Scleractinian corals of the suborders Dendrophylliina

ZOOTAXA 1952

Corals of the K/T-boundary: Scleractinian corals of the suborders Dendrophylliina, Caryophylliina, Fungiina, Microsolenina, and Stylinina ROSEMARIE C. BARON-SZABO

Magnolia Press Auckland, New Zealand

Rosemarie C. Baron-Szabo Corals of the K/T-boundary: Scleractinian corals of the suborders Dendrophylliina, Caryophylliina, Fungiina, Microsolenina, and Stylinina (Zootaxa 1952) 244 pp.; 30 cm. 5 Dec. 2008 ISBN 978-1-86977-303-8 (paperback) ISBN 978-1-86977-304-5 (Online edition)

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2 · Zootaxa 1952 © 2008 Magnolia Press

BARON-SZABO

Zootaxa 1952: 1–244 (2008) www.mapress.com / zootaxa/

ISSN 1175-5326 (print edition)

Copyright © 2008 · Magnolia Press

ISSN 1175-5334 (online edition)

ZOOTAXA

Corals of the K/T-boundary: Scleractinian corals of the suborders Dendrophylliina, Caryophylliina, Fungiina, Microsolenina, and Stylinina ROSEMARIE C. BARON-SZABO Smithsonian Institution, Department of Invertebrate Zoology, W-329, MRC-163, National Museum of Natural History, P.O. Box 37012, Washington, DC, 20013-7012, USA, and Senckenberg Research Institute, Senckenberganlage 25, D-60325 Frankfurt/Main, Germany. E-mails: [email protected] and [email protected]

Table of contents Abstract ......................................................................................................................................................................... 9 Introduction ....................................................................................................................................................................... 10 Material and methods ........................................................................................................................................................ 10 Results ....................................................................................................................................................................... 15 Systematic Palaeontology ................................................................................................................................................. 30 Order Scleractinia Bourne, 1900 ................................................................................................................................ 30 Suborder Dendrophylliina Vaughan & Wells, 1943 ........................................................................................... 30 Family Dendrophylliidae Gray, 1847 ........................................................................................................... 31 Genus Dendrophyllia Blainville, 1830 ................................................................................................. 31 Dendrophyllia candelabrum Hennig, 1899 ................................................................................... 31 Dendrophyllia dendrophylloides Milne Edwards & Haime, 1848 ................................................ 34 Genus Areopsammia Dietrich, 1917 ..................................................................................................... 34 Areopsammia alacca (Morren, 1828) ........................................................................................... 35 Genus Palaeopsammia Wanner, 1902 .................................................................................................. 35 Palaeopsammia collignoni (Alloiteau, 1958) ............................................................................... 36 Palaeopsammia zitteli (Vaughan, 1900) non Wanner, 1902 ......................................................... 38 Genus Wadeopsammia Wells, 1933 ...................................................................................................... 39 Wadeopsammia nodosa (Wade, 1926) .......................................................................................... 40 Genus Balanophyllia Searles Wood, 1844 ............................................................................................ 40 Balanophyllia besairiei (Alloiteau, 1958) .................................................................................... 44 Balanophyllia caulifera (Conrad, 1847) ....................................................................................... 43 Balanophyllia ponderosa Vaughan, 1900 ..................................................................................... 44 Suborder Caryophylliina Vaughan & Wells, 1943 ............................................................................................. 44 Family Caryophylliidae Dana, 1846 ............................................................................................................ 45 Subfamily Caryophylliinae Milne Edwards, 1857 ................................................................................... 45 Genus Caryophyllia Lamarck, 1801 ..................................................................................................... 45 Caryophyllia arcotensis (Forbes, 1846) ........................................................................................ 45 Caryophyllia konincki (Milne Edwards & Haime, 1848) ............................................................. 48 Caryophyllia mediavia Vaughan & Popenoe, 1935 ...................................................................... 51 Genus Bathycyathus Milne Edwards & Haime, 1848 .......................................................................... 52 Bathycyathus corneti (Alloiteau & Tissier, 1958) ......................................................................... 52 Bathycyathus lloydi (Vaughan, 1920) ........................................................................................... 53 Bathycyathus piveteaui (Alloiteau & Tissier, 1958) ..................................................................... 54 Genus Trochocyathus Milne Edwards & Haime, 1848 ......................................................................... 55 Trochocyathus speciosus (Gabb & Horn, 1860) ........................................................................... 56 Trochocyathus mitratus (Goldfuss, 1826) ..................................................................................... 57 Accepted by C. Wallace: 25 Sept. 2008; published: 5 Dec. 2008

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Trochocyathus lakii Duncan, 1880 ................................................................................................ 59 Genus Paracyathus Milne Edwards & Haime, 1848 ............................................................................ 62 Paracyathus altus Gregory, 1930 .................................................................................................. 62 Genus Deltocyathus Milne Edwards & Haime, 1848 ........................................................................... 63 Deltocyathus complanatus Squires, 1958 ..................................................................................... 63 Deltocyathus cupuliformis (Alloiteau, 1951) ................................................................................ 64 Deltocyathus whitei Durham, 1943 ............................................................................................... 64 Genus Stylocyathus d’Orbigny, 1850 .................................................................................................... 65 Stylocyathus besairiei Alloiteau, 1958 .......................................................................................... 65 Genus Stephanosmilia de Fromentel, 1862 .......................................................................................... 66 Stephanosmilia madagascariensis Alloiteau, 1958 ...................................................................... 66 Genus Stephanocyathus Seguenza, 1864 .............................................................................................. 66 Stephanocyathus sp. ...................................................................................................................... 66 Genus Cyathoceras Moseley, 1881 ....................................................................................................... 67 Cyathoceras embaensis Kuzmicheva, 1987 .................................................................................. 68 Cyathoceras mangyschlagensis Kuzmicheva, 1987 ..................................................................... 68 Genus Ceratotrochus Milne Edwards & Haime, 1848 ......................................................................... 69 Ceratotrochus supracretacea Hennig, 1899 ................................................................................. 69 Subfamily Parasmiliinae Vaughan & Wells, 1943 ................................................................................... 70 Genus Parasmilia Milne Edwards & Haime, 1848 .............................................................................. 70 Parasmilia centralis (Mantell, 1822)............................................................................................ 70 Parasmilia elongata Milne Edwards & Haime, 1848 ................................................................... 72 Genus Faksephyllia Floris, 1972 .......................................................................................................... 75 Faksephyllia faxoensis (Beck, 1835) ............................................................................................ 75 Genus Caryosmilia Wanner, 1902 ........................................................................................................ 75 Caryosmilia granosa Wanner, 1902 .............................................................................................. 76 Genus Asterosmilia Duncan, 1867........................................................................................................ 78 Asterosmilia alloiteaui (Alloiteau & Tissier, 1958) ...................................................................... 78 Genus Dasmosmilia Pourtalès, 1880 .................................................................................................... 79 Dasmosmilia kochii (Pratz, 1910) ................................................................................................. 79 Genus Smilotrochus Milne Edwards & Haime, 1851 ........................................................................... 80 Smilotrochus cornucopiae (Duncan, 1869) ................................................................................... 81 Smilotrochus galeriformis (Kner, 1848) ....................................................................................... 82 Smilotrochus ponderosus (Nielsen, 1922) ..................................................................................... 83 Smilotrochus milneri (Gregory, 1898) .......................................................................................... 84 Subfamily Desmophyllinae Vaughan & Wells, 1943 ............................................................................... 84 Genus Desmophyllum Ehrenberg, 1834 ................................................................................................ 84 Desmophyllum excavatum (Hagenow, 1839) ................................................................................ 85 Family Flabellidae Bourne, 1905 ................................................................................................................. 86 Genus Flabellum Lesson, 1831 ............................................................................................................ 86 ?Flabellum anderssoni (Felix, 1909) ............................................................................................ 86 Flabellum conoideum Vaughan, 1900........................................................................................... 87 Flabellum cuneiforme Lonsdale, 1845 .......................................................................................... 88 ?Flabellum fresnoense Durham, 1943 .......................................................................................... 90 Family Turbinoliidae Milne Edwards & Haime, 1848 ................................................................................ 90 Genus Turbinolia Lamarck, 1816 .......................................................................................................... 91 Turbinolia dickersoni Nomland, 1916 ........................................................................................... 91 Genus Bothrophoria Felix, 1909 ........................................................................................................... 92 Bothrophoria ornata Felix, 1909 ................................................................................................... 92 Genus Alveolocyathus Filkorn, 1994 ..................................................................................................... 93 Alveolocyathus felixi (Filkorn, 1994) ............................................................................................ 93 Genus Wellsotrochus Squires, 1960...................................................................................................... 93 Wellsotrochus cyathiformis (Squires, 1958) .................................................................................. 94 Wellsotrochus discus Squires, 1962 .............................................................................................. 94


BARON-SZABO

Genus Palocyathus Filkorn, 1994 ......................................................................................................... 96 Palocyathus seymourensis Filkorn, 1994 ...................................................................................... 96 Genus Platytrochus Milne Edwards & Haime, 1848 ............................................................................ 96 Platytrochus claibornensis de Gregorio, 1890 .............................................................................. 97 Suborder Fungiina Verrill, 1865 ......................................................................................................................... 97 Family Fungiidae Dana, 1846 ...................................................................................................................... 97 Genus Cycloseris Milne Edwards & Haime, 1849 ............................................................................... 97 Cycloseris (?) wellsi Baron-Szabo, 2002 ...................................................................................... 98 Family Fungiacyathidae Chevalier, 1987..................................................................................................... 99 Genus Fungiacyathus Sars, 1872........................................................................................................... 99 Fungiacyathus deltoidophorus (Felix, 1909) ................................................................................ 99 Family Acrosmiliidae Alloiteau, 1952 ....................................................................................................... 101 Genus Acrosmilia d’Orbigny, 1849 .................................................................................................... 101 Acrosmilia conica d'Orbigny, 1850 ............................................................................................. 101 Family Actinacididae Vaughan & Wells, 1943 ........................................................................................... 103 Genus Actinacis d’Orbigny, 1849 ....................................................................................................... 103 Actinacis barretti Wells, 1934 ..................................................................................................... 103 Actinacis haueri Reuss, 1854 ...................................................................................................... 104 Actinacis martiniana d’Orbigny, 1850......................................................................................... 105 Actinacis remesi Felix, 1903b ...................................................................................................... 106 Actinacis reussi Oppenheim, 1930 .............................................................................................. 108 Genus Actinhelia d'Orbigny, 1849 ...................................................................................................... 108 Actinhelia elegans (Goldfuss, 1826) ........................................................................................... 109 Genus Elephantaria Oppenheim, 1930 .............................................................................................. 109 Elephantaria lindstroemi Oppenheim, 1930 ............................................................................... 110 Family Haplaraeidae Vaughan & Wells, 1943 112 Subfamily Haplaraeinae Vaughan & Wells, 1943 .................................................................................. 112 Genus Astraraea Felix, 1900 .............................................................................................................. 113 Astraraea media (Sowerby, 1832) ............................................................................................... 113 Astraraea multiradiata (Reuss, 1854) ......................................................................................... 114 Genus Pleurocora Milne Edwards & Haime, 1848............................................................................. 115 Pleurocora arachnoides (Knorr & Walch, 1777) ........................................................................ 115 Pleurocora explanata Milne Edwards & Haime, 1848 .............................................................. 117 Pleurocora haueri Milne Edwards & Haime, 1848..................................................................... 117 Genus Pseudofavia Oppenheim, 1930 ................................................................................................ 118 Pseudofavia grandiflora (Reuss, 1854) ....................................................................................... 119 Genus Stiboriopsis Vaughan, 1899 ...................................................................................................... 119 Stiboriopsis jamaicaensis Vaughan, 1899.................................................................................... 120 Subfamily Meandrophylliinae Roniewicz, 1976..................................................................................... 121 Genus Meandrophyllia d’Orbigny, 1849 ............................................................................................. 121 Meandrophyllia oceani (de Fromentel, 1877) ............................................................................. 121 Meandrophyllia textilis (Goldfuss, 1826) .................................................................................... 122 Meandrophyllia velamentosa (Goldfuss, 1826) ........................................................................... 123 Genus Brachymeandra Alloiteau, 1957............................................................................................... 125 Brachymeandra leptophylla (Reuss, 1854).................................................................................. 125 Family Agariciidae Gray, 1847 ................................................................................................................... 126 Genus Cyathoseris Milne Edwards & Haime, 1849............................................................................ 126 Cyathoseris catadupensis (Vaughan, 1899) ................................................................................. 126 Cyathoseris formosa d’Achiardi, 1875 ........................................................................................ 127 Genus Pavona Lamarck, 1801............................................................................................................. 128 Pavona bronni (Haime, 1850)...................................................................................................... 128 Genus Morphastrea d’Orbigny, 1850 .................................................................................................. 130 Morphastrea escharoides (Goldfuss, 1826) ................................................................................ 130 Genus Heterogyra Reuss, 1868 .......................................................................................................... 132

SCLERACTINIAN CORALS OF THE K/T-BOUNDARY

Zootaxa 1952 © 2008 Magnolia Press ·

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Heterogyra murchisoni (d’Archiac & Haime, 1853) ................................................................... 132 Genus Anisoria Vidal, 1917................................................................................................................. 133 Anisoria vidali (Mallada, 1892) .................................................................................................. 133 Genus Valloria Vidal, 1874 ................................................................................................................. 134 Valloria egozcuei Vidal, 1874 ..................................................................................................... 134 Genus Trochoseris Milne Edwards & Haime, 1849 ........................................................................... 135 Trochoseris aperta Duncan, 1864 ............................................................................................... 135 Genus Vaughanoseris Wells, 1934 ...................................................................................................... 136 Vaughanoseris catadupensis Wells, 1934 ................................................................................... 136 Genus Ogilviastraea Oppenheim, 1930............................................................................................... 137 Ogilviastraea bigemmis (Felix, 1903) ........................................................................................ 137 Family Agathiphylliidae Vaughan & Wells, 1943 ..................................................................................... 138 Genus Agathiphyllia Reuss, 1864 ........................................................................................................ 138 Agathiphyllia bosniaca (Oppenheim, 1912) ............................................................................... 138 Genus Pattalophyllia D’Achiardi, 1867 ............................................................................................. 140 Pattalophyllia grumi (Catullo, 1852) .......................................................................................... 140 Family Poritidae Gray, 1842 ...................................................................................................................... 141 Genus Goniopora Blainville, 1830 ..................................................................................................... 142 Goniopora elegans (Leymerie, 1846) ......................................................................................... 143 Goniopora websteri (Bowerbank, 1840) ..................................................................................... 144 Goniopora imperatoris Vaughan, 1919 ....................................................................................... 144 Goniopora microscopica (Duncan, 1863) ................................................................................... 145 Goniopora reussiana (Duncan, 1865).......................................................................................... 146 Family Micrabaciidae Vaughan, 1905 ....................................................................................................... 146 Genus Micrabacia Milne Edwards & Haime, 1849 ............................................................................ 147 Micrabacia radiata (Goldfuss, 1827) ......................................................................................... 147 Micrabacia marylandica Stephenson, 1916 ................................................................................ 151 Genus Stephanophyllia Michelin, 1841 .............................................................................................. 152 Stephanophyllia indica Duncan, 1880 ........................................................................................ 152 Stephanophyllia lanquinei Alloiteau, 1936 ................................................................................. 153 Family Siderastreidae Vaughan & Wells, 1943 .......................................................................................... 153 Genus Siderastrea Blainville, 1830 .................................................................................................... 154 Siderastrea adkinsi (Wells, 1934) ............................................................................................... 154 Siderastrea conferta (Duncan, 1863) .......................................................................................... 155 Siderastrea vancouverensis Vaughan, 1923 ................................................................................ 155 Genus Siderofungia Reis, 1889............................................................................................................ 157 Siderofungia morloti (Reuss, 1864) ............................................................................................ 157 Genus Sideroseris Wells, 1945 ........................................................................................................... 158 Sideroseris durhami Wells, 1945 ................................................................................................ 158 Genus Pironastrea d'Archiardi, 1875 ................................................................................................. 159 Pironastrea discoides d'Archiardi, 1875 ..................................................................................... 160 Genus Hindeastraea White, 1888....................................................................................................... 160 Hindeastraea discoidea White, 1888 .......................................................................................... 160 Hindeastraea garloica (Tchéchmédjiéva, 1975) ......................................................................... 161 Suborder Microsolenina Morycowa & Roniewicz, 1995 ................................................................................. 162 Family Synastreaidae Alloiteau, 1952 ....................................................................................................... 162 Genus Synastrea Milne Edwards & Haime, 1848 .............................................................................. 162 Synastrea agaricites (Goldfuss, 1829) ........................................................................................ 162 Synastrea garumnica (Vidal, 1921) ............................................................................................ 163 Synastrea geometrica (Goldfuss, 1826) ....................................................................................... 164 Family Microsolenidae Koby, 1889 ........................................................................................................... 166 Genus Polyphylloseris de Fromentel, 1857 ........................................................................................ 166 Polyphylloseris microkothos n. sp. ............................................................................................. 166 Family Cunnolitidae Alloiteau, 1952 ......................................................................................................... 167


BARON-SZABO

Genus Cunnolites Alloiteau, 1957 ...................................................................................................... 168 Cunnolites cancellata (Goldfuss, 1826) ...................................................................................... 168 Cunnolites tenuiradiata (de Fromentel, 1863) ............................................................................ 169 Cunnolites scutellum (Reuss, 1854) ............................................................................................ 170 Cunnolites giganteus (d’Orbigny, 1850) ..................................................................................... 172 Cunnolites polymorphus (Goldfuss, 1826) ................................................................................. 173 Cunnolites angiostoma (Kühn, 1933) ......................................................................................... 176 Genus Aspidastraea Kühn, 1933 ........................................................................................................ 176 Aspidastraea clathrata (Goldfuss, 1826) .................................................................................... 178 Aspidastraea orientalis Kühn, 1933 ........................................................................................... 178 Aspidastraea semhae (Kossmat, 1907) ....................................................................................... 179 Genus Paracycloseris Wells, 1934 ..................................................................................................... 180 Paracycloseris nariensis (Duncan, 1880) ................................................................................... 180 Family Latomeandridae Alloiteau, 1952 ................................................................................................... 182 Genus Dimorphastrea d’Orbigny, 1850 .............................................................................................. 182 Dimorphastrea solida Umbgrove, 1925 ..................................................................................... 183 Genus Fungiastrea Alloiteau, 1952 .................................................................................................... 183 Fungiastrea crespoi (Felix, 1891) ............................................................................................... 183 Fungiastraea flexuosa (Goldfuss, 1826) ..................................................................................... 184 Genus Ellipsocoenia d’Orbigny, 1850................................................................................................. 186 Ellipsocoenia conferta (Umbgrove, 1925) .................................................................................. 186 Genus Baryphyllia Fromentel, 1857 .................................................................................................... 187 Baryphyllia maxima (Umbgrove, 1925) ..................................................................................... 187 Genus Gyrodendron Quenstedt, 1880 ................................................................................................. 188 Gyrodendron boltonae (Wells, 1934)........................................................................................... 188 Genus Loboseris M. Beauvais, 1982 ................................................................................................... 190 Loboseris abbreviata (Reuss, 1854) ........................................................................................... 190 Family Brachiphylliidae Alloiteau, 1952 ................................................................................................... 191 Genus Brachyphyllia Reuss, 1854 ...................................................................................................... 191 Brachyphyllia depressa Reuss, 1854 .......................................................................................... 191 Brachyphyllia felixi Baron-Szabo, 2000 ..................................................................................... 193 Genus Dermosmiliopsis Alloiteau, 1952 ............................................................................................ 194 Dermosmiliopsis orbignyi Alloiteau, 1952 ................................................................................. 194 Dermosmiliopsis tenuicosta (Reuss, 1854) ................................................................................. 195 Suborder Stylinina Alloiteau, 1952 .................................................................................................................. 195 Family Stylinidae d'Orbigny, 1851 ............................................................................................................ 195 Genus Stylina Lamarck, 1816 ............................................................................................................. 196 Stylina reussi Duncan, 1880 ........................................................................................................ 196 Genus Stylosmilia Milne Edwards & Haime, 1848 ............................................................................ 197 Stylosmilia kangpaensis Wu, 1975 .............................................................................................. 197 Genus Tubicora Latham, 1929............................................................................................................. 198 Tubicora aylmeri (Gregory, 1900) .............................................................................................. 198 Family Agatheliidae L. & M. Beauvais, 1975 ........................................................................................... 198 Genus Agathelia Reuss, 1854 ............................................................................................................. 200 Agathelia asperella Reuss, 1854 ................................................................................................. 200 Genus Multicolumnastraea Vaughan, 1899 ........................................................................................ 201 Multicolumnastraea cyathiformis (Duncan, 1865) ..................................................................... 201 Family Cladophylliidae Morycowa & Roniewicz, 1990 ........................................................................... 202 Genus Cladophyllia Milne Edwards & Haime, 1851 ......................................................................... 203 Cladophyllia cf. furcifera Roemer, 1888 .................................................................................... 203 Cladophyllia stewartae Wells, 1944 ........................................................................................... 203 Addendum to Corals of the K/T-Boundary: Scleractinian corals of the suborders Astrocoeniina, ................................ 204 Faviina, Rhipidogyrina, and Amphiastraeina (Baron-Szabo, 2006) ........................................................................ 204 Suborder Faviina Vaughan & Wells, 1943 ....................................................................................................... 204



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Family Faviidae Gregory, 1900 ................................................................................................................. 204 Genus Phacellocoenia Alloiteau & Tissier, 1958 ............................................................................... 204 Phacellocoenia thomkai (Eliášová, 1991) .................................................................................. 205 Family Mussidae Ortmann, 1890................................................................................................................ 206 Genus Circophyllia Milne Edwards & Haime, 1848 .......................................................................... 206 Circophyllia truncata (Goldfuss, 1826) ...................................................................................... 206 Family Meandrinidae Gray, 1847 ............................................................................................................... 207 Subfamily Euphyllinae Alloiteau, 1952 ................................................................................................. 207 Genus Euphyllia Dana, 1846 .............................................................................................................. 207 Euphyllia calyculata (Catullo, 1852) .......................................................................................... 207 Acknowledgments............................................................................................................................................................ 208 References ..................................................................................................................................................................... 209 Tables ....................................................................................................................................................................... 13 Appendix 1. Taxa mentioned from Maastrichtian and/or Paleocene strata without description or assigned doubtfully. 223 Appendix 2. Forms wrongly assigned to the Maastrichtian (newly assigned stratigraphic level indicated) or reported from strata of unknown/doubtful stratigraphic assignment (u) ......................................................................................... 224 Appendix 3. List of synonymies ..................................................................................................................................... 225


