101urn:lsid:arphahub.com:pub:DF2AD546-A4F0-51AC-B38C-88216742E553urn:lsid:zoobank.org:pub:9B96196F-9505-457E-8F01-0E12D644C289ZittelianaZITT2512-53382747-8106Pensoft Publishers10.3897/zitteliana.97.113796113796Research ArticleAnthozoaPalaeozoologyCretaceousAlpsMiddle Cenomanian coral fauna from the Roßsteinalmen (Northern Calcareous Alps, Bavaria, Southern Germany) – a revised and extended versionLöserHannes1Writing - original draftWernerWinfriedwerner@snsb.de2Writing - original draftDargaRobert3Writing - original draftEstación Regional del Noroeste, Instituto de Geología, Universidad Nacional Autónoma de México, Blvd. Luis Donaldo Colosio S/N y Madrid, 83250 Hermosillo, Sonora, MexicoUniversidad Nacional Autónoma de MéxicoHermosilloMexicoSNSB – Bayerische Staatssammlung für Paläontologie und Geologie and GeobioCenterLMU, Richard-Wagner-Strasse 10, D-80333 München, GermanySNSB – Bayerische Staatssammlung für Paläontologie und Geologie and GeobioCenterLMUMünchenGermanyNaturkunde- und Mammut-Museum Siegsdorf, Auenstraße 2, D-83313 Siegsdorf, GermanyNaturkunde- und Mammut-Museum SiegsdorfSiegsdorfGermany
Corresponding author: Winfried Werner (werner@snsb.de)
Academic editor: A. Nützel
20232012202397891470B2F9DF8-6A61-5518-B1D4-4DBB56689406D4564419-3213-4D38-96BB-E7CFE157E0F8104250370610202306112023Hannes Löser, Winfried Werner, Robert DargaThis is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.http://zoobank.org/D4564419-3213-4D38-96BB-E7CFE157E0F8
In the Northern Calcareous Alps, relics of a formerly widely distributed shallow marine facies belonging to the Branderfleck Formation (upper Albian to lower Turonian) crop out and contain locally abundant corals. The fauna described here derives from Middle Cenomanian sediments. This study complements a former revision. In total, the fauna includes 98 species in 46 genera, belonging to 16 scleractinian superfamilies and two octocorallian families. One species – Enallheliaoctasepta – is described as new. The fauna that was formerly located at the northern part of the Apulian plate (Austroalpine unit), south of the Penninic Ocean, shows palaeobiogeographic relationships to Cenomanian faunas from the Basque-Cantabrian Basin, the Prebetic zone, the Pelagonium, and the Quillan Basin, indicating stronger connections to Tethyan rather than Boreal faunas. The fauna also shares species with Aptian and Albian, but also with Late Cretaceous faunas of the Gosau Basin. Eight genera experienced a range extension; five genera have their last occurrence in the Middle Cenomanian fauna, and three genera have their first occurrence. Although the fauna presents numerous genera that became widespread in the Late Cretaceous, its generic composition is more closely related to late Early Cretaceous corals than to post-Cenomanian corals. The faunal turnover at the Cenomanian/Turonian boundary was not marked by the sudden appearance of new faunal elements, but rather by the disappearance of taxa. Some faunal elements that constitute post-Cenomanian faunas already existed in the Cenomanian, but they were very rare. The increase of taxa after the Cenomanian/Turonian boundary took place during the Coniacian and Santonian.
Only shortly after the appearance of the first publication on the corals from the Northern Calcareous Alps (Löser et al. 2013c), additional material from the same locality was discovered in the Bayerisches Landesamt für Umwelt - Geologie collections, and some further specimens were found in the collections of the Berliner Naturkundemuseum. The material in Berlin was collected by Josef Bruckmair and given to Werner Quenstedt (1893–1960) who worked as geologist and palaeontologist at the Friedrich-Wilhelms-Universität in Berlin. The Munich specimens come from the same collector, Otto Hölzl, who also sold and donated material to the Bayerische Staatssammlung für Paläontologie und Geologie. Surprisingly, the material recently discovered at the Bayerisches Landesamt für Umwelt - Geologie encompasses material that is taxonomically different from that described in Löser et al. (2013c). It is impossible to speculate about the reasons. One reason may be that some individual picked out the material (that is now at the Bayerisches Landesamt für Umwelt - Geologie, formerly Bayerisches Geologisches Landesamt) from a larger collection, and that the Bayerische Staatssammlung für Paläontologie und Geologie has received the rest of the material. Within the new collection, there are very attractive and well-preserved specimens, whereas, the material at the Bayerische Staatssammlung für Paläontologie und Geologie is, for a large part, represented by bulk samples. Among the new material there are multiple genera and species that are not yet reported. In this compilation, we include, therefore, all species, and also the material described in Löser et al. (2013c), but species described in the first paper are not figured, described and provided with dimensions. The discussion encompasses all species.
Our first contribution to the Middle Cenomanian coral fauna from the Northern Calcareous Alps was published more than ten years ago, and since then much more data about Cenomanian coral faunas became available. In Löser (2014b), the Boreal coral fauna from the Elbtal-Group (Saxony, Germany) of Cenomanian age was revised and completed with more data. In Löser (2015a), the Cenomanian coral fauna from the Le Mans area was briefly revised. Löser and Bilotte (2017) described a Cenomanian platy coral association from Southern France. Löser et al. (2018) report a lower Cenomanian coral fauna from Central Greece with nearly 80 species. Löser and Callapez (2022) present small coral faunas from the Cenomanian/Turonian boundary of Portugal. Löser and Wilmsen (2022, 2023) recently began publication of a lower Cenomanian coral fauna from Cantabria (Spain) that encompasses around 140 coral species. Another contribution that helps to link the Cenomanian coral faunas to other upper Cretaceous faunas constitutes the taxonomic revision of the Coniacian-Santonian coral fauna from the Gosau Basin (Löser et al. 2019), with descriptions and illustrations of 360 coral species. The new taxonomic data shed a different light on Cenomanian coral faunas and the evolution of this group during the Cretaceous.
Geological setting
The Roßsteinalmen locality is situated in the Alpine Mountains about 50 km south of Munich, 9 km SE of the Lenggries community and 5 km W of the Kreuth community (Fig. 1). At this locality, a small series of mainly siliciclastic Cretaceous sediments crop out, which were for a long time generally called ”Cenoman Serie” (e.g., Boden 1935). Outcrops of this lithostratigraphic unit are, generally, restricted to relatively small areas in the Northern Calcareous Alps. They represent relics of a formerly more widely distributed facies, which have been protected from erosion by their tectonic position within synclinal structures. The ”Cenoman Serie” is subdivided into two different formations, (1) the Losenstein Formation of Middle Albian to Lower Cenomanian age, consisting of silty marls, turbitic sandstones and deep water conglomerates with exotic pebbles, and (2) the Branderfleck Formation, consisting mainly of marls, calcareous sandstones, and breccias (?Upper Albian to Turonian; Gaupp 1980, 1982; Weidich 1984a, b; Faupl and Wagreich 2000). The Roßsteinalmen area belongs, tectonically to the northern part of the Lechtal nappe and here the Branderfleck Formation overlies Upper Jurassic radiolarites and limestones with slight angular unconformity (Steinberg 1980; Weidich 1984b).
Location of the section Roßsteinalmen and tectonic units. NCA, Northern Calcaerous Alps, F + H, Flysch and Helvetic unit, MOL, Molasse. From Löser et al. (2013c).
https://binary.pensoft.net/fig/955649
The section at the Roßsteinalmen is about 150 m thick and consists of sandy marls and sandstones alternating with thick carbonaceous fine breccias (Fig. 2; Steinberg 1980; Weidich 1984b). The upper part is characterised by thick, coarse chaotic breccias that are interpreted by Weidich (1984b) as channel deposits cutting the marly series. The components of the breccias are mainly Triassic and Jurassic carbonates, as well as ”Cenomanian” breccias and, therefore, they reflect synorogenetic reworking of local material (e.g. Boden 1935; Kuhn 1991; Steinberg 1980). For the base, micropalaeontological data, based on planctonic foraminifera (e.g., Rotaliporacushmani (Morrow, 1934)), indicate a Middle to lower Upper Cenomanian age. For the upper part (centre of the synclinal fold), the upper Cushmani zone and thus, an Upper Cenomanian age is confirmed (Weidich 1984b). The marls and sandstones of the section are partly rich in tests of orbitolinid foraminifera (Steinberg 1980). This important index fossil family did not reach the Late Cenomanian (Schroeder and Neumann 1985). The only species known from the Middle Cenomanian is Conicorbitolinaconica (d‘Archiac, 1837), which was also indicated in the Branderfleck Fm by Schlagintweit and Wagreich (2005). Orbitolinids were collected in the marls but thin sections revealed their poor state of conservation. The marls and sandstones are, therefore, of a Middle Cenomanian age.
Palaeogeographically, the Branderfleck Formation represents sediments deposited at the northern margin of the Austroalpine unit and, thus, at the southern border of the Penninic Ocean (see Dercourt et al. 2000; Faupl and Wagreich 2000; Pfiffner 2010). For the clastics of the lower Branderfleck Formation, palaeocurrent data give evidence of a source area located toward the south (Gaupp 1980). The uplift of the northern border of the Austroalpine unit may be linked to the formation of an accretionary wedge in the context of the nascent subduction of the Penninic Ocean (Pfiffner 2010). According to Weidich (1984b), the sediments at the Roßsteinalmen were deposited in water ranging from about 50 m deep on the inner shelf (base of the section) to about 100 m on a middle shelf area (upper part of the section).
The corals described in the first part (Löser et al. 2013c) come from the lower part of the section (Fig. 2). Only specimen 2012 X 1 originates from a higher level and probably has a lower Upper Cenomanian age. The labels of the new coral material described and depicted for the first time in this study do not indicate the exact position within the Roßsteinalmen section; however, based on the preservation and a general rareness of fossils in the upper part of the section, these corals certainly do all come from the lower part of the section.
Material and methods
In total, 240 coral specimens were included in this extended study; 220 of them could be identified at the species level. Nearly 200 thin sections in both transversal and longitudinal orientation were prepared. The coral material varies in its state of preservation, with exceptionally well-preserved specimens and others that are strongly recrystallised or fragmented.
Section at the Roßsteinalmen. Planktonic foraminifera stratigraphy after Weidich (1984a, b), occurrence of corals and orbitolinid foraminifers after Steinberg (1980), Weidich (1984b), and personal observations. Scale only valid for the lower part of section. From Löser et al. (2013c).
https://binary.pensoft.net/fig/955650
Thin sections were scanned by passing light through them using a flatbed scanner with an optical resolution of 6,400 dpi. Scanned images were then transferred to grey scale bit maps. Their quality was amended by histogram contrast manipulation (contrast stretching) where possible.
To gain more insight into the intraspecific variation of fossil corals and to obtain a better strategy for comparing species, corallite dimensions of each specimen were systematically measured. To achieve statistical significance, the largest number of possible measurements was taken. This number was mainly controlled by the size and quality of the thin section and the size of the single corallites in relation to the size of the thin sections. Septa were counted for numerous corallites where the septal symmetry was not regular (in corals with a regular symmetry the number of septa is the same in all corallites). For each type of measurement (corallite diameter and distance, width and distance of corallite row) and count (principally septal counts) in one thin section, the following values were obtained:
n number of measurements or counts;
min–max lowest and highest measured or counted values (mm for measurements);
µ arithmetic mean (average);
s standard deviation;
cv coefficient of variation according to K. Pearson
µ±s first interval.
Measurements are always in millimetre. Thin sections were measured and values were calculated using the Palaeontological Database System PaleoTax, module PaleoTax/Measure (https://www.paleotax.de/measure); for details on the mathematical background, see Löser (2012b). Morphometric data of the corals were compared against the morphometric data of specimens in worldwide fossil coral collections, and an associated image database. The database encompasses approximately 28,920 coral specimens from Triassic to modern forms. Approximately 8,200 of them are type specimens, and 16,800 specimens are illustrated. The database is located in the Estación Regional de Noroeste (Instituto de Geología, UNAM), Sonora, Mexico. Data storage and processing were carried out using the PaleoTax database program (Löser 2004).
To compare the studied fauna with other coral faunas outside the study area, a computer database of about 3,100 worldwide coral localities with coral indications was used (Löser et al. 2002, 2005). To simplify the analysis, localities of the same age, belonging to the same basin, on the same continental margin or the same interoceanic platform, were grouped together into one palaeo-province (a type of large faunule, sensu Johnson 2007). Altogether, this produced 440 provinces, reaching from the Jurassic into the Palaeogene. Only firmly dated localities were assigned to a province in order to ensure that the subsequent analysis was valid, and that the studied locality was not included in any existing province. For the study area, an independent province was created to allow a clear comparison between the existing provinces and the new material. Interregional comparisons were carried out between the new province and existing provinces having at least three species in common with the fauna of the studied area. The comparisons were carried out using, exclusively, specimens that were available to the first author (H.L.), but not the indications in the literature. The literature does not offer precise morphometric data, that are necessary to separate species and to compare them to each other. For details, see also Löser (2008), and Löser and Minor (2007). Data analysis, statistics, and the creation of charts were all carried out using the Database System PaleoTax and the graphic module PaleoTax/Graph (www.paleotax.de). The material is kept in the Bayerisches Landesamt für Umwelt - Geologie/Paläontologie (Hof), the Bayerische Staatssammlung für Paläontologie und Geologie (Munich), and the Museum für Naturkunde der Humboldt-Universität (Berlin), all of which are located in Germany.
Systematic description
The distribution data (as reflected in the synonymy lists) are almost entirely based on material examined by the first author (H.L.). Material only mentioned in the literature, material not available, or insufficiently described and/or poorly illustrated in the literature, was not taken into account. To obtain better insight into the distribution patterns of the studied coral fauna, much unpublished material was included. Therefore, distribution data indicated under ‘Other occurrences’ could also be provided for species remaining in open nomenclature. As mentioned above, we include here all species, and also those species that are described in detail in Löser et al. (2013c), but measurements, descriptions, remarks, and illustrations are not repeated.
In scleractinian corals, the separation of superfamilies, families, and genera is based on qualitative characteristics, whereas the species are separated on the basis of morphometric data (corallite dimensions, septal counts, and septal density). The species of one genus are, more or less, alike and differ only in their measurements. For this reason descriptions of species are not given here. We provide for the taxonomic levels from superfamily to genus detailed descriptions, without repeating characteristics in lower levels. When a superfamily is characterised by large trabeculae, perforate septa, and the presence of synapticulae, this is also the case for all families and all genera. There are rare exceptions, but these are mentioned in the descriptions. When a superfamily encompasses only one family, the description is not repeated with the family.
The abbreviations used in the synonymy lists follow Matthews (1973): *, earliest valid publication of the species name; non, the citation is excluded from the species; p, the described material belongs only in part to the species concerned; v, the specimen was observed by the first author (H.L.). The abbreviation cf indicates that the present species is similar to the species indicated in the concerned reference, but the assignment remains uncertain. A year in italics indicates that the quotation is provided with neither a description nor an illustration.
The following abbreviations are used to describe the dimensions of the corals: c, corallite diameter (outer diameter); ccd, distance between corallite centres; clmax, large lumen; clmin, small lumen; cmax, larger outer corallite diameter; cmin, smaller outer corallite diameter; crd, distance of corallite series; crw, width of corallite series; md, distance between crests in a hydnophoroid colony.
Collection abbreviations are as follows: BSPG, Bayerische Staatssammlung für Paläontologie und Geologie, München, Germany; LFU, Bayerisches Landesamt für Umwelt - Geologie, Hof, Germany; NHMW, Naturhistorisches Museum, Wien, Austria; MNHN, Muséum National d’Histoire Naturelle, Paris, France; PU, Museo di Geologia e Paleontologia dell’ Università di Torino, Torino, Italy; MB, Museum für Naturkunde der Humboldt-Universität, Berlin, Germany.
Class Anthozoa Ehrenberg, 1834
Subclass Hexacorallia Haeckel, 1866
Order Scleractinia Bourne, 1900
Fossil scleractinian corals are described since the end of the 18th century. Since the late 20th century, they are examined using thin sections, leading to the discovery of hitherto unknown morphological elements and to changes in the classification. During former systematic revisions (Vaughan and Wells 1943; Alloiteau 1952; Wells 1956; Alloiteau 1957), the order Scleractinia was subdivided into suborders and families. The subdivision into suborders is now, up to 80 years later, not considered practical for various reasons, as explained by Löser et al. (2018), Löser and Callapez (2022), and Löser and Wilmsen (2023). Therefore, the classification system that was introduced by Löser (2016c) does not apply suborders, but rather applies superfamilies which group families together. This practical approach has the advantage that superfamilies are covered by the International Code of Zoological Nomenclature (ICZN 1999) and follow the type principle. This is not the case for the taxonomic level of orders, including suborders. Superfamilies clearly refer to a family, genus, type species, and a type. The characteristics of a superfamily are thus connected to a physical specimen that more precisely limits the characteristics of this higher rank taxon. Practically, suborders are, at the present time, replaced by superfamilies. Currently, 27 superfamilies with 56 families (or informal groups) are distinguished, that range from the Middle Jurassic to the Palaeogene, partly to the Neogene, and extant (Löser 2016c). Contrary to the former classification system based on suborders, the superfamilies may constitute monophyletic groups. The basic characteristics for the distinction of the superfamilies is the size of the trabeculae relative to the septa. Further distinction is based on the presence or absence of synapticulae and septal perforation.
