. Calicular outline circular. Septa compact. Symmetry of septa radial and regularly hexa- Remarks meral. Cycles of septa regular. Septal cycles differ in The species has very short septa. Septal cycles cannot almost length and thickness. Septa not connected to each other. not be distinguished. The costae are short and non-confluent. Septal lateral face smooth, inner margin smooth. Pali occurrence and paliform lobes absent. Costae present, confluent or See Löser (2013b) for details. sub-confluent. Synapticulae and columella absent. En- dotheca consists of numerous tabulae and some dis- Cryptocoenia annae (Volz, 1903) sepiments. Wall compact, parathecal. Coenosteum Figs. 5.4-6 narrow, consists of costae and tabulae. Budding extra- Material calicinal. The various species do differ much in their BSPG 2014 XVIII 20; 2 thin sections. general morphology, only in their dimensions. There- Synonymy fore detailed descriptions are not provided for all 1857 Cyathophora antiqua – Fromentel, p. 42 species. *1903 Cyathophora Annae Volz, p. 26 [18], pl. 4, figs. 9-13 v 1963 Cyathophora atempa Felix 1891 – Reyeros Navarro, Cryptocoenia aguilerai (Reyeros Navarro, 1963) p. 8, pl. 4, figs. 7, 8 Figs. 5.1-3 v 1981 Pseudocoenia annae (Volz 1903) – Turnšek and Mi- Material hajlovic, p. 15, pl. 10, fig. 5 BSPG 2014 XVIII 11, 2014 XVIII 26; 3 thin sections. v 2003 Cyathophora haysensis Wells, 1932 – Baron-Szabo et Synonymy al., p. 208, pl. 37, fig. 7, pl. 39, fig. 2

Revista de la Sociedad Geológica de España, 29(1), 2016 Hannes Löser and Patrick Zell 15

Dimensions Material (20) n min-max µ σ cv µ±σ BSPG 2014 XVIII 29; 2 thin sections. cl min 16 1.79-2.74 2.22 0.28 12.6 1.94-2.51 Synonymy cl max 16 2.28-3.41 2.91 0.38 13.1 2.53-3.29 *v 1891 Cyathophora atempa Felix, p. 155, pl. 25, figs. 7, 8 ccd 12 3.03-4.14 3.62 0.36 10.1 3.25-3.98 v 2010 Cryptocoenia atempa (Felix, 1891) – Löser, p. s 24 591, fig. 3.4 [detailed synonymy here]

Remarks Dimensions The septa are very short and the septal cycle cannot be (29) n min-max µ σ cv µ±σ differentiated. The costae are unknown. cl min 11 1.52-2.32 1.83 0.26 14.3 1.57-2.10 occurrence cl max 11 1.72-2.54 2.19 0.29 13.4 1.89-2.48 Early Hauterivian (Acanthodiscus radiatus Zone) of ccd 20 2.36-4.03 3.05 0.40 13.1 2.65-3.45 France (Haute-Marne) Saint Dizier. France (Yonne) Gy- s 24 l'Evêque, fields SW Gy-l'Evêque (BSPG 2003 XX 6401). Barremian of Mexico (Puebla) Tehuacán, San Antonio Remarks Texcala (ERNo L-4412). Aptian of Mexico (Puebla) San The septa are very short and the septal cycle cannot be Juan Raya; Serbia (East Serbia) Zljebine. Early Aptian differentiated. The costae are unknown. (Palorbitolina? lenticularis Zone) of Romania (Suceava) occurrence Pojorîta area, Cîmpulung-Moldovenesc, Valea Seaca. Late See Löser (2010) for details. Aptian to Albian of Iran (Esfahan) Esfahan Basin, Dizlu. Early Cenomanian (Mantelliceras dixoni zone) of Spain Cryptocoenia miyakoensis (Eguchi, 1936) (Cantabria, Santander) Cobreces, Luaña playa (BSPG Figs. 7.1-2 2007 V 366). Material BSPG 2014 XVIII 12; 2 thin sections. Cryptocoenia atempa (Felix, 1891) Synonymy Figs. 5.7-9 *v 1936 Miyakopora miyakoensis Eguchi, p. 70, figs. 4, 4 a v 1976 Cyathophora miyakoensis (Eguchi 1936) – Turnšek Fig. 8.- Stratigraphic distribution and abundance of species. The and Buser, p. 11, 38, pl. 1, figs. 3-5 thickness of the bars indicates the number of localities in which v 1992 Cyathophora pygmaea Volz 1903 – Turnšek et al., the concerned species was found (see scale). Grey bar indicates p. 213, pl. 5, figs. 1, 2 the stratigraphic range of the study area.