BARON-SZABO

Abstract This study is the second of two parts of a taxonomic review of the scleractinian corals of the Maastrichtian and Paleocene periods. The first part (Baron-Szabo, 2006) focused on the scleractinian suborders Astrocoeniina, Faviina, Rhipidogyrina, and Amphiastraeina. The second part deals with the remaining five suborders: Dendrophylliina, Caryophylliina, Fungiina, Microsolenina, and Stylinina. The two parts together represent the first extensive compilation of coral species of the K/T-(Cretaceous/Tertiary) boundary, and deal with more than 2500 records of 550 nominal taxa, of which 259, belonging to 149 genera (including Lazarus taxa=taxa that disappeared before the Maastrichtian and re-occurred after the Paleocene) are considered valid. In the five suborders evaluated in this paper, 136 valid species (including in an addendum 3 taxa belonging to the suborders of the first part) of 81 genera can be reliably documented as occurring in the Maastrichtian and/or the Paleocene. For the following taxa, new combinations are proposed: Palaeopsammia collignoni (Alloiteau, 1958), Palaeopsammia zitteli (Vaughan, 1900) non Wanner, 1902, Bathycyathus corneti (Alloiteau & Tissier, 1958), Bathycyathus lloydi (Vaughan, 1920), Bathycyathus piveteaui (Alloiteau & Tissier, 1958), Trochocyathus speciosus (Gabb & Horn, 1860), Deltocyathus cupuliformis (Alloiteau, 1951), Asterosmilia alloiteaui (Alloiteau & Tissier, 1958), Dasmosmilia kochii (Pratz, 1910), Desmophyllum excavatum (Hagenow, 1839), Smilotrochus cornucopiae (Duncan, 1869), Smilotrochus milneri (Gregory, 1898), Alveolocyathus felixi (Filkorn, 1994), Pleurocora arachnoides (Knorr & Walch, 1777), Meandrophyllia textilis (Goldfuss, 1826), Meandrophyllia velamentosa (Goldfuss, 1826), Cyathoseris catadupensis (Vaughan, 1899), Heterogyra murchisoni (d’Archiac & Haime, 1853), Pattalophyllia grumi (Catullo, 1852), Stephanophyllia cribraria (Stephenson, 1916), Siderofungia morloti (Reuss, 1864), Hindeastraea garloica (Tchéchmédjiéva, 1975), Aspidastraea clathrata (Goldfuss, 1826), Paracycloseris nariensis (Duncan, 1880), Fungiastraea flexuosa (Goldfuss, 1826), Ellipsocoenia conferta (Umbgrove, 1925), Baryphyllia maxima (Umbgrove, 1925), Tubicora aylmeri (Gregory, 1900), Phacellocoenia thomkai (Eliášová, 1991), and Euphyllia calyculata (Catullo, 1852). One species is newly described: Polyphylloseris microkothos n. sp.. In addition to the re-examination and re-evaluation of described forms, this study also includes the first description of the largest Maastrichtian coral assemblage known (consisting of about 4000 specimens from Jamaica), as well as new material from the Campanian-Maastrichtian of Argentina, Lower Maastrichtian of Mexico (Cerralvo), and the Paleocene of Austria (Kambühel-Kalke). Furthermore, lost or “forgotten “coral collections were discovered and illustrated for the first time, including the type and original material of d’Achiardi (1875, Eocene of Italy), Wanner (1902, Maastrichtian-Danian of Egypt), Berryhill, in Berryhill et al. (1960, Danian of Puerto Rico), and Schlotheim (1820, Mesozoic-Recent). A diagnosis is provided for each species, as well as for each higher level taxonomic category, and issues concerning new taxonomic assignments are discussed in detail. The descriptions are accompanied by illustrations of representatives of each species, and in many cases, include illustrations of type or original material. Also included is the first comprehensive overview of the stratigraphical and geographical ranges of each taxon. The largest number of species occurring at the K/T-boundary are in the suborders Faviina (79), Fungiina (51), and Caryophylliina (41). In all of the nine suborders 259 valid species are known from the Maastrichtian and/or Paleocene, of which 204 occurred before the K/T-event and 106 species (52 %) crossed the K/T boundary. In the Paleocene 55 new species appeared. While species of all suborders crossed the K/T-boundary, no new species of the suborders Rhipidogyrina, Amphiastraeina, and Microsolenina appeared in the Paleocene. On the genus level 96 of the 131 genera (=73.3%) that occurred before the K/T-event crossed the K/T-boundary. Thirty-five genera went extinct and 18 genera have their first occurrence in the Paleocene. A generic extinction rate of 26.7% across the K/T-boundary, as estimated here, is considerably less than the rates of around 60% previously stated, but is quite similar to recently reported results for other macroinvertebrate groups after taxonomic revision (e.g., echinoids). Key words: Maastrichtian, Paleocene, taxonomy, survivorship, stratigraphical and geographical distribution



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Introduction The current paper deals with the taxa of scleractinian corals of the suborders Dendrophylliina, Caryophylliina, Fungiina, Microsolenina, and Stylinina that occurred during the Maastrichtian and/or the Paleocene time periods (Tables 1 and 2). It represents the second and final part of a study of corals of the K-T-boundary, the first of which dealt with the suborders Astrocoeniina, Faviina, Rhipidogyrina, and Amphiastraeina (Baron-Szabo 2006). Both parts together form the taxonomical basis of the previously published results concerning the extinctions and recovery patterns of scleractinian corals at the K-T-boundary (Kiessling & Baron-Szabo 2004) (Tables 3 and 4). As pointed out earlier (Baron-Szabo 2006), although Maastrichtian and/or Lower Tertiary corals have been reported from numerous localities worldwide, the vast majority of these papers document only very small faunas of no more than 10 taxa, or only list corals without giving any further taxonomical information, or, in the rare cases of publications that contain taxonomic documentation of large scleractinian coral faunas of the K/T-boundary, they deal with the fauna of a single geological period. The present paper utilizes the data provided by these earlier studies, as well as new data on the Maastrichtian and Paleocene corals from various localities (see chapter Material and Methods). In order to do this, around 2500 records of over 550 nominal coral taxa described for this period had to be re-evaluated. Because of the large amount of information to be presented, the study has been divided into two parts. The first part dealt with 123 valid species belonging to 65 genera. The current work adds 3 more taxa to the suborder Faviina (see Addendum), increasing the number of valid taxa to 126 belonging to 67 genera. The second part deals with 131 valid species belonging to 81 genera. In recent years, a small number of marine macroinvertebrate groups have been the subject of studies evaluating their fate at the K-T-boundary. Besides recently carried out studies of scleractinian corals (Kiessling & Baron-Szabo 2004; Baron-Szabo 2006 and this paper), bivalves (Raup & Jablonski 1993) and echinoids (Smith & Jeffery 2000) were analyzed and these results were compared to data compiled by Sepkoski (2002) who gave a generic extinction rate for macroinvertebrate groups of over 90%. Although the generic extinction for bivalves was shown to be around 63%, the extinction rate in echinoids was not more than 36%, thus very closely corresponding to the results of 26.7% for scleractinian corals (as documented in the current paper). The analysis carried out for bivalves differed in that the extinction patterns for echinoids and scleractinians were analyzed by only one author (e.g., in Kiessling & Baron-Szabo 2004, Baron-Szabo was responsible for the taxonomic evaluation) or one group of authors, respectively, the evaluation of bivalves, however, was carried out in a way that included consultation with an unidentified number of specialists; consequently a number of different taxonomic views may have been included (see Raup & Jablonski 1993, p. 971). If a single taxonomic model had been used, a different result may have been obtained. In the current work, the approach has been to evaluate taxa morphologically, regardless of their geographic and stratigraphic occurrences. The author was not restricted by the assumption that taxa were confined within major geological time periods. Some workers have assumed that it would be unlikely for scleractinian corals to cross these boundaries because of the inplied long time intervals. However, as a matter of fact, this taxonomic revision of scleractinian species occurring worldwide at the K/T-boundary has revealed that the vast majority (238 out 259 species = 91.9 %) span time ranges that are often significantly shorter than those of many species known from Lower to Upper Cretaceous strata (Table 1).

Material and methods The illustrated specimens are housed in the type collections of the following institutions: AUG

University of Auckland, Department of Geology, New Zealand;


BARON-SZABO

BMNH BSP BSPG CAS CO GBA IPB MB MCZ MGSB MNHN NHMW NMNH NMNZ PMB NRM PMU RMNH SAZU SMF SNM UANL

The Natural History Museum, London, UK; Geologisches Institut, Universität Karlsruhe, Germany; Bayerische Staatssammlung Munich, Germany; California Academy of Sciences, San Francisco, USA; New Zealand Geological Survey, Wellington, New Zealand; Geologische Bundesanstalt, Vienna, Austria; Geologisch-Paläontologisches Institut der Rheinischen Friedrich-Wilhelms Universität, Bonn, Germany; Natural Museum of the Humboldt University, Berlin; Museum of Comparative Zoology, Harvard University, Cambridge; Museu Geològico del Seminari de Barcelona; Institut de Paléontologie du Museum d'Histoire Naturelle de Paris, France; MPUR Museo di Paleontologia, Università di Roma, Italy; Naturhistorisches Museum, Vienna, Austria; National Museum of Natural History, Smithsonian Institution, Washington, DC (formerly USNM); National Museum of New Zealand, Wellington, New Zealand; Natural History Museum Beograd, Serbia; Naturhistoriska Riksmuseet, Stockholm, Sweden; Paleontologiska Museet, Uppsala, Sweden; Nationaal Natuurhistorisch Museum, Leiden, Netherlands (formerly Rijksmuseum van Natuurlijke Historie); Slovenska akademija znanosti in umetnosti, Ljubljana, Slovenia; Forschungsinstitut Senckenberg, Senckenberg Museum, Frankfurt, Germany; Slovak National Museum, Bratislava, Slovakia; Universidad Autónoma de Nuevo León; USNM now NMNH.

In addition to the re-examination and re-evaluation of described forms, this study also includes the first assessment of the largest Maastrichtian coral assemblage (consisting of about 4000 specimens from Jamaica, known as the ‘Coates collection’), as well as new material from the Campanian-Maastrichtian of Argentina, Lower Maastrichtian of Mexico (Cerralvo), and the Paleocene of Austria (Kambühel-Kalke). In most cases thin sections (cross and longitudinal cuts) were used to identify the new coral material. To get a comprehensive overview and to examine type material to verify identifications, collections at the following Institutions or Museum were consulted: Paris (NMHN, collections of Alloiteau, Arnaud, Basse & Menard, Besaire, Bührer, Chevalier, Collignon, de Fromentel, Jacob, Michelin, Milne Edwards & Haime, d‘Orbigny, Tissier, Thomas, and Villette); Rome (MPUR, collections of d’Angelis d’Ossat and Michelotti); Pisa (Paleontological Museum, collection of d‘Achiardi); Berlin (MB, collections of Ehrenberg, Felix, Oppenheim, and Schlotheim), Frankfurt (SMF, collection of Esper); London (BMNH, collections of Blagrove, Duncan, Felix, Gregory, Heneken, Kühn, Latham, Oppenheim, and Trechmann); Leiden (RMNH, collection of Umbgrove); Washington, DC (NMNH, collections of Budd, Cairns, Dana, Hoffmeister, Nomland, Sokolov, Squires, Squires & Demetrion, Stephenson, Vaughan, Wade, Weisbord, and Wells); Wellington, NZ (CO, collection of Squires); Ljubljana (SAZU, collection of Turnšek); Vienna (GBA and NHMW, collections of Beauvais, Felix, Kühn, and Reuss); Barcelona (MGSB, collections of Alloiteau, Bataller, and Reig Oriol); Bonn (IPB, collections of Goldfuss and Roemer); Cambridge, Mass. (MCZ, collection of Vaughan); Uppsala (PMU, collections of Cleve, Cleve & Molander, and Goers). Descriptions were supplemented by digital scans of published material. The Maastrichtian material from Jamaica was collected during the years 1966 and 1972 by A. Coates, J. Jackson, and E. Kauffman. The Campanian-Maastrichtian material from Argentina and Lower Maastrichtian



11

corals from Mexico were sampled by W. Stinnesbeck and colleagues; the Paleocene corals of Austria were made available to me by H. Lobitzer (Bad Ischl, Austria), F. Schlagintweit (Munich, Germany), and G. Schubert (GBA, Vienna, Austria). Noteworthy for their relevance to the current work is the re-discovery of four collections of type and/or original material: 1) Schlotheim’s (1820) original collection of scleractinian corals of Paleozoic to Recent age that is housed at the Natural Museum of the Humboldt University Berlin, Germany, was long lost and forgotten. The present paper provides the first photographic illustrations of some specimens of this collection that are relevant to this paper. 2) D’Achiardi’s (1875) type collection of corals of the Eocene of Italy (Friul) was lost for over a century and re-discovered in its original wrappings in the collections of the Paleontological Museum, Pisa, by the author in 2003. The present paper gives the first photographic illustrations of some specimens of this collection that are relevant to this work. 3) Berryhill’s (1960) collection of corals from the Danian of Puerto Rico that is housed at the Smithsonian Institution (all of them are catalogued as “PR-7”) was never described or illustrated but taxa from it were mentioned in the publication by Kaye (1956). The present work provides the first photographic illustrations of some of the corals. 4) Wanner’s (1902) type collection of Maastrichtian-Paleocene scleractinian corals from Egypt was originally housed in the Bayerische Staatssammlung Munich, Germany, and destroyed during WWII. However, in 2004 the author found 64 specimens of the type collection in the inventory of the Rijksmuseum Stockholm, Sweden. It is assumed that the specimens had been given to the Rijksmuseum as part of an exchange programme between museums and institutions early last century. The specimens represent the only known corals of the Wanner collection. Because the labels were signed by the former director of the Bayerische Staatssammlung Munich, Prof. von Zittel, they are undoubtedly extractions from the type collection, and therefore, they all represent original material, most of which are syntypes. The present paper gives the first re-description and photographic documentation of this material. In addition to the examination of the above mentioned material, more than 2500 species records were studied. The data and information retrieved from this procedure resulted in the synonymy lists presented. For further explanation of the ideas on which the synonymies were based see Remarks sections for individual species of e.g., Caryophyllia konincki (Milne Edwards & Haime, 1848) and Parasmilia centralis (Mantell, 1822). In contrast to the extinction/survivorship pattern of species (Tables 1), the survivorship estimation for genera requires a different approach. Based on the currently available data, several genera are represented by only one species at the K/T-boundary, some of which became extinct by the end of the Maastrichtian. However, from the palaeontological record it is known that some of these genera re-occurred in strata younger than the Paleocene and are therefore not taxonomically described in the current paper which focuses solely on scleractinian corals of the Maastrichtian-Paleocene period. Therefore, the genera that re-appeared in postPaleocene strata are not being considered as having gone extinct. In addition to the genera already mentioned in the first part (Baron-Szabo 2006), these genera (“Lazarus taxa”) include: Platygyra calzadai (Reig Oriol, 1989) (holotype: MGSB 47189), from the Campanian of northern Spain); Platygyra wellsi (Durham, 1942) from the Middle Eocene of California; Agathelia Reuss, 1854, from the Upper Maastrichtian (Baron-Szabo 2000 and this paper) and from the Oligocene of Libya (“Agathelia sp.” in Hladil et al. 1992); Pironastrea d’Achiardi, 1875, from the Paleocene and Upper Cretaceous (Campanian and older, represented by taxa formerly ascribed to the genus Koilomorpha Alloiteau, 1952, this paper); Dasmosmilia Pourtalès, 1880, form the Upper Maastrichtian (this paper) and various Tertiary and Recent localities, e.g. D. vaughanianus (Wells 1945) (holotype USNM 68334) from the Eocene of Barbados as well as the type species of this genus Dasmosmilia lymani Pourtalès, 1880, Recent, West Indies;


BARON-SZABO

Cycloseris Milne Edwards & Haime, 1849a, form the Maastrichtian (Baron-Szabo 2002 and this paper) and from various Tertiary and Recent localities (e.g., C. aegyptiaca Pratz, 1883, from the Eocene of Egypt); Cyathoceras Moseley, 1881, from the Maastrichtian (Kuzmicheva 1987; and this paper) and from the Teriary e.g., C. periallus Squires, 1962, from the Oligocene of New Zealand and the type species C. cornu Moseley, 1881, Recent, southern Atlantic; Heterogyra Reuss, 1868, from the Paleocene and from various Cretaceous localities, e.g., H. zitteli (Felix, 1903a), Upper Santonian of Austria (Baron-Szabo 2002); in addition to its Cretaceous occurrences (e.g., Maastrichtian, this paper);



13

Ellipsocoenia d’Orbigny, 1850, from e.g., the Eocene of Spain (E. bauzai Mallada, 1885); and from Mesozoic occurrences (e.g., Maastrichtian, this paper); Baryphyllia Fromentel, 1857, from European Cretaceous strata and Asian Teriary sediments (B. gemmans [Fritsch, 1878], Eocene of Borneo, this paper) and European Tertiary localities (B. variabilis (Reuss, 1872), Oligocene of Italy; Stylina Lamarck, 1816, from Jurassic and Cretaceous localities, including the type species Stylina echinulata Lamarck, 1816, from the Jurassic of France.


BARON-SZABO

Results A total of 259 species belonging to 149 genera of nine scleractinian suborders (Astrocoeniina, Faviina, Rhipidogyrina, Amphiastraeina, Dendrophylliina, Fungiina, Microsolenina, Caryophylliina, and Stylinina) occurred at the K/T-boundary (Tables 3 and 4). The majority of the taxa (including Lazarus taxa) are colonial: 181 species (=69.9%) belonging to 103 genera (=69.1 %); 78 species (30.1 %) belonging to 46 genera (30.9 %) are solitary forms.



15

Twenty-one (8.1 %) of the species that occurred at the K/T-boundary were also reported from the Lower Cretaceous, nine (3.5 %) of which also crossed into the Tertiary, including one species of Faviina (solitary), four species of the Caryophylliina (solitary), two species of the Fungiina (both colonial), and two species of the Microsolenina (both solitary). Nine species (3.5 %) survived into the Neogene, seven (2.7 %) of which had also crossed the K/T-boundary: four species of the suborder Faviina (all colonial), two species of the Fungiina (both colonial), and one species of the suborder Microsolenina (solitary). Not one of the taxa that had its first occurrence in the Lower Cretaceous passed into the Neogene. Taxa which disappeared after the Maastrichtian (98 species) show only close affinities to Upper Cretaceous coral assemblages of adjacent regions, e.g. in the European regions to the Western Atlantic area, or


BARON-SZABO

Eastern Africa to Central-South Asia/Pacific area but hardly correspond to Lower Cretaceous coral associations (Baron-Szabo 2006, Tables 1 and 2, and Tables 1 and 2 this paper). Forms which first appeared in the Paleocene (55 species) show greatest affinities to European and Asian assemblages of the Lower Tertiary but, with the exception of two taxa, have not been reported from younger strata (Baron-Szabo 2006: Tables 1 and 2, and Tables 1 and 2 this paper). The only species that had their first occurrence in the Paleocene and passed into the Neogene are the meandroid form Heterogyra murchisoni (d’Archiac & Haime, 1853) and the cerioid species Siderastrea conferta (Duncan, 1863). Both of these taxa passed into the Neogene in regions from which they had been reported for the Lower Tertiary.