Cerioid, phaceloid, or plocoid colonies. Septa compact, mostly in a regular radial symmetry. Septa often connected to each other. Lateral faces with thorns, upper margin granulated. Microstructure of septa of medium-sized trabeculae. Lonsdaleoid septa present in one genus, main septa absent. Synapticulae absent. Pali in some genera present, columella in most genera. Endotheca varies, generally made of thin tabulae. Wall compact or subcompact, mainly septothecal. Marginarium in one genus. Coenosteum generally present, consists of isolated trabeculae or costae. Budding extracalicinal and intracalicinal.
Family <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Actinastreidae</tp:taxon-name-part></tp:taxon-name> Alloiteau, 1952AnimaliaScleractiniaActinastreidae32921E1F-0D14-59B2-BD88-C7C7B384921DActinastread’Orbigny, 1849Type species.
Actinastreagoldfussi d’Orbigny, 1850.
Description.
Plocoid colony with narrow coenosteum that is made of large isolated trabeculae. Corallites circular or polygonal. Septa in a regular radial symmetry, rarely bilateral. Septa of the first and second generation can be connected to the columella, younger septa can be connected to older septa. Columella styliform and large. No pali.
AnimaliaScleractiniaActinastreidae68BA5BCB-1D57-5357-939A-99047AE3F7D6ActinastrealimbataAlloiteau, 1954b*1954bActinastrealimbata nov. sp. – Alloiteau: 89, text-fig. 17, pl. 2, fig. 5, pl. 10, fig. 7. v2013Stelidioserisminima (de Fromentel, 1857) – Löser, Werner and Darga: 44, pl. 1, figs 3, 4. v2019Actinastrealimbata Alloiteau, 1954 – Löser, Heinrich and Schuster: 42, figs 50a–c. Material.
BSPG 1947 XVI 26, 1947 XVI 61; two thin sections.
Remarks.
The material does not belong to the genus Stelidioseris because it shows isolated trabeculae in the coenosteum as it is typical for Actinastrea. Therefore, the species name has been changed, compared to our publication from 2013.
Other occurrences.
Upper Turonian of the Western Tethys (France), Coniacian to Santonian of the Central Tethys (Austria), lower Campanian of the Western Tethys (France), lower Maastrichtian of the Western Atlantic (Mexico).
AnimaliaScleractiniaActinastreidaeBFBA24FE-7124-569B-8B65-2B434CBDDA2FActinastreapolygonataAlloiteau, 1954bPlate 1: figs 1–3*v1954bActinastreapolygonata nov. sp. – Alloiteau: 43, pl. 4, fig. 11, pl. 7, fig. 5. v1989Actinastreaschizoformis nov. sp. – Reig Oriol: 21, pl. 2, fig. 4, pl. 6, fig. 2. v2019Actinastreapolygonata Alloiteau, 1954 – Löser, Heinrich and Schuster: 45, figs 54a–c. Material.
LFU 8336SG015004#1; one thin section.
Dimensions.
(LFU 8336SG015004#1).
n
min–max
µ
s
cv
µ±s
clmin
20
1.25–1.60
1.44
0.09
6.6
1.35–1.54
clmax
20
1.53–2.06
1.70
0.14
8.7
1.56–1.85
ccd
27
1.37–2.02
1.73
0.18
10.6
1.54–1.91
septa
10+10
Other occurrences.
Upper Turonian of the Western Tethys (France), Coniacian to Santonian of the Central Tethys (Austria), Santonian of the Western Tethys (Spain).
AnimaliaScleractiniaActinastreidaeD4C7A2CE-6537-5D08-853F-42EB456DF47EActinastreasubdecaphylla(Oppenheim, 1930)v*1930Astrocoeniasubdecaphylla n. sp. – Oppenheim: 460, pl. 15, fig. 9. 1982Actinastraeasubdecaphylla (Oppenheim) 1930 – Beauvais: (1), 13, figs 1, 2. v2013Actinastrearegularis (de Fromentel, 1887) – Löser, Werner and Darga: 43, pl. 1, figs 1, 2. v2019Actinastrea?subdecaphylla Oppenheim, 1930 – Löser, Heinrich and Schuster: 48, figs 58a–d. Material.
BSPG 1947 XVI 60; one thin section.
Remarks.
In Löser et al. (2013c) this specimen was assigned to Actinastrearegularis (de Fromentel, 1887). After examining the type material of Actinastreasubdecaphylla this assignation was corrected. Both species are similar but differ in the corallite dimensions (see Löser et al. 2019: 40 for comparison).
Other occurrences.
Coniacian to Santonian of the Central Tethys (Austria), upper Santonian of the Western Tethys (France), upper Campanian to lower Maastrichtian of the Arabian Peninsula (United Arab Emirates).
In the Cretaceous only solitary corals. Septa compact, mostly in a subregular radial symmetry. Septa in places connected to each other. Lateral faces with thorns, upper margin smooth. Microstructure of septa of small trabeculae. Lonsdaleoid and main septa absent. Synapticulae present, mainly in the wall. Pali absent, columella by septal fusion. Endotheca generally made of thin tabulae. Wall compact, mainly septothecal with synapticulae. Marginarium absent. Coenosteum generally present, consists of costae. Budding intracalicinal.
Family <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Agariciidae</tp:taxon-name-part></tp:taxon-name> Gray, 1847AnimaliaScleractiniaAgariciidaeABD7BE93-C8F7-511A-AAAB-F67BC134E303AntilloserisVaughan, 1905Type species.
Turbinoseriseocaenica Duncan, 1873.
Description.
Solitary elliptical coral. Septa compact, in a regular radial symmetry. Septa of the first three cycles thicker in the corallite centre and free, septa of further cycles more regular in thickness and connected to each other. Septa lateral faces with thorns, directing to the corallite center. No pali. Columella styliform and small. Enthodeca absent. Wall perforated, with synapticulae.
The assignation of the present specimen to this genus is preliminary. It shares with Antilloseris the arrangement of septa, with the difference that the septa of the first three cycles are, in the present specimen, not thickened in the corallite centre. The columella is parietal and not styliform as in Antilloseris.
Turbinate solitary coral with an elliptical or circular outline. The septa are in a regular radial symmetry, regularly connected to each other, and many of them are fused together in the centre of the corallite to form the columella.
AnimaliaScleractiniaAgariciidae0F3C168E-02D8-51A8-A43F-C50E043FEBF9TrochoseropsisettalensisSöhle, 1897*v1897TrochoseropsisEttalensis – Söhle: 45, pl. 7, figs 2, 2ab. v1958Smilotrochustarraconensis nov. sp. – Alloiteau: 84, pl. 2, figs 7, 8. v2013aTrochoseropsisettalensis Söhle, 1897 – Löser: 16, figs 6b–i. v2013Trochoseropsisettalensis Söhle, 1897 – Löser, Werner and Darga: 50, pl. 3, figs 7–9. Material.
BSPG 1947 XVI 15, 1947 XVI 16, 1947 XVI 24, 1947 XVI 28, 1947 XVI 29, 1947 XVI 73, 1947 XVI 74, 1991 X 63, LFU 8336SG015145, MB K2984#2, K2984#3; ten thin sections.
Other occurrences.
Lower Albian of the Western Tethys (France), lower Cenomanian of the Central Tethys (Germany), Campanian of the Western Tethys (Spain).
Mainly solitary, rarely colonial (phaceloid) corals. Septa compact, regular thickness, mostly in a regular radial symmetry. Septal upper margins smooth, lateral faces with few small thorns or granulae. Lonsdaleoid septa and main septum absent. Microstructure of probably very small trabeculae that can only be traced by a dark line. Synapticulae absent. Pali and columella can be present. The endotheca is variable, but generally poorly developed in solitary forms. Marginarium absent. Wall septothecal. Coenosteum absent.
https://binary.pensoft.net/fig/955651Family <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Caryophylliidae</tp:taxon-name-part></tp:taxon-name> Dana, 1846
The classification of the family is difficult. Their fossil members are generally poorly known, and particularly this is the case for many type species. The reason is that solitary corals have been – in contrast to colonial corals – much less studied using thin sections. Whereas large coral colonies often provide pieces to prepare thin sections, the preparation of thin sections from a solitary coral generally results in its complete loss. The study of solitary corals always demands specimen-rich populations. These populations are not always available. Therefore, not all material shown here can be assigned to a genus.
Coral with an elliptical outline. Septa free, in a regular hexameral symmetry. Twelve septa with very strongly inflated inner margins (that can be interpreted as pali). Columella very large, styliform. Endotheca poorly developed.
LFU 8336SG015099#2, 8336SG015132#1; four thin sections.
Dimensions.
(LFU 8336SG015099#2).
c
14.1×14.6
septa
48
Description.
The solitary coral has a regular septal symmetry with four septal cycles. Only the third cycle bears occasionally pali. The columella consists of some small elements.
Turbinate solitary coral with circular outline. A columella is present, but difficult to distinguish from the septa. No pali. The endotheca is poorly developed.
The coral Parasmiliacentralis is far more than one hundred times cited in the literature. Since most of these citations are not accompanied by proper illustrations and/or corallite measurements and septal counts, a synonymy list cannot be compiled. The distribution data are based on material observed by the first author (H.L.).
Other occurrences.
Cenomanian of the European Boreal (UK), Coniacian to lower Santonian of the Central Tethys (Austria), upper Coniacian to Maastrichtian of the European Boreal (UK, Germany).
Solitary and (cerioid, phaceloid, plocoid) colonial corals. Septa compact and with regular thickness. Septal symmetry radial, regular or sub-regular. Septa often connected to each other. Septal lateral faces with thorns and/or granulae, septal upper margin with fine granulations. Lonsdaleoid septa and main septum absent. Microstructure of medium-sized trabeculae. Synapticulae absent. Pali present in some genera. Columella present in most genera, styliform, lamellar, parietal, or by septal fusion. Endotheca generally present. Marginarium absent. Wall present, septothecal (by septal thickening) or as tabulotheca. Coenosteum present in plocoid genera. Budding extracalicinal (cerioid, plocoid) or intracalicinal (phaceloid).
Phaceloid colony. Corallites with a small diameter (< 6 mm). Septa in a sub-regular radial symmetry; but systems can be recognised. The number of septa increases with the corallite diameter. The septa of the first two cycles have pali. The columella is lamellar.
AnimaliaScleractiniaCladocoridae71FF9D11-5232-5E2F-901C-ED00C619E4E5Procladocorasimonyi(Reuss, 1854)Plate 2: figs 4–6*v1854Cladocora Simonyi – Reuss: 112, pl. 12, figs 5–7. v1930Cladocoralibidinum n. sp. – Oppenheim: 362, pl. 38, fig. 14. v1936Cladocora jamaicaënsisVaughan 1899 – Hackemesser: 38, pl. 5, fig. 3. v2000Procladocorajamaicaensis (Vaughan 1899) – Löser: 52, pl. 3, figs 1–5. Material.
LFU 8336SG015082#3, 8336SG015162; three thin sections.
Dimensions.
(LFU 8336SG015082#3).
c
4.1×5.7
septa
45
Other occurrences.
Lower Albian of the Western Tethys (Spain), upper Cretaceous of the Central Tethys (Greece, Austria).
AnimaliaScleractiniaCladocoridae4CBB3B65-098E-5BEA-B9F1-F19C1D06E080Procladocorasp.Plate 2: fig. 3v1997Pleurocora cf. alternans Milne-Edwards and Haime, 1849 – Baron-Szabo: 77, pl. 10, figs 4, 6. Material.
LFU 8336SG015140#1; four thin sections.
Dimensions.
(LFU 8336SG015140#1).
c
3.8×4.1
septa
28
Other occurrences.
Upper Cenomanian of the Western Tethys (France), lower Coniacian of the Central Tethys (Austria).
Plocoid colonies. Septal symmetry regular radial and mostly hexameral. Pali present in some genera. Columella varies: styliform, styliform and double, lamellar, or absent. Coenosteum with costae.
Plocoid coral with septa in a regular hexameral or decameral symmetry. Columella lamellar or small and styliform. Irregular pali at the first septal cycle, not very pronounced. Coenosteum narrow.
The septal symmetry is particular in this specimen. Whereas in Eocolumastrea the septal symmetry is normally a multiple of six or ten, the present specimen shows varying systems such as eleven and ten.
Plocoid colony with circular or elliptical corallites. Symmetry regular hexameral. The first two septal cycle bear pali; the columella is lamellar and small.
AnimaliaScleractiniaColumastreidaeA3D41761-913D-5A1A-8E2D-69C1CD2313ABNeocoeniaexsculpta(Reuss, 1854)*v1854Astraeaexsculpta – Reuss: 114. v1957 StephanaxophylliaCasterasi nov. sp. – Alloiteau: 73, figs 20, 21, pl. 9, fig. 8, pl. 16, fig. 1. vp2013Neocoenia cf. casterasi (Alloiteau, 1957) – Löser, Werner and Darga: 44. v2019Neocoeniaexsculpta (Reuss, 1854) – Löser, Heinrich and Schuster: 77, figs 107a, b. Material.
BSPG 1947 XVI 33, 1947 XVI 65, 1991 X 73, 1991 X 77, LFU 8336SG015079#1, 8336SG015079#3; nine thin sections.
Remarks.
This material was formerly assigned to Neocoenia cf. casterasi (Alloiteau, 1957). After studying the type material of Neocoeniaexsculpta, the determination could be improved.
Other occurrences.
Upper Turonian to Santonian of the Central Tethys (Austria), Santonian of the Western Tethys (France).
https://binary.pensoft.net/fig/955652AnimaliaScleractiniaColumastreidae1DC63AC5-1C5A-5263-89DA-F4241CFFDB6ANeocoeniakuehnii(Oppenheim, 1930)v*1930Pleurocorakuehnii n. sp. – Oppenheim: 371, pl. 38, fig. 13, pl. 40, fig. 9. 1982Barycorakuehnii (Oppenheim) 1930 – Beauvais: (1), p. 99. vp2013Neocoenia cf. casterasi (Alloiteau, 1957) – Löser, Werner and Darga: 44, pl. 1, figs 7–9. Material.
BSPG 1991 X 48, 1991 X 74, 1991 X 75, 1991 X 76, 1991 X 78, 2012 X 4, LFU 8336SG015005#1; four thin sections.
Remarks.
This material was assigned to Neocoenia cf. casterasi by Löser et al. (2013c). In 2016, it was possible to study the type material of Oppenheim (1930) and to clarify the taxonomy of the present material.
Other occurrences.
Santonian of the Central Tethys (Austria).
AnimaliaScleractiniaColumastreidae3A047321-1716-50F9-99D9-78CA15A501BDNeocoeniarenziHackemesser, 1936v*1936Neocoeniarenzi n. g. n. sp. – Hackemesser: 24, pl. 3, figs 4–6. v2013Neocoeniarenzi (Hackemesser, 1936) – Löser, Werner and Darga: 46, pl. 1, figs 10–12. v2016cNeocoeniarenzi Hackemesser, 1936 – Löser: 471, figs N4a, b. v2019Neocoeniarenzi (Hackemesser, 1936) – Löser, Heinrich and Schuster: 77, fig. 105, figs 108a–c. Material.
BSPG 1947 XVI 10, 1947 XVI 46, 1947 XVI 8, LFU 8336SG015077, 8336SG015079#2; four thin sections.
Other occurrences.
Middle Turonian to Santonian of the Central Tethys (Greece, Austria), upper Campanian of the Western Tethys (Spain), undefined Cretaceous of the Central Tethys (Greece).
Solitary and colonial corals. Septa either with regularly distributed perforations, with perforations only in certain parts of the septa, or almost compact. Septa generally thick, without notable symmetry (except Negoporitidae). Septa often connected to each other. Septal lateral faces with pennulae and thorns. Septal distal margin with large granulae. Both lonsdaleoid septa and main septum absent. Microstructure of large trabeculae. Synapticulae present. Pali in some genera probably present but difficult to distinguish from the perforated inner margins of the septa. Columella poorly defined. Endotheca present or absent. When present, mostly thin tabulae. Marginarium absent. Wall poorly developed. Coenosteum varies depending on the organisation type. Budding varies.