Revista de la Sociedad Geológica de España, 29(1), 2016 16 LATE APTIAN CoRALS v 1998 Adelocoenia fontserei (Bataller 1947) – Schöllhorn, Cryptocoenia steinmanni (Fritzsche, 1924) p. 74, pl. 18, fig. 7, pl. 27, figs. 2-4 Figs. 5.10-12 Material Dimensions BSPG 2014 XVIII 3, 2014 XVIII 8; 3 thin sections. (12) n min-max µ σ cv µ±σ Synonymy cl min 15 0.97-1.44 1.19 0.12 10.7 1.07-1.32 *v 1924 Cyathophora steinmanni Fritzsche, p. 316, pl. 3, cl max 15 1.08-1.74 1.43 0.18 13.0 1.24-1.61 fig. 8, pl. 4, fig. 3 ccd 20 1.26-1.99 1.62 0.24 15.0 1.38-1.86 v 2013b Cryptocoenia almerai (d'Angelis d'ossat, 1905) – s 6 Löser, p. 31, fig. 10l [here detailed synonymy] v 2014 Cryptocoenia steinmanni (Fritzsche, 1924) – Löser, Remarks p. 47, fig. 7j The species presents only one septal cycle with rather long septa. The costae are not visible. The specimen is very Dimensions small and did not allow to provide a longitudinal thin sec- (8) n min-max µ σ cv µ±σ tion. cl min 25 1.14-1.69 1.42 0.13 9.2 1.29-1.55 occurrence cl max 25 1.35-1.75 1.59 0.12 7.8 1.46-1.71 Cretaceous of Slovenia (West Slovenia) Banjska ccd 30 1.51-2.59 1.97 0.27 13.7 1.70-2.24 planota, Levpa. Early Valanginian of Spain (Valencia, s 6 Alicante) oliva, Elca (BSPG 2003 XX 4768). Early late Aptian of Spain (Cataluñnya, Lérida) Comarca del Alt Remarks Urgell, Municipio de Cabó, Senyús section; Spain The species present only one septal cycle. The septa are (Catalunya, Lérida) Comarca de La Noguera, Municipio short, but well visible. The costae are short and non-confluent. de Vilanova de Meià, Montsec de Rubies, section NW occurrence La Cabrua quarry (BSPG 2003 XX 6312). Late Aptian of See Löser (2013b) for details. Spain (Murcia) Jumilla, Solana del Sopalmo (MGSB 74393). Late Aptian to middle Albian of Slovenia (South Holocystis Lonsdale, 1849 Slovenia) Kocevje region, Slovenski vrh, horizon A-C. Description Latest Aptian of Japan (Iwate-ken) Miyako-shi. Earliest Plocoid colony. Calicular outline irregular circular. Albian (Leymeriella tardefurcata Zone) of Spain Septa compact. Symmetry of septa radial and regularly (Catalunya, Barcelona) Comaca del Alt Penedès, Cas- tetrameral. Cycles of septa subregular. Septal cycles dif- tellvi de la Marca, Can Pascual, section loc. 2 (ERNo fer in length and thickness. Septa not connected to each L-6295). Early Albian of Mexico (Sonora) Municipio other. Septal lateral face smooth, inner margin smooth. Cucurpe, Cucurpe, La Mesa (ERNo L-4278). Early Pali and paliform lobes absent. Costae present, confluent Cenomanian (Mantelliceras mantelli Zone) of Germany or sub-confluent. Synapticulae and columella absent. En- (Nordrhein/Westfalen) Mülheim/Ruhr, Kassenberg dotheca consists of numerous tabulae and some dissepi- (BSPG 2003 XX 1181). ments. Wall compact, parathecal. Coenosteum narrow,

Fig. 9.- Former stratigraphic distribution of genera of the studied fauna. Ranges according to Löser (2005) and unpublished data. Ju- rassic not shown because of uncertainties about the ranges. Grey bar indicates the stratigraphic range of the studied fauna.