17

Suborders: Species of all nine scleractinian suborders, Astrocoeniina, Faviina, Rhipidogyrina, Amphiastraeina, Dendrophylliina, Caryophylliina, Fungiina, Microsolenina, and Stylinina, survived into the Tertiary (Tables 1–4). Five suborders show species survival rates below the overall average number of 52%: Astrocoeniina (44.4%), Rhipidogyrina (12.5 %), Amphiastraeina (40 %), Microsolenina (33.3%), and Stylinina (20%). As previously pointed out by Kiessling & Baron-Szabo (2004), the K/T-extinction involved predominantly colonial taxa of thamnasterioid and meandroid growth forms and/or corals of restricted geographic distribution, mainly or exclusively occurring in the reefal environments of the tropics. Genera and species: A total of 259 species belonging to 149 genera were analyzed (Baron-Szabo 2006: Tables 1 and 2, and Tables 3 and 4 this paper). On the generic level, 96 out of 131 genera, corresponding to 73.3 % of taxa, crossed the K/ T-boundary (Table 4). On the species level, 106 species out of 204 (52 %) crossed the K/T-boundary. 55 species in 19 genera had their first occurrence in the Paleocene. While taxa of all nine suborders crossed the K/Tboundary, the suborders Rhipidogyrina, Amphiastraeina, and Microsolenina have no new genera or species in the Paleocene. Although two new species of the suborder Dendrophylliina occurred in the Paleocene, no new genus was recorded. A detailed discussion on the extinctions and recovery patterns of scleractinian corals at the K/T-boundary was recently provided (Kiessling & Baron-Szabo 2004). It should be noted however, that the survival/extinction rates slightly differ in the former work due to a slightly different approach used e.g., rarefaction statistics


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19


BARON-SZABO



21


BARON-SZABO



23


BARON-SZABO



25


BARON-SZABO

were applied. In addition, some species could not be studied at that time, and/or have only recently been rediscovered (see chapter Material and Methods). That led to a slight decrease in the extinction rates in the present work, especially affecting the genus level. Because the results presented in Kiessling & Baron-Szabo (2004) were directly derived from the taxonomic work presented in the two “K/T parts” (Baron-Szabo 2006, and this paper), they are briefly summarized here. The following four key points were evaluated: 1 the intensity of the end-Cretaceous coral extinction and pace of recovery; 2 the ecological selectivity of coral extinction; 3 the impact of coral extinctions on the global reef ecosystem; 4 the geographic patterns of coral extinction and recovery. TABLE 3. Number of suborders and species that occurred at the K/T-boundary. Suborders

no. of spe- K/T-occurrences of species: number of species having their surviving cies at K/T occurrence in Maas- first occurrence in occurrence in Maastrich- species (%) Paleocene trichtian tian and Paleocene (=survivors)

Astrocoeniina

34

18

16

8

44.4

Faviina

79

62

17

35

56.5

Rhipidogyrina

8

8

-

1

12.5

Amphiastraeina

5

5

-

2

40

Dendrophylliina

9

7

2

4

57.1

Caryophylliina

41

30

11

17

56.7

Fungiina

51

44

7

29

65.9

Microsolenina

25

27

-

9

33.3

Stylinina

7

5

2

1

20

Total

259

204

55

106

52

TABLE 4. Number of suborders and genera that occurred at the K-T-boundary (including Lazarus taxa). Suborders

no. of genera at K/T-occurrences of genera: number of genera having their K/T

surviving genera

(including Lazarus taxa)

occurrence Maastrichtian

Astrocoeniina

11

6

5

5

83.3

Faviina

52

47

5

37

78.7

Rhipidogyrina

4

4

-

2

50

Amphiastraeina

1

1

-

1

100

Dendrophylliina

5

5

-

3

60

Caryophylliina

26

21

5

16

76.2

Fungiina

31

29

2

21

72.4

Microsolenina

13

13

-

8

61.5

Stylinina

6

5

1

3

60

Total

149

131

18

96

73.3


in first occurrence in occurrence in Maastrich- (%) tian and Paleocene Paleocene (=survivors)


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TABLE 5. Skeletaldimensions of species of Cunnolites occurring at the K/T-boundary. Species of Cunnolites

Density of septa (s/mm) in

Ratio of min/max diameter

peripheral areas

around calicinal pit

throughout ontogeny

17–25/5

22–25/5

0.9–1

Cunnolites tenuiradiata (de Fromentel, 15–20 (22)/5 1863)

12–14/5

0.67–1 (0.7–0.85 most common)

Cunnolites scutellum (Reuss, 1854)

14–20/5

15–22/5

0.8–1

Cunnolites giganteus (d’Orbigny, 1850)

8–15/5

8–15/5

0.73–0.97

Cunnolites polymorphus (Goldfuss, 1826)

17–20 (25)/5

17–20/5

0.7–0.98

Cunnolites angiostoma (Kühn, 1933)

20–25/5

25/5

0.85–0.89

Cunnolites cancellata (Goldfuss, 1826)

1. The intensity of the end-Cretaceous coral extinction and pace of recovery 1.1 Extinction intensities Genus-level: 30 (±8) % (previously reported to be around 60 %) (26.1 % in present work). Species-level: 45 (±5) % (previously reported to be around 95 %) (48 % in present work). 1.2 Survivorship and abundance (including Lazarus taxa) — Abundance was not insurance against extinction — No significant differences in extinction risk between species-poor and species-rich genera

1.3 Origination rates — Evolution of new genera gradual but concentrated in the Danian 2. The ecological selectivity of coral extinction — Z-like corals (most likely to have been zooxanthellate, term sensu Rosen & Turnšek 1989) were more affected than az-like corals (most likely to have been azooxanthellate, term sensu Rosen & Turnšek 1989) (extinction of z-like ca. 40 %, az-like ca. 27 %), that means end-Cretaceous extinction was selective against photosymbiosis — Extinction risk was higher for corals with high corallite integration (meandroid, thamnasterioid) — “Larval strategy shows to be no selective criterion” (nearly equal number of brooders and broadcasters survived) — mean skeleton size of species within genera had no significant influence on survivorship, mean size species of Maastrichtian and Paleocene genera was nearly identical. Therefore, extinction did not act against species large size genera — Extinction rate did not vary significantly between habitats:


BARON-SZABO

Reefs=32 (±11) % Coastal environments=24 (±11) % Carbonate platforms=30 (±9) % 3. The impact of coral extinctions on the global reef ecosystem — Low number of coral reefs recorded for latest Cretaceous and Paleocene (± 50 for each interval) but their globally preserved volume peaked in the Paleocene (in previous works “Paleocene was time of depressed reef building”) — K/T-boundary represents the only mass extinction event without significant drop in reefal carbonate production at stage level: Reef corals either little affected by extinctions or recovered more rapidly than at other extinction intervals Coral occurrences from reefs: — Latest Cretaceous: 16 (±3) % genera — Paleocene: 36 (±5) % genera These data contrast with the partitioning of az-like and z-like corals in both intervals because the occurrence of z-like corals corresponded to: — Late Cretaceous assemblages: 59 (±4) % genera — Paleocene assemblages: 37 (±5) % genera Z-like corals vs. reef building — Several az-like corals are involved in Paleocene reef building but played no role in the latest Cretaceous — Paleocene z-like more common in reefal associations than the late Cretaceous ones (rather due to evolutionary radiation of encrusting algae than due to reef building potential of corals) 4. The geographic patterns of coral extinction and recovery — Highest extinction rates in low paleolatitudes, drop off toward higher paleolatitudes — Extinction patterns are strongly controlled by occurrence of the extinction resistant az-like corals: latitude had no direct influence on extinction rates No extinction hot spots Extinction in: ? North America: 33 (±11) % ? Europe: 35 (±10) % ? Africa and India: 30 (±11) % ? Same applies for endemics to regions (40-50 %) — Wide geographic distributions of clades in the Maastrichtian was insurance against extinction — New z-like genera in the Paleocene were concentrated in low paleolatitudes: tropics represented a source SCLERACTINIAN CORALS OF THE K/T-BOUNDARY


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of evolutionary novelty.

Systematic Palaeontology The classification system used in this work for assigning and arranging genera is the one created by BaronSzabo (2002), which was based on a combination of several different proposed classifications, including those of Vaughan & Wells (1943), Alloiteau (1952a, 1957), and Wells (1956). In addition, microstructural features as proposed by these authors are given for each family. When other concepts are adopted, supplementary remarks are included. Further information on microstructure is incorporated in each generic diagnosis if it differs from the family character. For different and additional microstructural concepts see e.g. Beauvais (1981, modified classification according to microstructure), Gill (1981, synapticulae and related microstructural features), Roniewicz (1982, synapticulae and related microstructural features; 1996, septal and thecal microstructure), Morycowa & Roniewicz (1995, modified classification according to microstructure), and Stolarski & Roniewicz (2001, discussion of microstructural and microarchitectural characters). Abbreviations for the dimensions in the text are as follows: d =corallite diameter; dl =diameter of lumen; d (max) =maximum diameter of corallite; d (min) =minimum diameter of corallite; c-c =distance between centres of corallites; c-c =distance between corallite centres; c-c (series) =distance between corallite centres of same series or adjacent series (as further specified in text);c-c (wall) =width of calicinal series; s =number of septa; s/mm =density of septa; diss/mm =density of dissepiments; h =height of corallum; S1, S2, etc. =septa of 1st cycles, septa of 2nd cycles, etc.; C1, C2, etc. =costae corresponding to 1st cycle septa, costae corresponding to 2nd cycle septa, etc. Taxonomic terms and abbreviations: v =material studied by the author; (v) =material documented by very good and detailed photographs; publication date in italics =taxon only mentioned without any description or illustration; * =first description of taxon to which the assignment of specimen refers.

Order Scleractinia Bourne, 1900 Diagnosis. Solitary or colonial Zoantharia with calcareous external skeleton secreted by the ectodermal body layer, consisting essentially of radial partitions or septa, which are intermesenterial in position and formed within upward infoldings of the basal part of the polyp column wall, and attendant supporting structures: basal plate, epitheca, dissepiments, synapticulae, and mural structures; septa developed in ontogeny following pattern of mesenteries, additional septa after first 6 being inserted in all 6 primary mesenterial exocoeles in successive cycles of six, 12, 24, 48, and so on, in dorsoventral order.

Suborder Dendrophylliina Vaughan & Wells, 1943 Diagnosis. Solitary and colonial. Wall synapticulothecal, irregularly porous. Septa structurally similar to the caryophylliids but usually secondarily thickened, more or less porous, with margins wholly smooth or partly dentate. Septa usually following the Pourtalès plan. Remarks. This suborder was proposed for the reception of a single large family, the Dendrophylliidae. The microscopic structure of the septa, the morphology of the polyps, and the corallum habit are all character-


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istic of the Caryophylliina, but the porosity and thickening of the septa, due to irregular trabecular growth, and the perforate synapticulothecal wall are conditions of specialization unknown in that group; the latter features are more characteristic of the fungiid group. It is suggested that the group originated from a caryophylliid-like ancestor in which very irregular trabecular growth leads to septal thickening and perforation and replacement of septotheca by synapticulotheca, and these in turn broke up extratentacular mesenterial extensions into canaliculate prolongations of the coelenteron through and over the wall. Septa usually following the Pourtalès plan.

Family Dendrophylliidae Gray, 1847 Diagnosis. Solitary and colonial. Colony formation by intra- and extratentacular budding. Wall formed by trabecular outer ends of septa and simple but very irregular synapticulae, irregularly porous, usually thick, irregularly costate or covered by interrupted costae or irregularly scattered granulations, occasionally epithecate. Porous, layered coenenchyme developed in some colonial genera. Septa composed of one fan system of simple trabeculae, but the trabeculae tend to be very irregular in development, often not closely united in the plane of the septum and vertically discontinuous because of sclerodermites bending outwards from the plane of the septum in unpaired fashion, especially peripherally in the mural region and near the columella, leaving pores. Laterally the septa are strongly granulated, marginally they are usually smooth except proximally and distally where there are irregular dentations or granulations, especially costally; and in some species the septa are more or less wholly dentate. Insertion of the septa usually following the Pourtalès plan. Columella, when present, spongy. Endotheca thin, poorly developed. Remarks. Cairns (2001) gave detailed discussions of the taxonomic history of the Dendrophylliidae and provided a generic revision and phylogenetic analysis of the genera of this family.

Genus Dendrophyllia Blainville, 1830 Type species. Madrepora ramea Linnaeus, 1758, Recent, Mediterranean Sea.

Diagnosis. Colonial, dendroid, but polyps remain organically connected. Extensive development of coenosteum may embed branches, resulting in plocoid to fasciculate appearances. Gemmation extracalicinal. Costosepta arranged according to the Pourtalès plan, subcompact with irregularly occurring pores, granular laterally. Columella irregular, spongy. Endothecal dissepiments thin, subtabulate. Wall thick, septothecal-synapticulothecal, irregularly porous.

Dendrophyllia candelabrum Hennig, 1899 Pl. 1, Figs 1a, b v1828

Caryophyllia fasciculata Lamarck: Morren, p. 48.

v*1899

Dendrophyllia candelabrum, n.sp.: Hennig, p. 8, pl. 1, figs 6–12 (topotypes studied).

v1925

Galaxea fasciculata (Morren spec. gen. nov. nom.): Umbgrove, p. 106, pl. 10, fig. 15.

v1972

Dendrophyllia candelabrum Hennig, 1899: Floris, p. 92, pl. 6, figs 35–37, pl. 7, figs 1–6, pl. 8, figs 1A–B



31

(older synonyms cited therein)(topotypes studied). 1987

Dendrophyllia candelabrum Hennig, 1899: Kuzmicheva, p. 149, pl. 26, figs 2a, b.

v1988

Dendrophyllia candelabrum Hennig: Drobne et al., p. 185, pl. 29, figs 4–6.

1990

Dendrophyllia candelabrum Hennig, 1899: Bernecker & Weidlich, p. 113, pl. 26, figs 1, 2, pl. 29, figs 1–4, pl. 30, figs 1, 2.

?1993

Dendrophyllia candelabrum Hennig, 1899: Carbone et al., p. 227.

1997

Denddrophyllia candelabrum Hennig, 1902: Vecsei & Moussavian, p. 128, pl. 35, fig. 3.

v1998

Dendrophyllia candelabrum Hennig, 1899: Turnšek, in Turnšek & Drobne, p. 135, pl. 4, figs 1, 2.

parsv2004 Placocoenia macrophthalma (Goldfuss, 1826): Leloux, p. 319, pls 1–3, non pl. 4. v2005

Dendrophyllia candelabrum Hennig, 1899: Bernecker & Weidlich, p. 11ff., figs 5C, D, 6 (1), A (9) (topotypes studied).

Dimensions. d=3.5–5 mm, juvenile around 2 mm, up to 12 mm in latest adult stages; s=24–60, latest adult stages up to 96. Description. Colonial. dendroid; corallites generally 3.5–5 mm in diameter, bud off in any direction at angles around 90º; costosepta, up to 12 of which reach the center of the calice, arranged clearly or indistinctly according to the Pourtalès plan; columella weakly developed; endothecal dissepiments sparse, subtabulate; wall septothecal-synapticulothecal, often thick with irregularly occurring pores. Remarks. The taxonomic position of Caryophyllia (Galaxea) fasciculata sensu Morren, 1828, from the Maastrichtian of The Netherlands has long been discussed. Leloux (2004) assigned this specimen to Placocoenia macrophthalma (Goldfuss, 1826). However, in a recent evaluation Baron-Szabo (2006, p. 46) pointed out that Morren’s form showed the presence of porous septa and synapticulae (in ‘steinkern’ preservation), and therefore differed from the concept of the placocoeniid group. In fact, Morren’s specimen very closely corresponds to the parts of steinkern preservation that are present in Dendrophyllia as shown on Pl. 1, fig. 2 of the current work. Therefore, the taxon Caryophyllia (Galaxea) fasciculata sensu Morren, 1828, is grouped with the genus Dendrophyllia. In the dimensions of its skeletal elements it corresponds well with the ones observed in D. candelabrum. Type locality of species. Danian of Denmark (Fakse limestone). Distribution. Danian of Denmark, Slovenia, and Sweden (Limhamn), Paleocene of Italy and Russia, Upper Paleocene-Lower Eocene of ?Somalia. PLATE 1: Fig. 1 Dendrophyllia candelabrum Hennig, 1899, topotype, Bayerische Staatssammlung Munich, BSP 1981-I-11, Danian of Denmark (Fakse limestone). 1a, upper surface of colony, scale bar: 6 mm; 1b, basal view of colony, scale bar: 6 mm. Fig. 2 Dendrophyllia dendrophylloides Milne Edwards & Haime, 1848d, topotype, upper surface of colony, MNHN, M-01166, Eocene of England, scale bar: 3 mm. Fig. 3 Dendrophyllia dendrophylloides Milne Edwards & Haime, 1848d, cross thin section, NMNH, Coates coll., no. 397, Middle-Upper Maastrichtian of Jamaica, scale bar: 2.3 mm. Fig. 4 Palaeopsammia zitteli (Vaughan, 1900) (holotype of Kumbiopsammia besairiei Alloiteau, 1958), MNHN, Mo-5080, Maastrichtian of Madagascar, scale bar: 3 mm. 4a, cross view; 4b, longitudinal view. Fig. 5 Areopsammia alacca (Morren, 1828), topotype (type material of Areopsammia maestrichtensis Dietrich, 1917, and Porosmilia alacca [Morren, 1828] of Umbgrove, 1925), RGM 76599, Umbgrove coll., upper surface of specimen, steinkern preservation, Maastrichtian of The Netherlands, scale bar: 4.5 mm.


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33

Dendrophyllia dendrophylloides Milne Edwards & Haime, 1848 Pl. 1, Figs 2, 3 v*1848d

Dendrophyllia dendrophylloides: Milne Edwards & Haime, vol. III, p. 102 (topotypes studied).

v1850a

Dendrophyllia dendrophylloides: Milne Edwards & Haime, p. 36, pl. 6, figs 2–2c (topotypes studied).

1925

Dendrophyllia dendrophylloides: M. Edw. et J. Haime 1848: Felix, pars 28, p. 162 (older synonyms cited therein).

(v)1975

Dendrophyllia dendrophylloides Edwards et Haime, 1850: Kuzmicheva, p. 28, pl. 4, figs 2, 3.

1987

Dendrophyllia weselovi Kuzmicheva, sp. nov.: Kuzmicheva, p. 149, pl. 26, figs 3a, b.

v1988

Dendrophyllia dendrophylloides Milne Edwards & Haime 1850: Drobne et al., p. 185, pl. 30, figs 1, 2.

1997

Dendrophyllia new sp.: Stemann, in Bryan et al., p. 36, text-fig. 2B, Table 1.

v1998

Dendrophyllia dendrophylloides Milne Edwards & Haime, 1850: Turnšek, in Turnšek & Drobne, p. 135, pl. 4, fig. 5.

v2000

Dendrophyllia nodosa Reuss, 1868: Baron-Szabo, p. 112, pl. 7, figs 2–4.

v2002

Dendrophyllia nodosa Reuss, 1868: Baron-Szabo, p. 77, pl. 55, figs 1–3.

Dimensions. d (lumen, max.)=3–5.5 mm, juvenile around 2 mm, late adult stages 8.5 mm; d (lumen, min.): 2.5–5 mm; s=40–48+s5, in latest adult stages up to around 60, in juveniles up to 24. Description. Colonial, dendroid to subfasciculate-subplocoid; polyps connected by a vesicular coenosteum; costosepta arranged according to the Pourtalès plan; endothecal dissepiments thin, subtabulate; wall septothecal-synapticulothecal, often thickened. Remarks. The type material of D. dendrophylloides Milne Edwards & Haime (see Pl. 1, Fig. 2, upper surface view of colony) from the Eocene of England and the material from the Middle-Upper Maastrichtian of Jamaica (see Pl. 1, Fig. 3, cross thin section) show corallites that are connected by well-developed coenosteum and have diameters ranging from 2 to around 8 mm (juvenile and adult corallites, including wall). In addition, in both specimens the number of septa is the same in corallites of the same diameter, e.g., ranging between 50 and 60 in corallites of lastest adult stages of around 8 mm in diameter (numbers include septa that are short and extend beyond the corallite wall as short spines), in juvenile corallites of around 2.5 mm in diameter around 24 septa are present. Therefore, the specimens from the Eocene of England and the MiddleUpper Maastrichtian of Jamaica are considered the same taxon. Type locality of species. Eocene of England (Bracklesham Bay). Distribution. Middle-Upper Maastrichtian of Jamaica (this paper) and the UAE/Oman border region, Paleocene of Russia, Slovenia, Ukraine, and the USA (Salt Mountain Limestone, Alabama), Eocene of England and Ukraine. New Material. Middle-Upper Maastrichtian of Jamaica, NMNH, Coates coll., sample no.: 397 (=Jerusalem Mountain Inlier).

Genus Areopsammia Dietrich, 1917 (=Porosmilia Umbgrove, 1925; Type species. Cyclolites alacca Morren, 1828, Maastrichtian of the Netherlands [Maastricht]) Type species. Cyclolites alacca Morren, 1828 (=Areopsammia maestrichtensis Dietrich, 1917), Maastrichtian of the Netherlands (Maastricht).


BARON-SZABO

Diagnosis. Solitary, turbinate or bowl-shaped. Costosepta irregularly porous, strongly granulated laterally, basally reduced to granulations. Septa arranged in normal insertion (not following the Pourtalès plan). Columella feebly developed, papillose. No epitheca. Endothecal dissepiments thin and poorly developed. Wall synapticulothecal, thick and porous.

Areopsammia alacca (Morren, 1828) Pl. 1, Fig. 5 1799 v*1828 1914 v1917 v1925 1943 1952a 1956 1957 1999 2000 2000 v2001 v2002

Fungite: Faujas-Saint-Fond, p. 199, pl. 38, fig. 6. Cyclolites alacca: Morren, p. 49 (topotypes studied). Cyclolites alacca Morren 1828: Felix, pars 7, p. 185. Areopsammia maestrichtensis n. sp.: Dietrich, p. 304ff., text-fig. on p. 305. Porosmilia alacca, gen. nov. Morren spec.: Umbgrove, p. 113, pl. 11, fig. 17. Areopsammia maestrichtensis Dietrich, 1917: Vaughan & Wells, p. 235. Areopsammia maestrichtensis Dietrich, 1917: Alloiteau, p. 680. Areopsammia maestrichtensis Dietrich, 1917: Wells, p. F433. Areopsammia maestrichtensis Dietrich, 1917: Alloiteau, in table on p. 352. Areopsammia alacca: Leloux, p. 193, fig. 2. Areopsammia alacca (Morren, 1828): Löser, p. 9. Areopsammia maestrichtensis, Dietrich 1917: Löser, p. 9. Areopsammia alacca (Morren, 1828): Cairns, p. 11, pl. 1, figs b–d. Areopsammia alacca (Morren, 1828)(=Areopsammia maestrichtensis Dietrich, 1917): Baron-Szabo, p. 77, pl. 56, figs 1–2.

Dimensions. d=18 x 22 mm; s=96. Description. Solitary, bowl-shaped; septa crowded, arranged in 5 complete cycles in 6 systems; fossa deep. Remarks. Umbgrove (1925) based his new genus Porosmilia on the same specimen that Dietrich (1917) used for his genus Areopsammia. Because Umbgrove's taxon is a junior synonym, Dietrich's Areopsammia has priority. Moreover, the generic name Porosmilia was previously used by de Fromentel (1860). Therefore, Umbgrove's taxon is a junior homonym. Type locality of species. Maastrichtian of The Netherlands (St. Pietersberg). Distribution. Maastrichtian of The Netherlands.

Genus Palaeopsammia Wanner, 1902 (=Kumbiopsammia Alloiteau, 1958; Type species. Kumbiopsammia besairiei Alloiteau, 1958, Maastrichtian of Madagascar); (=Diegosmilia Alloiteau, 1958; Type species. Microseris complanata Collignon, 1931, Cenomanian of Madagascar); (=Sakalavicyathus collignoni Alloiteau, 1958, Type species. Sakalavicyathus collignoni Alloiteau, 1958, Maastrichtian of Madagascar). Type species. Palaeopsammia multiformis Wanner, 1902, Danian of Egypt (subsequent designation by Wells 1936).

Diagnosis. Solitary, conical to patellate. Costosepta not arranged in Pourtalès plan but higher cycle septa (S4) frequently anastomose with those of lower cycle (S3). Pali present, variable in number and shape. Costae



35

poorly developed, but represented by linear arrangement of coarse granules. Columella generally well-developed, parietal-spongy-papillose. Endothecal dissepiments vesicular. Wall (parathecal-) synapticulothecal, (?pseudo-) septothecal when secondaryly thickened. Epithecal wall present or absent.

Palaeopsammia collignoni (Alloiteau, 1958) Pl. 2, figs 4a, b v1958 2000 v2002

Sakalavicyathus Collignoni nov. sp.: Alloiteau, p. 195, pl. 27, figs 10–17, 19. Sakalavicyathus collignoni, Alloiteau 1958: Löser, p. 72. Sakalavicyathus collignoni Alloiteau, 1958: Baron-Szabo, p. 172, text-fig. 70.