The family encompasses numerous solitary and colonial coral genera that show septa with perforations which are concentrated on the inner margin of the septa. The septa are thinner than in the Synastraeidae and less perforated than in the Microsolenidae. The thickness of septa and the space between them are similar.
Thamnasterioid colony, with mostly only at the inner margin perforated septa, that barely differ in length and thickness. The corallites are regularly distributed, generally not in rows, or only when juvenile. Costae run between all corallites. There is no wall.
AnimaliaScleractiniaLatomeandridae10F0DFAF-B60F-539D-B506-CEC3945C1D07Astraeofungiadecipiens(Michelin, 1846)*1846Astreadecipiens – Michelin: 200, pl. 50, fig. 13. v1891Thamnastraea Crespoi – Felix: 146, pl. 22, fig. 5. v1951Thamnasteriajezoensis Eguchi, n.sp. – Eguchi: 54, pl. 18, figs 5, 6. v1957Astreadecipiens Michelin – Alloiteau: 213, figs 153–155, pl. 3, fig. 3, pl. 14, fig. 5, pl. 18, fig. 6. v1963Thamnasteriacrespoi (Felix) – Reyeros Navarro: 4, pl. 1, figs 1, 4. v1994Thamnasteriacotteaui Fromentel – Liao and Xia: 127, pl. 32, figs 6, 7. v1996Synastrea cf. dubia Fromentel, 1861 – Baron-Szabo and Steuber: 25, pl. 14, figs 1, 7. v2013bAstraeofungiatenochi (Felix, 1891) – Löser: 20, fig. 3.2. v2013Astraeofungiatenochi (Felix, 1891) – Löser, Werner and Darga: 58, pl. 7, figs 4–6. v2016cAstreadecipiens Michelin, 1841 – Löser: 181, figs A50a–c. v2023Astraeofungiadecipiens (Michelin, 1841) – Samaniego-Pesqueira et al.: 123, fig. 5E. Material.
BSPG 1947 XVI 27; two thin sections.
Remarks.
In Löser et al. (2013c) this specimen was assigned to Astraeofungiatenochi (Felix, 1891). Improved and more systematic measurements show that Astraeofungiatenochi has larger dimensions and higher septal counts than the present specimen.
Other occurrences.
Valanginian to Aptian of the Western Atlantic (Mexico), lower Hauterivian of the European Boreal (Germany, France), Barremian of the Western Atlantic (Mexico), lower Aptian of the Western Tethys (France) and Central Tethys (Greece), upper Aptian of the Western Pacific (Japan), Aptian to lower Albian of the Central Tethys (Greece, Hungary), lower Albian of the Western Tethys (Spain) and Western Atlantic (Mexico), upper Aptian to Albian of the Eastern Tethys (Iran), middle Albian of the Western Atlantic (Mexico), middle Cenomanian of the European Boreal (France).
Valanginian to Aptian of the Western Atlantic (Mexico), lower Hauterivian of the European Boreal (France), and the Western Tethys (France), upper Barremian to lower Aptian of the Central Tethys (Switzerland), upper Aptian of North Africa (Algeria), and the Western Tethys (Spain), lower Albian of the Western Atlantic (USA, Mexico), Albian of the Western Tethys (Spain), lower Cenomanian of the Western Tethys (Spain), middle Cenomanian of the European Boreal (France), and upper Cenomanian of the European Boreal (Germany).
Thamnasterioid colony with corallites arranged in concentric rows. The colony surface is plane, the corallite centres may be slightly depressed. Septa are more often connected between corallites of neighboured rows than with corallites of the same row. No wall. Dimorphastrea is similar to Astraeofungia and juvenile colonies of Astraeofungia and Dimorphastrea are difficult to distinguish.
AnimaliaScleractiniaLatomeandridaeB1652154-FC80-5566-BEE6-582ACA8EB2B4Dimorphastreacf.hiraigaensis(Eguchi, 1951)cf1951Meandraraeahiraigaensis Eguchi, n. sp. – Eguchi: 37, pl. 6, figs 5, 7, pl. 7, fig. 7. v2013bDimorphastreainsignis (Fromentel, 1887) – Löser: 20, fig. 3.3. v2013Dimorphastrearegularis (de Fromentel, 1857) – Löser, Werner and Darga: 60, pl. 6, figs 5–6. v2015Dimorphastrea sp. – Löser, Arias and Vilas: 56, figs 7a–c. Material.
BSPG 1947 XVI 11, 1947 XVI 9; two thin sections.
Remarks.
The present material differs from Dimorphastreahiraigaensis by a clearly higher number of septa.
Other occurrences.
Lower Hauterivian of the European Boreal (France), Albian to lower Cenomanian of the Western Tethys (Spain), middle Cenomanian of the European Boreal (Germany).
Solitary cylindrical coral. Corallite outline elliptical, centre slightly depressed. Septa irregularly perforated, more common in the central part of the septal blade. Symmetry of septa irregular radial. Septa of younger generations are with their inner margins occasionally connected to septa of older generations. Synapticulae abundant. Columella absent or as some small elements, presumably trabecular extensions of septal inner margins.
AnimaliaScleractiniaLatomeandridaeBE337FE6-EF97-5DA6-A25C-053E2898DE2ELeptophyllaraeacf.granulata(de Fromentel, 1863b)cf1863Leptophylliagranulata – de Fromentel: 303, pl. 61, fig.1. v2013Leptophyllaraea cf. granulata (de Fromentel, 1863) – Löser, Werner and Darga: 60, pl. 7, figs 7–9. Material.
Solitary cylindric coral. Corallite outline circular or elliptical, centre slightly depressed. Symmetry of septa irregular radial. Synapticulae moderately common. Columella absent or developed as some small elements, presumably trabecular extensions of septal inner margins. Endotheca consists of numerous dissepiments. Wall absent or epithecal.
Remarks.
As already explained in Löser et al. (2021b), in the historic literature the genus Placoseris was considered synonymous with Acrosmilia d’Orbigny, 1849. Acrosmilia is a conceptual genus; the type specimen of the type species is available but so poorly preserved that important diagnostic features, such as the presence or absence of pennulae or the amount of septal perforation, cannot be observed. For this reason, the genus Leptophyllia Reuss, 1854 was applied (Löser et al. 2019) in place of Acrosmilia. Leptophyllia was for a long time considered to be a junior synonym of Acrosmilia. The study of type specimens and topotypical material has shown that Leptophyllia belongs to the mainly Late Cretaceous family Synastraeidae and is restricted to the Late Cretaceous, whereas Placoseris belongs to the Jurassic and mainly Lower Cretaceous family Latomeandridae. Leptophyllia has thicker and less perforate septa, whereas in Placoseris the septa are thinner and more perforate at the inner margin. Moreover, the septa are often connected to each other in the latter, a characteristic that is less common in Leptophyllia (see Löser et al. 2019 for details).
AnimaliaScleractiniaLatomeandridae07DD0080-8F60-59BD-8C2D-8171365D8C9FPlacoseriseturbensis(de Fromentel, 1857)Plate 3: figs 1, 2*v1857 TrochoserisEturbensis – de Fromentel: 19, pl. 1, fig. 8. v1897Leptophylliapatellata – Söhle: 44, pl. 6, fig. 5. v1941Thecoseriscenomanensis n.sp. – Alloiteau: 22, pl. 1, figs 18, 19. v1989Acrosmiliapatellata (Michelin 1845) – Löser: 131, text-fig. 34, pl. 26, fig. 1. v2015aAcrosmilia sp. – Löser: appendix. v2018Placoseriseturbensis (Fromentel, 1857) – Löser, Steuber and Löser: 42, pl. 4, figs 1–3. v2023Placoseriseturbensis (Fromentel, 1857) – Löser and Wilmsen: 293, figs 10.7–10.9. Material.
LFU 8336SG015085; six thin sections.
Dimensions.
(LFU 8336SG015085).
c
26×31.8
septa
154
Other occurrences.
Valanginian to Aptian of the Western Atlantic (Mexico), lower Hauterivian of the European Boreal (France), lower Albian of the Western Tethys (Spain), lower Cenomanian of the Central Tethys (Greece), the Western Tethys (Spain), and the European Boreal (Germany), middle Cenomanian of the European Boreal (France), upper Cenomanian of the European Boreal (Germany).
AnimaliaScleractiniaLatomeandridae399EBA44-6FA4-58D8-884E-4999C70736DAPlacoseriscf.eturbensis(de Fromentel, 1857)Plate 3: figs 3–5cf1857 TrochoserisEturbensis – de Fromentel: 19, pl. 1, fig. 8. v2013Acrosmilia sp. – Löser, Werner and Darga: 58, pl. 7, figs 1–3. v2014bAcrosmiliabaumbergeri (Koby, 1898) – Löser: 34, fig. 5d. Material.
BSPG 1947 XVI 4, 1947 XVI 5, 1991 X 64, 1991 X 65, 1991 X 66, 1991 X 68, 1991 X 69, LFU 8336SG015091; four thin sections.
Dimensions.
(LFU 8336SG015091).
c
19.4×30.1
septa
202
Remarks.
The present material differs from Placoseriseturbensis by a higher number of septa.
Other occurrences.
Hauterivian to Cenomanian of the European Boreal (France, Germany), upper Cenomanian of the Western Tethys (France) and the European Boreal (Germany).
Thamnasterioid-cerioid colony. Corallite outline irregular with corallite centres slightly depressed. Symmetry of septa irregular. Costae confluent or sub-confluent. Synapticulae occasional, mainly in the space between corallites. Columella consists of isolated trabeculae or one more solid element. Wall subcompact, made of synapticulae. Polyastreopsis differs from Thalamocaeniopsis by a poorly defined corallite outline, and a more incomplete wall.
AnimaliaScleractiniaLatomeandridaeB741A84A-2DC9-54FE-A7B7-E0697CFB8E44Polyastropsiscf.fascigera(Felix, 1909)cf1909Isastraeafascigera – Felix: 172, pl. 7, fig. 2. v2013Thalamocaeniopsis sp. – Löser, Werner and Darga: 63, pl. 8, figs 7–9. Material.
BSPG 1947 XVI 56; two thin sections.
Remarks.
In Löser et al. (2013c) this specimen was assigned to the genus Thalamocaeniopsis. This is changed here. Thalamocaeniopsis has polygonal corallites and a marked wall, even if thin and incomplete, whereas Polyastreopsis has no wall at all and the corallite outline is irregular. The specimen differs from Polyastropsisfascigera by having smaller distances of the corallites and a higher number of septa.
Other occurrences.
Upper Aptian of the Western Tethys (Spain), lower Albian of the Western Atlantic (Mexico), lower Cenomanian of the European Boreal (Germany).
Cerioid colony. Corallite outline polygonal with centres slightly depressed. Symmetry of septa irregular. Synapticulae occasional, mainly in the space between corallites. No costae. Columella consists of isolated trabeculae or one more solid element. Wall subcompact, made of synapticulae. Budding extracalicinal.
AnimaliaScleractiniaLatomeandridae4E6C743B-5DD0-5F29-BB1B-51A15EE54584Thalamocaeniopsiscf.taramellii(d’Achiardi, 1880)Plate 3: figs 6–8cf1880Latimeandrataramellii – d’Achiardi: 249: pl. 17, fig. 7. v2004Isastreaminima Prever, 1909 – Löser and Mohanti: 583, fig. 2c. v2018Thalamocaeniopsis sp. – Löser, Steuber and Löser: 44, pl. 4, figs 10, 11. Material.
BSPG 1947 XVI 75; two thin sections.
Dimensions.
(BSPG 1947 XVI 75).
n
min–max
µ
s
cv
µ±s
clmin
10
1.55–2.03
5.71
1.17
20.5
4.54–6.88
clmax
10
5.65–10.3
8.00
1.69
21.1
6.31–9.69
ccd
10
3.84–8.71
5.82
1.58
27.2
4.24–7.40
septa
10
36–63
45.80
8.57
18.7
37–54
Remarks.
The specimen differs from Thalamocaeniopsistaramellii by a lower number of septa.
Other occurrences.
Lower Aptian of the Western Tethys (Spain), lower Cenomanian of the Central Tethys (Greece), Cenomanian of the Southern Tethys (India).
Valanginian to lower Aptian of the Central Tethys (Bulgaria, Italy), lower Aptian of the European Boreal (UK) and the Central Tethys (Greece), lower Albian of the Western Tethys (Spain) and the Western Atlantic (Mexico), lower Cenomanian of the Western Tethys (Spain).
Solitary (not in the Cretaceous) and colonial (cerioid, hydnophoroid, meandroid, phaceloid, plocoid, thamnasterioid) colonies. Septa completely and regularly perforated. Interseptal space larger than or equal to septal thickness.
Thamnasterioid colony with corallites arranged in rows. The corallites are well separated from each other. Septa run mostly between neighboured corallite rows, less between adjacent corallites. A central corallite may be present.
AnimaliaScleractiniaMicrosolenidaeA455EC80-3E9B-5C41-A339-24EA697DD630Dimorpharaeawilliamsonensis(Wells, 1944)*v1944Microsolenawilliamsonensis Wells, n.sp. – Wells: 100, pl. 20, figs 1–4. v2013Dimorpharaeajaponica Eguchi, 1951 – Löser, Werner and Darga: 63, pl. 8, figs 10–12. v2017Dimorpharaeajaponica Eguchi, 1951 – Löser and Bilotte: 9, figs 7g–i. Material.
BSPG 1991 X 80; two thin sections.
Remarks.
In Löser et al. (2013c), this specimen was assigned to Dimorpharaeajaponica Eguchi, 1951. Dimorpharaeajaponica and Dimorpharaeawilliamsonensis have very similar dimensions but differ in the septal counts.
Other occurrences.
Upper Aptian of the Western Tethys (Spain), Albian to lower Cenomanian of the Western Atlantic (Mexico, USA), lower Cenomanian of the European Boreal (Germany), upper Cenomanian of the Western Tethys (France).
AnimaliaScleractiniaMicrosolenidaeE94552B1-0A12-56F6-A240-21104FC82C5DEocomoserisMelnikova et al., 1993Type species.
Astreoid colony with small corallites. Symmetry of septa irregular, but two size orders can be distinguished. Number of septa up to 30. Septa occasionally connected to each other. Costae sub-confluent to non-confluent. Synapticulae fairly common. Columella styliform. Endotheca and wall absent. Coenosteum narrow.
AnimaliaScleractiniaMicrosolenidae8D9F981D-BA14-5421-BB72-129CC457EACBEocomoserissp.Plate 4: figs 3, 4v2012Eocomoserisraueni Melnikowa et al., 1993 – Bover Arnal, Löser and Moreno Bedmar: 58, figs 11I–K. v2013EocomoserisraueniMelnikova et al. 1993 – Löser, Castro and Nieto: 25, pl. 8, figs 11, 12. v2021Eocomoseris sp. 1 – Löser, Nieto, Castro and Reolid: 25, figs 27.1–3. Material.
LFU 8336SG015094#3; one thin section.
Dimensions.
(LFU 8336SG015094#3).
n
min–max
µ
s
cv
µ±s
clmin
6
1.64–2.16
1.90
0.19
9.8
1.71–2.08
clmax
6
1.84–2.32
2.15
0.18
8.4
1.97–2.33
ccd
10
1.82–3.89
2.65
0.64
24.3
2.01–3.30
septa
7
19–28
23.9
3.63
15.2
20–27
Remarks.
The difficulty of species separation was already discussed in Löser et al. (2021b).
Other occurrences.
Lower Valanginian of the Western Tethys (Spain), Valanginian to Aptian of the Western Atlantic (Mexico), lower Aptian to middle Cenomanian of the Western Tethys (Spain, Portugal), Albian to lower Cenomanian of the European Boreal (UK).
Plocoid colony. Corallite outline circular, corallites small (generally below 3 mm in diameter). Septa irregularly perforated. Symmetry of septa bilateral. Septal cycles differ in length, but hardly at all in thickness. Septa of younger cycles often and regularly connected to the septa of preceding cycles. Not well-separated pali irregularly present. Costae non-confluent. Synapticulae fairly common, mainly in the wall. Columella small, substyliform. Endotheca consists of numerous thin tabulae. Wall subcompact, synapticulothecal. Coenosteum moderately broad, consists of trabeculae and tabulae.
AnimaliaScleractiniaNegoporitidaeEF648F8F-1BBF-5FC7-BD2B-0F379C8B0DB5Negoporitescf.quartusEliášová, 1995cf1995Negoporitesquartus sp. n. – Eliášová: 28, pl. 2, fig. 2, pl. 3, figs 1, 2. v2013Negoporites cf. quartus Eliášová, 1995 – Löser, Werner and Darga: 62, pl. 8, figs 1–3. Material.
BSPG 1947 XVI 45; one thin section.
Other occurrences.
Lower Cenomanian of the Western Tethys (Spain).
AnimaliaScleractiniaNegoporitidae62781B1C-32B3-5DB5-8228-5DE657F43097Negoporitessp.v2013Negoporites sp. – Löser, Werner and Darga: 62, pl. 8, figs 4–6. Material.