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Fig. 10.- Correlation of provinces with joint species of the study area. Provinces with less than two joint species are suppressed. The Correlation Ratio coefficient was applied. For details of calculation see Löser and Minor (2007). consists of costae and tabulae. Budding extracalicinal. Holocystis dupini (orbigny, 1850) The various species do not differ much in their general Figs. 6.4-6 morphology, only in their dimensions. Therefore detailed Material descriptions are not provided for all species. BSPG 2014 XVIII 30; 2 thin sections. Synonymy Holocystis cieszynica (Kolodziej and Gedl, 2000) * 1850 Tetracoenia Dupiniana orbigny, (2), p. 121 Figs. 6.1-3 1988 Holocystis bukowinensis Voltz, 1903 – Kuzmicheva Material and Aliev, p. 157, pl. 1, fig. 4 BSPG 2014 XVIII 23; 1 thin section. Synonymy Dimensions *v 000 Nowakocoenia cieszynica Kolodziej and Gedl, p. (30) n min-max µ σ cv µ±σ 187, figs. 3, 8, 9 cl min 25 1.40-2.36 1.92 0.20 10.8 1.71-2.12 v 2003 Holocystis elegans (Lonsdale, 1849) – Baron-Szabo cl max 25 1.71-2.81 2.19 0.25 11.7 1.94-2.45 and González León, p. 204, figs. 6A, B ccd 35 1.94-2.93 2.39 0.23 9.7 2.15-2.62 s 10 8-14 11.70 1.82 15.6 10-14 Dimensions (23) n min-max µ σ cv µ±σ Remarks cl min 9 1.69-2.69 2.18 0.30 13.7 1.88-2.48 The septa of the first cycle are rather long and reach cl max 6 2.23-2.97 2.59 0.28 10.9 2.31-2.87 about one third of the calicular diameter. ccd 4 2.47-2.87 2.71 0.18 6.8 2.53-2.90 occurrence s 16 Hauterivian to early Aptian of France (?). Barremian of Azerbaijan (Kubatlinskij) Alikuliushagi. occurrence Late Barremian to early Aptian (Paleorbitolina lenti- Holocystis elegans (Lonsdale, 1849) cularis Zone) of Mexico (Sonora) Municipio Ures, Cerro Figs. 7.3-4 de oro. Late Barremian to early Aptian (Colchidites Material sarasini to Deshayesites forbesi zones) of Poland BSPG 2014 XVIII 33; 2 thin sections. (Slaskie) Bielsko-Biala, Rudzica. Synonymy

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*v 1849 Cyathophora ? elegans Lonsdale, p. 83, pl. 4, figs. 12-15 Remarks v 2006 Holocystis elegans (Lonsdale, 1849) – Löser, p. In comparison to the other species of the study area, H. 294, figs. 1e-g nomikosi has rather thick and well developed septa. occurrence Dimensions Late Barremian to early Aptian (Palorbitolina lenticu- σ σ (33) n min-max µ cv µ± laris Zone) of Mexico (Sonora) Municipio Ures, Cerro de cl min 12 2.14-2.87 2.38 0.19 8.1 2.19-2.58 oro (ERNo L-4378). Early Aptian of Spain (Murcia) cl max 11 2.32-3.35 2.81 0.37 13.2 2.43-3.18 Sierra de la Muela. Early Aptian of Tanzania (Tanganyika, ccd 13 2.21-3.36 2.84 0.37 13.1 2.46-3.21 Mtwara) Nambawala plateau, Pilepile. Early Aptian s 5 8-14 10.80 2.28 21.1 9-13 (Dufrenoyia furcata Zone) of Spain (Vascongadas, Viz- caya) Bilbao, Peñascal. Late Aptian of Greece (Viotía) occurrence Aliartos, Chiarmena. Late Barremian to early Aptian (Palorbitolina lenticula- ris Zone) of Mexico (Sonora) Municipio Ures, Cerro de oro Discussion (ERNo L-4313). Early Aptian (Prodeshayesites fissicostatus and Deshayesites forbesi zones) of UK (Isle of Wight) Ather- The coral fauna shows a relatively low diversity which field point. Early Aptian of Spain (Murcia) Jumilla, Solano is probably due to the reduced thickness of the coral bea- del Sopalmo (MGSB 73675). Early Albian of Mexico ring bed and the low number of samples available. High (Sonora) Municipio Ures, Cerro de oro (ERNo L-4890). Corg.- and increased siliciclastic contents of beds over - and underlying the coral bearing horizon as well as the Holocystis magnicanaliculata (Prever, 1909) small-sized architecture of the assemblage suggest that Figs. 6.7-9 the biostrome was formed during a short period of in- Material creased oxygen availability and lowered deltaic input of BSPG 2014 XVIII 2; 2 thin sections. siliciclastics and organic matter. Deltaic influenced, sha- Synonymy llow marine environments with lowered oxygenation and *v 1909 Polytremacis magnicanaliculata Prever, p. 68, pl. high siliciclastic input represent difficult living condi- 1, figs. 20, 22 a tions especially for corals, which are sensitive to external physico-chemical changes (e.g., Dupraz and Strasser, Dimensions 2002). With 13 species, the fauna is dominated by plo- (2) coid corals with generally low calicular diameter (less cl min 1.2-1.6 than 4 mm) of the genera Cryptocoenia, Holocystis and s 4 (8) Nudacolumastrea. Ceroid (?Thalamocaeniopsis), flabe- lloid (Diplocteniopsis), meandrinoid (Diplogyra), Remarks phaceloid (Actinastraeopsis, Montlivaltoides), and soli- The only specimen is very small and did not provide tary corals (Angelismilia, Aulastraeopora, Plesiosmilia) large thin sections that would allow systematic measure- are subordinated. ments. The reported species have a limited stratigraphical dis- occurrence tribution that ranges from the Valanginian to the Ceno- Early Aptian of Italy (Abruzzi, L'Aquila) Monti d'ocre, manian (Fig. 8). Except for one, all species are also Fossa Cerasetti. Early late Aptian of Spain (Catalunya, reported from the early Aptian, and the majority of them Lérida) Comarca de La Noguera, Montsec de Rubies, sec- also in the early Albian. The genera have a slightly wider tion NW La Cabrua quarry (BSPG 2003 XX 4200). range, most being known from the Aptian to early Albian (Fig. 9). Except for Plesiosmilia, no genus reaches the Holocystis nomikosi Löser, 2006 Turonian. So the fauna has a typical Early Cretaceous com- Figs. 6.10-12 position. The genus Angelismilia that was hitherto endemic Material to the Montmell Formation and restricted to the early Al- BSPG 2014 XVIII 9, 35, 38; 6 thin sections. bian (Leymeriella tardefurcata Zone) experienced a range Synonymy extension. v 1926 Cyathophora sp. – Dietrich, p. 66, pl. 6, fig. 3 The palaeobiogeographic relationships are poor *v 2006 Holocystis nomikosi Löser, p. 295, figs. 1j-l (Fig. 10). Although the present fauna shares 18 species v 2013c Holocystis dupini – Löser et al., figs. 5kl with other provinces, the number of species that it v 2013a Holocystis nomikosi Löser, 2006 – Löser, p. 110, shares with each province, is low (see Löser and Minor figs. 6.10-6.12 2007 for more explication). It shares most species with the early Albian of East Iberia, mainly encompassing Dimensions the lower Albian Montmell Formation W and SW of (38) n min-max µ σ cv µ±σ Barcelona, the early Albian of the Bisbee Basin in cl min 25 1.48-2.13 1.90 0.18 9.7 1.71-2.08 Northern Mexico, the Aptian of the Puebla Basin in cl max 25 1.78-2.35 2.05 0.12 5.9 1.93-2.18 Central Mexico, and early Aptian of the Pelagonium ccd 30 2.06-3.10 2.65 0.26 9.9 2.38-2.91 (Greece). s 8-10