Dimensions. d=2.5–4 mm; s=32–42; h=up to 10 mm. Description. Subceratoid-subcylindrical, circular in outline; costosepta arranged in 6 irregular systems; columella spongy-trabecular. Remarks. Alloiteau (1958, p. 195–196) described the solitary genus Sakalavicyathus and stated that it was characterized by a subceratoid corallum, compact and radially arranged costosepta, with granulated lateral sides, the presence of pali, that are variable in number and shape, the presence of a columella formed by a small number of trabeculae, thin endothecal dissepiments, and a parathecal-synapticulothecal wall with intercostal pores. However, re-examination of the holotype revealed that higher cycle septa frequently merged with lower cycle ones, that the trabecular columella was well-developed, and that costae were rather represented by a linear arrangement of coarse granules. Therefore, together with the above mentioned charatceristics, Sakalavicyathus Alloiteau is considered a junior synonymy of the genus Palaeopsammia. Type locality of species. Upper Maastrichtian of Madagascar. Distribution. Upper Maastrichtian of Madagascar.

PLATE 2: Fig. 1 Palaeopsammia zitteli (Vaughan, 1900), NMNH, Mo158169 (1st hypotype), Danian of California, cross view, polished surface, scale bar: 3 mm. Fig. 2 Palaeopsammia zitteli (Vaughan, 1900), NMNH, Mo158169, Danian of California, longitudinal view, scale bar: 2.5 mm. Fig. 3 Palaeopsammia zitteli (Vaughan, 1900), NMNH, Mo158169 (2nd hypotype), Danian of California, cross view, polished surface, scale bar: 3 mm. Fig. 4 Palaeopsammia collignoni (Alloiteau, 1958), holotype (originally assigned by Alloiteau to Sakalavicyathus collignoni), MNHN, Alloiteau coll., Mo5166, Upper Maastrichtian of Madagascar, scale bar: 1 mm. 4a, cross view; 4b, longitudinal view. Fig. 5 Palaeopsammia zitteli (Vaughan, 1900), (syntype of Palaeopsammia multiformis Wanner, 1902), NRM-PZ, Wanner coll., Cn 47549, Danian of Egypt, scale bar: 3 mm. 5a, cross view, upper surface; 5b, longitudinal view. Fig. 6 Palaeopsammia zitteli (Vaughan, 1900), (syntype of Palaeopsammia multiformis Wanner, 1902), NRM-PZ, Wanner coll., Cn 47548, Danian of Egypt, cross view, upper surface, scale bar: 3 mm. Fig. 7 Palaeopsammia zitteli (Vaughan, 1900), Vaughan coll., NMNH, I32379 (figured as holotype of Steriphonotrochus leithensis Vaughan, 1920, in Vaughan 1920), Paleocene of California, scale bars: 1 mm; 7a, longitudinal view; 7b, cross view. Fig. 8 Palaeopsammia zitteli (Vaughan, 1900), (syntype of Palaeopsammia multiformis Wanner, 1902), NRM-PZ, Wanner coll., Cn 47519, Danian of Egypt, scale bar: 3 mm. 8a, cross view, upper surface; 8b, longitudinal view.


BARON-SZABO



37

Palaeopsammia zitteli (Vaughan, 1900) non Wanner, 1902 Pl. 1, figs 4a, b, Pl. 2, figs 1–3b, 5a–8b, Pl. 3, figs 2a, b v1897 v*1900 v1902 v1902 1902 v1902 1902 1903 v1920 1925 v1931 v1933 ?1955 v1958 v1958 v1958 1958 1958 v1958 1970 1970 1971 1981 1987 (v)1995 1995 non1996 (v)1996 (v)1996 (v)1996 1996 v1997 2000 2000 v2000 2000 2000 2000 2000 2001 v2001 v2002 2002 2002 v2002

Placosmilia n. sp.: Merriam, p. 770. Trochocyathus zitteli Merriam: Vaughan, p. 102, pl. 8, figs 4–6a, fig. 7. Thecocyathus aegyptiacus n. sp.: Wanner, p. 99, pl. 14, figs 1–1a. Palaeopsammia zitteli n. sp.: Wanner, p. 104, pl. 15, figs 1–4. Caryophyllia Jasmundi n. sp.: Wanner, p. 100, pl. 14, figs 2–4. Palaeopsammia multiformis n. sp.: Wanner, p. 105, pl. 15, figs 5–9. Palaeopsammia zitteli Wanner: Quaas, p. 161. Palaeopsammia zitteli Wanner: Oppenheim, p. 448, pl. 7, figs 17–18a. Steriphonotrochus leithensis Vaughan, n. sp.: Vaughan, p. 64, pl. 10, figs 6–6b. Trochocyathus zitteli Vaughan 1900: Felix, pars 28, p. 204. Microseris complanata: Collignon, pl. V (I), figs 4–5b. Palaeopsammia fastigiata nov. sp.: Kühn, p. 183, pl. 17, fig. 9. Trochocyathus cf. epicharis Wann.: Rossi Ronchetti, p. 108. Microseris complanata Collignon: Alloiteau, p. 153. Diegosmilia complanata Collignon sp.: Alloiteau, p. 153, pl. 34, figs 5, 6, text-fig. 25. Kumbiopsammia Besairiei nov. sp.: Alloiteau, p. 201, pl. 34, figs 1–4. Palaeopsammia multiformis Wanner: Alloiteau, p. 203. Palaeopsammia zitteli Wanner: Alloiteau, p. 203. Dasmosmilia ? spissa n. sp.: Squires, p. 41, pl. 6, figs 15–16. Palaeopsammia multiformis Wanner: Hassan & Salama, p. 92, pl. 3, figs 1a–j. Kumbiopsammia cf. besairiei Alloiteau: Hassan & Salama, p. 94, pl. 3, figs 2a–i. Palaeopsammia multiformis Wanner: Salama, p. 86, pl. 10, figs 1–5. Palaeopsammia multiformis Wanner: Barthel & Herrmann-Degen, p. 162. Palaeopsammia multiformis Wanner, 1902: Kuzmicheva, p. 146, pl. 15, figs 7–9. Palaeopsammia multiformis Wanner 1902: Abdel-Gawad & Gameil, p. 27, pl. 20,figs 5–6. Bathypsammia cleopatrae Youssef and Salama, 1969: Abdel-Gawad & Gameil, p. 28, pl. 20,figs 9–10. Caryophyllia jasmundi Wanner, 1902: Schuster, p. 80. Notocyathus (Notocyathus) tineidensis n. sp.: Schuster, p. 81, pl. 19, figs 3a, b, text-fig. 43. Palaeopsammia multiformis Wanner, 1902: Schuster, p. 83, text-fig. 46. Kumbiopsammia salamae n. sp.: Schuster, p. 84, text-fig. 48. Kumbiopsammia cf. besairiei Alloiteau, 1958: Schuster, listed in appendix. Palaeopsammia multiformis Wanner, 1902: Cairns, p. 11 (older synonyms cited therein). Dasmosmilia spissa Squires: Löser, p. 27. Diegosmilia complanata (Collignon 1931): Löser, p. 28. Diegosmilia complanata (Collignon, 1931): Baron-Szabo, p. 126, pl. 12, fig. 1. Caryophyllia jasmundi, Wanner 1902: Löser, p. 17. Kumbiopsammia besairiei, Alloiteau 1958: Löser, p. 44. Palaeopsammia fastigiata, Kühn 1933: Löser, p. 58. Palaeopsammia zitteli, Wanner 1902: Löser, p. 58. Palaeopsammia aegyptiaca (Wanner, 1902): Cairns, p. 13. Kumbiopsammia besairiei Alloiteau, 1958: Cairns, p. 40, pl. 14, figs g–i. Diegosmilia complanata (Collignon, 1931): Baron-Szabo, p. 16, pl. 119, figs 2–3, 6. Palaeopsammia fastigiata Kühn, 1933: Baron-Szabo, p. 78. Palaeopsammia zitteli Wanner, 1902: Baron-Szabo, p. 78. Kumbiopsammia besairiei Alloiteau, 1958: Baron-Szabo, p. 79, pl. 57, fig. 2

Dimensions. d (min)=5–23 mm; d (max)=6–30 mm; d (min)/d (max)=0.68–1; s=48–96; h=up to 58 mm (bro-


BARON-SZABO

ken piece). Description. Solitary, variably conical or patellate; costosepta arranged in 4–5 cycles in 6 systems in corallites of around 15 mm and larger; S1–S3 nearly equal in adult corallites, reaching the axial region; S1 often branch near the outer edge; oldest septa often straight, younger ones tend to be irregularly wavy; columella generally well-developed, spongy, fasciculate or formed by twisted segments, sometimes appearing net-like; columella projecting or deeper in corallum. Remarks. The specimens documented as Bathypsammia cleopatrae Youssef & Salama, 1969, in AbdelGawad & Gameil (1995) from the Paleocene of Egypt differ from the generic concept of Bathypsammia Marenzeller, 1906, in having a rather patellate shaped, free corallum, and lacking the septal arrangement according to the Pourtalès plan, thus closely agree with Palaeopsammia Wanner, 1902. In his revision on dendrophylliids Cairns (2001, p. 13) merged the species Caryophyllia jasmundi and Trochocyathus aegyptiacus and transferred them to the genus Palaeopsammia. The material documented in Schuster (1996) however, seems to closely correspond to the genus Caryophyllia. The specimens in the above list are interpreted to belong to the same species based on the occurrence of identical septal arrangement in corallites of the same size. The fact that in some specimens all of the septa are very wavy and flexuous whereas in others the septa seem rather straight is interpreted as a taxonomically irrelevant factor since in some specimens of the Egyptian type collection both types of septa appear, sometimes they are straight only in their uppermost margins but are flexuous a few mm below (see e.g., Pl. 2, Fig. 5a, upper part of image). In addition, in the Egyptian specimens the corallite walls are often very poorly preserved and can only be observed in a few places, which can result in appearances that seem to be different from the specimens with well-preserved wall structures. Moreover, in some corallites the columellar structures are more bulging than in others, e.g., the specimen figured on Pl. 2, Fig. 7b shows axial developments that seem to be different from the ones seen in the specimen figured on Pl. 2, Fig. 6. However, in both corals the columella is well-developed spongy-papillose. Moreover, in the Egyptian type material, in addition to specimens having a bulging columella, forms showing a flat or depressed columella occur (e.g., specimen figured on Pl. 2, Fig. 8a has a rather flat, in some places even depressed, columella). Therefore, these specimens are considered synonymous to the same species. Type locality of species. Danian of the USA (Martinez Group, California). Distribution. Cretaceous of Egypt, (?)Upper Cretaceous-(?)Eocene of New Zealand, Cenomanian and Maastrichtian of Madagascar, Maastrichtian of Mexico, ?Libya, Iran (Neyriz, Esfahan), Tunisia, and Turkmenistan, Maastrichtian-Paleocene of Egypt, Middle-Upper Maastrichtian of the UAE/Oman border region, Paleocene of Libya and the USA (Cannonball Marine Formation, South Dakota; Martinez Group, California). New Material. Danian of Egypt, NRM-PZ, sample nos.: Cn 47505–47515 and 47518–47565 (Babel Jasmund).

Genus Wadeopsammia Wells, 1933 Type species. Trochosmilia nodosa Wade, 1926, Maastrichtian of the USA (Tennessee, Ripley Formation).

Diagnosis. Corallum simple, subconical, curved, with a small or slightly expanded base which may enclose a small tabular cavity. Upper surface broad and nearly flat, covered by radiating rows of low nodules corresponding to the principal septa. Wall thin, cellular, composed of a porous tangle of synapticulae and disconnected distal portions of the septa. Exterior devoid of epitheca, covered with small rounded granules, scattered



39

toward the base but arranged in rows forming costae near the calice. Rows of small pores present in the corallite walls between costae, usually filled near the base of the corallum. Upper edges of septa beaded or dentate, covered by long granules laterally. Septa laminar and imperforate in structure. Columella formed by cemented axial ends of septa, large and solid, but not styliform. Dissepiments absent.

Wadeopsammia nodosa (Wade, 1926) Pl 3, figs 1a, b v*1926 v1933 2000 v2001 v2002

Trochosmilia nodosa n. sp.: Wade, p. 26, pl. 1, figs 1, 2, 5. Wadeopsammia nodosa (Wade): Wells, p. 228, pl. 12, figs 14–15, pl. 15, figs 23–25. Wadeopsammia nodosa (Wade 1926): Löser, p. 86. Wadeopsammia nodosa: Cairns, p 20–21, figs 3f–3i. Wadeopsammia nodosa (Wade, 1926): Baron-Szabo, p 79, pl. 57, figs 3–4.

Dimensions. d=3.3 mm; s=36; h=3 mm. Description. Having the characters of the genus; septa arranged in 3 complete cycles with the beginning of a fourth one. According to Cairns (2001, p. 21) the holotype represents a juvenile corallum. In excluding a second specimen reported for this genus by Wells (1933, p. 228), that has a papillose columella and lacks thecal porosity, Cairns states that the genus has been known from only the holotype, and therefore remains a poorly known taxon. Type locality of species. Maastrichtian of the USA (Tennessee, Ripley Formation). Distribution. Campanian-Maastrichtian of the USA (Ripley and Navarro Formations).

Genus Balanophyllia Searles Wood, 1844 (=Eupsammia Milne Edwards & Haime, 1848d, Type species. Madrepora trochiformis Pallas, 1766, Eocene of France); (=Ceratopsammia Alloiteau, 1958, Type species. Ceratopsammia besairiei Alloiteau, 1958, Campanian-Maastrichtian of Madagascar) Type species. Balanophyllia calyculus Searles Wood, 1844, Lower Pliocene of England (Norfolk).

PLATE 3: Fig. 1 Wadeopsammia nodosa (Wade, 1926), holotype, NMNH, I32703, Maastrichtian of the USA, scale bar: 1.5 mm. 1a, longitudinal view; 1b, cross view (photographs courtesy S. Cairns). Fig. 2 Palaeopsammia zitteli (Vaughan, 1900), BMNH, K⎫hn coll., R.30280 (holotype of Palaeopsammia fastigiata Kühn, 1933), Maastrichtian of Iran, scale bar: 3 mm. 2a, cross view, polished surface; 2b, longitudinal view. Fig. 3 Balanophyllia ponderosa Vaughan, 1900, syntype, NMNH, Vaughan coll., I158364, Paleocene of the USA. 3a, cross view, scale bar: 3 mm; 3b, longitudinal view, scale bar: 4 mm. Fig. 4 Balanophyllia caulifera (Conrad, 1847) (paratype of Balanophyllia schlosseri Traub, 1938), polished surface, cross view, BSP, Kch6, Middle Paleocene of Austria, scale bar: 5 mm. Fig. 5 Balanophyllia caulifera (Conrad, 1847) (holotype of Eupsammia narindensis Alloiteau, 1936), MNHN, Mo5211, Upper Maastrichtian of Madagascar. 5a, longitudinal view, scale bar: 5 mm; 5b, cross view, scale bar: 4.5 mm. Fig. 6 Balanophyllia besairiei (Alloiteau, 1958), syntype, MNHN, Mo5028, Campanian-Maastrichtian of Madagascar, scale bar: 5 mm. 6a, longitudinal view; 6b, cross view.


BARON-SZABO



41

Diagnosis. Solitary, variable conical, often trochoid, ceratoid, or turbinate. Base broad or narrow, attached or unattached. Asexual budding may occur from edge zone. Costae short, costosepta subcompact, Pourtalès plan present. Columella elongate, spongy. Wall synapticulothecal. Epitheca present or absent. Remarks. The validity of the taxon Eupsammia and its possible synonymy with Balanophyllia have been discussed since the early 20th century. In the generic revision and phylogenetic analysis of dendrophylliid genera, Cairns (2001) gave a detailed discussion of the taxonomic history of these two forms and provided descriptions for Balanophyllia and Eupsammia. While Cairns acknowledged the arguments that support the grouping of Balanophyllia with Eupsammia as its junior synonym (e.g., structurally they are identical, cooccurrence of the two forms in many regions since at least the Eocene), he separated the two forms on the basis of their attached or unattached living mode and differences in their early ontogenetical development, the latter idea of which had been proposed earlier by Durham (1949). According to Durham (1949, p. 139ff.) the taxon Balanophyllia shows a polycentric development (=”...formation of the adult corallum through two or more distinct stages....”) in its very first ontogenetical stages, whereas Eupsammia is considered to show a monocentric development in its earliest ontogenetical stages (=”....formation of the adult corallum by direct conical enlargement of the prototheca...”). However, recent investigations regarding ontogenetical stages of solitary and colonial scleractinians from Austria (Gosau Group, Upper Cretaceous) by Baron-Szabo (2003 and unpublished data) indicate that different septal and wall developments in the earliest ontogenetical stages are the result of whether the polyp was attached or was free-living (see remarks regarding the presumed characteristic of Cunnolites to be “free in ephebic stage” in the chapter of the Family Cunnolitidae Alloiteau). The sediments of the Upper Cretaceous part of the Gosau Group are characterized by muddy, soft-bottom paleoenvironments that often contain bioclastics. As a result, unattached and attached corals occur side by side. In numerous outcrops, the author of this paper has observed specimens of the same taxa of, e.g., Cunnolites, that always showed the development of a single septal cycle in the earliest ontogenetical stage with no or only one poorly developed thecal ring, whereas in attached forms at least 2 or 3 cycles and generally well-developed thecal structures were observed in the earliest stages that could be studied. This observation corresponds to the statement by Durham (1949, p. 143) regarding the attached form of Balanophyllia elegans: “No specimens representing the earliest stages of the corallum of Balanophyllia elegans were found....” Therefore, it is assumed that Balanophyllia and Eupsammia represent only one taxon that has the ability to settle on soft and hard grounds, living attached and unattached, and that, resulting from this, the terms monocentric and polycentric have no taxonomic value for this genus. With the exception of terms monocentric and polycentric, which are considered to have no taxonomic value for this genus, the diagnosis given for Balanophyllia represents a combination of the two descriptions provided by Cairns (2001, pp. 14 and 17) for Balanophyllia and Eupsammia.

Balanophyllia besairiei (Alloiteau, 1958) Pl. 3, figs 6a, b v*1958 ?1958 1987 v2001 2000 2002

Ceratopsammia Besairiei nov. sp.: Alloiteau, p. 200, pl. 34, figs 7–9. Eupsammia lehmani nov. sp.: Alloiteau & Tissier, p. 263, pl. 1, figs 10a–b, 10a’. Balanophyllia subcylindrica (Philippi, 1851): Kuzmicheva, p. 148, pl. 26, figs 1a–b. Balanophyllia (=Eupsammia) besairiei (Alloiteau, 1958): Cairns, p. 17ff., pl. 2, figs b, f. Ceratopsammia besairiei, Alloiteau 1958: Löser, p. 18. Balanophyllia besairiei (Alloiteau, 1958): Baron-Szabo, p. 80.


BARON-SZABO

Dimensions. d=4–6 (? up to 7.9) mm; d (min)/d (max)=0.75–0.83; s=48–60 (? up to 98); h=10–15 (? up to 16.5) mm. Description. Trochoid to subcylindrical, elliptical or slightly elliptical in outline; septa arranged in 3 cycles in 12 systems with the beginning of a fourth cycle. Remarks. In forming a long and slim corallum that is elliptical in outline, and having 98 septa in a maximum corallite diameter of 7.9 mm, the species Eupsammia lehmani Alloiteau & Tissier, 1958, from the Paleocene of France most likely represents an ontogenetically older image of Balanophyllia besairiei (Alloiteau, 1958). However, because the development of the septal apparatus could not be identified, the species is only provisionally grouped with B. besairiei. Type locality of species. Campanian-Maastrichtian of Madagascar (Analavelona). Distribution. Campanian-Maastrichtian of Madagascar, Paleocene of Ukraine and ?France.

Balanophyllia caulifera (Conrad, 1847) Pl. 3, figs 4–5b *1847 v1900 v1900 parsv1935 v1936 v1938 1941a v1951b v1951b ?1951b 1955 v1958 v1967 1974 1974 1984 1984 1987 nonv1996 2000 2000 2000 2000 2000 2001 2001 2001 2001 2002 2002

Turbinolia caulifera: Conrad, p. 296. Balanophyllia caulifera (Conrad): Vaughan, p. 176, pl. 20, figs 15–18 (older synonyms cited therein)(topotypes studied). Balanophyllia caulifera var. multigranosa var. nov.: Vaughan, p. 177, pl. 20, figs 19–20. Balanophyllia ponderosa var. texana new variety: Vaughan & Popenoe, p. 341, pl. 4, fig. 11, non figs 6–10. Eupsammia narindensis (nov. sp.): Alloiteau, p. 14, pl. 6, figs 24–27. Balanophyllia schlosseri: Traub, p. 37, pl. 1, figs 4a–c, text-fig. 1. Balanophyllia pinraensis, n. sp.: Wells, p. 16 (316), pl. 2, figs 6–6a. Eupsammia narindensis All.: Alloiteau, p. 82, pl. 13, fig. 9. Eupsammia narindensis All. var. elongata nov. var. (Alloiteau): Alloiteau, p. 82, pl. 13, figs 10–11. Eupsammia Collignoni nov. sp.: Alloiteau, p. 83, pl. 13, figs 12–14. Eupsammia pozaryskii n. sp.: Rozkowska, p. 257, pl. 1, fig. 7, text-figs 11–14. Palaeopsammia mitsinjoensis nov. sp.: Alloiteau, p. 204, pl. 37, figs 14, 16–18. Balanophyllia schlosseri Traub: Kühn & Traub, p. 16, pl. 1, figs 7–10. Balanophyllia (Balanophyllia) santaelenensis n. sp.: Frost & Langenheim, p. 308, pl. 120, figs 1–3, pl. 121, figs 1–7. Balanophyllia schlosseri Traub, 1938: Kuzmicheva, p. 65, pl. 10, figs 4a–b. Balanophyllia aff. B. (Eupsammia) varians Stol: Pal, et al., p. 57. Balanophyllia (E) ladakhensis sp. nov.: Pal, et al., p. 57, pl. 1, figs 1–2. Balanophyllia schlosseri Traub, 1938: Kuzmicheva, p. 147, pl. 25, figs 10a–b. Palaeopsammia mitsinjoensis Alloiteau, 1958: Baron-Szabo & Steuber, p. 20, pl. 10, fig. 1, 3. Eupsammia collignoni, Alloiteau 1951: Löser, p. 35. Eupsammia ladakhensis, Pal, et al 1984: Löser, p. 35. Eupsammia narindensis, Alloiteau 1936: Löser, p. 35. Eupsammia narindensis var. elongata, Alloiteau 1936: Löser, p. 35. Palaeopsammia mitsinjoensis, Alloiteau 1958: Löser, p. 58. Balanophyllia (B.) schlosseri Traub, 1938: Cairns, p. 15. Balanophyllia (B.) santaelensis Frost, 1975: Cairns, p. 15. Balanophyllia (E.) caulifera (Conrad, 1847): Cairns, p. 18. Balanophyllia (E.) ponderosa var. texana Vaughan and Popenoe, 1935: Cairns, p. 18. Palaeopsammia mitsinjoensis Alloiteau, 1958: Baron-Szabo, p. 78. Balanophyllia narindensis (Alloiteau, 1936): Baron-Szabo, p. 80.