Astreoid colony with strong, at the inner margin slightly perforated septa, that stand in a bilateral symmetry. The septal lateral faces are ornamented with pennulae. The columella is small and styliform. The wall is synapticulothecate, in places septothecal, subcompact. Endotheca with tabulae.
AnimaliaScleractiniaNegoporitidae340216F6-4B7E-51BC-BB5E-0EE7692C9479ParactinacisuliaeLöser & Heinrich, 2018Plate 4: figs 5–7v2018Paractinacisuliae spec. nov. – Löser and Heinrich: 131, fig. 4. v2019Paractinacisuliae Löser and Heinrich, 2018 – Löser, Heinrich and Schuster: 92, figs 126, 129a–c. Material.
BSPG 1947 XVI 69, LFU 8336SG015157; three thin sections.
Dimensions.
(LFU 8336SG015157).
n
min–max
µ
s
cv
µ±s
cmin
7
2.58–3.69
3.20
0.36
11.4
2.84–3.57
cmax
6
2.96–4.04
3.46
0.51
14.7
2.95–3.97
ccd
9
2.60–4.07
3.07
0.51
16.5
2.56–3.58
septa
24–32
Other occurrences.
Coniacian to Santonian of the Central Tethys (Austria).
Solitary and (astreoid, meandroid, thamnasterioid) colonial corals. Septa almost compact with few perforations at the inner margins. Interseptal space smaller than septal thickness.
Thamnasterioid colony, where – in contrast to Synastrea – the corallites are clearly marked and slightly elevated. The septa are bent outside the corallite.
AnimaliaScleractiniaSynastreidae54DDFC34-1196-59B4-8938-7A68BE810066Brachycoeniaaff.composita(Sowerby, 1832)Plate 4: figs 8–10v2019Brachycoeniaaff.composita (Sowerby, 1832) – Löser, Heinrich and Schuster: 95, figs 135a–c. Material.
BSPG 1955 XIX 34; two thin sections.
Dimensions.
(BSPG 1955 XIX 34).
n
min–max
µ
s
cv
µ±s
cmin
10
6.50–8.36
7.26
0.62
8.5
6.64–7.87
cmax
10
6.32–8.58
7.24
0.80
11.1
6.43–8.04
ccd
10
8.33–10.1
9.11
0.65
7.1
8.46–9.76
septa
10
36–49
42.3
3.40
8.0
39–46
Remarks.
The specimen has smaller corallite dimensions compared to Brachycoeniacomposita. Since the type material of Brachycoeniacomposita is not available and a correct comparison is not possible, a new species cannot be established for this specimen.
Other occurrences.
Coniacian to Santonian of the Central Tethys (Austria).
Turbinate or trochoid solitary coral with circular, elliptical or irregular outline. The septa are numerous, thick, slightly perforated at the inner margin and rarely connected to each other. The endotheca is well developed.
AnimaliaScleractiniaSynastreidae022A0124-0EF7-5925-A687-132B82F16041Leptophylliasp.Plate 5: figs 1–3vp1854Leptophylliaclavata – Reuss: 101. v2019Leptophyllia sp. 2 – Löser, Heinrich and Schuster: 137, figs 203a–c. Material.
LFU 8336SG015155#1; two thin sections.
Dimensions.
(LFU 8336SG015155#1).
c
10.6×12.9
septa
140
Remarks.
This specimen marks the first stratigraphical occurrence of the genus.
Other occurrences.
Coniacian to Santonian of the Central Tethys (Austria).
Thamnasterioid colony with strong, only at the inner margin perforated septa, that barely differ in length and thickness. The corallites are regularly distributed, often slightly depressed. Costae run between all corallites.
AnimaliaScleractiniaSynastreidaeBF250C0D-4A9A-53DB-AAAD-271CFE6AD150Synastreaagaricites(Goldfuss, 1826)Plate 5: figs 4–6*v1826Astraea Agaricites – Goldfuss: 66, pl. 22, fig. 9. v1854Thamnastraeaprocera – Reuss: 120, pl. 5, figs 1, 2. v2019Synastreaagaricites (Goldfuss, 1826) – Löser, Heinrich and Schuster: 142, fig. 212, figs. 214a–c. Material.
LFU 8336SG015099#1; two thin sections.
Dimensions.
(LFU 8336SG015099#1).
n
min–max
µ
s
cv
µ±s
ccd
30
3.76–5.75
4.73
0.63
13.3
4.10–5.36
septa
10
49–61
53.9
3.81
7.1
50–58
Other occurrences.
Upper Turonian to Santonian of the Central Tethys (Austria).
AnimaliaScleractiniaSynastreidae23FDF69F-D69F-56C2-82CA-D2F130F129CBSynastreacf.catadupensis(Vaughan, 1899)Plate 5: figs 7–9cf1899Mesomorphacatadupensis, n. sp. – Vaughan: 246, pl. 41, figs 1-3. v2019Synastrea cf. catadupensis Vaughan, 1899 – Löser, Heinrich and Schuster: 144, figs 216a–c. Material.
BSPG 1947 XVI 70, LFU 8336SG015088; six thin sections.
Dimensions.
(BSPG 1947 XVI 70)
n
min–max
µ
s
cv
µ±s
ccd
25
4.29–7.38
5.98
0.79
13.2
5.19–6.76
septa
12
38–56
45.25
5.51
12.2
40–51
Remarks.
Synastreacatadupensis has higher septal counts compared to the present material.
Other occurrences.
Coniacian to Santonian of the Central Tethys (Austria).
AnimaliaScleractiniaSynastreidae2059B6F9-EC12-5326-ACCD-034FC0D237F9Synastreaexaltata(Reuss, 1854)Plate 5: figs 10–12*1854Thamnastraeaexaltata – Reuss: 118, pl. 19, figs 5, 6. v2019Synastrea?exaltata (Reuss, 1854) – Löser, Heinrich and Schuster: 146, figs 220a–c. Material.
Coniacian to Santonian of the Central Tethys (Austria), Campanian to Maastrichtian of the Western Atlantic (Jamaica).
AnimaliaScleractiniaSynastreidae1BB66B39-4FE9-5AD1-ACE9-AEE6CBBE0DC8Synastreaheberti(Alloiteau, 1952)Plate 6: figs 1–3*v1952UxacalcaraeaHeberti All. – Alloiteau: 665. v1957UxacalcaraeaHeberti nov. gen., nov. sp. – Alloiteau: 221, pl. 8, fig. 7, pl. 9, fig. 12. v1997Thamnaraeacladophora Felix, 1903 – Baron-Szabo: 80, pl. 13, fig. 1. v2012aAstraeofungiasiva (Stoliczka, 1873) – Löser: 28, figs 2.11–2.12. v2016cUxacalcaraeaheberti Alloiteau, 1952 – Löser: p. 681, figs U1a–c. v2019Synastreaheberti Alloiteau, 1952 – Löser, Heinrich and Schuster: 147, figs 221a–c. Material.
BSPG 1947 XVI 67; two thin sections.
Dimensions.
(BSPG 1947 XVI 67).
n
min–max
µ
s
cv
µ±s
ccd
15
3.71–5.20
4.41
0.47
10.7
3.93–4.88
septa
8
31–46
38.6
5.18
13.4
33–44
Other occurrences.
Upper Turonian of the Western Tethys (France), upper Turonian to Santonian of the Central Tethys (Austria), lower Coniacian of the Western Tethys (Spain), lower Campanian of the Central Tethys (Austria, Turkey).
AnimaliaScleractiniaSynastreidaeF17094D9-DB5A-5F0E-AE97-5174D9B2DEEASynastrea?salisburgensis(Beauvais, 1982)Plate 6: figs 4, 5?*1982Fungiastraeasalisburgensis nov. sp. – Beauvais: (2), p. 75, pl. 27, fig. 5, pl. 28, fig. 1. v2018Synastrea sp. – Löser, Steuber and Löser: 46, pl. 6, figs 1–3. v2019Synastreasalisburgensis (Beauvais, 1982) – Löser, Heinrich and Schuster: 151, figs 226a–c. Material.
LFU 8336SG015227#2, 8336SG015228#3, 8336SG015228#5; three thin sections.
Dimensions.
(LFU 8336SG015228#5).
n
min–max
µ
s
cv
µ±s
ccd
20
4.33–6.90
5.69
0.85
14.9
4.85–6.54
septa
10
29–42
34.6
5.25
15.2
29–40
Remarks.
As already mentioned in Löser et al. (2019), the type material of the species is not available and the morphometric data provided in the literature do not allow a clear separation from other species.
Other occurrences.
Aptian to Santonian of the Central Tethys (Greece, Austria), Campanian to Maastrichtian of the Western Atlantic (Jamaica).
AnimaliaScleractiniaSynastreidae20ED378C-FA32-5DD3-AAA3-F10ACD73CFC0Synastreasp.v2013Synastrea sp. – Löser, Werner and Darga: 63, pl. 9, fig. 1. Material.
Colonial corals. Septa strong, compact. Symmetry irregular. Septa often connected to each other. Septal distal margins smooth, lateral faces smooth or with thorns, inner margins slightly swollen. Lonsdaleoid septa absent. Main septum absent. Microstructure of very small trabeculae, only marked by a dark line. Synapticulae and pali absent. Columella by septal fusion. Endotheca mostly present, consisting of tabulae or dissepiments. Marginarium absent. Wall generally present, septothecal. Coenosteum present in some genera. Budding varies.
Family <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Dendrogyridae</tp:taxon-name-part></tp:taxon-name> Alloiteau, 1952AnimaliaScleractiniaSynastreidae1FA2B407-832F-52D0-B5FA-D53F00C0427FHydnophoraraeaOppenheim, 1930Type species.
Monticulariastyriaca Michelin, 1847.
Description.
Hydnophoroid colony. Corallite centres can be recognised in places. Septa straight, free, often alternating in length and thickness. Their lateral faces bear thorns or are smooth. The columella is rudimentarily developed; it is difficult to assign the elements between the crests to any morphological unit. Endotheca well developed.
Remarks.
Formerly, this genus was assigned to the family Eugyridae (Löser 2016c, Löser et al. 2019). But it differs from this family by the less regular endotheca and thinner septa. Moreover, all other members of the Eugyridae became extinct before the Turonian. Hydnophoraraea shows certain similarities to the family Dendrogyridae, but there is also material that differs from this concept such as, for instance, by having a dissepimental instead of a tabular endotheca, or showing the lateral faces ornamented with thorns instead of being smooth. It is also possible that Hydnophoraraea encompasses heterogeneous material and it is also possible that further studies could reveal that the material assigned to Hydnophoraraea may belong to different genera. Further, it is also problematic that the type material of Monticulariastyriaca Michelin, 1847, type species of Hydnophoraraea, is not available (despite being registered in the catalogue of the MNHN in Paris).
The present specimen differs from H.digitata by larger distances between the crests. Moreover, the crests are polygonal in H.digitata, but conical in the present specimen.
AnimaliaScleractiniaSynastreidae14434EDC-409C-534F-B556-3EF12A5FDF2CHydnophoraraeaobliqua(Reig Oriol, 1992)v*1992Hydnophoraobliqua n. sp. – Reig Oriol: 26, pl. 1, fig. 6, pl. 5, figs 4, 5. vp2012aHydnophoraobliqua Reig Oriol, 1992 – Löser: 26. vp2013Hydnophora cf. obliqua Reig Oriol, 1992 – Löser, Werner and Darga: 50, pl. 3, figs 4–6. Material.
BSPG 1947 XVI 21, 1947 XVI 22, 1963 VI 190, 1991 X 59, 1991 X 60; two thin sections.
Other occurrences.
Middle Santonian of the Central Tethys (Austria), Early Campanian of the Central Tethys (Turkey), Campanian of the Western Tethys (Spain).
AnimaliaScleractiniaSynastreidae3983153A-AD97-55FA-A6D8-B4B771DBF954HydnophoraraeaparviconusOppenheim, 1930*v1930Hydnophoraraeaparviconus Oppenheim n. sp. – Oppenheim: 227, pl. 18, figs 3–5, 8. v1930Hydnophoraraeaaconus n. sp. – Oppenheim: 232, pl. 18, fig. 2, pl. 19, fig. 4. v1989Hydnophoraminima nov. sp. – Reig Oriol: 17, pl. 5, fig. 4. vp2012a –Hydnophoraobliqua Reig Oriol, 1992 – Löser: 26, figs 3.1–3.3. vp2013Hydnophora cf. obliqua Reig Oriol, 1992 – Löser, Werner and Darga: 50. v2013Hydnophora sp. – Löser, Werner and Darga: 48, pl. 4, figs 8, 9. v2019Hydnophoraraeaparviconus Oppenheim, 1930 – Löser, Heinrich and Schuster: 159, figs 235, 236, 240a–c. Material.
BSPG 1963 VI 189, 1991 X 106; two thin sections.
Remarks.
In Löser et al. (2013c) this material was assigned to Hydnophora cf. obliqua Reig Oriol, 1992 and, respectively, Hydnophora sp. After having been able to study the type material of Oppenheim (1930) in Jerusalem in 2016, it was possible to correct this assignation.
Other occurrences.
Coniacian to Early Campanian of the Central Tethys (Austria, Turkey), Campanian of the Western Tethys (Spain).
AnimaliaScleractiniaSynastreidae96828023-AA85-5799-B103-F20C8415FDAEHydnophoraraearapulumOppenheim, 1930Plate 6: figs 8, 9v1877Hydnophora Styriaca – de Fromentel: 468, pl. 120, fig. 2. [nonMonticulariastyriaca Michelin, 1847] *v1930Hydnophoraraearapulum n. sp. – Oppenheim: 230, pl. 14, fig. 3, pl. 18, fig. 7. 1992Hydnophoradissimilis n. sp. – Reig Oriol: 27, pl. 1, figs 4, 5, pl. 5, fig. 6. v2019Hydnophoraraearapulum Oppenheim, 1930 – Löser, Heinrich and Schuster: 160, figs 242a, b. Material.
LFU 8336SG015087#1; two thin sections.
Dimensions.
(LFU 8336SG015087#1).
n
min–max
µ
s
cv
µ±s
md
15
1.97–2.55
2.31
0.20
8.5
2.11–2.51
septa
13
8–12
9.92
1.55
15.6
8–11
Other occurrences.
Upper Cenomanian of the Western Tethys (France), Coniacian to Santonian of the Central Tethys (Austria), upper Santonian of the Western Tethys (France), Santonian to Campanian of the Western Tethys (Spain).
AnimaliaScleractiniaSynastreidae19985FEF-A0E6-5439-94BC-AB01A555C127Hydnophoraraeaaff.rapulumOppenheim, 1930Plate 7: figs 1–3v2018Hydnophoraraeastyriaca (Michelin, 1847) – Löser, Steuber and Löser: 47, pl. 6, figs 4–6. Material.
LFU 8336SG015012#1; two thin sections.
Dimensions.
(LFU 8336SG015012#1).
n
min–max
µ
s
cv
µ±s
md
15
1.85–2.57
2.23
0.26
11.8
1.97–2.49
septa
20
10–14
12.20
1.47
12.1
11–14
Remarks.
Compared to Hydnophoraraearapulum, the distances between crests are lower but the septal counts are higher.
Colonial (cerioid, flabelloid, hydnophoroid, meandroid, phaceloid, plocoid) corals. Septa compact. Septal symmetry regular and in various systems, in size orders or irregular. Septa poorly ornamented. Septal microstructure of small trabeculae. Lonsdaleoid septa only in the Felixigyrids; main septa absent. Synapticulae and pali absent. Columella rare. Endotheca well-developed, generally as dense, thick and regular tabulae. Marginarium absent. Wall compact and tabulothecal or septothecal by thickening of septa. Coenosteum and budding varies.
Colonial (cerioid, flabelloid, hydnophoroid, meandroid or hybrid) corals. Septa in size orders. Columella rare. Wall compact and septothecal by thickening of septa.
Hydnophoroid-cerioid colony with distinct corallites. Corallite outline irregular, centres depressed. Symmetry of septa radial and irregularly hexameral. Pali or paliform lobes absent. Costae absent. Columella styliform or by septal fusion in the centre of the corallite. Endotheca consists of regular tabulae and occasional dissepiments. Wall compact, septothecal. Coenosteum absent.
AnimaliaScleractiniaEugyridae60608221-894F-5280-9340-50E5970A2633ColumellophoravelimensisEliášová, 1989*v1989Columellophoravelimensis n.sp. – Eliášová: 114, pl. 1, fig. 1, pl. 3, fig. 2. v2013Columellophora cf. velimensis Eliášová, 1989 – Löser, Werner and Darga: 48, pl. 2, figs 1–3. v2016cColumellophoravelimensis Eliášová, 1989 – Löser: 247, figs C38a–c. Material.