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Acknowledgements gion central de la provincial de Valencia y noreste de la provin- cial de Albacete. Seminarios de Estratigrafía, Serie Mono- The authors acknowledge the late Stefan Götz, Yvonne grafías, 1: 1-334. Spychala and Enric Pascual-Cebrian for their support du- Gregory, J.W. (1899): New species of Cladophyllia, Prionastraea ring fieldwork and Felix Schlagintweit for microfossil iden- and Stylina. Annals and Magazine of natural History, (7), 4, 19: 457-461. tification. Financial support for this research was provided by the Heidelberg University Graduate Academy (LGFG He, Xinyi and Xiao, Jin-dong (1990): Jurassic and Cretaceous hexacorals of Ngari area. In: Paleontology of Ngari, Tibet 2012-9; to P.Z.). A grammatical check was carried out by (Xizang), (Yang Zunyi and Nie Zetong, Eds). China Univer- Proof-Reading-Service (Letchworth Garden City, En- sity Geoscience Press, Beijing, 146-159. gland). Preparation of thin sections in the ERNo labora- Koby, F. 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(1849): Notes on fossil Zoophytes found in the de- Angelis d'ossat, G. de (1905): Coralli del Cretacico inferiore della posits described by Dr.Fitton in his Memoir entitled "A strati- Catalogna. Palaeontographia Italica, 9: 169-251. graphical account of the section from Atherfield to Rocken Arias, C., Mas, J.R., García, A., Alonso, A., Vilas, L., Rincón, R. End". Quarterly Journal of the Geological Society of London, and Meléndez, N. (1979): Les faciès urgoniens et leurs varia- 5: 55-103. tions pendant la transgression aptienne occidentale de la Löser, H. (2005): Stratigraphy of Cretaceous coral genera. Neues Chaîne Ibérique (Espagne). Geobios Mémoire Spécial, 3: 11- Jahrbuch für Geologie und Paläontologie, Abhandlungen, 23. 238: 231-277. Baron-Szabo, R.C. and González León, C. M. (2003): Late Ap- Löser, H. 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