43

Dimensions. d (min)=5.3–24 mm; d (max)=7.5–34 mm; d (min)/d (max)=0.63–0.9; s=48–96; h=8.5–50 mm. Description. Trochoid to subceratoid, nearly circular to elliptical in outline; septa arranged in 5 cycles in 6 systems, complete in late adult stages; S1 and S2 equal or nearly equal; synapticulae abundant peripherally. Remarks. The forms given in the synonymy list above are interpreted to represent various ontogenetical stages of the same species according to which 48 septa arranged in 4 complete cycles are generally present in a corallite diameter of 5 to 8 mm, septa of the beginning fifth cycle occur in a corallite diameter of 9 mm and larger. In corallites having reached a diameter of around 25 mm, septa arranged in 5 complete cycles are present. In juvenile ontogenetical stages the corallite shape is nearly circular (ratio of minimum diameter/maximum diameter is generally around 0.8–0.9) and becomes more oval in later stages of ontogeny. Type locality of species. Lower Oligocene of the USA (Vicksburgian). Distribution. Campanian-Maastrichtian of India (Ladakh), Upper Maastrichtian of Madagascar, Upper Maastrichtian-Lower Danian of Ukraine, Paleocene of the USA (Texas), Middle Paleocene of Austria (Kroisbach, Grabung Kch 7), Upper Danian of Poland, Middle Eocene of Peru (Negritos), Middle-Upper Eocene of Mexico (San Juan Formation and Ixtaclum shale), Lower Oligocene of the USA.

Balanophyllia ponderosa Vaughan, 1900 Pl. 3, figs 3a, b v*1900 parsv1935 v1938 v1967 2001 2001

Balanophyllia ponderosa sp. nov.: Vaughan, p. 173, pl. 20, figs 1–2. Balanophyllia ponderosa var. texana new variety: Vaughan & Popenoe, p. 341, pl. 4, figs 6–10, non fig. 11. Balanophyllia sp.: Traub, p. 38, pl. 1, fig. 5, text-fig. 2. Balanophyllia traubi nov. spec.: Kühn & Traub, p. 17, pl. 2, fig. 3. Balanophyllia (E.) ponderosa Vaughan, 1900: Cairns, p. 18. Balanophyllia (E.) ponderosa var. texana Vaughan and Popenoe, 1935: Cairns, p. 18.

Dimensions. d (min)=9–20 mm; d (max)=10–24 mm; d (min)/d (max)=0.71–0.9; s=120 up to around 160 in the larger specimens; h=12–45 mm. Description. Trochoid to subturbinate, subpatellate during some ontogenetical stages; circular in outline in juvenile specimens, elliptical in outline in later adult stages; septa thin, numerous, arranged in 5 cycles in 6 systems, with the beginning of a sixth cycle in late adult stages; in corallum of 9–10 mm in diameter about 80 septa present, in corallite diameter of around 18 mm the fifth septal cycle complete; columella often large, spongy or net-like; synapticulae very abundant peripherally, forming a dense vesicular wall. Type locality of species. Paleocene of the USA (Alabama, Midwayan). Distribution. Paleocene of Austria and the USA (Alabama and Texas).

Suborder Caryophylliina Vaughan & Wells, 1943 Diagnosis. Solitary and colonial. Septa laminar, composed of one fan system of simple, very small trabeculae; septal margins completely smooth. Dissepiments usually not developed. Polyps small to moderate in size, with ridged stomodaea, and tentacles arranged in 1 to 3 rings.


BARON-SZABO

Family Caryophylliidae Dana, 1846 Diagnosis. Solitary and colonial. Colony formation by extracalicinal budding (rarely intracalicinal) budding, forming phaceloid or dendroid colonies. Costae commonly covered by stereome or epitheca. Septa exsert. Columella absent or formed by curled trabecular laths, solid, spongy. Pali or paliform lobes common. Endothecal dissepiments developed in some groups. Remarks. Gray (1847) is commonly referred to as the first author to have described the family Caryophylliidae (e.g. Milne Edwards 1857; Vaughan & Wells 1943; Alloiteau 1952a). However, as pointed out by Cairns (1989) one year prior to Gray’s work the family Caryophylliidae was created by Dana (1846: 364), which gives Dana priority of authorship.

Subfamily Caryophylliinae Milne Edwards, 1857 Diagnosis. Mostly solitary, fixed or free. Ahermatypic. Phaceloid or dendroid colonies formed by extratentacular budding from the edge zone. Septothecate, with strong costae. Endothecal dissepiments rare.

Genus Caryophyllia Lamarck, 1801 Type species. Madrepora cyathus Ellis and Solander, 1786, Recent, Mediterranean Sea.

Diagnosis. Solitary, turbinate or cylindrical, fixed or free. Costosepta laminar, compact. septal margins smooth or nearly smooth. Pali opposite third cycle of septa or before 2nd group of septa where hexameral symmetry is lost. Columella formed by twisted trabecular segments. Endothecal dissepiments few in number. Wall septothecal.

Caryophyllia arcotensis (Forbes, 1846) Pl. 4, figs 1a, b, Text-Fig. 1 v*1846 1850 1851b non1873 1873 non1922 non1936 (v)1951b (v)1951b 1955 non1958 non1995 2000 2000

Turbinolia arcotensis, sp. nov.: Forbes, p. 163, pl. 19, figs 9a–b. Ellipsosmilia arcotensis: d’Orbigny, vol. 2, p. 276. Trochosmilia arcotensis: Milne Edwards & Haime, p. 17. Caryophyllia arcotensis, Stoliczka: Stoliczka, p. 7, pl. 1, figs 1–10. Turbinolia arcotensis, Stoliczka, p. 8, pl. 1, figs 11–12. Caryophyllia arcotensis, Forbes: Cottreau, p. 169, pl. 1, figs 6–6b. Caryophyllia arcotensis Forbes: Alloiteau, p. 9. Caryophyllia arcotensis Stol., var. regularis nov. var.: Alloiteau, p. 81, pl. 13, figs 1–1a. Caryophyllia antonibensis, nov. sp.: Alloiteau, p. 80, pl. 13, figs 2–4. Caryophyllia antonibensis All.: Rossi Ronchetti, p. 113, pl. 11, figs 5–6. Protrochocyathus arcotensis Forbes, sp.: Alloiteau, p. 136, pl. 17, figs 6a–g. Caryophyllia arcotensis (Forbes, 1846): Tchéchmédjiéva, p. 55, pl. 10, figs 11, 12. Caryophyllia antonibensis, Alloiteau 1951: Löser, p. 17. Caryophyllia arcotensis (Forbes 1846): Löser, p. 17.



45


BARON-SZABO

PLATE 4: Fig. 1 Caryophyllia arcotensis (Forbes, 1846), holotype, BMNH, R.29211, Maastrichtian of India, scale bar: 4.5 mm. 1a, cross view; 1b, longitudinal view. Fig. 2 Caryophyllia mediavia Vaughan & Popenoe, 1935, holotype, NMNH, M371041, Danian of the USA, scale bar: 3.4 mm. 2a, cross view; 2b, longitudinal view. Fig. 3 Bathycyathus lloydi (Vaughan, 1920) (holotype of Steriphonotrochus manorensis, Wells, 1933), NMNH, Wells coll., I75132, Upper Maastrichtian of the USA. 3a, upper surface, cross view, scale bar: 5 mm; 3b, polished surface, juvenile stage of corallum, scale bar: 1 mm; 3c, longitudinal view, scale bar: 2 mm. Fig. 4 Bathycyathus lloydi (Vaughan, 1920), holotype, NMNH, Vaughan coll., I32376, Paleocene of the USA, scale bar: 2.5 mm. 4a, upper surface, cross view; 4b, longitudinal view. Fig. 5 Trochocyathus speciosus (Gabb & Horn, 1860) (syntype of Trochocyathus epicharis Wanner, 1902), NRM-PZ, Wanner coll., Cn 47525, Danian of Egypt, scale bar: 1.5 mm. 5a, cross view, upper surface; 5b, longitudinal view; 5c, side view.

Dimensions. d (min)=16.5–24 mm; d (max)=17–25 mm; d (min)/d (max)=0.8–1; s=40–48+s5; h=up to 35 mm. Description. Solitary, variable conical (e.g., turbinate, cylindrical), slightly elliptical or circular in outline; costosepta straight, developed in 4 complete cycles in 6 systems, regularly or irregularly alternating in length and thickness; septa laterally smooth or finely granulated; pali seem to be axial to the second set of septa.

Text-Fig. 1 Caryophyllia arcotensis (Forbes, 1846) as figured in Alloiteau (1951b) (as holotype of Caryophyllia arcotensis var. regularis Alloiteau, 1951b). A, B. Upper Maastrichtian of Madagascar, scale bar: 10 mm; A, upper surface longitudinal view; B, cross view of corallum.

Remarks. The specimens described from the Albian of Madagascar as Caryophyllia arcotensis (Forbes, 1846) by Alloiteau (1936), which were later transferred to the genus Protrochocyathus in Alloiteau, 1958 (Protrochocyathus is interpreted here to be the junior synonym of Trochocyathus), have calicinal diameters ranging from 8–10 mm with 48 septa arranged in 6 systems. Therefore, they most likely represent junior synonyms of Trochocyathus antsiranensis Collignon, 1931, from the Cenomanian of Madagascar. The latter species is characterized by a calicinal diameter of around 8 mm and 48 septa arranged in 6 systems. SCLERACTINIAN CORALS OF THE K/T-BOUNDARY


47

The specimens described as Caryophyllia arcotensis (Forbes) from the Upper Campanian of Bulgaria by Tchéchmédjiéva (1995) differ from the genus Caryophyllia in lacking pali. They most likely belong to the genus Parasmilia. Type locality of species. Maastrichtian of India (Pondicherry). Distribution. Maastrichtian of India and Libya (Sofeggin), Upper Maastrichtian of Madagascar.

Caryophyllia konincki (Milne Edwards & Haime, 1848) Text-Fig. 2 v*1848d v1850a v1850a 1850a v1857 v1857 v1857 v1863 1869 1873 1880 1880 1880 pars1914 1914 1914 1914 1914 1914 1914 1922 1925 1925 v1933 v1933 1952b 1955 1955 (v)1955 1955 1970 1970 1970 1970 1972 1975 1984 1987 1987 1987

Cyathina Koninckii: Milne Edwards & Haime, p. 290. Cyathina cylindrica: Milne Edwards & Haime, p. 45. Cyathina Bredae: Milne Edwards & Haime, p. 46. Cyathina Debeyana: Milne Edwards & Haime, p. 46. Caryophyllia cylindrica: Milne Edwards, vol. 2, p. 17. Caryophyllia Konincki: Milne Edwards, vol. 2, p. 17. Caryophyllia Bredai: Milne Edwards, vol. 2, p. 18. Caryophyllia decemplex: de Fromentel, p. 168, pl. 21, figs 2–2b. Caryophyllia decemeris de Fromentel: Duncan, p. 16. Caryophyllia arcotensis Stoliczka, p. 7, pl. 1, figs 1–10 (non Forbes 1846). Caryophyllia compressa, Duncan: Duncan, p. 17, pl. 1, figs 1–4. Caryophyllia Indica, Duncan: Duncan, p. 17, pl. 1, figs 5–7. Caryophyllia Feddeni, Duncan: Duncan, p. 18, pl. 1, figs 8–10. Caryophyllia Arcotensis Forbes sp. 1846: Felix, pars 7, p. 209. Caryophyllia Bredai E. H. sp. 1850: Felix, pars 7, p. 209. Caryophyllia compressa Duncan 1880: Felix, pars 7, p. 210. Caryophyllia cylindrica E. H. sp. 1850: Felix, pars 7, p. 210. Caryophyllia Feddeni Duncan 1880: Felix, pars 7, p. 210. Caryophyllia Indica Duncan 1880: Felix, pars 7, p. 210. Caryophyllia Konincki E. H. sp. 1848: Felix, pars 7, p. 210. Caryophyllia danica, n. sp.: Nielsen, p. 221, pl. 2, figs 19, 20. Caryophyllia Bredai E. H. sp.: Umbgrove, p. p. 114, pl. 11, figs 20, 25–26. Caryophyllia Konincki E. H. sp.: Umbgrove, p. 114. Caryophyllia stephensoni n. sp.: Wells, p. 124, pl. 12, figs 6–7, pl. 14, figs 15–18. Caryophyllia mississippiensis n. sp.: Wells, p. 126, pl. 14, figs 19–20. Cylindrocyathus popenguinensis nov. sp.: Alloiteau, p. 11, pl. 1, figs 3–4. Caryophyllia arcotensis (Forb.): Rossi Ronchetti, p. 114. Caryophyllia cupuliformis Stol.: Rossi Ronchetti, p. 114. Caryophyllia calcitrapai (v. Koenen): Rozkowska, p. 246, pl. 1, figs 3, 4, pl. 2, figs 1, 2, text-figs 4–6. Caryophyllia kongieli n. sp.: Rozkowska, p. 251, pl. 1, figs 1, 2, pl. 2, fig. 4, text-figs 7, 8. Caryophyllia aegyptiaca sp. nov.: Hassan & Salama, p. 82, pl. 2, figs 11a–c. Caryophyllia phanerocosta sp. nov.: Hassan & Salama, p. 82, pl. 2, figs 11a–c. Caryophyllia quadragenaria var. vigentipali Hassan & Salama, p. 84, pl. 2, figs 4a, b. Asterosmilia decapali sp. nov.: Hassan & Salama, p. 89, pl. 3, figs 5a–c. Caryophyllia andreasi sp. n.: Floris, p. 40, pl. 2, figs 1A–5. Caryophyllia matesovae sp. nov.: Kuzmicheva, p. 23, pl. 2, figs 8–9. Caryophyllia arcotensis Stoliczka, 1873: Pal. et al., p. 60, pl. 2, fig. 20. pl. 3, fig. 21. Caryophyllia jasmundi Wanner, 1902: Kuzmicheva, p. 114, pl. 18, figs 4a–5b. Caryophyllia kongieli Rozkowska, 1955: Kuzmicheva, p. 115, pl. 18, figs 6a, b. Caryophyllia danica Nielsen, 1922: Kuzmicheva, p. 117, pl. 19, figs 2a, b.


BARON-SZABO

1987 1995 1995 ?1995 non1995 1995 1995 1995 1996 ?1997 1999 2000 2000 2000 2000 2000 2000 2000 2000 2000 2000 2000 2002 2002 2002 2002 v2002 2002 2002 2002 2002 2002 v2002 (v)2005

Caryophyllia matesovae Kuzmicheva, 1975: Kuzmicheva, p. 117, pl. 19, figs 3–4. Caryophylliidae sp. indet. 2: Abdel-Gawad & Gameil, p. 20, pl. 20, figs 7–8. Caryophyllia decemplex Fromentel, 1862: Tchéchmédjiévaa, p. 54, pl. 10, fig. 9. Caryophyllia konincki (Edwards & Haime, 1847): Tchéchmédjiéva, p. 55, pl. 10, fig. 10. Caryophyllia agatdalensis Floris, 1972: Tchéchmédjiéva, p. 55, pl. 10, fig. 10. Caryophyllia cylindrica (Edwards & Haime, 1850): Tchéchmédjiéva, p. 56, pl. 11, fig. 7, pl. 12, fig. 1. Caryophyllia debeyana (Edwards & Haime, 1850): Tchéchmédjiéva, p. 57, pl. 12, figs 2, 3. Caryophyllia bredae (Edwards & Haime, 1850): Tchéchmédjiéva, p. 57, pl. 12, fig. 4. Caryophyllia jasmundi Wanner, 1902: Schuster, p. 80. Caryophyllia cf. konincki Milne-Edw. & H. 1848: Löser, p. 104. Caryophyllia bredai: Leloux, p. 193, figs 2. Caryophyllia arcotensis, Stoliczka 1873: Löser, p. 17. Caryophyllia bredai (Milne-Edwards & Haime 1851): Löser, p. 17. Caryophyllia compressa, Duncan 1880: Löser, p. 17. Caryophyllia cylindrica (Milne-Edwards & Haime 1851): Löser, p. 17. Caryophyllia decemplex, de Fromentel 1862: Löser, p. 17. Caryophyllia feddeni, Duncan 1880: Löser, p. 17. Caryophyllia indica, Duncan 1880: Löser, p. 17. Caryophyllia konincki (Milne-Edwards & Haime 1848): Löser, p. 17. Caryophyllia mississippiensis, Wells 1933: Löser, p. 17. Caryophyllia stephensoni, Wells 1933: Löser, p. 17. Cylindrocyathus popenguinensis, Alloiteau 1952: Löser, p. 27. Caryophyllia arcotensis Stoliczka, 1873: Baron-Szabo, p. 157. Caryophyllia bredai (Milne Edwards & Haime, 1850): Baron-Szabo, p. 157. Caryophyllia compressa Duncan, 1880: Baron-Szabo, p. 157. Caryophyllia cylindrica (Milne Edwards & Haime, 1850): Baron-Szabo, p. 157. Caryophyllia decemplex Fromentel, 1863: Baron-Szabo, p. 157, pl. 117, figs 4, 7. Caryophyllia feddeni Duncan, 1880: Baron-Szabo, p. 157. Caryophyllia indica Duncan, 1880: Baron-Szabo, p. 157. Caryophyllia konincki (Milne Edwards & Haime, 1850): Baron-Szabo, p. 157. Caryophyllia mississippiensis Wells, 1933: Baron-Szabo, p. 157. Caryophyllia popenguinensis (Alloiteau 1952): Baron-Szabo, p. 157. Caryophyllia stephensoni Wells, 1933: Baron-Szabo, p. 157, pl. 117, figs 5–6. Caryophyllia danica: Bernecker & Weidlich, p. 16ff, fig. 7A, figs 7 (2, 3).

Dimensions. d (min)=1.5–16 mm; d (max)=2–18 mm; d (min)/d (max)==0.81–0.96; s (adult)=48+s5; h=up to 45 mm. Description. Turbinate to variably conical, circular to slightly elliptical in outline; costosepta generally thin, arranged in 4 cycles in 6 systems, complete in specimens that are 8 mm in diameter and larger, incomplete in younger forms (3–7 mm in diameter); in specimens 2–4 mm in diameter the number of septa is 24 or slightly larger (around 30), in a corallite diameter of 6 mm the number of septa is around 36, in a corallite of 7 to 8 mm in diameter 4 complete or nearly complete cycles (44–48 septa) are developed; in corallites thar are larger than 8 mm in diameter, septa of an imcomplete 5th cycle are present (e.g., in d: 14–18 mm, 68–72 septa present); septa S1 and S2 more dominant, becoming equal in later ontogenetical stages; columella made of a small number of twisted segments, weakly developed; pali elongate, often thick. Remarks. The uniting of the species in the synonymy list above is based on the idea that they form different ontogenetical stages of the same species. The varying shape of the calice from almost circular to slightly



49

elliptical can be observed in all of the investigated specimens. Both larger and smaller specimens have been reported from Denmark, Egypt, Pakistan, and Ukraine. In the original description of the species Caryophyllia popenguinensis (Alloiteau, 1952b) from the Maastrichtian of Senegal, Alloiteau (1952b) stated that 48 septa were present in a corallite of 4.5 mm in diameter. However, the illustration of the type specimen (Alloiteau 1952b, pl. 1, fig. 4) seems to only show around 34 septa, thus closely corresponding with very juvenile stages of the species C. konincki. The material from the Maastrichtian of Libya assigned to Caryophyllia cupuliformis by Rossi Ronchetti (1955, p. 114) is slightly larger (10 mm) than the species C. konincki (=7 mm), has exactly 4 complete cycles or additional septa (48–54) of the beginning fifth cycle, and is nearly circular in outline. It assumed that these specimens represent a slightly later ontogenetical stage than the specimens described as C. konincki.

Text-Fig. 2 Caryophyllia konincki (Milne Edwards & Haime, 1848d). A, B, syntype of Caryophyllia cylindrica (Milne Edwards & Haime, 1850a), MNHN, Mo1151, Maastrichtian of Belgium, scale bar: 5 mm; A, cross view of corallum; B, upper surface longitudinal view; C, type material of Caryophyllia kongieli Rozkowska, 1955, as figured in Rozkowska (1955), Upper Danian of Poland, cross view, scale bar: 2 mm; D, G, syntypes of Caryophyllia konincki (Milne Edwards & Haime, 1848d), MNHN, Mo1301, Milne Edwards & Haime coll., Maastrichtian of Belgium; D, longitudinal view of corallum, slightly oblique, scale bar: 3 mm; G, cross view, polished surface, scale bar: 3.5 mm; E, F, syntype of Caryophyllia bredai (Milne Edwards & Haime, 1850a), MNHN, Mo 1150, Maastrichtian of Belgium, scale bar: 5.5. mm; E, cross view of corallum, oblique; F, upper surface longitudinal view; H, Caryophyllia indica (Duncan, 1880), type material as figured in Duncan (1880), Paleocene of Pakistan, cross view of corallum showing part of septal apparatus, scale bar: 3.5 mm; I, Caryophyllia feddeni (Duncan, 1880), type material as figured in Duncan (1880), Paleocene of Pakistan, cross view of corallum showing part of septal apparatus, scale bar: 4 mm.


BARON-SZABO

The species Caryophyllia andreasi Floris, 1972, was described from the Danian of Denmark by Floris (1972, p. 40ff) as having 72 septa in a corallite 17 x 15.3 mm in diameter. However, on page 42 he combined specimens in an illogical ontogenetical manner in that corals are grouped which 1) have the same number of septa in coralla significantly differing in diameter, and 2) larger corallites that have a smaller number of septa are put together with smaller corallites that have a distinctly larger number of septa (e.g., coralla that are 7–8 mm in diameter with 64–66 septa are grouped with corals that are 9 mm in diameter but only have 44 septa). Therefore, it is assumed that these specimens represent different species. Among the specimens described by Floris are corals that very closely agree with the species C. konincki (d: 7–8 mm with 48 septa). Moreover, forms having 52–60 septa in corallites 9–11 mm in diameter are documented which are here interpreted to represent intermediate ontogenetical of C. konincki. In addition, there are corals that are believed to belong to the ontogenetically latest stages (d: 14–18 mm with 68–72 septa). Besides the dimensions of skeletal elements, specimens assigned to C. andreasi closely agree with the original description of the species C. konincki in having 1) rather thin septa, 2) S1 and S2 nearly equal, 3) a rather weakly developed columella, and 4) elongate pali. From the Upper Campanian of Bulgaria Tchéchmédjiéva (1995) described the form Caryophyllia agatdalensis Floris, 1972. However, the specimen illustrated shows a septal arrangement according to the Pourtalès plan, therefore representing a dendrophylliid form. Type locality of species. Maastrichtian of Belgium (Ciply). Distribution. Senonian of England (Northfleet), Upper Campanian of Bulgaria and Germany, Maastrichtian Belgium (Ciply), the Netherlands, Libya (Sofeggin), India (Arrialoor Group), ?Germany, and Senegal, Upper Maastrichtian of the USA (Navarro Formation, Mustang River, Texas), Campanian-Maastrichtian (Ladakh) and Danian of Pakistan (Laki Range, Sind), Paleocene of Egypt and Ukraine, Danian of Denmark (Greenland and Fakse), Upper Danian of Poland and Turkmenistan.