BSPG 1947 XVI 54, 1991 X 105; four thin sections.
Other occurrences.
Upper Cenomanian of the Western Tethys (France), upper Cenomanian to lower Turonian of the European Boreal (Czech Republic).
AnimaliaScleractiniaEugyridae5F7B71C4-C579-59C7-9473-AFD7C060EF83Columellophorasp.v2013Columellophora sp. – Löser, Werner and Darga: 48, pl. 2, figs 4–6. Material.
BSPG 1947 XVI 51; two thin sections.
Other occurrences.
Cenomanian of the European Boreal (Czech Republic).
Hydnophoroid colony. Crests conical, thick, often connected to each other. Corallites distinct. No septal symmetry, but size orders can be distinguished. Pali or paliform lobes absent. Costae unknown. Columella absent. Endotheca consists of thin tabulae and dissepiments. Wall compact, septothecal. Coenosteum absent.
AnimaliaScleractiniaEugyridaeAE62EF01-F39C-5BD7-9EEF-36FEDF1D3EC9FelixigyradeangelisiPrever, 1909*v1909FelixigyraDeangelisi – Prever: 118, pl. 12, figs 7, 8. v2013Felixigyradeangelisi Prever, 1909 – Löser, Werner and Darga: 54, pl. 3, figs 1–3 [= with detailed synonymy]. Material.
BSPG 1947 XVI 62, 1991 X 81, 1991 X 98, 1991 X 99; three thin sections.
Cerioid colony. Corallite outline irregular. Septa very short, with a triangular outline. Symmetry of septa irregular. Pali, costae, synapticulae, and columella absent. Endotheca consists of numerous and regular tabulae. Wall compact, probably tabulothecal.
AnimaliaScleractiniaSolenocoeniidae1E94DB88-28B3-58AA-8584-844F66F62A0FConfusaformaweyeriLöser, 1987Plate 7: figs 4–6*v1987Confusaformaweyeri n.sp. – Löser: 234, pl. 1, figs 1–3. v2016cConfusaformaweyeri Löser, 1987 – Löser: 254, figs C49abc. v2018Confusaformaweyeri Löser, 1987 – Löser, Steuber and Löser: 48, pl. 6, figs 10–12. [here more detailed synonymy] Material.
BSPG 2016 XXII 1; two thin sections.
Dimensions.
(BSPG 2016 XXII 1).
n
min–max
µ
s
cv
µ±s
clmin
20
0.54–0.99
0.75
0.13
16.9
0.62–0.87
clmax
20
0.87–1.18
1.02
0.11
10.4
0.91–1.12
Remarks.
The only specimen is poorly preserved and did not allow septal counts.
Other occurrences.
Lower Aptian of the Central Tethys (Slovenia, Italy), lower Albian of the Western Atlantic (Mexico), lower Cenomanian of the Central Tethys (Greece) and the Western Tethys (Spain), upper Cenomanian of the European Boreal (Germany, Czech Republic).
Plocoid colony. Corallite outline circular. Symmetry of septa radial and regularly hexameral or decameral. Septa very short, free. Pali or paliform lobes absent. Costae present, sub-confluent to non-confluent. Columella absent. Endotheca consists of regular tabulae and occasional dissepiments. Wall compact, as tabulotheca. Coenosteum moderately broad, consists of costae and tabulae.
AnimaliaScleractiniaSolenocoeniidaeCDCE5B83-E58D-5DD7-8B37-8CF22213C51FCryptocoeniaaguilerai(Reyeros Navarro, 1963)*v1963Procyathophoraaguilerai n.sp. – Reyeros Navarro: 8, pl. 3, figs 3, 5. v2013Cryptocoeniaaguilerai (Reyeros Navarro, 1963) – Löser, Werner and Darga: 64, pl. 9, figs 4–6 [with more detailed synonymy]. v2016Cryptocoeniaaguilerai (Reyeros Navarro, 1963) – Löser and Zell: 14, figs 5.1–3. Material.
BSPG 1991 X 70; two thin sections.
Other occurrences.
Tithonian to lower Berriasian of the European Boreal (Czech Republic), Valanginian to Aptian of the Western Atlantic (Mexico), upper Barremian to lower Aptian of the Central Tethys (Germany, Greece), lower Aptian of the Western Tethys (Spain) and the Central Tethys (Greece), upper Aptian of the Western Tethys (Spain), upper Aptian to lower Albian of the Western Tethys (Spain, France), lower Albian of the Western Tethys (Spain, France), upper Albian of the European Boreal (UK).
AnimaliaScleractiniaSolenocoeniidaeAAAB8EDE-27F3-5AC7-8844-12D6E212C3B1Cryptocoeniaantiquad’Orbigny, 1850*v1850Cryptocoeniaantiqua – d’Orbigny: (2), p. 92. v1964Cyathophorasteinmanni Fritzsche 1924 – Morycowa: 24, pl. 3, fig. 2, pl. 5, figs 2, 3. v1996Pentacoeniaelegantula d’Orbigny, 1850 – Baron-Szabo and Steuber: 8, pl. 3, fig. 3. v1996Pseudocoeniaannae (Volz, 1903) – Baron-Szabo and Steuber: 8, pl. 2, fig. 1. v2010Cryptocoeniaatempa (Felix, 1891) – Löser: 591, fig. 3.4. v2013Cryptocoeniabulgarica (Toula, 1884) – Löser, Werner and Darga: 64, pl. 9, figs 2–3. v2016Cryptocoeniaatempa (Felix, 1891) – Löser and Zell: 15, figs 5.7–9. Material.
BSPG 1947 XVI 48, 1991 X 71; two thin sections.
Remarks.
The material was in Löser et al. (2013c) identified as Cryptocoeniabulgarica (Toula, 1884). After this publication the type of Cryptocoeniaantiqua became available and could be measured. The present material is much closer to Cryptocoeniaantiqua; Cryptocoeniabulgarica has smaller dimensions.
Other occurrences.
Lower Hauterivian of the European Boreal (France), upper Barremian of the Western Tethys (France), upper Barremian to lower Aptian of the Central Tethys (Poland, Greece), upper Aptian of the Western Tethys (Spain).
Lower Callovian of the Southern Tethys (Madagascar), Valanginian of the Central Tethys (Bulgaria), upper Barremian of the Western Tethys (France), upper Barremian to lower Aptian of the Central Tethys (Poland, Greece), Aptian to Lower Albian of the Western Atlantic (Mexico), lower Albian of the Western Tethys (France), Cenomanian of the Central Tethys (Greece) and the Southern Tethys (India), Middle Cenomanian of the European Boreal (Germany, Czech Republic).
AnimaliaScleractiniaSolenocoeniidaeC58CC178-C2F6-5FE9-B704-F75C7B5FFC4FCryptocoeniawaltoni(Milne Edwards & Haime, 1851)v*1851Convexastreawaltoni – Milne Edwards and Haime: 109, pl. 23, figs 5, 6. v1873Astrocoenia Reussiana, Stoliczka – Stoliczka: 27, pl. 5, figs 3, 4. v1947Cyathophora Fontserei Bataller 1944 – Bataller: 48, text-fig. v1966Stylinaelegans Beauvais – Beauvais: 121, pl. 1, fig. 1. v1974Cyathophorapygmaea Volz – Turnšek and Buser: 12, 33, pl. 4, fig. 1. v1981Cyathophorapygmaea Volz 1903 – Turnšek and Mihajlovic: 18, pl. 13, figs 1, 2. v1994Adelocoeniapygmaea (Volz 1903) – Löser: 10, text-figs 4, 5, pl. 12, figs 1, 2. v2013Cryptocoeniafontserei (Bataller, 1947) – Löser, Werner and Darga: 66, pl. 9, figs 7–9. v2015cCryptocoeniareussiana (Stoliczka, 1873) – Löser: 21, figs 3D–F. Material.
BSPG 1947 XVI 12, 1947 XVI 20, 1947 XVI 23, 1947 XVI 49, 1947 XVI 6, 1947 XVI 77, 1991 X 43, LFU 8336SG015013#1, 8336SG015107, 8336SG015228#4; eight thin sections.
Remarks.
In Löser et al. (2013c) this material was assigned to Cryptocoeniafontserei Bataller, 1947. Afterwards, it was possible to examine the types of Cryptocoeniawaltoni and Cryptocoeniafontserei. It transpired that both species are synonymous.
Other occurrences.
Aalenian to Callovian of the European Boreal (UK), Callovian of the Central Tethys (Tunisia), Kimmeridgian of the European Boreal (Germany), upper Barremian to lower Aptian of the Central Tethys (Bulgaria, Serbia, Slovenia), lower Aptian of the Western Tethys (Spain), lower Albian of the Western Atlantic (Mexico), lower Albian to lower Cenomanian of the Western Tethys (Spain, France), upper Albian of the Southern Tethys (India) and the European Boreal (UK), lower Cenomanian of the Western Tethys (Spain), middle Cenomanian of the European Boreal (Belgium, Germany).
Plocoid colony. Corallite outline circular. Symmetry of septa radial and regularly trimeral. Septa very short, free. Pali or paliform lobes absent. Costae present, sub-confluent to non-confluent. Columella absent. Endotheca consists of regular tabulae and occasional dissepiments. Wall compact, as tabulotheca. Coenosteum moderately broad, consists of costae and tabulae.
Solitary and colonial (astreoid, meandroid, phaceloid) corals. Septa compact or perforated. Younger septal cycles have more perforations than older septal cycles. Septa very thick, those of the first cycles being the thickest. Septal symmetry regular or subregular radial. Septal upper margins with granulae, lateral faces with thorns. Lonsdaleoid septa and main septum absent. Microstructure of large trabeculae. Synapticulae common. Pali absent, columella parietal. Endotheca mostly absent. Marginarium and wall absent. Coenosteum in some genera. Budding extracalicinal.
Family <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Felixaraeidae</tp:taxon-name-part></tp:taxon-name> Beauvais, 1982AnimaliaScleractiniaSolenocoeniidaeDCF02328-2D77-5DF9-B293-E0320A788F97FelixaraeaBeauvais, 1982Type species.
Felixaraearennensis Beauvais, 1982.
Description.
Turbinate solitary coral with a circular outline. The septa are perforated, younger (thinner) septa more than older (thicker) septa.
Remarks.
The present material of Felixaraea marks the first occurrence of the genus.
The specimen is larger than Felixaraeaagassizi and has a slightly higher septal count.
Other occurrences.
Campanian to Maastrichtian of the Western Atlantic (Jamaica).
AnimaliaScleractiniaSolenocoeniidae0C37C9EC-0408-5280-9C3B-70E31A6BFA26FelixaraearennensisBeauvais, 1982Plate 8: figs 4–6vp1930Haplaraeareticularis n. sp. – Oppenheim: 35, pl. 27, figs 11, 12. [non figs 8–10] v1952Haplaraearennensis All. – Alloiteau: pl. 2, fig. 4, text-fig. 110. [without description] v1957Haplaraearennensis n.sp. – Alloiteau: figs 278–280. [without description] *1982Felixaraearennensis Alloiteau – Beauvais: (2), 25. v2013cFelixaraearennensis Beauvais, 1982 – Löser: 752, figs 1G–I. v2016cFelixaraearennensis Beauvais, 1982 – Löser: 339, figs F3a, b. v2019Felixaraearennensis (Beauvais, 1982) – Löser, Heinrich and Schuster: 165, figs 247, 248, figs 253a, b. Material.
LFU 8336SG015135#2; one thin section.
Dimensions.
(LFU 8336SG015135#2).
c
16.3×21.2
septa
100
Remarks.
Felixaraearennensis was only illustrated but not described by Alloiteau (1952, 1957). Illustrations alone do not constitute a valid first description of a species. Beauvais (1982) was the first to describe and illustrate the species.
Other occurrences.
Coniacian to Santonian of the Central Tethys (Austria), upper Santonian to middle Campanian of the Western Tethys (France, Spain).
Solitary turbinate coral with an elliptical outline. Septa made of large trabeculae. They are perforated, younger septa more than older septa. Septa rarely connected to each other. Symmetry irregular radial. Columella parietal. Wall with various rings of synapticulae. Endotheca absent.
Remarks.
Even the position of the specimen within the family Felixaraeidae is uncertain. In the Felixaraeidae, the trabeculae are still larger, the septa are stronger and synapticulae are more frequent.
Solitary and colonial (cerioid, phaceloid, and plocoid) corals. Septa compact, thick, with ornamented lateral faces. Symmetry radial and bilateral. Lonsdaleoid septa may occur. Septal microstructure with small trabeculae, visible as a medium dark line. Synapticulae absent, pali absent. Columella rarely developed. Endotheca well-developed. Marginarium present in some genera. Wall trabecular or septothecal. Budding extracalicinal.
Solitary corals or (phaceloid, plocoid) colonies. Septa in a higher number as in the other families, generally in a radial symmetry with septal cycles generally regular, except for large corallites. No costae. Lonsdaleoid and main septa absent. Columella weak. Marginarium absent.
AnimaliaScleractiniaAgatheliidaeF4EE53F9-86BB-5417-9FAB-9AA8B4D625DCEotheliaLöser, Werner & Darga, 2013Type species.
Eotheliahoelzli Löser, Werner & Darga, 2013.
Description.
Plocoid coral colony with circular corallites. Thick, compact septa in a regular septal symmetry. Septa in a regular hexameral symmetry. Septa of the first cycle attached to each other in the centre of the corallite forming the columella, septa of the second cycle shorter. Endotheca made of few tabulae, coenosteum consists of dissepiments and rare trabeculae.
AnimaliaScleractiniaAgatheliidaeFCA01DED-19F1-5F75-A2C9-78E427D25DA9EotheliabavaricaLöser, Werner & Darga, 2013v2013Eotheliabavarica n. gen., n. sp. – Löser, Werner and Darga: 52, pl. 5, figs 1–4. Material.
BSPG 1947 XVI 13, 1947 XVI 52, 1947 XVI 57, 1947 XVI 58, 1947 XVI 59, 1991 X 93; four thin sections.
AnimaliaScleractiniaAgatheliidaeF321DF88-2EC9-5624-A350-FDE1F687B184EotheliahoelzliLöser, Werner & Darga, 2013v2013Eotheliahoelzli n. gen., n. sp. – Löser, Werner and Darga: 52, pl. 4, figs 1–7. Material.
BSPG 1980 XIII 13, 1991 X 103, 1991 X 42; five thin sections.
Only (phaceloid, plocoid) colonies. Symmetry radial, cycles subregular. A larger septum is often present. Lonsdaleoid septa and a marginarium can be present.
Plocoid colony with circular corallites. Septal symmetry regular in varying systems. A main septum can be present. A trabecular wall always exists. The coenosteum is granulated at its surface.
Plocoid colony. Corallite outline irregular circular. Symmetry of septa radial and in various symmetries. Cycles of septa regular. Septa not connected to each other. Costae non-confluent, with pali-like outgrowths (costal pali). Endotheca consists of numerous and regular tabulae. Wall subcompact, septothecal. The Coenosteum is moderately broad and it consists of tabulae and costal pali.
AnimaliaScleractiniaHeterocoeniidae63E6F258-05CD-5AD6-961E-D0F398B0DB89Styloheterocoeniasp.v2013Heterocoenia sp. – Löser, Werner and Darga: 54, pl. 3, figs 10–12. Material.
BSPG 1991 X 101; three thin sections.
Remarks.
The material was formerly assigned to the genus Heterocoenia. Löser et al. (2018) established for this coral type the new genus Styloheterocoenia.
BSPG 1947 XVI 82, LFU 8336SG015148, 8336SG015152#1, 8336SG015152#2; four thin sections.
Dimensions.
(LFU 8336SG015152#2).
c
8.4×9.5
septa
28
Description.
Plocoid to sub–phaceloid small colonies. Septa in a subregular hexameral symmetry. No wall.
Remarks.
The small colonies very probably represent a new genus within the family Heterocoeniidae, but the low number of specimens and their small size do not allow to establish a new species and genus.
(?) Phaceloid coral. Corallites densely packed, with a polygonal outline. The compact septa are low in number. A main septum can be present. No septal symmetry. Endotheca probably with dissepiments.
Remarks.
The material is questionable and its systematic position uncertain.
Solitary and colonial (phaceloid, plocoid) corals. Septa compact, with varying thickness, in a subregular radial symmetry. Septa not connected to each other (Rayasmiliidae) or connected only in the centre of the corallite (Misistellidae). Lateral faces with fine granulae or smooth, upper margin smooth. Lonsdaleoid and main septa absent. Microstructure of septa of small trabeculae. Synapticulae absent. Pali absent, columella present in most genera, lamellar or by septal fusion. Endotheca present. Marginarium absent. Wall absent, but an epitheca is often present (when preserved). Coenosteum varies. Budding intracalicinal.
Solitary and phaceloid corals. The septa are always free. A lamellar columella is present in some genera. One or two septa may be connected to the columella.
Solitary turbinate coral. Corallite outline circular. Symmetry of septa radial and irregularly hexameral. Septa rarely connected to each other. Pali absent. Costae present. Columella absent.
AnimaliaScleractiniaRayasmiliidae96B1E3B4-3B3D-5C16-B372-4CDEF4F0FCD7CeratosmiliaarnaudiAlloiteau, 1957*v1957CeratosmiliaArnaudi nov. sp. – Alloiteau: 116, 421, fig. 74, pl. 3, fig. 7. v2013Ceratosmiliaarnaudi Alloiteau, 1957 – Löser, Werner and Darga: 56, pl. 5, figs 7–9. Material.
BSPG 1947 XVI 38, 1947 XVI 39, 1947 XVI 44, 1991 X 104, 1991 X 47, 1991 X 52, 1991 X 89, 1991 X 90, 1991 X 91, 2012 X 3, LFU 8336SG015082#1, 8336SG015137#2, 8336SG015153; two thin sections.
Solitary turbinate coral. Septa compact, not connected to each other, in a regular radial symmetry. Septa can be connected with the columella. Wall absent. Epitheca present. Endotheca well developed. Columella lamellar.
Solitary turbinate coral with an elliptical outline. Septa made of small trabeculae, in a regular hexameral symmetry, not connected to each other. Septal lateral faces strongly dentated. Columella parietal. Wall septothecal. Endotheca unknown.
Remarks.
The assignation of this specimen to the Rayasmiliidae is preliminar. It differs from this family by strongly ornamented lateral septal faces and the septothecal wall, but coincides in the septal outline, septal microstructure, and regular symmetry.
Cylindric or turbinate solitary coral with circular or slightly elliptical outline. No columella, no pali. Endotheca made of large dissepiments. No wall, just a thin epitheca that is often not present.
AnimaliaScleractiniaRayasmiliidaeB57D96D4-E9AC-59FA-B956-0188ECE79160Trochophylliaaprutina(Prever, 1909)v*1909Coelosmiliaaprutina – Prever: 109, text-fig. 14, pl. 10, fig. 25. v2013Paramontlivaltiaruvida (Prever, 1909) – Löser, Werner and Darga: 56, pl. 6, figs 10–12. Material.
BSPG 1947 XVI 30, 1947 XVI 31, 1947 XVI 32; two thin sections.
Remarks.
This material was described as Paramontlivaltiaruvida (Prever, 1909) by Löser et al. (2013c). This species is very similar to Trochophylliaaprutina and differs only by smaller corallite dimensions. It is possible that the species are synonyms.
Other occurrences.
Lower Aptian of the Central Tethys (Italy), lower Cenomanian of the Western Tethys (Spain, France).
BSPG 2016 XXII 4, LFU 8336SG015109; three thin sections.
Dimensions.
(LFU 8336SG015109).
c
36.9×42.15
septa
92
Other occurrences.
Middle Oxfordian of the Western Tethys (France).
AnimaliaScleractiniaRayasmiliidaeC7A31FF5-5F1F-55FE-8DF2-AAEE3E2F82CETrochophylliaruvida(Prever, 1909)*v1909Epismiliaruvida – Prever: 113, text-fig. 19. v1909Trochosmiliapolymorpha – Prever: 108, text-figs 12, 13, pl. 10, figs 5–23. vnon2013Paramontlivaltiaruvida (Prever, 1909) – Löser, Werner and Darga: 56, pl. 6, figs 10–12. Material.
LFU 8336SG015137#3; one thin section.
Remarks.
The specimens assigned in Löser et al. (2013c) to Paramontlivaltiaruvida belong to Trochophylliaaprutina. Both species are similar in their number of septal cycles but T.aprutina has a larger corallite diameter (13–15 mm) compared to T.ruvida (8–11 mm).
Other occurrences.
Aptian to Santonian of the Central Tethys (Italy, Greece, Austria).
Solitary and (astreoid, cerioid, flabelloid, meandroid, phaceloid, thamnasterioid) colonial corals. Septa compact. Septal thickness regular, septa in an irregular radial symmetry, but septal generations can be distinguished. Septa not connected to each other. Septal lateral faces with vertical keels, upper margins with granulae. Lonsdaleoid septa and main septum absent. Microstructure of large trabeculae. Synapticulae absent. Pali rarely present. Columella in some genera, generally lamellar. Endotheca well-developed. Marginarium absent. Wall poorly defined, generally only with an epitheca. Coenosteum present, depending on the organisation form. Budding generally extracalicinal.
Solitary and (astreoid, flabelloid, meandroid, phaceloid, plocoid) colonial corals. Septa compact. First septal generation thicker than all others. Symmetry irregular or subregular radial. Septa in some genera connected to each other. Septal distal margins smooth, lateral faces with fine thorns, inner margins often swollen or T-shaped. Lonsdaleoid septa very rare. Main septum absent. Microstructure of very small trabeculae, only marked by a dark line. The costae are made by medium-sized trabeculae that result in granulated costal surfaces. Synapticulae and pali absent. Columella present or absent. Endotheca mostly present. Marginarium absent. Wall generally present, septothecal. Coenosteum present in some genera. Budding varies.
Family <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Phyllosmiliidae</tp:taxon-name-part></tp:taxon-name> Felix, 1903bAnimaliaScleractiniaMontlivaltiidae72D2C9DC-DD18-5F2B-BE44-41080FA93CBEAulosmiliaAlloiteau, 1952Type species.
Trochosmiliaarchiaci de Fromentel, 1863b.
Description.
Solitary coral with an elliptical outline. Costae are not pronounced. The septal symmetry is quite regular, but not all septa reach the upper margin. The columella is lamellar, but generally deep in the corallite. The endotheca is poorly developed.
AnimaliaScleractiniaMontlivaltiidae6BB0ABC5-7841-53C9-9815-E45C5C5F38E7AulosmiliacristataBeauvais, 1982Plate 10: figs 10, 11vp1921Placosmiliabofilli, n. sp. – Vidal: 4, pl. 4, figs 6, 7. *1982Aulosmiliacristata nov. sp. – Beauvais: (1), 224, pl. 19, fig. 9. v1997Peplosmiliafromenteli Angelis d’Ossat, 1905 – Baron-Szabo: 72, pl. 8, fig. 1. v2019Aulosmiliacristata Beauvais, 1982 – Löser, Heinrich and Schuster: 239, figs 378a, b. Material.
LFU 8336SG015006#1; two thin sections.
Dimensions.
(LFU 8336SG015006#1).
c
20.4×28
septa
133
sd
9/5mm
Remarks.
Placosmiliabofilli Vidal, 1921 is based on two syntypes that belong to different species: Aulosmiliainconstans and Aulosmiliacristata. Only one specimen was illustrated. A lectotype has so far not been selected.
Other occurrences.
Coniacian to Santonian of the Central Tethys (Austria), Santonian of the Western Tethys (Spain).
AnimaliaScleractiniaMontlivaltiidae7412AA0D-7CE3-5F4C-AD8C-8A3BB87CDCD7Aulosmiliainconstans(de Fromentel, 1862b)v*1862bTrochosmiliainconstans – de Fromentel: 266, pl. 30, fig. 1, pl. 33, fig. 1. v1862bTrochosmiliaheterophyllia – de Fromentel: 272, pl. 35, fig. 1. vp1921Placosmiliabofilli, n. sp. – Vidal: 4. v1952Aulosmiliaarchiaci de From. 1862 – Alloiteau: 636, fig. 92, pl. 7, fig. 2. v1952Strotogyra (Rhipidogyra) granulata All. – Alloiteau: pl. 3, fig. 3. v1957Phragmosmiliacrassa nov. sp. – Alloiteau: 86, figs 35, 36, pl. 2, fig. 1, pl. 19, fig. 9. v2013Aulosmilia?inconstans (de Fromentel, 1862) – Löser, Werner and Darga: 55, pl. 6, figs 7–9. v2016cTrochosmiliainconstans Fromentel, 1862 – Löser: 522, figs P45a, b. v2019Aulosmiliainconstans Fromentel, 1862 – Löser, Heinrich and Schuster: 240, figs 373–375, figs 379a–c. Material.
BSPG 1947 XVI 36, 1947 XVI 37, 1991 X 41, 1991 X 53, 1991 X 54, 1991 X 55, 1991 X 56, 1991 X 57, 1991 X 58, 1991 X 85, 1991 X 86, 1991 X 87; four thin sections.
Remarks.
During the systematic revision of the Coniacian and Santonian corals of the Gosau Group, the genus Aulosmilia was profoundly investigated (Löser et al. 2019) and much Upper Cretaceous coral type material was studied. The present material can now clearly be assigned to Aulosmiliainconstans.
Other occurrences.
Upper Cenomanian to Santonian of the Western Tethys (France), Coniacian to Santonian of the Central Tethys (Austria), Santonian to middle Campanian of the Western Tethys (Spain, France).
The specimen differs from Aulosmiliamagnifica by much higher septal counts.
Other occurrences.
Lower Cenomanian of the Western Tethys (Spain, France).
AnimaliaScleractiniaMontlivaltiidaeACDFF5CB-B159-5F8A-A484-A5CAEECB55ABAulosmiliaparkinsoni(Milne Edwards & Haime, 1848c)*v1848cPlacosmilia Parkinsonii – Milne Edwards and Haime: 235. v1854Placosmiliaconsobrina – Reuss: 84, pl. 5, figs 17–19. v1862aPlacosmiliaangulata – de Fromentel: 225, pl. 18, fig. 3. v1903bPlatysmiliamulticincta Felix (Reuss sp.) – Felix: 285, pl. 20, figs 2–5. v1903bTrochosmiliachondrophora nov. sp. – Felix: 327, pl. 24, fig. 12. v1921Trochosmiliaosensis, n. sp. – Vidal: 4, pl. 3, figs 4–6. v1930Trochosmiliachondrophora Felix – Oppenheim: 483, pl. 31, fig. 8. v2006Aulosmiliacuneiformis (Milne-Edwards and Haime, 1848) – Baron-Szabo: 69, pl. 14, figs 2, 4. v2013Aulosmilia?bipartita (Reuss, 1854) – Löser, Werner and Darga: 54, pl. 6, figs 1, 2. vp2013Aulosmilia?consobrina (Reuss, 1854) – Löser, Werner and Darga: 55. v2019Aulosmiliaparkinsoni (Milne Edwards and Haime, 1848) – Löser, Heinrich and Schuster: 241, figs 381a, b. Material.
BSPG 1947 XVI 18, 1947 XVI 19, 1991 X 82, 1991 X 83, 1991 X 84; three thin sections.
Remarks.
In Löser et al. (2013c) this material was assigned to Aulosmilia?bipartita and Aulosmilia?consobrina. More recent research on the very variable genus (e.g. Löser et al. 2019) has shown that species must be understood in a much wider concept. Hence, the material is now nearly entirely assigned to Aulosmiliaparkinsoni.
Other occurrences.
Lower Coniacian of the Western Tethys (France), upper Turonian to Santonian of the Central Tethys (Austria), Coniacian to Santonian of the Western Tethys (France), Santonian of the Western Tethys (Spain, France), Maastrichtian of the Western Atlantic (Jamaica), Paleocene to Eocene base of the Central Tethys (Austria).
AnimaliaScleractiniaMontlivaltiidae266E2F70-3B65-5E0D-ABCD-9F529C95D58DAulosmiliasp.vp2013Aulosmilia?consobrina (Reuss, 1854) – Löser, Werner and Darga: 55. v2019Aulosmilia sp. – Löser, Heinrich and Schuster: 241, figs 383a, b. Material.
BSPG 1947 XVI 17; two thin sections.
Other occurrences.
Upper Turonian to Santonian of the Central Tethys (Austria), middle Campanian of the Western Tethys (Spain).
Meandroid colony with long, straight or sinuous corallite rows, that are separated by a mostly wide coenosteum. Individual corallites cannot be distinguished. The corallite rows can be in the same level as the coenosteum, or can be elevated. In the latter case, the coral appears flabelloid. Columella lamellar.
AnimaliaScleractiniaMontlivaltiidae946F2C05-7392-5170-9926-F4E9D34D6D51Pachygyracucullata(Zuffardi-Comerci, 1930)v*1930Eugyracucullata n. sp. – Zuffardi-Comerci: 15, pl. 3, figs 3, 8. v2013Pachygyrakrameri Oppenheim, 1930 – Löser, Werner and Darga: 55, pl. 5, figs 5, 6. v2019Pachygyracucullata (Zuffardi-Comerci, 1930) – Löser, Heinrich and Schuster: 274, figs 442a, b. Material.
BSPG 2012 X 1; five thin sections.
Remarks.
In Löser et al. (2013c) this specimen was assigned to Pachygyrakrameri Oppenheim, 1930. In 2016 it was possible to study the type specimen of this species and it transpired that it has much smaller dimensions as discernable from the publication of Oppenheim (1930).
Other occurrences.
Turonian to Santonian of the Central Tethys (Italy, Austria), upper Santonian of the Western Tethys (France).
Colonial (astreoid, plocoid) corals. Septa irregularly perforated. Symmetry of septa irregular radial or bilateral. Septa connected to each other. Septal distal margin coarsely dentated, lateral face with rare granulae, inner margin slightly swollen in places. Lonsdaleoid septa and main septum absent. Microstructure of medium-sized trabeculae. Synapticulae abundant. Pali present in some genera. Columella and endotheca present. Marginarium absent. Wall varies. Coenosteum generally present. Budding extracalicinal.
Astreoid colony. The corallites are small, elliptical, well marked, and with large distances to each other. Septal symmetry is between radial and bilateral. Septa often and regularly connected to each other. Pali present but because of the septal perforations not well visible.
AnimaliaScleractiniaActinacididae8C746D98-2BB4-5C96-BF80-5D875656E957Actinacis?remesiFelix, 1903aPlate 11: figs 3–5?*1903aActinacisRemesi – Felix: 567, text-figs 1, 2. v1911ActinacisRemesi – Trauth: 75, text-fig. 4, pl. 4, fig. 1. v1921ActinacisRemesi Felix – Zuffardi-Comerci: 11, pl. 2, figs 1, 2. v1957Actinacisvignyensis n.sp. – Alloiteau: pl. 18, fig. 9 [= nom. nud.]. v2000Actinacisremesi Felix, 1903 – Baron-Szabo: 119, pl. 10, figs 3, 6. v2013ActinacismagnaAlloiteau 1958 – Löser, Castro and Nieto: 27, pl. 9, figs 5, 6. v2019Actinacis sp. 2 – Löser, Heinrich and Schuster: 306, figs 499a, b. v2022Actinacis?remesi Felix, 1903 – Löser and Callapez: 157, figs 7.j–l. Material.
LFU 8336SG015228#2; one thin section.
Dimensions.
(LFU 8336SG015228#2).
n
min–max
µ
s
cv
µ±s
clmin
5
1.53–2.05
1.76
0.21
12.1
1.55–1.98
clmax
5
1.78–2.54
2.16
0.29
13.5
1.86–2.45
septa
5
24–31
28.2
2.77
9.8
25–31
Remarks.
The material can clearly be assigned to the genus Actinacis, but could not be assigned to an existing species with certainty. Closely related is material labelled as ”Actinacisvignyensis Alloiteau, 1957” (MNHN; but never formally described) and material labelled as Actinacisremesi Felix, 1903 (NHMW, coll. Trauth; PU, coll. Zuffardi-Comerci) from the Eocene of Klokocov (Moravia, Czech Republic) and the Turonian of Libya, but the type of Actinacisremesi itself is unknown.
Other occurrences.
Upper Albian of the Western Tethys (Spain), Turonian of North Africa (Libya), Turonian to lower Coniacian of the Western Tethys (Portugal, Spain), middle Santonian of the Central Tethys (Austria), upper Campanian to Maastrichtian of the Arabian Peninsula (United Arab Emirates), Palaeocene to Eocene of the European Boreal (France, Czech Republic).
Solitary and colonial (astreoid, cerioid, plocoid, reptoid, and thamnasterioid) corals. Septa almost not perforated. Normal septal thickness, septal symmetry regular radial or irregular, septa often and regularly connected to each other. Upper septal border and lateral faces with granulations. No lonsdaleoid septa, no main septum. Microstructure of medium-sized trabeculae. Synapticulae present. Pali absent. Columella present, styliform or by septal fusion. Endotheca present. Marginarium absent. Wall absent or present. Coenosteum depending on the colony type. Budding extracalicinal.
Only cyclolitid forms. Septa (younger more than older) at the inner margins perforated. Symmetry radial and irregular hexameral, often connected to each other. Synapticulae not common.
Astreoid, cerioid, plocoid, reptoid, and thamnasterioid colonies. The symmetry is regular or subregular radial in varying systems, or irregular. The columella is formed by septal fusion.