Caryophyllia mediavia Vaughan & Popenoe, 1935 Pl. 4, figs 2a, b v*1935 v1935 1970 1972 1987 ?1993

Caryophyllia mediavia new species: Vaughan & Popenoe, p. 333, pl. 3, figs 12, 13. Caryophyllia dumblei new species: Vaughan & Popenoe, p. 334, pl. 3, figs 14–16. Asterosmilia aliquantula Squires, 1958: Hassan & Salama, p. 89, pl. 3, figs 6a–c. Caryophyllia agatdalensis sp. n.: Floris, p. 49, pl. 2, figs 8a–9. Caryophyllia agatdalensis Floris, 1972: Kuzmicheva, p. 116, pl. 19, figs 1a, b. Caryophyllia dumblei ? Vaughan and Popenoe: Hansen et al., p. 257, Table 1.

Dimensions. d (max)=9–12 mm; d (min)=8.3–10 mm; s=40–48 (+?s5); h=up to 16.5 mm. Description. Variably conical corallum (trochoid, cylindrical, cornute); costosepta thin, fragile, arranged in 4 compete or incomplete cycles in 6 systems in a corallite of 9–12 mm; 40–44 septa are present in coralla of 7–8 mm diameter; wall rather thick. Type locality of species. Danian of the USA (Texas, Falls County). Distribution. Lower Maastrichtian of Mexico (Cerralvo, this paper), Maastrichtian-Danian of Denmark (Greenland), Danian of Azerbaijan and the USA (Texas), Paleocene of Egypt. New Material. Lower Maastrichtian of Mexico (Cerralvo), sample nos: UANL CE MAAS-211, -215, 218.



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Genus Bathycyathus Milne Edwards & Haime, 1848 (=Monticyathus Alloiteau & Tissier, 1958, Type species. Monticyathus lehmani Alloiteau & Tissier, 1958, Danian of Belgium); (=Coronocyathus Alloiteau &Tissier, 1958, Type species. Coronocyathus goffinti Alloiteau & Tissier, 1958, Danian of Belgium); (=Frescocyathus Barta-Calmus, 1969, Type species. Frescocyathus tessieri Barta-Calmus, 1969, Paleocene of Ivory Coast); (=Steriphonotrochus Vaughan, 1900, Type species. Steriphonotrochus pulcher Vaughan, 1900, Oligocene of the USA [Red Bluff, Mississippi]). Type species. Bathycyathus chilensis Milne Edwards & Haime, 1848c, Recent, Pacific Ocean (off the coast of Chile).

Diagnosis. Solitary, turbinate or variably conical, fixed or free. Often circular in outline in juvenile stages, becoming compressed in later ontogenetical stages. Costosepta laminar, compact. Septal margins smooth or nearly smooth. Pali not distinct from columellar laths. Columella formed by twisted trabecular segments. Endothecal dissepiments few in number. Wall septothecal, septoparathecal when not properly thickened. Remarks. Alloiteau & Tissier (1958) stated that their newly created genus Coronocyathus was characterized by 1) a parathecal and septothecal wall, 2) laterally finely dentate compact costosepta, 3) pali opposite S1 and S2, 4) a spongy columella forming twisted segments, and 5) the lack of both endothecal developments and synapticulae. However, in contrast to this statement the illustrations of the type material (Alloiteau & Tissier, 1958, pl. 1, figs 5 a–e) indicate that the specimens are characterized by a rather septothecal wall throughout their ontogenetical development, show a small number of endothecal dissepiments, and have paliform structures which, especially in younger ontogenetical stages, are not distinct from columellar developments. Therefore, the assignment to the genus Bathycyathus is suggested. Alloiteau & Tissier (1958) created the genus Monticyathus. The illustrations of the type specimen differ from the description of the holotype in that there are dissepiments and rather septothecal (septoparathecal in places) developments present, whereas the authors stated that an endotheca was absent and a parathecal wall was present. In addition, the illustrations of the holotype show a turbinate corallum, that has a spongy-net-like columella, and pali that are generally fused with columellar segments. Therefore, the assignment to the genus Bathycyathus is suggested. Barta-Calmus (1969) created the genus Frescocyathus from the Paleocene of Ivory Coast, stating that it differed from Bathycyathus in having a trochoid-shaped corallum and a weakly developed columella. Because in Bathycyathus the corallum shape is most likely largely controlled by its mode of attachment, due to which the corallum later becomes fixed or free, and the size of the columella that can be strongly or weakly developed, the assignment of Frescocyathus to the genus Bathycyathus is suggested.

Bathycyathus corneti (Alloiteau & Tissier, 1958) Text-Fig. 3 (v)*1958 (v)1958 (v)1958

Kionotrochus corneti nov. sp.: Alloiteau & Tissier, p. 279, pl. 1, figs 9a–b’. Monticyathus lehmani nov. sp.: Alloiteau & Tissier, p. 284, pl. 1, figs 8a–b’. Coronocyathus goffinti nov. sp.: Alloiteau & Tissier, p. 287, pl. 1, figs 5 a–e.

Dimensions. d (max., adult)=6.7 mm; d (min., adult)=3.8 mm; s (adult)=48 + s4 (12s1+12s2+24s3+s4); h=up to about 45 mm.


BARON-SZABO

Description. Corallum compressed, conical, elliptical in outline; septal arrangement in 4 cycles in 12 systems in late adult stages, in irregularly developed 6 systems in younger ontogenetical stages; laterally granulate. Dimensions of skeletal elements during ontogeny see Fig. 3. Remarks. The species Bathycyathus lehmani (Alloiteau & Tissier, 1958) from the Danian of Belgium seems to completely correspond to the slightly more juvenile stage of Bathycyathus goffinti (Alloiteau & Tissier, 1958) illustrated on pl. 1, fig. 5c of Alloiteau & Tissier (1958) from the same locality. Therefore, it is assumed that they are synonymous. Type locality of species. Danian of Belgium. Distribution. Danian of Belgium.

Text-Fig. 3 Bathycyathus corneti (Alloiteau & Tissier, 1958). A, B, holotype of Bathycyathus corneti (Alloiteau & Tissier, 1958), based on the illustration in Alloiteau & Tissier (1958), figured as Kionotrochus corneti Alloiteau & Tissier, 1958, Danian of Belgium, A, cross view of corallum; B, upper surface longitudinal view; C, D, holotype of Bathycyathus goffinti (Alloiteau & Tissier, 1958), based on the illustration in Alloiteau & Tissier (1958), figured as Coronocyathus goffinti Alloiteau & Tissier, 1958, Danian of Belgium, C, cross view of corallum; D, upper surface longitudinal view; E, F, G, H, paratype of Bathycyathus goffinti (Alloiteau & Tissier, 1958), based on the illustrations in Alloiteau & Tissier (1958), figured as Coronocyathus goffinti Alloiteau & Tissier, 1958, Danian of Belgium, cross view of corallum, polished surface, showing different ontogenetical stages; I, J, holotype of Bathycyathus lehmani (Alloiteau & Tissier, 1958), based on the illustration in Alloiteau & Tissier (1958), figured as Monticyathus lehmani Alloiteau & Tissier, 1958, Danian of Belgium, I, cross view of corallum; J, upper surface longitudinal view. A, B, C, D, E: scale bar 5 mm; F, G, H: scale bar 3 mm; I, J: scale bar: 4 mm.

Bathycyathus lloydi (Vaughan, 1920) Pl. 4, figs 3a–4b v*1920 v1920 v1920 v1933

Paracyathus lloydi, n. sp.: Vaughan, p. 62, pl. 10, figs 3–3b. Paracyathus thomi, n. sp.: Vaughan, p. 63, pl. 10, figs 4–4d. Paracyathus kayserensis, n. sp.: Vaughan, p. 68, pl. 10, figs 5–5b. Steriphonotrochus ? manorensis n. sp.: Wells, p. 123 (205), pl. 14, fig. 21.



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(v)1972 2000

Paracyathus kangliaensis sp. n.: Floris, p. 60, pl. 3, figs 6A–9. Steriphonotrochus manorensis, Wells 1933: Löser, p. 74.

Dimensions. d (adult)=8 x 14.5 mm, (juvenile)=4.5 mm; s (adult)=64, s (juvenile): 36; costae (adult)=64, (juvenile)=around 30; h=13.5–18 mm. Description. Corallum elongated conical, elliptical in outline in later adult stages; septal arrangement rather irregular in juvenile stages, with about 30 extending to the columella in adult image; in late adult stages number of costae equal number of septa, in juvenile stages number of costae often smaller than number of septa; septa laterally granulate, marginally smooth; in corallite of around 7 mm in diameter 48 septa developed; around 60 septa are present in corallite of d (max) 10 mm. Remarks. The species B. kayserenesis (Vaughan, 1920), B. lloydi (Vaughan, 1920), and B. thomi (Vaughan, 1920), all of which were described from the Danian of the USA (Cannonball Marine Formation), are assumed to represent a more immature stage of the form B. manorensis (Table 4, figs 3a–c ). The species B. kangliaensis from the lower Danian of Greenland closely corresponds to the more immature ontogenetical stage. Type locality of species. Paleocene of the USA (Cannonball Marine Formation). Distribution. Upper Maastrichtian-Paleocene of the USA (Navarro Formation, Texas; Cannonball Marine Formation), Lower Danian of Denmark (Greenland).

Bathycyathus piveteaui (Alloiteau & Tissier, 1958) Text-Fig. 4 (v)*1958 (v)1969 (v)1969 (v)1969

Kionotrochus piveteaui nov. sp.: Alloiteau & Tissier, p. 275, pl. 1, figs 4–4b’. Platycyathus eburnensis n. sp.: Barta-Calmus, p. 822, pl. 1, figs 1–2a. Frescocyathus tessieri n. sp.: Barta-Calmus, p. 823, pl. 1, fig. 3, text-figs 1a–c. Frescocyathus nagagreboensis n. sp.: Barta-Calmus, p. 823, pl. 1, figs 4, 5, 10, 11.

Dimensions. d (max.)=6.4–18.7 mm; d (min.)=5.5–19 mm; d (min)/d (max)=0.61–0.96; s (adult)=96+s6. Description. Variably conical-trubinate, slightly elliptical to compressed in outline; costosepta developed in 5 complete cycles in 6 systems in corallites that have a maximum diameter of around 13 mm and larger; columella formed by a small number of twisted segments.

Text-Fig. 4 Bathycyathus piveteaui (Alloiteau & Tissier, 1958), holotype based on the illustration in Alloiteau & Tissier (1958), Danian of Belgium, scale bar: 4 mm; A, cross view of corallum; B, upper surface longitudinal view.


BARON-SZABO

Remarks. The merging of the above taxa is based on the idea that they represent different ontogenetical stages of the same species, according to which the forms B. eburnensis, B. tessieri, and B. nagagreboensis are more adult stages (corallite diameters ranging between d (min) of 10–18 and d (max) of 12 and 18 mm, 96 septa) and the species B. piveteaui represents a juvenile image (corallite diameter of 5.5 x 6.4 mm and 48 septa). Type locality of species. Danian of Belgium. Distribution. Paleocene of Ivory Coast, Danian of Belgium.

Genus Trochocyathus Milne Edwards & Haime, 1848 (=Protrochocyathus Alloiteau, 1958, Type species. Protrochocyathus madagascariensis Alloiteau, 1958, Albian of Madagascar); (=Paratrochocyathus Alloiteau, 1958, Type species. Paratrochocyathus collignoni Alloiteau, 1958, Albian of Madagascar); (=Cyrtocyathus Alloiteau, 1958, Type species. Cyrtocyathus collignoni Alloiteau, 1958, Maastrichtian of Madagascar); (=Elasmosmilia M. Beauvais, 1961, Type species. Elasmosmilia padernensis M. Beauvais, 1961, Upper Santonian of France); (=Tethocyathus Kühn, 1933, Type species. Thecocyathus microphyllus Reuss, 1871, Tortonian of Austria). Type species. Turbinolia mitrata Goldfuss, 1826, Upper Cretaceous of Germany (Aachen) (see Milne Edwards & Haime 1848c).

Diagnosis. Solitary, variably conical, often turbinate to ceratoid, or discoidal, fixed or free. Costosepta compact, finely granulated laterally. Pali or paliform lobes in 2 crowns opposite all but last cycle. Columella fascicular or spongy. Wall septothecal. Endothecal dissepiments vesicular. Epithecal wall absent or present. Remarks. The genus Trochocyathus has long been considered as lacking an epithecal wall. Re-examination of the type material of this genus, which consists of two syntypes, showed that, in general, an epithecal wall is not present (in the syntype figured in Baron-Szabo 2002, pl. 118, fig. 1, no epithecal wall seems to be present), but in the lower part of one of the two syntypes (the one figured in Baron-Szabo 2002, pl. 118, fig. 2) short strips of incomplete epithecal rings were observed. These short structures can be found only in a very few places, but they are wall developments sitting on top of the costae, which excludes them from being parts of the septothecal developments in this genus. Therefore, the generic diagnosis of Trochocyathus was emended here in adding the statement “Epithecal wall absent or present” to the generic diagnosis. The genus Tethocyathus Kühn, 1933, had been distinguished from Trochocyathus by the presence of both a narrow edge zone and an extensive epitheca (Wells 1956). Because the development of an epithecal wall is dependent on the presence of a narrow edge zone (pers. comm. S. Cairns 2008), it first of all means that the two genera are actually only distinguished by a single characteristic (=presence of a narrow edge zone). In addition, while it seems questionable whether the feature of a wide or narrow edge zone could be used to separate genera (pers. comm. S. Cairns 2008), this distinguishing characteristic (= “epithecal wall absent in Trochocyathus but present in Tethocyathus”) does not exist because the presence of an epithecal wall in Trochocyathus was observed (see remarks above). Subgenus. Platycyathus Fromentel, 1862 (Type species. Trochocyathus terquemi Milne Edwards & Haime, 1857, Lower Santonian of France): like Trochocyathus but corallum discoidal, free.



55

Trochocyathus speciosus (Gabb & Horn, 1860) Pl. 4, figs 5a–c *1860 1902 v1902 v1933 parsv1933 v1938 1941a 1955 v1958 1958 ?1958 v1967 1970 1970 1970 (v)1972 1977 (v)1987 (v)1987 2000 2000 2002 2002

Platytrochus speciosus n. sp.: Gabb & Horn, p. 399, pl. 69, figs 15–17. Trochocyathus speciosus (Gabb and Horn): Vaughan, p. 207–209, figs 1–2a. Trochocyathus epicharis n. sp.: Wanner, p. 99, pl. 14, figs 5–7. Trochocyathus mississippiensis n. sp.: Wells, p. 134 (216), pl. 14, figs 22–23. Trochocyathus gardnerae n. sp.: Wells, p. 135 (217), pl. 14, figs 8–10, non p. 135, paratype no. 4, non specimen no. 3 on page 136. Trochocyathus kroisbachensis n. sp.: Traub, p. 36, pl. 1, figs 2a, b. Trochocyathus sp. cf. T. mississippiensis Wells: Wells, p. 295 (13). Deltocyathus staszici n. sp.: Rozkowska, p. 243, pl. 1, fig. 6, pl. 2, fig. 5, text-figs 1–3. Trochocyathus lemniscatus n. sp.: Squires, p. 48, pl. 10, figs 11–15. Paratrochocyathus epicharis Wanner 1902 sp.: Alloiteau, p. 190. Paratrochocyathus Jasmundi Wanner 1902: Alloiteau, p. 190. Trochocyathus(Protrochocyathus) kroisbachensis Traub: Kühn & Traub, p. 12, pl. 1, figs 5, 6, pl. 2, figs 4, 5. Paratrochocyathus epicharis (Wanner): Hassan & Salama, p. 84, pl. 1, figs 5a, b. Paratrochocyathus lemniscatus (Squires): Hassan & Salama, p. 85, pl. 1, figs 6a–c. Paratrochocyathus deniseptatus sp. nov.: Hassan & Salama, p. 85, pl. 2, figs 1a–g. Trochocyathus johannesi sp. n.: Floris, p. 54, pl. 3, figs 4A–5. Trochocyathus gardnerae Wells, 1933: Wolleben, p. 396, pl. 3, fig. 1. Paratrochocyathus epicharis (Wanner 1902): Kuzmicheva, p. 124, pl. 20, figs 7a, b. Tethocyathus kopetdagensis Kuzmicheva, sp. nov.: Kuzmicheva. 124, pl. 20, figs 8a-9b, ?pl. 21, figs 1, 2. Paratrochocyathus epicharis (Wanner 1902): Löser, p. 60. Trochocyathus, gardnerae Wells 1933: Löser, p. 82. Trochocyathus epicharis Wanner, 1902: Baron-Szabo, p. 158. Trochocyathus gardnerae Wells, 1933: Baron-Szabo, p. 158.

Dimensions. d (max)=8–16 mm; d (min)=6–10 mm; d(min)/d(max)=0.5–1; s (in juvenile corallites of 4–6 mm) up to around 40; s (adult)=48+s5; h=3–20 mm. Description. Subturbinate to variably conical; corallites circular in outline; costosepta arranged in 4 complete cycles in 6 systems; S 1 and S2 nearly equal, remaining septa alternate in length and thickness; paliform structures elongate. Remarks. Vaughan (1902) transferred the species Platytrochus speciosus to the genus Trochocyathus. In his re-examination of the type material, Vaughan reported that this taxon was characterized by slightly elliptical, variably conical corallites with diameters ranging between 11.5 and 13.5 mm (greater diameter) and 10–11 mm (lesser diameter) and 48 septa arranged in 6 systems. The same year, Wanner (1902) published the description of the species Trochocyathus epicharis from the Maastrichtian-Danian of Egypt which, according to his description, was characterized by short, turbinate-subpatellate (he called it “umbrella-like”) coralla of 7–12 mm in diameter and 48 septa arranged in 6 systems. Re-examination of recently re-discovered type material of the Wanner collection at the Rijksmuseum Stockholm revealed however, that 48 septa were only present in the largest corallites (=10–12 mm in diameter), and that trochoid-ceratoid coralla were present in addition to the short turbinate type. Moreover, in corallites that were smaller than 10 mm in diameter, the number of septa was 40 or less. Therefore, T. epicharis is considered a junior synonym of T. speciosus. A very similar situation was found in Trochocyathus lemniscatus Squires from the Middle Eocene of New Zealand. Squires (1958, p. 48) stated that the number of “usually 48 septa” was present in corallites of 10–16 mm in


BARON-SZABO

diameter. Re-examination of the type material showed that 48 septa were present in corallites that were around 12 mm in diameter. In corallites of around 15 or 16 mm in diameter, the number of septa was about 60, which corresponds to a septal development that would be expected in a slightly later ontogenetical stage. Therefore, Trochocyathus lemniscatus is considered a junior synonym of Trochocyathus speciosus. In the original description, Traub (1938, p. 36) stated that his new species Trochocyathus kroisbachensis had the corallite diamter of 13 x 17 mm and 48 septa. Later, in Kühn & Traub (1967, p. 12) this statement was corrected in that the holotype was reported to be only 7.5 x 9.5 mm (and the paratype 9 x 11 mm). In addition, re-examination of the type specimen by the author of the current work revealed that the number of septa in the holotype was slightly less than 48 (around 44), and therefore very closely agreed with the dimensions of T. speciosus. The species Trochocyathus johannesi Floris is practically identical to the species T. epicharis, a statement which was even made by Floris (1972, p. 56) himself. According to Floris, the only difference from T. epicharis (and also Trochocyathus mississippiensis Wells, 1941a, see synonmy list above) is in the bowl-shaped corallum of the latter. As already noted above, the type collection of Trochocyathus epicharis consists of specimens that have various corallum types, which is also one of the characteristics of T. speciosus. Because the species Deltocyathus staszici Rozkowska from the Upper Danian of Poland seems to be lacking pali before the last cycles, it differs from the genus Deltocyathus, but agrees with the genera Trochocyathus and Paracyathus Milne Edwards & Haime. In the latter genus the pali are often indistinguishable from the columella, whereas in Deltocyathus staszici they can be well distinguished; therefore, the latter species seems to be more closely related to Trochocyathus. With a skeletal structure of 36 septa developed in 6 systems in a corallite 4.2 mm in diameter, it closely corresponds to the juvenile stages of the forms listed in the synonymy of T. speciosus. The species Tethocyathus kopetdagensis Kuzmicheva from the Upper Danian of Turkmenistan is characterized by 48 septa in corallites 10–12 mm in diameter. Because the generic separation of Tethocyathus from Trochocyathus does not seem to be valid (see discussion regarding the genus Tethocyathus in Remarks above), the species is considered to be identical to T. speciosus. Based on measurements carried out by the author, it can be stated that all of the above listed forms are characterized by a remarkably large number of septa, ranging between 36 and 40, in corallites that are between 4 and 6 mm in diameter. Type locality of species. Paleocene (Midwayan) of the USA (Hardeman County, Tennessee). Distribution. Campanian of Cuba, Maastrichtian-Danian of Egypt, Santonian-Maastrichtian (Mississippi, Texas, Navarro Formation) and Paleocene (Hardeman County, Tennessee) of the USA, Maastrichtian of Mexico (Parras Basin), Maastrichtian of Tunisia (see Alloiteau, 1958, p. 190), ?Upper Maastrichtian of Madagascar, Paleocene of Austria, Danian of Denmark (Greenland), Upper Danian of Poland and Turkmenistan, Middle Eocene of New Zealand. New Material. Danian of Egypt, NRM-PZ, sample nos.: Cn 47522 and 47525–47531 (Babel Jasmund).

Trochocyathus mitratus (Goldfuss, 1826) Pl. 5, figs 1–4b v*1826 1848d 1857

Turbinolia mitrata nobis: Goldfuss, pl. 15, figs 5a–c. Trochocyathus Konincki: Milne Edwards & Haime, p. vol. 2, p. 305. Trochocyathus Konincki: Milne Edwards. vol. 2, p. 30.