Astreoid colony with large corallites. Septal symmetry irregularly radial, in various systems. Septa are quite regularly connected to each other. In the corallite centre, septa of the first cycles fuse to form the columella.
AnimaliaScleractiniaRhizangiidae78E989D1-F8E8-5A88-86D0-49D83EA9C2CBEosiderastreacf.glomerata(Reuss, 1854)cf1854Brachyphylliaglomerata – Reuss: 104, pl. 2, figs 11, 12. v2013 ?Diploastrea sp. – Löser, Werner and Darga: 48, pl. 1, figs 5, 6. v2016bEosiderastrea sp. 5 – Löser: 397, pl. 7, figs 1–3. Material.
BSPG 1991 X 97; two thin sections.
Remarks.
This specimen is very closely related to Eosiderastreaglomerata in showing a decameral septal symmetry and four septal cycles, but differs from this species clearly in smaller corallites.
Other occurrences.
Lower Cenomanian of the Central Tethys (Greece).
AnimaliaScleractiniaRhizangiidae6A986846-A009-5BEC-82F3-E3C4F7E9FAB2EosiderastreastefaniLöser, 2016bv2013 ?Diploastreatanohataensis (Eguchi, 1951) – Löser, Werner and Darga: 46, pl. 2, figs 7–9. 2014Diploastreacrassa Kuzmicheva, 1980 – Baron-Szabo: pl. 69, figs 2, 3. v2015a “Diploastrea” tanohataensis (Eguchi, 1951) – Löser: 281, fig. B. v*2016bEosiderastreastefani n. sp. – Löser: 394, pl. 2, figs 7–9. v2018Eosiderastreastefani Löser, 2016 – Löser, Steuber and Löser: 69, pl. 19, figs 10, 11. v2023Eosiderastrea cf. stefani Löser, 2016c – Samaniego-Pesqueira et al.: 142, figs 14H, I. Material.
BSPG 1947 XVI 47; two thin sections.
Remarks.
In Löser et al. (2013c), this specimen was assigned to ?Diploastreatanohataensis. In Löser (2016a) the new genus Eosiderastrea was established and in Löser (2016b) systematically revised.
Other occurrences.
Upper Barremian to lower Aptian of the Central Tethys (Germany), upper Aptian to lower Albian of the Western Tethys (Spain), middle Albian of the Western Atlantic (Mexico), lower Cenomanian of the Western Tethys (France, Spain) and the Central Tethys (Greece), middle Cenomanian of the European Boreal (France). Lower Cretaceous without specification of the Central Tethys (Israel).
The present specimen has a decameral septal symmetry with three septal cycles. All other Eosiderastrea with such a symmetry and large corallite dimensions (as the present specimen) have four septal cycles. Hence, the present specimen is very probably a new species.
AnimaliaScleractiniaRhizangiidaeC7B6357F-90D5-5C4E-A11A-0C03F51AF602SideroheliaLöser et al., 2021аType species.
Sideroheliaaquilai Löser et al., 2021а.
Description.
Phaceloid. Corallite circular to elliptical. Septa in a regular radial symmetry and varying systems. Younger septa attached to older septa in a regular plan. Costae short. Columella by septal fusion. Synapticulae rare, only close to the wall. Wall compact, septothecal. Endotheca absent.
Solitary and colonial corals. Septa compact. Septal symmetry mainly regular, radial, or bilateral. First septal cycle (or cycles) generally thicker than later cycles. Septal lateral faces ornamented. Septal inner margins with auriculae. Lonsdaleoid septa present in two families. No main septa. Microstructure of small trabeculae. Synapticulae and pali absent. Columella present in two families. Endotheca well-developed. Marginarium absent. Wall compact, septothecal. Coenosteum present in some genera. Budding varies depending on the colony type.
Astreoid colony with small corallites. No septal symmetry. The wall is incomplete, septothecal. Endotheca well developed with numerous small dissepiments.
Remarks.
The assignation of the unique specimen to a genus is difficult. It can be compared to Preverastraea Beauvais, 1976. Although Preverastraea varies in its corallite arrangement, it has always a wall that is incomplete in the present material.
Meandrinoid colony with straight and parallel rows. Rows very narrow, coenosteum wide. In places large isolated corallites are present. Septa in rows short and irregular, in isolated corallites well developed, in a hexameral symmetry. Endotheca well developed.
Remarks.
The single specimen does not allow to establish a new genus and species.
Cerioid colony. Corallite outline irregularly polygonal, with a small diameter. Symmetry of septa radial and irregularly hexameral. Septa not connected to each other. Pali absent. Costae hardly present, confluent. Endotheca consists of numerous tabulae. Wall subcompact, septothecal. Coenosteum absent.
Plocoid colony growing in the form of branches which show corallites only on one face. Corallite outline circular. Symmetry of septa radial and regularly hexameral, rarely octameral. Pali absent. Costae present, non-confluent. Columella styliform. Endotheca consists of thin tabulae. Coenosteum broad.
The new species has a remarkable octameral symmetry of septa.
Holotype.
LFU 8336SG015215#1 with one thin section.
Locus typicus.
Germany, Bavaria, community of Lenggries, Roßsteinalmen.
Stratum typicum.
Cretaceous, middle Cenomanian, Branderfleck Fm.
Paratype.
LFU 8336SG015215#2 with one thin section.
Material.
Holotype and paratype.
Diagnosis.
Enallhelia with an octameral septal symmetry, two septal cycles, a small corallite diameter of 1.1–1.2 mm and a larger corallite diameter of 1.4 -1.7 mm.
Description.
Plocoid colony growing in the form of branches which show corallites only on one face. Corallite outline slightly elliptical. Symmetry of septa radial and regularly octameral, two septal cycles. Septa of the first cycle are thicker and longer than septa of the second cycle. Septa compact, with occasional auriculae at their inner margins. Pali absent. Costae present, non-confluent. Columella styliform. Endotheca consists of thin tabulae. Wall compact, probably septothecal. Coenosteum broad.
Remarks.
The genus has, to date, a stratigraphical range from the Oxfordian to the lower Albian. It is more common in the Upper Jurassic than the Lower Cretaceous. The present two specimens constitute the youngest occurrence of the genus.
Relationship.
Enallhelia has generally a hexameral septal symmetry. There are no formally established species with an octameral septal symmetry as in the present material.
BSPG 1991 X 44, LFU 8336SG015223; three thin sections.
Dimensions.
(LFU 8336SG015223).
n
min–max
µ
s
cv
µ±s
clmin
15
0.92–1.28
1.10
0.13
11.7
0.97–1.23
clmax
15
1.10–1.48
1.24
0.09
7.4
1.14–1.33
ccd
20
2.09–3.32
2.68
0.33
12.3
2.35–3.01
septa
10
14–17
15.70
0.82
5.2
15–17
Remarks.
The septal symmetry is octameral with two septal cycles. The species is, to date, only known from the middle Jurassic to the Valanginian.
Other occurrences.
Bathonian of the Southern Tethys (Madagascar), Aalenian to lower Valanginian of the Western Tethys (Spain), Kimmeridgian of the European Boreal (Germany), lower Tithonian of the Central Tethys (Italy).
AnimaliaScleractiniaStylinidae50FF1FA4-1589-5DD1-9BE9-66CA656B8586Stylinainwaldensis(Ogilvie, 1897)v1880Stylinabernardana ? – d’Achiardi: 288. [nonStylinabernardana Etallon, 1859] v*1897Diplocoeniainwaldensis – Ogilvie: 165, pl. 18, figs 7, 8. v2013Stylinainwaldensis (Ogilvie, 1897) – Löser, García-Barrera et al.: 396, figs 4g–i . [here more detailed synonymy] v2015Stylinainwaldensis (Ogilvie, 1897) – Löser, Arias and Vilas: 59, figs 3h, i. v2019Stylinainwaldensis (Ogilvie, 1897) – Löser, Arias and Vilas: 281, figs 11.1–3. Material.
BSPG 1947 XVI 42, 1991 X 45, 1991 X 46, 2010 VI 1; three thin sections.
Other occurrences.
Lower Tithonian of the Central Tethys (Italy), Tithonian to lower Berriasian of the European Boreal (Czech Republic), Berriasian of the Central Tethys (Ukraine), upper Valanginian of the Western Tethys (Spain), lower Hauterivian of the Central Tethys (Ukraine) and the European Boreal (France), upper Barremian to lower Aptian of the Western Atlantic (Mexico, Venezuela), Aptian of the Central Tethys (Greece), upper Albian to lower Cenomanian of the Western Tethys (Spain).
The specimen represents a holdfast of an octocoral with an approximate diametre of 15 mm.
Order <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="order">Scleralcyonacea</tp:taxon-name-part></tp:taxon-name><xref ref-type="bibr" rid="B125">McFadden et al., 2022</xref>AnimaliaScleractiniaHelioporidae4FD23590-357B-56E6-A58B-CAE307BE42BEHelioporidaeMoseley, 1876Description.
The family encompasses genera that form small plocoid colonies. The corallites are small (generally below 2 mm in diameter). The structure is simple because it is only made up of trabeculae and tabulae. The so-called septa are just short wing-like extensions of the trabeculae that reach into the corallites. Pali, a columella or synapticulae do not exist. The tabulae cross corallites and coenosteum in the same level. The coenosteum is formed by trabeculae and is granulated on its surface.
The genus forms small spheric, fingerlike or incrusting colonies. The corallites are circular and very regular. The septa are short. The coenosteum appears cellular with very fine pores and tiny spines, or is rarely vermiculate.
Remarks.
As explained in Hernández Morales and Löser (2018), the genus Polytremacis is a junior synonym of the extant genus Heliopora. The corallite dimensions of the species of the study area are compared in Table 1.
The type material of this species is very probably based on specimens that were first reported by Roemer (1862) as ”Chaetetesradians” from Carboniferous glacial drift material. The name ”Chaetetesradians” was solely mentioned, without giving a description or illustration. Remeš (1898) has, therefore, the priority.
Other occurrences.
Upper Albian of the European Boreal (UK), Turonian of the European Boreal (Poland), Coniacian of the Central Tethys (Austria), Danian of the European Boreal (France).
Measurements of the Heliopora species in the studied fauna.
https://binary.pensoft.net/fig/955664AnimaliaScleractiniaHelioporidae8433A5DB-0A50-575E-871A-7D434F2CE939Helioporaradiata(d’Orbigny, 1850)v1850Centrastrearadiata – d’Orbigny: (2), p. 207. v2013Polytremacisvermiculata (Felix, 1903) – Löser, Werner and Darga: 67, pl. 10, figs 7–9. v2018Helioporaradiata (Orbigny, 1850) – Löser, Steuber and Löser: 69, pl. 20, figs 7–9. v2019Helioporaradiata Orbigny, 1850 – Löser, Heinrich and Schuster: 324, figs 529a–c. Material.
BSPG 1947 XVI 43, 1947 XVI 55; two thin sections.
Remarks.
In Löser et al. (2013c), these specimens were assigned to Polytremacisvermiculata (Felix, 1903b), but afterwards it was possible to examine the type material of Helioporaradiata (d’Orbigny, 1850) and to take more detailed measurements in the (only) syntype of Helioporavermiculata that show that both species differ in their dimensions.
Other occurrences.
Upper Aptian to lower Cenomanian of the Western Tethys (Spain, France), lower Cenomanian of the Central Tethys (Greece) and the Western Tethys (Spain, France), Coniacian to Santonian of the Central Tethys (Austria), upper Santonian of the Western Tethys (France), Maastrichtian of the Arabian Peninsula (United Arab Emirates).
AnimaliaScleractiniaHelioporidaeFA2E9142-036E-5493-8984-6C1620978E75Helioporaramosa(d’Orbigny, 1849)Plate 15: figs 1–3*v1849Dactylacisramosa – d’Orbigny: p. 11. v1850Dactylacisramosa – d’Orbigny: (2), p. 183. v2016cDactylacisramosa Orbigny, 1849 – Löser: 276, fig. D1. v2018Helioporaramosa (Orbigny, 1849) – Hernández Morales and Löser: 357, fig. 2.6. v2019Helioporaramosa (Orbigny, 1849) – Löser, Heinrich and Schuster: 325, figs 530a–c. Material.
LFU 8336SG015104; two thin sections.
Dimensions.
(LFU 8336SG015104).
n
min–max
µ
s
cv
µ±s
clmin
30
0.48–0.67
0.57
0.05
8.8
0.52–0.62
clmax
30
0.55–0.72
0.61
0.05
7.4
0.56–0.65
septa
20
12–16
14.4
1.10
7.6
13–16
Other occurrences.
Lower Albian of the Western Atlantic (Mexico), Cenomanian to lower Coniacian of the Western Tethys (France, Spain), Coniacian to Santonian of the Central Tethys (Austria).
Lower Hauterivian of the European Boreal (France), upper Barremian of the Western Tethys (France), upper Barremian to lower Aptian of the Central Tethys (Bulgaria, Serbia), lower Aptian of the Central Tethys (Greece, Serbia) and the Western Tethys (Spain), upper Aptian of the Western Pacific (Japan), lower Albian of the Western Atlantic (Mexico), upper Cenomanian of the European Boreal (Czech Republic), Turonian of the Central Tethys (Somalia), Coniacian to Santonian of the Central Tethys (Austria), lower Eocene of the Central Tethys (Somalia) and the European Boreal (Czech Republic).
AnimaliaScleractiniaHelioporidae51647E5A-F95E-500D-B1F1-578E3B5439C6Helioporaurgonensis(Koby, 1898)v*1898Polytremacisurgonensis – Koby: 87, pl. 21, fig. 5. v1936HelioporaedwardsanaStoliczka 1873 – Hackemesser: 76, pl. 6, fig. 11. Material.
BSPG 1947 XVI 40, 1947 XVI 41, 1947 XVI 7, 1991 X 62; two thin sections.
Dimensions.
(BSPG 1947 XVI 7).
n
min–max
µ
s
cv
µ±s
clmin
25
0.56–0.78
0.67
0.07
9.9
0.60–0.74
clmax
25
0.63–0.86
0.74
0.07
8.9
0.68–0.81
septa
22
11–17
14.86
1.58
10.6
13–16
Other occurrences.
Barremian of the Central Tethys (France), lower Albian of the Western Atlantic (Mexico), Coniacian to Santonian of the Central Tethys (Austria), Cretaceous without specification of the Central Tethys (Greece).
AnimaliaScleractiniaHelioporidaeDCFE559F-2FEB-579E-8A18-F91143808405Helioporasp. 1v2013Polytremacis sp. – Löser, Werner and Darga: 68, pl. 10, figs 10–12. Material.
BSPG 1947 XVI 63; one thin section.
Other occurrences.
Coniacian of the Central Tethys (Austria), upper Eocene of the Western Atlantic (Panama).
LFU 8336SG015021#1, 8336SG015021#2; three thin sections.
Dimensions.
(LFU 8336SG015021#2).
n
min–max
µ
s
cv
µ±s
clmin
9
1.11–1.36
1.27
0.09
6.9
1.18–1.35
clmax
9
1.22–1.52
1.38
0.11
7.6
1.27–1.48
septa
3
19–23
21.33
2.08
9.8
19–23
Other occurrences.
Lower Aptian of the Central Tethys (Italy), upper Cenomanian of the Western Tethys (France), upper Turonian to Santonian of the Central Tethys (Austria), Palaeocene of the European Boreal (Germany).
https://binary.pensoft.net/fig/955665AnimaliaScleractiniaHelioporidae368B9115-0B48-5F17-B957-809A1A2EA16EHelioporasp. 3v2013Polytremacisbofilli (Bataller, 1936) – Löser, Werner and Darga: 67, pl. 10, figs 1–3. vp2019Helioporapartschi Reuss, 1854 – Löser, Heinrich and Schuster: 324. Material.
BSPG 1947 XVI 14, 1947 XVI 25, 1947 XVI 50, 1991 X 40, 1991 X 61, LFU 8336SG015015#1, 8336SG015015#2, 8336SG015096; seven thin sections.
Remarks.
In Löser et al. (2013c), the material was assigned to Polytremacisbofilli. After more detailed measuring it transpires that the dimensions are larger than in Helioporabofilli. Helioporabofilli itself is probably a junior synonym of Helioporablainvilleana Michelin, 1841. This is difficult to decide; the type material of Helioporablainvilleana comes from Uchaux (Vaucluse, France) and is very poorly preserved. Therefore, Helioporablainvilleana probably should be discarded. The type material of Helioporapartschi is available but did not deliver well-constrained morphometric data.
Other occurrences.
Coniacian to Santonian of the Central Tethys (Austria), lower Santonian of the Western Tethys (France), Campanian to Maastrichtian of the Western Atlantic (Jamaica), Maastrichtian of the Western Tethys (France).