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1880 v1900 1901 v1903a 1914 1914 ?1916 v1920 v1920 1925 1925 v1930a pars v1933 v1934 (v)1938 ?1951b v1958 1967 (v)1975 ?(v)1975 (v)1975 (v)1987 1993 1994 2000 2000 2000 2000 2000 v2000 2001 2002 2002 2002 2002 v2002 v2002 2002

Trochocyathus corbicula, Duncan: Duncan, p. 27, pl. 7, figs 12–14. Trochocyathus clarkeanus (Vaughan): Vaughan, p. 100, pl. 7, figs 20–23 (paratypes studied). Trochocyathus clarkeanus Vaughan: Clarke & Martin, p. 81. Trochocyathus microphyes: Felix, p. 354, pl. 18, figs 9, 10 (topotypes studied). Trochocyathus Konincki E. H. 1848: Felix, pars 7, p. 208. Trochocyathus microphyes Felix 1903: Felix, pars 7, p. 209. Trochocyathus imperialis, n. sp.: Nomland, p. 63, pl. 4, figs 11–12. Trochocyathus dakotaensis Vaughan, n. sp.: Vaughan, p. 61, pl. 10, figs 1–1b. Trochocyathus ? neumani Vaughan, n. sp.: Vaughan, p. 62, pl. 10, figs 2–2c. Trochocyathus Clarkeanus Vaughan 1895: Felix, pars 28, p. 196. Trochocyathus imperialis Nomland 1916: Felix, pars 28, p. 199. Trochocyathus micraphye Felix: Oppenheim, p. 548 (topotypes studied). Trochocyathus gardnerae n. sp.: Wells, p. 135ff. (217ff.), non pl. 14, non figs 8–10. Trochocyathus Matleyi, new species: Wells, p. 74, pl. 2, figs 5–6. Trochocyathus haunsbergensis n. sp.: Traub, p. 36, pl. 1, figs 2a, b. Asterosmilia sakalavensis nov. sp. (Alloiteau): Alloiteau, p. 82, pl. 13, figs 7–8. Protrochocyathus septempartitus nov. sp.: Alloiteau, 189, pl. 31, figs 9–11. Trochocyathus(Protrochocyathus) haunsbergensis Traub: Kühn & Traub, p. 12, pl. 1, figs 5, 6, pl. 2, Paracyathus kangpaensis (sp. nov.): Wu, p. 109, pl. 7, figs 6–9, pl. 9, figs 7–8, pl. 10, fig. 2. Cyathoceras ellipticus (sp. nov.): Wu, p. 109, pl. 9, fig. 9. Notocyathus morozi Kuzmicheva, sp. nov.: Kuzmicheva, p. 22, pl. 2, figs 5a–6. Notocyathus morozi Kuzmicheva, sp. nov.: Kuzmicheva, p. 139, pl. 24, figs 9a–b. Paracyathus kangpaensis Wu: Liao & Xia, p. 209, fig. 3.8. Paracyathus kangpaensis Wu: Liao & Xia, p. 186, pl. 55, figs 4–9. Paracyathus kangpaensis, Wu 1975: Löser, p. 58. Protrochocyathus septempartitus, Alloiteau 1958: Löser, p. 68. Sakalavicyathus microphyes (Felix 1903): Löser, p. 72. Trochocyathus konincki, Milne Edwards & Haime 1848: Löser, p. 82. Trochocyathus matleyi, Wells 1934: Löser, p. 82. Trochocyathus microphyes Felix, 1903: Baron-Szabo, p. 126, pl. 10, figs 5, 7, pl. 12, fig. 3. Paracyathus kangpaensis Wu 1975: Löser & Liao, p. 666. Trochocyathus kangpaensis (Wu, 1975): Baron-Szabo, p. 158. Trochocyathus konincki Milne Edwards & Haime, 1848: Baron-Szabo, p. 158. Trochocyathus matleyi Wells, 1934: Baron-Szabo, p. 158. Trochocyathus mitratus (Goldfuss, 1826): Löser, p. 688 (older synonyms cited therein). Trochocyathus mitratus (Goldfuss, 1826): Baron-Szabo, p. 158, pl. 118, figs 1, 2. Trochocyathus microphyes Felix, 1903: Baron-Szabo, p. 158, pl. 118, figs 3, 6, 9–10. Trochocyathus septempartitus (Alloiteau, 1958): Baron-Szabo, p. 158.

Dimensions. d (max)=9–41 mm; d (min)=8–30 mm; s=50, up to around 200 (12s1+12s2+24s3+48s4+96s5+s6, in adult stages); h=25–65 mm; d (min)/d (max)=0.5–0.9; d (juvenile stages)=3.5–8 mm; s (juvenile stages)=up to 48 (6s1+6s2+12s3+24s4). Description. Turbinate, trochoid, or ceratoid corallum; corallite slightly elliptical or compressed in adult stages, circular to slightly elliptical in juvenile stages; costosepta compact, thin or moderate, nearly equal in juvenile stages, arranged in 5 complete with the beginning of a sixth cycle in 12 systems in adult stages; septal development of 4 complete cycles in 6 systems present in juvenile corallites of around 5–6 mm in diameter; pali very thick and elongate in juvenile stages, becoming thinner and more obsolete in later ontogenetical stages; columella very variably shaped: spongy-papillose, or fused papillae may form elongated segments, or very weakly developed.


BARON-SZABO

Remarks. The 2 syntypes of Trochocyathus mitratus (Goldfuss) have corallites that are 10 and 11.5 mm in diameter and nearly 60 septa in their latest ontogenetical stage. In their juvenile stages there are 48 septa in 4–5 mm diameter. These skeletal dimensions are were found in juvenile stages of the specimens from the UAE/ Oman border region (Baron-Szabo 2000), are chararcteristic of the type material of Trochocyathus matleyi Wells, and Trochocyathus konincki Milne Edwards & Haime, as well as in Trochocyathus microphyes Felix, which all have 48 septa in 4 cycles in a calicular diameter of around 4–5 mm. In addition, it is suggested that the larger specimens of e.g. the UAE/ Oman represent ontogenetically older stages of T. mitratus. A very similar kind of ontogenetical development seems to be present in the form Trochocyathus kangpaensis Wu from the Campanian-Maastrichtian of Tibet (see Wu 1975, pl. 7, figs 6–9, pl. 9, figs 7–8, pl. 10, fig. 2; Liao & Xia, 1994, pl. 55, figs 4–9) and in the species Trochocyathus corbicula, Duncan from the Paleocene of Pakistan. Therefore, all these species are considered junior synonyms of T mitratus. Re-examination of the type specimens described as Trochocyathus septempartitus (Alloiteau, 1958), revealed that they represent slightly more juvenile ontogenetical stages of Trochocyathus mitratus with 56 septa and a septal arrangement in 9 or 10 irregular systems in a corallite of 6 x 8 mm in diameter (in contrast to Alloiteau’ description in which he stated the presence of a septal development in 14 systems). The transformation during ontogenetical development from septal orders in 6 systems in juvenile stages to systems higher than 6 in adult stages has recently been discovered in solitary taxa like e.g. Aulosmilia aspera (Sowerby) (Baron-Szabo 2003). The species T. septempartitus matches the development of an ontogenetical stage intermediate between a juvenile corallite with 6 systems and the development of 12 systems in adult stages. Some of the specimens assigned to Trochocyathus gardnerae Wells from the Maastrichtian of the USA (Texas, Navarro Formation) by Wells (1933, p. 135, paratype no. 4 and specimen no. 3 on p. 136) closely correspond to juvenile stages of the species T. mitratus in that they have nearly circular corallites with diameters ranging between 5–7 mm, as well as 48 septa arranged in 12 systems. Type locality of species. Campanian of Germany (Aachen, Zevenwegen-Schichten). Distribution. Santonian of Austria (Gosau Group), Campanian of Germany, Campanian-Maastrichtian of Tibet (Gamba County, Houshan) and Jamaica (Blue Mountain Peak), Middle-Upper Maastrichtian of the UAE/Oman border region, Upper Maastrichtian of Madagascar, Maastrichtian (Texas, Navarro Formation)Danian (Cannonball marine Member) of the USA, Paleocene of Austria and Belgium (Obourg, Mons), Upper Danian of Ukraine, Eocene of the USA (Maryland [Aquia and Nanjemoy Formations], ?California [Lower Tejon Group]).

Trochocyathus lakii Duncan, 1880 Text-Fig. 5 *1880 v1900 1925 1955 1955 v1958 v1958 (v)1958 (v)1977 2000

Trochocyathus Lakii, Duncan: Duncan, p. 18, pl. 1, figs 11–13. Trochocyathus hyatti sp. nov: Vaughan, p. 93, pl. 6, figs 19–21, pl. 7, fig. 1. Trochocyathus hyatti Vaughan 1900: Felix, pars 28, p. 199. Trochocyathus desioi n. sp.: Rossi Ronchetti, p. 106, pl. 11, figs 2–3. Trochocyathus libycus n. sp.: Rossi Ronchetti, p. 110, pl. 11, figs 4a–b. Cyrtocyathus collignoni nov. sp.: Alloiteau, p. 193, pl. 19, fig. 6, pl. 26, figs 9–15, pl. 28, fig. 3. Cyrtocyathus collignoni nov. sp. var. mixtus: Alloiteau, p. 194, pl. 19, figs 6a–b. Asterosmilia aliquantula n. sp.: Squires, p. 61, pl. 11, figs 4–8. Trochocyathus hyatti Vaughan: Toulmin, p. 146, pl. 1, figs 9–11. Cyrtocyathus collignoni, Alloiteau 1958: Löser, p. 27.



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Cyrtocyathus collignoni var. mixtus, Alloiteau 1958: Löser, p. 27. ?Trochocyathus collignoni (Alloiteau, 1958): Baron-Szabo, p. 158 (pro Cyrtocyathus Alloiteau, 1958).


BARON-SZABO

PLATE 5: Fig. 1 Trochocyathus mitratus (Goldfuss, 1826), syntype, IPB, Goldfuss coll., no. 179, Campanian of Germany, scale bar: 4 mm. 1a, cross view; 1b, longitudinal view. Fig. 2 Trochocyathus mitratus (Goldfuss, 1826) (holotype of Trochocyathus dakotaensis Vaughan, 1920), NMNH, Vaughan coll., I32374, Danian of the USA, scale bar: 4 mm. 2a, cross view; 2b, longitudinal view. Fig. 3 Trochocyathus mitratus (Goldfuss, 1826), syntype, IPB, Goldfuss coll., no. 179, Campanian of Germany. 3a, cross view, scale bar: 4 mm; 3b, longitudinal view, scale bar: 1.5 mm. Fig. 4 Trochocyathus mitratus (Goldfuss, 1826), BMNH, AZ 792, Middle-Upper Maastrichtian of the UAE/Oman border region. 4a, cross view, thin section, juvenile stage, scale bar: 2.5 mm; 4b, longitudinal view, thin section, adult stage, scale bar: 3.5 mm. Fig. 5 Trochocyathus speciosus (Gabb & Horn, 1860) (holotype of Trochocyathus gardnerae Wells, 1933), NMNH, Wells coll., I75143, Maastrichtian of the USA. 5a, longitudinal view, scale bar: 3 mm; 5b, cross view, scale bar: 4 mm. Fig. 6 Stylocyathus besairiei Alloiteau, 1958, holotype, MNHN, Mo5185, Upper Maastrichtian of Madagascar. 6a, cross view, scale bar: 2 mm; 6b, longitudinal view, scale bar: 2.5 mm. Fig. 7 Deltocyathus whitei Durham, 1943, holotype, CAS, 69418, Upper Paleocene of the USA, scale bar: 5 mm. 7a, upper surface, cross view; 7b, base of corallum. Fig. 8 Stephanosmilia madagascariensis Alloiteau, 1958, holotype, MNHN, Mo5176, Upper Campanian-Maastrichtian of Madagascar. 8a, longitudinal view, scale bar: 2 mm; 8b, cross view, upper surface, scale bar: 1.8 mm. Fig. 9 Deltocyathus complanatus Squires, 1958 (holotype of Discosmilia analavelonensis Alloiteau, 1958), MNHN, Mo5060, Upper Campanian-Maastrichtian of Madagascar, cross view, upper surface, scale bar: 5.5 mm. Fig. 10 Parasmilia centralis (Mantell, 1822), (topotype of Parasmilia lindstroemi Hennig, 1899), topotype, MB, Schlotheim coll., K.3631.1, Upper Danian of Denmark (Fakse), scale bar: 3 mm. 10a, longitudinal view; 10b, cross view. Fig. 11 Deltocyathus complanatus Squires, 1958 (paratype of Discosmilia analavelonensis Alloiteau, 1958), MNHN, in same box with holotype Mo5176 Upper Campanian-Maastrichtian of Madagascar, longitudinal view, upper surface, scale bar: 3.5 mm.

Dimensions. d (min)=6–13.5 mm; d (max)=7–21 mm; d (min)/d (max)=0.59–0.96; s=30–48+s5; h=8–23 mm. Description. Solitary, trochoid, nearly circular in younger ontogenetical stages, becoming more elliptical in later stages; costosepta developed in 4 complete or nearly complete in a corallite of around 10 mm in diameter; septa of first 3 cycles nearly equal in thickness, alternating in length; spongy columella well-developed. Remarks. In the original description of Trochocyathus desioi Rossi Ronchetti (1955, p. 106) stated that the septa formed sets of 13. However, the illustrations of the type material (Rossi Ronchetti 1955, pl. 11, figs 2–3) show a septal development in sets of 12 nearly equal septa, which are irregularly influenced by the development of a beginning fifth septal cycle. The specimens of the species T. collignoni are interpreted to be juvenile stages (the specimens which have corallite diameters ranging between 6–7 mm and 30–34 septa) and early adult stages (the specimens which are characterized by corallite diameters of around 8 mm and around 40 septa). The type material of the species Trochocyathus hyatti from the Danian of the USA (Alabama) very closely corresponds to Trochocyathus lakii in having four incomplete septal cycles in 6 systems (around 44 septa) in a syntype 10 mm in diameter (the diameter the type of Trochocyathus lakii is 10 mm and has around 40 septa) and thin, delicately granular septa. Type locality of species. Paleocene of Pakistan. Distribution. Maastrichtian of Libya (Sofeggin), Upper Maastrichtian of Madagascar, Paleocene of Pakistan and the USA (Alabama), Eocene of New Zealand.



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Text-Fig. 5 Trochocyathus lakii Duncan, 1880. A, B, D, F, USNM M158134 (syntypes of Trochocyathus hyatti Vaughan, 1900), Paleocene of the USA (Alabama); A, cross view of corallum (larger syntype); B, upper surface longitudinal view (larger syntype); D, cross view of corallum (smaller syntype); F, upper surface longitudinal view (smaller syntype); C, E, G, type specimens of Trochocyathus lakii Duncan, 1880 based on the illustrations as provided by Duncan (1880), Paleocene of Pakistan; C, cross view of corallum showing part of septal apparatus; E, longitudinal view of part of corallum; G, longitudinal view of whole corallum. A, B, D, F: scale bar 5 mm; C, G: scale bar 4 mm; E: scale bar 2 mm.

Genus Paracyathus Milne Edwards & Haime, 1848 Type species. Paracyathus procumbens Milne Edwards & Haime, 1848c, Middle Eocene of France.

Diagnosis. Solitary, turbinate, fixed or free. Septotheca costate. Paliform lobes often bi- or trilobated, opposite all but last cycle. Columella papillose, often indistinguishable from the inner paliform lobes. Paracyathus altus Gregory, 1930 Text-Fig. 6 *1930 1993

Paracyathus altus, sp. nov: Gregory, p. 100, pl. 11, figs 8–9b. Paracyathus cf. altus (Gregory, 1930): Carbone et al., p. 227.

Dimensions. d (max., adult)=12 mm; d (min., adult)=10 mm; s (adult)=48 (6s1+6s2+12s3+24s4); h=14–18 mm. Description. Corallum elliptical in outline; septal arrangement in 4 complete cycles in 6 systems in late adult stages, a few of which unite in triplets; septa of an incomplete fifth cycles can be present; there are 36 septa in a corallite diameter of 7 mm; S1–S3 nearly equal; pali in three crowns, a few are bifid.


BARON-SZABO

Remarks. Because no Paleocene material of this taxon was available, an image of the Eocene as given in Gregory (1900) is presented here. Type locality of species. Eocene of India (Upper Ranikot Series). Distribution. Upper Paleocene of Somalia, Eocene of India.

Text-Fig. 6 Paracyathus altus Gregory, 1930, based on the illustration of holotype in Gregory (1930), Eocene of India, cross view, polished surface, scale bar: 4 mm.

Genus Deltocyathus Milne Edwards & Haime, 1848 (=Discosmilia Alloiteau, 1958, Type species. Discosmilia analavelonensis Alloiteau, 1958, Upper Campanian-Maastrichtian of Madagascar). Type species. Turbinolia italicus Michelotti, 1838, Tertiary of Italy (see Milne Edwards & Haime, 1848c).

Diagnosis. Solitary, discoid to patellate. Costosepta laminar, exsert, compact. Septal margins smooth or nearly smooth. Pali or paliform lobes opposite all cycles, forming elongate segments. Columella papillose. Endothecal dissepiments few in number. Wall septothecal. Remarks. Alloiteau (1958, p. 196ff.) stated that the presence of a parathecal-synapticulothecal wall was one of the generic characteristics of the genus Discosmilia Alloiteau, 1958. Therefore, based on the original description, Baron-Szabo (2002, p. 172) concluded that there was a close resemblance to the genus Sakalavicyathus. However, investigation of the type material of Discosmilia revealed the absence of a parathecal-synapticulothecal wall but the existence of a rather (parathecal-) septothecal wall. Moreover, the septal arrangement seems to correspond well with that of Deltocyathus. Therefore, the assignment to the latter genus is suggested. Deltocyathus complanatus Squires, 1958 Pl. 5, figs 9, 11 *1958 v1958 non1970 non1996 1997 2000 2000 2002 2002

Deltocyathus ? complanatus n. sp.: Squires, p. 50, pl. 10, figs 21–22. Discosmilia analavelonensis nov. sp.: Alloiteau, p. 198, pl. 33, figs 6–12, text-figs 34–35. Discosmilia cf. analavelonensis Alloiteau: Hassan & Salama, p. 87, pl. 1, figs 8a–b. Discosmilia cf. analavelonensis Alloiteau, 1958: Schuster (listed in appendix). Deltocyathus ? complanatus: Stilwell, p. 104, tab. 1. Deltocyathus complanatus, Squires 1958: Löser, p. 27. Discosmilia analavelonensis, Alloiteau 1958: Löser, p. 32. Deltocyathus complanatus Squires, 1958: Baron-Szabo, 159. Sakalavicyathus analavelonensis (Alloiteau, 1958): Baron-Szabo, p. 172.



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Dimensions. d=4–8 mm; s=40–48+s5; h=1.3–2.5 mm. Description. Discoid-subpatellate, circular in outline; base flat; costosepta arranged in 3 complete cycles with the beginning of a fourth cycle in 6 systems in corallites that are around 4 mm in diameter; costosepta arranged in 4 complete cycles in 6 systems in corallites having a diameter of around 7 mm, nearly complete in corallites that are around 6 mm in diameter; S1 and S2 equal, straight, reaching the axial region; lower cycle septa fuse with higher cycle ones in the centre of corallum. Type locality of species. Campanian-Maastrichtian of New Zealand. Distribution. Campanian-Maastrichtian of New Zealand, Upper Campanian-Maastrichtian of Madagascar.

Deltocyathus cupuliformis (Alloiteau, 1951) Text-Fig. 7 (v)*1951b 1987 2000 2002

Trochocyathus cupuliformis nov. sp. (Alloiteau): Alloiteau, p. 82, pl. 13, figs 5–6. Deltocyathus (Levipalifer) arctous Kuzmicheva, sp. nov.: Kuzmicheva, p. 123, pl. 20, figs 6a–b. Levipalifer arctous, Kuzmicheva 1987: Löser, p. 48. Deltocyathus arctous (Kuzmicheva, 1987): Baron-Szabo, p. 159.

Dimensions. d=up to 35 mm; s=48+s5; h=up to 20 mm. Description. Subdiscoid-patellate, circular to subcircular in outline; costosepta straight, arranged in 4 complete cycles in 6 systems in a corallite ranging between 25–28 mm, regularly alternating in length and thickness; S1 reach the axial region; columella rather weakly developed. Type locality of species. Upper Maastrichtian of Madagascar. Distribution. Maastrichtian of Russia, Upper Maastrichtian of Madagascar.

Deltocyathus whitei Durham, 1943 Pl. 5, figs 7a, b v*1943 (v)1969

Deltocyathus whitei Durham, n. sp.: Durham, p. 200, pl. 32, figs 13, 16, text-fig. 1. Platycyathus ghiroboensis n. sp.: Barta-Calmus, p. 820, pl. 1, figs 6–7.

Dimensions. d (max)=11–14 mm; s=84–96; h=3.8–6.8 mm. Description. Discoid, circular in outline; costosepta thin, wavy, arranged in 4 complete cycles in 6 systems with a small number of septa of a beginning fifth cycle, regularly alternating in length and thickness; S1–S3 reach the axial region; character of columella uncertain. Remarks. Barta-Calmus (1969) described the species Platycyathus ghiroboensis from the Paleocene of Ivory Coast. In having only a small number of endothecal dissepiments (absence of endotheca according to Barta-Calmus [1969] but type specimens seem to show a small number of endothecal dissepiments in the illustrations on Plate 1, Figs 6a and 7), a corallum that is discoidal in shape, and pali that seem to be positioned before all septal cycles, the specimens rather correspond to the genus Deltocyathus, and closely resemble the species D. whitei. Type locality of species. Upper Paleocene of the USA (California, Panoche Quadrangle). Distribution. Paleocene of Ivory Coast, Upper Paleocene of the USA.


BARON-SZABO

Text-Fig. 7 Deltocyathus cupuliformis (Alloiteau, 1951b), based on the illustration of holotype in Alloiteau (1951b), Upper Maastrichtian of Madagascar, cross view of lower part of corallum, scale bar: 10 mm.

Genus Stylocyathus d’Orbigny, 1850 (=Stylotrochus de Fromentel, 1862, Type species. Stylotrochus arcuatus de Fromentel, 1862, Senonian of France [Var]). Type species. Stylocyathus dentalinus d’Orbigny, 1850, Cenomanian of France.