Discussion
In the first description of the coral fauna from the Roßsteinalm area, 39 species in 25 genera were described. This amount has more than doubled, now reaching 98 species in 46 genera. The new material from the Bayerisches Landesamt für Umwelt - Geologie/Paläontologie (Hof) and the Museum für Naturkunde der Humboldt-Universität (Berlin) is taxonomically very different from the formerly studied material kept at the Bayerische Staatssammlung für Paläontologie und Geologie (Munich). We can only speculate about the reasons. Most material (Hof and Munich) was sold or donated by the private collector Otto Hölzl, as aforementioned. It is possible that the Bayerisches Landesamt für Umwelt - Geologie/Paläontologie first picked out attractive material from his collection and the remaining material went to the Bayerische Staatssammlung für Paläontologie und Geologie. Another possibility is that the material was collected at different times and at slightly different places or horizons at the Roßsteinalmen, or that the attractive specimens were the first to be collected and later all of the rest were collected. For example, all of the material of the genus Eothelia (that is unattractive for collectors) is housed at the Bayerische Staatssammlung für Paläontologie und Geologie (Munich). No specimen was found in the other two collections. On the other hand, the (attractive) specimens of the genus Felixarea are all from the Bayerisches Landesamt für Umwelt - Geologie/Paläontologie (Hof).
The new material has enriched the fauna considerably but various taxonomic assignments – genera and species – of the first revision have changed (Table 2). This has different reasons:
After the publication, numerous collections in Europe and the USA were visited by the first author (H.L.) and much type material was studied. Of importance for the present revision was the study of the collections of Johannes Felix (Leipzig), Paul Oppenheim (Jerusalem), and August Emanuel Reuss (Vienna).
With the publication of the systematic revision of the Cretaceous corals on the genus level (Löser 2016c) an improved classification system was proposed. Therefore, the systematic position of many genera was modified, and many genera became synonyms.
The revision of the corals of the Conacian and Santonian corals of the area of Rußbach and Gosau (Austria) by Löser et al. (2019) has improved the knowledge on Late Cretaceous corals that also had influence on the taxonomy of Cenomanian corals.
Finally, the improved methods of recording and analysing morphometric data in Scleractinian corals changed the viewpoint at the species concept level and the way in which species are distinguished. Many type specimens were measured with more details and much more morphometric data became available overall.
Faunal composition
The present middle Cenomanian coral fauna encompasses 98 coral species, with 88 species belonging to the subclass Hexacorallia, and ten species belonging to the subclass Octocorallia. Compared to other Cretaceous coral faunas, this is a comparably high value which is so far only nearly reached or even exceeded by two Cenomanian coral faunas: Kozani (Greece; Löser et al. 2018) and Cantabria (Spain; Löser and Wilmsen 2022). There are two further upper Cretaceous species-rich faunas: the Turonian to Santonian coral faunas from Austria (Löser et al. 2019), and a Campanian fauna from Spain (e.g. Bataller 1937). Both coral faunas are allochthonous, in that the material was brought together from different areas and may also differ slightly in age. This is not the case for the present coral fauna.
With the exception of one specimen originating from the upper part of the Roßsteinalmen section, all other specimens certainly come from the marly facies of the lower part of this section. However, an attribution to single horizons and thus a separation of different palaeocommunities is not possible.
In our first compilation, published in 2013, we stated that solitary corals dominate the collection in number of specimens (80%), whereas colonial forms dominate in number of genera. The new material includes coral genera of various growth forms and although colonial coral genera still dominate the association, the number of solitary coral genera increased (Fig. 3). The new specimens - solitary and colonial corals - are also small, rarely exceeding 5 cm in their largest dimension. This may suggest - as already discussed in Löser et al. (2013c) - unfavourable substrate conditions and limited light. All of those limiting factors had no influence on the species richness.
Distribution of the corallite integration types in the studied fauna. The total number of genera (50) is larger because all corals where included, also those not assigned to any genus.
https://binary.pensoft.net/fig/955666
Changes in the taxonomy for the Roßsteinalmen coral fauna published in 2013.
Leptophyllaraea cf. granulata (de Fromentel, 1863)
Leptophyllaraea cf. granulata (de Fromentel, 1863)
Microphyllia cf. oldhamiana (Stoliczka, 1873)
Microphyllia cf. oldhamiana (Stoliczka, 1873)
Negoporites cf. quartus Eliášová, 1995
Negoporites cf. quartus Eliášová, 1995
Negoporites sp.
Negoporites sp.
Neocoenia cf. casterasi (Alloiteau, 1957)
Neocoeniaexsculpta (Reuss, 1854)
Neocoenia cf. casterasi (Alloiteau, 1957)
Neocoeniakuehnii (Oppenheim, 1930)
Neocoeniarenzi (Hackemesser, 1936)
Neocoeniarenzi (Hackemesser, 1936)
Pachygyrakrameri Oppenheim, 1930
Pachygyracucullata (Zuffardi-Comerci, 1930)
Paramontlivaltiaruvida (Prever, 1909)
Trochophylliaaprutina (Prever, 1909)
Paramontlivaltiaruvida (Prever, 1909)
Trochophylliaruvida (Prever, 1909)
Polytremacisbofilli (Bataller, 1936)
Heliopora sp. 3
Polytremacis sp.
Heliopora sp. 1
Polytremacisvermiculata (Felix, 1903)
Helioporaradiata (d’Orbigny, 1850)
Stelidioserisminima (de Fromentel, 1857)
Actinastrealimbata Alloiteau, 1954
Synastrea sp.
Synastrea sp.
Thalamocaeniopsis sp.
Polyastropsis cf. fascigera (Felix, 1909)
Trochoseropsisettalensis Söhle, 1897
Trochoseropsisettalensis Söhle, 1897
Twenty-seven superfamilies of the order Scleractinia were distinguished in Löser (2016c) for the Cretaceous. Another superfamily is introduced here (Dendrogyroidea) and one superfamily (Ficariastraeoidea) is put into synonymy with the Phyllosmilioidea. Of the resulting 27 superfamilies, 22 occur in the Middle Cenomanian. In the studied fauna, 15 of those superfamilies are present. For the remaining seven superfamilies Dendrophyllioidea, Fungioidea, Haplaraeoidea, Madreporoidea, Micrabacioidea, Orbicelloidea, and Stylophoroidea an occurrence in the Middle Cenomanian is confirmed, but they were not found in the study area.
Fig. 4 shows the comparison of the number of genera per superfamily for the Albian to Turonian on a global scale to the number of genera per superfamily in the study area (only order Scleractinia). Both global and local data are very similar, with the difference that in the present fauna the Caryophyllioidea, Eugyroidea, Montlivaltioidea and Phyllosmilioidea have less genera compared to the global data, whereas the Cyclolitoidea and Stylinoidea are represented by slightly more genera. The lower number of Caryophyllioidea genera in the present shallow water fauna can be explained by the preference of this superfamily for deeper marine environments. On the whole the faunal composition is comparable to other coral faunas of Cenomanian age.
Distribution of genera of the studied fauna among superfamilies (order Scleractinia) based on verified stratigraphical ranges. These ranges were obtained through the observation of material and do not come from the literature. A comprehensive compilation of the ranges were published in Löser (2016c). The left bar shows the distribution of (192) genera worldwide that have an occurrence (at least partly) in the Cenomanian. Only those superfamilies are shown that occur also in the study area, resulting in 124 genera. The right bar shows the distribution of (50) genera of the studied fauna.
https://binary.pensoft.net/fig/955667Stratigraphic distribution
Fig. 5 is an alphabetical chart that shows the stratigraphical distribution of the species of the study area in other areas. It can be observed that the species of the same genus have a comparable stratigraphical distribution. Some genera have their principal distribution in the Lower Cretaceous and others in the Upper Cretaceous. This is obvious for the lower Cretaceous genera Astraeofungia, Cryptocoenia, Stylina and the upper Cretaceous genera Actinastrea, Aulosmilia, Hydnophoraraea, Synastrea. Only a few genera are occurring throughout the whole Cretaceous, such as the octocoral Heliopora.
When looking directly at the stratigraphical distribution of the genera (Fig. 6), this polarisation is even more obvious: 15 genera occur in the Lower Cretaceous but have their last occurrence in the Cenomanian; 14 genera occur before and continue after the Cenomanian; eight genera occur only in the Cenomanian; nine genera originate in the Cenomanian and continue into the Upper Cretaceous.
The summarised data in Fig. 7 for both species and genera show that more taxa of this study have their stratigraphic distribution in the Lower Cretaceous than in the Upper Cretaceous. There are many genera that had their last occurrence in the Cenomanian, some were present in the whole Cretaceous and many others had their first occurrence in the Cenomanian. Some genera experienced a range extension with the present fauna; the most striking are Cyathophoropsis, Enallhelia and Felixaraea.
Palaeobiographic relationships
Fig. 8 shows the correlation of palaeo-provinces (faunules) based on (76) species of the study area that have a distribution in other areas. The number of joint species is generally low and, therefore, no clear pattern can be observed. The highest number of species (16 out of 76) shares the present fauna with the coral fauna from Cóbreces (Cantabria, Spain; Löser and Wilmsen 2022). Generally, the present fauna tends to correlate more with Tethyan faunas; Boreal faunas are in a separate cluster. Even if not shown in Fig. 8, the present fauna shares many species with the well-studied Coniacian to Santonian coral fauna of the area of Rußbach and Gosau (Austria); out of the 76 species that were indicated at other places, 27 occur also in the Rußbach/Gosau area. When comparing only coral faunas with a range in the Cenomanian (Fig. 9), the fauna from Cóbreces again shows the highest number of joint species. The correlation with other areas does not allow a clear conclusion; very probably because the number of joint species is too low.
Evolutionary aspects
Almost ten years have passed since the publication of the first part of the coral fauna from the Northern Calcareous Alps . At the same time or afterwards, more large Albian to Cenomanian coral faunas were published (e.g. Löser 2013a, 2014b, 2015a; Löser et al. 2015; Löser et al. 2013a; Löser and Bilotte 2017; Löser et al. 2018; Löser and Wilmsen 2022, 2023). The Coniacian to Santonian coral fauna of the Gosau area was revised (Löser et al. 2019). All of these revisions have extended or limited the stratigraphical ranges of the coral genera. When correlating the palaeo-provinces based on genera (Fig. 10), two major groups can be observed, the Lower Cretaceous plus Cenomanian and the Turonian to Santonian. There is one exception; the Quillan Basin of Cenomanian age is grouped together with the Turonian to Santonian provinces. The Quillan Basin province encompasses, principally, the locality of Prat-Périe (Sougraigne, Les Corbières, Aude, France) with an Uppermost Cenomanian age (juddii-zone; Bilotte 1985; Michel Bilotte personal communication). It is shown again that the faunal turnover (see also Löser and Callapez 2022, fig. 9) from the Lower Cretaceous plus Cenomanian to the Turonian and younger strata was transitional and can only be partly connected to the sea level highstand and Ocean Anoxic Event 2 at the Cenomanian/Turonian boundary. When looking at the stratigraphic distribution of the superfamilies (that may represent monophyletic groups), the faunal turnover is less obvious (Fig. 11). Only three superfamilies became extinct during the Cenomanian, and three superfamilies originated.
Distribution of species of the studied fauna in localities outside of the study area. The thickness of the horizontal bars corresponds to the number of localities where the species was found. The vertical green bar marks the age of the investigated coral fauna. Only Tithonian to Palaeocene is shown.
Verified stratigraphic ranges of the coral genera in the study area, sorted according to their occurrence. Ranges as above after Löser (2016c), but improved by newer data. Red lines mark range extensions as a result of this study. The vertical green bar indicates the age of the studied fauna. Only Tithonian to Palaeocene is shown; some genera may have longer ranges.
During the Cenomanian, the general trend of a rising sea level continued (Haq et al. 2014). Whereas the lower Cenomanian is characterised by a sea level increase, the middle Cenomanian experienced a slight decrease. During the upper Cenomanian, the sea level increased again, reaching a high stand at the Cenomanian/Turonian boundary. Globally, nearly all precisely dated large Cenomanian coral faunas can be assigned either to the lower or the upper Cenomanian (for example, for the Boreal regions see Löser 1994, for the Western Tethys Löser and Wilmsen 2022, for the Central Tethys Löser and Bilotte 2017, for the Eastern Tethys Löser et al. 2018, and literature cited therein). Shallow marine middle Cenomanian coral faunas are poorly reported. The classical Cenomanian outcrop area near Le Mans (Sarthe, France) can only be assigned to the middle to upper Cenomanian (Morel ed. 2015). Middle to Upper Cenomanian corals are mentioned from the Batna area in Algeria (Vila 1977). Some of these outcrops could be restricted to the middle Cenomanian. Unrevised coral faunas of a lower to middle Cenomanian age are reported from Tournai beds in Hainaut (Belgium; d’Archiac 1847) The Boreal coral fauna of the Meißen Fm (Saxon, Germany) has a lower middle Cenomanian age (Löser 2014b).
Summarised ranges of genera and species. The chart of genera shows the verified distribution of the genera of the study area (based on Löser 2016c with later precision). The chart of species shows the number of faunules where the species from the study area occurred (not localities). The new data obtained in the present study are not shown in both charts. The vertical green bar marks the time interval of the study area.
Correlation of the palaeo-provinces where species of the studied fauna occur. Only provinces with more than two species were included, and only the time period Albian to Turonian is considered. The Correlation Ratio coefficient was applied. Abbreviations: Ap, Aptian; Al, Albian; Ce, Cenomanian, Tu, Turonian, Co, Coniacian. The number 1. indicate lower, the number 2. middle, and the number 3. upper. The numbers in brackets are the numbers of joint species. The stratigraphy of the area Pelagonium S margin is uncertain. The study area is marked in bold letters.
Correlation of the palaeo-provinces with a range in the Cenomanian where species of the studied fauna occur. Only provinces with more than one species were included. The Correlation Ratio coefficient was applied. Abbreviations: Ap, Aptian; Al, Albian; Ce, Cenomanian, Tu, Turonian, Co, Coniacian. Number 1. indicates lower, number 2. indicates middle and number 3. indicates upper. The numbers in brackets are the numbers of joint species. The stratigraphy of the area Pelagonium S margin is uncertain, but also clearly includes Cenomanian corals. The study area is marked in bold letters.
Correlation of the palaeo-provinces where genera of the studied fauna occur. Only provinces with more than two genera were included, and only the time period Hauterivian to Santonian was considered. The Correlation Ratio coefficient was applied. Abbreviations: Va, Valanginian; Ha, Hauterivian; Ap, Aptian; Al, Albian; Ce, Cenomanian, Tu, Turonian, Co, Coniacian; Sa, Santonian. Number 1. indicates lower, number 2. indicates middle and number 3. indicates upper. The numbers in brackets are the numbers of joint genera. The stratigraphy of the area Pelagonium S margin is uncertain. The study area is marked in bold letters. The dotted line separates Hauterivian to Cenomanian and Turonian to Santonian palaeo-provinces (with one exception).
Verified stratigraphic distribution of the superfamilies. The ranges are based on the verified distribution data for the genera, as explained above. Only Albian to Turonian is shown. The vertical green bar marks the age of the studied fauna. All superfamilies are shown, as well as those superfamilies that do not occur in the studied fauna. Superfamilies that do not occur in the study area are shown as light grey bars.
https://binary.pensoft.net/fig/955674
The present large coral fauna that can clearly be restricted to the middle Cenomanian is, therefore, an exceptional record. The duration of the lower, middle and upper Cenomanian is too short to allow the recognition of evolutionary changes, but the present fauna links together, even more solidly, the typical upper Lower Cretaceous coral faunas (Barremian to Albian) and the lower Upper Cretaceous (Turonian to Santonian). One reason may be the marly facies; other Cenomanian coral faunas are found in coastal transgressive areas with a considerable siliciclastic input, or in deposits of carbonate ramps. Other Cenomanian coral faunas in a marly facies such as from the study area are hardly known.
With the present study it can be confirmed that the shallow marine coral faunas experienced a transition that already started in the Lower Cretaceous and continued during the Cenomanian. The gap in the fossil coral record during the Turonian is just the lack of shallow marine sediments (and therefore outcrop areas; Löser 2016c, Fig. 6.3.5.3) due to the regression during the Turonian and the subsequent erosion. Coniacian coral faunas differ from Cenomanian faunas which concerns particularly the genus richness of vicariant families. As, for instance, Latomeandridae became reduced in the number of genera, Synastraeidae increased, or Montlivaltiidae were reduced in genera but Phyllosmiliidae were increased (cf. Löser 2016c, Fig. 6.2.2).
Acknowledgements
We are grateful to Erwin Geiss from the Bayerisches Landesamt für Umwelt and Dieter Korn from the Berliner Museum für Naturkunde for providing access to new material from the Roßsteinalmen. Preparation of thin sections by Aimée Orcí (Hermosillo) is kindly acknowledged. English text correction by Matthew Copley (Barcelona). Reviews by Ricardo Garberoglio (Buenos Aires) and Jacob Leloux (Leiden) helped to improve the manuscript.
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