Diagnosis. Solitary corallum, turbinate or ceratoid, fixed or free. Costosepta compact. Pali in one irregular crown opposite first 2 cycles. Columella trabecular, appearing substyliform or formed by 2 or 3 twisted laths. Wall septothecal. Endothecal dissepiments rare. Affinities. The genus Stylocyathus d’Orbigny shows close affinities to the genus Stephanosmilia de Fromentel but differs from the latter in having both a different corallum shape and develoment of axial structures (trochoid to subcylindrical, fixed, and pali in two crowns in front of S1 and S2 in Stephanosmilia).

Stylocyathus besairiei Alloiteau, 1958 Pl. 5, figs 6a, b v*1958 2000

Stylocyathus Besairiei nov. sp.:Alloiteau, p. 188, pl. 32, fig. 11. Stylocyathus besairiei, Alloiteau 1958: Löser, p. 76.



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Stylocyathus besairiei Alloiteau, 1958: Baron-Szabo, p. 160.

Dimensions. d=4.5 x 5.5 mm; s=28 (7s1+7s2+14s3); h=6 mm. Description. Fragment of a solitary, ?turbinate, corallum; costosepta compact, developed in 3 cycles in 7 systems; columella fasciculate; elongate pali present before S1 and S2. Type locality of species. Upper Maastrichtian of Madagascar. Distribution. Upper Maastrichtian of Madagascar.

Genus Stephanosmilia de Fromentel, 1862 Type species. Stephanosmilia perlata de Fromentel, 1862, Aptian of France.

Diagnosis. Solitary, trochoid to subcylindrical, fixed. Septa compact, bilaterally to radially arranged, with coarse granules laterally. Columella sublamellar to substyliform. Pali in two crowns in front of S1 and S2. Endothecal dissepiments sparse. Wall septothecal.

Stephanosmilia madagascariensis Alloiteau, 1958 Pl. 5, figs 8a, b 1952b v*1958 2000 2002

?Stylocyathus sp.: Alloiteau, p. 9, pl. 1, figs 9–10. Stephanosmilia madagascariensis nov. sp.: Alloiteau, p. 191, pl. 37, figs 7–8. Stephanosmilia madagascariensis Alloiteau 1958: Löser, p. 74. Stephanosmilia madagascariensis Alloiteau, 1958: Baron-Szabo, p. 161.

Dimensions. d=2–3 mm; s=24–48. Description. Ceratoid corallum; costosepta richly granulated, arranged in 3–4 cycles in 6 systems; S4 very short, complete or incomplete; columella trabecular, formed by a twisted, sublamellar segment. Type locality of species. Upper Campanian-Maastrichtian of Madagascar. Distribution. Upper Campanian-Maastrichtian of Madagascar, Maastrichtian of Senegal.

Genus Stephanocyathus Seguenza, 1864 Type species. Stephanocyathus elegans Seguenza, 1864, Recent, Mediterranean Sea, off Sicily.

Diagnosis. Solitary, patellate, free. Costosepta compact, thin, finely granulated laterally. Paliform lobes very elongate, present on all cycles, or absent. Columella trabecular, papillose, or trabeculae fused on surface.

Stephanocyathus sp. Text-Fig. 8 1972 1972

Stephanocyathus (Stephanocyathus) sp.: Floris, p. 63, pl. 3, fig. 10. Stephanocyathus (Stephanocyathus) ? sp. no. 1: Floris, p. 64, pl. 3, figs 11A–12B.


BARON-SZABO

1972

Stephanocyathus (Stephanocyathus) ? sp. no. 2: Floris, p. 65, pl. 4, fig. 1.

Text-Fig. 8 Stephanocyathus sp., (?Campanian-) Maastrichtian-Upper Danian of Greenland, as figured in Floris (1972), cross view, upper surface, scale bar: 4 mm.

Description. Floris (1972) described several poorly preserved specimens of solitary, apparently free, subcircular coralla; one set of specimens consists of corallites having diameters of 7–12 mm, number of septa probably ranging between (?) 56–64; a second set of specimens, possibly representing more juvenile stages of the former set, consists of coralla that are 6–9 mm in diameter and have 24–48+ septa, probably 2 crowns of pali before S2 and S3; wall septothecal. Distribution. (?Campanian-) Maastrichtian-Upper Danian of Greenland.

Genus Cyathoceras Moseley, 1881 Type species. Cyathoceras cornu Moseley, 1881, Recent, southern Atlantic.

Diagnosis. Solitary, turbinate or cylindrical, fixed or free. Costosepta laminar, compact. Septal margins smooth or nearly smooth. Columella formed by twisted trabecular segments. Endothecal dissepiments few in number. Wall septothecal.

Cyathoceras embaensis Kuzmicheva, 1987 Text-Fig. 9 *1987

Cyathoceras embaensis Kuzmicheva sp. nov.: Kuzmicheva, p. 118, pl. 19, figs 5a–b.



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2000 2002

Cyathoceras embaensis, Kuzmicheva 1987: Löser, p. 23. Cyathoceras embaensis Kuzmicheva, 1987: Baron-Szabo, p. 162.

Dimensions. d (min)=24–28 mm; d (max)=28–32 mm; d (min)/d (max)=0.86–0.88; s=70–72. Description. Turbinate; slightly elliptical in outline; costosepta arranged in 4 complete cycles in 6 systems; fifth cycle incomplete; septa straight and regularly alternate in length and thickness; columella irregularly trabecular, weakly developed. Type locality of species. Maastrichtian of Kazakhstan (Emby basin). Distribution. Maastrichtian of Kazakhstan.

Text-Fig. 9 Cyathoceras embaensis Kuzmicheva, 1987. A, B, holotype, based on the illustration in Kuzmicheva (1987), Maastrichtian of Kazakhstan; A, cross view, part of corallum, scale bar: 5 mm; B, longitudinal view, part of corallum, oblique, scale bar: 10 mm.

Cyathoceras mangyschlagensis Kuzmicheva, 1987 Text-Fig. 10 *1987 2000 2002

Cyathoceras mangyschlagensis Kuzmicheva sp. nov.: Kuzmicheva, p. 118, pl. 19, figs 6–7b. Cyathoceras mangyschlagensis, Kuzmicheva 1987: Löser, p. 23. Cyathoceras mangyschlagensis Kuzmicheva, 1987: Baron-Szabo, p. 162.

Dimensions. d (min)=22–23 mm; d (max)=25–27 mm; d (min)/d (max)=0.85–0.88; s=90–96. Description. Turbinate; slightly elliptical in outline; costosepta arranged in 5 complete cycles in 6 systems in late adult stage; septa straight or slightly bent and irregularly alternate in length and thickness; columella composed of twisted segments, well-developed. Type locality of species. Maastrichtian of Kazakhstan (Mangyschlak). Distribution. Maastrichtian of Kazakhstan.


BARON-SZABO

Text-Fig. 10 Cyathoceras mangyschlagensis Kuzmicheva, 1987. A, B, holotype, based on the illustration in Kuzmicheva (1987), Maastrichtian of Kazakhstan, scale bar: 10 mm; A, cross view, slightly oblique; B, longitudinal view.

Genus Ceratotrochus Milne Edwards & Haime, 1848 Type species. Turbinolia multiserialis Michelotti, 1838, Tertiary of Italy (see Milne Edwards & Haime, 1848c).

Diagnosis. Solitary, trochoid, fixed or free. Costosepta compact, columella papillose, large. Pali absent. Endothecal dissepiments sparse. Wall septothecal. Ceratotrochus supracretacea Hennig, 1899 Text-Fig. 11 *1899 1927 1955 1987 2000 2002

Ceratotrochus supracretacea, n. sp.: Hennig, p. 21, pl. 2, figs 37–40. Ceratotrochus polonicus n. sp.: Siemiradzki, p. 317, pl. 1, figs 5a–c. Axocyathus punctatus (de From.): Rossi Ronchetti, p. 115, pl. 12, figs 1–2. Ceratotrochus rengarteni Kuzmicheva, sp. nov.: Kuzmicheva, p. 127, pl. 21, figs 6–7. Ceratotrochus polonicus, Siemiradzki 1927: Löser, p. 18. Ceratotrochus polonicus Siemiradzki, 1927: Baron-Szabo, p. 163.

Dimensions. d=9–25 mm; s=36–96; h=up to 28 mm. Description. Trochoid to ceratoid, circular in outline; costosepta thin, developed in 5 cycles in 6 systems in late adult stages; they slightly alternate in thickness, more clearly alternate in length; columella as observed in C. punctatus and C. rengarteni rather small. Remarks. The species listed in the synonymy list above are believed to form an ontogenetical series in which: 1) the forms assigned to the species C. punctatus from the Maastrichtian of Libya and the species C. supracretacea from the Danian of Denmark represent early to intermediate ontogenetical stages (d ranging between 9–13.5 mm, number of septa 36–48 in C. punctatus; d ranging between 12.5 mm (calculated) and 16 SCLERACTINIAN CORALS OF THE K/T-BOUNDARY


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mm, number of septa 48 in C. supracretacea); 2) the species C. rengarteni from the Danian of Turkmenistan, Ukraine, Kazakhstan, and Russia form intermediate stages (d up to18 mm, s=67–75); 3) the species C. polonicus from the Maastrichtian of Ukraine corresponds to late ontogenetical stages (d=25 mm, s=96). However, because the ontogenetical stages could not be studied by sectioning the material, the synonymy listed above is only provisional. Type locality of species. Danian of Denmark. Distribution. Maastrichtian of Libya, Maastrichtian-Danian of Ukraine, Danian of Denmark, Turkmenistan, Kazakhstan, and Russia.

Text-Fig. 11 Ceratotrochus supracretacea Hennig, 1899, based on the illustration of the type material in Hennig (1899), Danian of Denmark. A, upper surface longitudinal view, scale bar: 5 mm; B, upper surface cross view, scale bar: 5 mm; C, longitudinal cut, scale bar: 2.5 mm.

Subfamily Parasmiliinae Vaughan & Wells, 1943 (=Family Parasmiliidae Alloiteau, 1952a p.p.) Diagnosis. Solitary and colonial, ahermatypic. Colony formation by intra- and extracalicinal budding. Coenosteum, epitheca, and stereome rarely developed. Endothecal dissepiments sparse. Columella trabecular or absent.

Genus Parasmilia Milne Edwards & Haime, 1848 Type species. Madrepora centralis Mantell, 1822, Campanian-Maastrichtian of England (Sussex) (see Milne Edwards & Haime, 1848d).

Diagnosis. Solitary, trochoid, fixed. Columella spongy to irregularly trabecular. Costosepta compact. Septal margins smooth or slightly granular. Endothecal dissepiments few in number, developed deep in corallum. Wall septothecal to septoparathecal.

Parasmilia centralis (Mantell, 1822) Pl. 5, figs 10a, b, Pl. 6, figs 1a–4 v1820 *v1822 1829

Hippurites elongatus (conf. Parkins): Schlotheim, p. 354. Madrepora centralis: Mantell, p. 159, pl. 16, figs 15–16. Caryophyllia centralis: Mantell, vol. 3, p. 204.


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1829 v1848d v1848d 1863 1863 1866 1866 1866 v1869 v1899 1899 1904 1909 1914 1914 1914 1914 v1922 1927 1955 1955 1955 1955 non1958 1986 1987 1987 1987 1987 1995 1995 2000 2000 v2002 2002 v2002 2002 2002 2002 2002 2002 (v)2005

Caryophyllia centralis Mantell: Phillips, vol. 1, p. 119, pl. 1, fig. 13. Parasmilia centralis: Milne Edwards & Haime, vol. 4, p. 244. Parasmilia Gravesiana: Milne Edwards & Haime, vol. 4, p. 245. Parasmilia Gravesi: de Fromentel, p. 212, pl. 22, fig. 1. Parasmilia cylindracea: de Fromentel, p. 214, pl. 29, fig. 1. Parasmilia cylindrica M. Edw. u. Haime: Bölsche, p. 465, pl. 8, figs 2, 3. Parasmilia Gravesiana M. Edw. u. Haime: Bölsche, p. 466, pl. 8, fig. 4. Parasmilia laticostata n. sp.: Bölsche, p. 466, pl. 8, fig. 5. Parasmilia granulata, n. sp.: Duncan, p. 13, pl. 6, figs 5–8. Parasmilia lindströmi n. sp.: Hennig, p. 15, pl. 2, figs 18–33 (topotypes studied). Parasmilia scanica n. sp.: Hennig, p. 20, pl. 2, figs 34–36. Parasmilia centralis var. gravesiana: Jukes-Browne, vol. 3, p. 524. Parasmilia centralis (Mantell): Lang, p. 294, text-fig. 40. Parasmilia centralis: Mantell sp. 1822: Felix, pars 7, p. 227. Parasmilia granulata Duncan 1869: Felix, pars 7, p. 229. Parasmilia Gravesi E. H. 1849: Felix, pars 7, p. 229. Parasmilia laticostata Bölsche 1866: Felix, pars 7, p. 229. Parasmilia lindströmi Hennig: Nielsen, p. 28 (topotypes studied). Parasmilia Fittoni M. Edw. & Haime: Siemiradzki, p. 318, pl. 1, fig.2a–c. Parasmilia cf. cylindracea de From.: Rossi Ronchetti, p. 117, pl. 12, figs 3a–b. Parasmilia granulata Duncan: Rossi Ronchetti, p. 118, pl. 12, figs 4a–b. Caryosmilia granosa Wann.: Rossi Ronchetti, p. 119, pl. 12, figs 5a–c. Parasmilia helenae n. sp.: Rozkowska, p. 254, pl. 1, fig. 5, pl. 2, fig. 3, text-figs 9, 10. Parasmilia aff. lindströmi Hennig: Alloiteau & Tissier, p. 245, pl. 2, figs 2a–b’. Parasmilia centralis: Tchéchmédjiéva, p. 58ff. Parasmilia centralis (Mantell, 1822): Kuzmicheva, p. 134, pl. 23, figs 5–7. Parasmilia cylindrica Edwards et Haime, 1850: Kuzmicheva, p. 135, pl. 23, figs 8–10. Parasmilia fittoni Edwards et Haime, 1851: Kuzmicheva, p. 135, pl. 23, fig. 11, pl. 24, fig. 1. Parasmilia helenae Rozkowska, 1955: Kuzmicheva, p. 138, pl. 24, figs 6a, b. Parasmilia centralis (Mantell, 1822): Tchéchmédjiéva, p. 58, pl. 12, figs 7, 8. Parasmilia gravesana Edwards & Haime, 1848: Tchéchmédjiéva, p. 59, pl. 12, figs 9, 10. Parasmilia granulata, Duncan 1869: Löser, p. 60. Parasmilia laticostata, Bölsche 1866: Löser, p. 60. Parasmilia centralis (Mantell, 1822): Baron-Szabo, p. 165, pl. 119, figs 1, 7. Parasmilia cylindrica Milne Edwards & Haime, 1850: Baron-Szabo, p. 165. Parasmilia granulata Duncan, 1869: Baron-Szabo, p. 165, pl. 119, figs 4–5. Parasmilia gravesi Milne Edwards & Haime, 1848: Baron-Szabo, p. 165. Parasmilia laticostata Bölsche, 1866: Baron-Szabo, p. 165. Parasmilia centralis (Mantell 1822): Löser et al., p. 485 (older synonyms cited therein). Parasmilia cylindrica Milne-Edwards & Haime 1851: Löser et al., p. 488 (older synonyms cited therein). Parasmilia gravesi Milne Edwards & Haime, 1848: Löser et al., p. 491 (older synonyms cited therein). Parasmilia lindströmi: Bernecker & Weidlich, p. 16ff, fig. 7A (6).

Dimensions. d (adult)=10–13 mm; s (adult):=48; h=up to 60 mm. Description. Trochoid-ceratoid, circular to slightly elliptical in outline; compressed or cylindrical in juvenile stages; the ratio of minimum/maximum diameter ranges between 0.8–1 in adult stages, can be as small as 0.49 in juvenile stages; costosepta thin, slightly flexuous, arranged in 3 complete cycles in 6 systems in early ontogenetical stages (corallite diameter of 2–3.5 mm); 4 complete cycles in 6 systems present in adult stages of corallite diameter ranging between 10 and 13 mm; in the holotype of the species P. centralis the latest adult



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stage is reached at a corallite diameter of 12–13 mm; around 12 septa reach the axial region; columella generally well-developed. Remarks. The synonymy list given above is based on the idea that the specimens used to create the taxa P. cylindracea, P. gravesiana, P. helenae, P. scanica, P. lindstroemi, P. laticostata, and P. granulata represent different ontogenetical stages of the species P. centralis. The species P. centralis, P. cylindracea, P. laticostata, and P. gravesiana are characterized by skeletal dimensions of d=10–13 mm, s=48, and h=up to 40 mm. In the species P. helenae and P. scanica the number of septa ranges between 30 and 40 (24+s) in a corallite diameter of around 5 mm. The specimens upon which the latter species was established closely correspond to the juvenile part of the holotype of the species P. centralis. The species P. lindstroemi was based on a series of syntypes which include larger coralla of 11 mm in diameter and 48 septa, as well as small coralla of e.g., 7 mm in diameter having around 24 septa. Therefore, the type series of the species P. lindstroemi forms an ontogenetical succession uniting the above mentioned taxa. Type location of species. Campanian-Maastrichtian of England (Sussex). Distribution. Cretaceous of Germany, Upper Cretaceous of France, Cenomanian-Turonian of the Czech Republic (Bohemia), Turonian of Bulgaria and Great Britain, Santonian of Tunisia, Senonian of Great Britain and Poland, Lower Campanian of Bulgaria, Upper Campanian of Ukraine, Lower Maastrichtian of Mexico (Cerralvo, this paper) and Bulgaria, Maastrichtian of Libya (Sofeggin), Ukraine, Belgium, and The Netherlands, Upper Danian of Denmark (Fakse, Stevns Klint) and Poland. New Material. Lower Maastrichtian of Mexico (Cerralvo), sample nos: UANL CE MAAS-212–214.

Parasmilia elongata Milne Edwards & Haime, 1848 Pl. 6, figs 5a, b

PLATE 6: Fig. 1 Parasmilia centralis (Mantell, 1822), holotype, BMNH, R.5462, Campanian-Maastrichtian of England. 1a, cross view, upper surface, scale bar: 4 mm; 1b, longitudinal view, scale bar: 3 mm. Fig. 2 Parasmilia centralis (Mantell, 1822), UANL CE MAAS-213, Lower Maastrichtian of Mexico, scale bar: 1.5 mm. 2a, cross view, upper surface; 2b, longitudinal view. Fig. 3 Parasmilia centralis (Mantell, 1822) (holotype of Parasmilia granulata Duncan, 1869), BMNH, R.6612, Senonian of Great Britain. 3a, cross view, upper surface, scale bar: 1.7 mm; 3b, longitudinal view, scale bar: 1.5 mm. Fig. 4 Parasmilia centralis (Mantell, 1822), (topotype of Parasmilia lindstroemi Henning, 1899), MB, Schlotheim coll., K.3631.2, Upper Danian of Denmark (Fakse), lateral view, ‘steinkern preservation’, scale bar: 3 mm. Fig. 5 Parasmilia elongata Milne Edwards & Haime, 1848d, topotype (?syntype), MNHN, Milne Edwards & Haime coll., M-549, Lower Maastrichtian of Belgium. 5a, longitudinal view, scale bar: 3 mm; 5b, cross view, scale bar: 3 mm. Fig. 6 Faksephyllia faxoensis (Beck, 1835), topotype, Bernecker & Weidlich coll., University of Erlangen, M.68, Middle Danian of Denmark (Fakse limestone), cross view of colony, scale bar: 2.5 mm (photograph courtesy M. Bernecker & O. Weidlich). Fig. 7 Faksephyllia faxoensis (Beck, 1835), topotype, BSP, 1967-XII-1, Middle Danian of Denmark (Fakse limestone), upper surface of colony, scale bar: 1.5 mm. Fig. 8 Faksephyllia faxoensis (Beck, 1835), topotype, MB, Schlotheim, coll., K. K.2743.1, upper surface of colony, scale bar: 4 mm.


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v*1848d 1848d 1850a v1850a 1850 v1857 v1857 non1866 1914 1914 1914 1922 1922 1922 non1927 1930 1987 1987 non1987 ?1987 2000 2000 2000 2002 2002 2002 2002 2002 2005

Parasmilia elongata: Milne Edwards & Haime, vol. 4, p. 246. Parasmilia punctata: Milne Edwards & Haime, vol. 4, p. 246. Parasmilia cylindrica: Milne Edwards & Haime, p. 50, pl. 8, fig. 5. Parasmilia Fittoni: Milne Edwards & Haime, p. 50, pl. 9, figs 2-2b (topotypes studied) Cyclosmilia elongata: d’Orbigny, vol. 2, p. 276. Parasmilia ? elongata: Milne Edwards, vol. 2, p. 174. Parasmilia Fittoni: Milne Edwards, vol. 2, p. 174 (topotypes studied). Parasmilia cylindrica M. Edw. u. Haime: Bölsche, p. 465, pl. 8, figs 2, 3. Parasmilia cylindrica E. H. 1850: Felix, pars 7, p. 228. Parasmilia elongata E. H. 1849: Felix, pars 7, p. 228. Parasmilia Fittoni E. H.: Felix, pars 7, p. 228. Parasmilia biseriata, M. U. H.: Nielsen, p. 26, pl. 4, figs 5–9. Parasmilia cincta, n. sp.: Nielsen, p. 27, pl. 4, figs 10–12. Parasmilia danica, n. sp.: Nielsen, p. 28, pl. 4, figs 17–18. Parasmilia Fittoni M. Edw. & Haime: Siemiradzki, p. 318, pl. 1, fig.2a–c. cf. Parasmilia danica Brünnig-Nielsen: Kühn, p. 547. Parasmilia elongata Edw. et Haime: Kuzmicheva, p. 64. Parasmilia cylindrica Edwards et Haime, 1850: Kuzmicheva, p. 135, pl. 23, figs 8–10. Parasmilia fittoni Edwards et Haime, 1851: Kuzmicheva, p. 135, pl. 23, fig. 11, pl. 24, fig. 1. Parasmilia biseriata (Forchhammer et Steenstrup, 1850): Kuzmicheva, p. 137, pl. 24, figs 3a–b. Parasmilia elongata, Milne-Edwards & Haime 1848: Löser, p. 60. Parasmilia fittoni, Milne-Edwards & Haime 1848: Löser, p. 60. Parasmilia punctata, Milne-Edwards & Haime 1848: Löser, p. 60. Parasmilia elongata Milne Edwards & Haime, 1848: Baron-Szabo, p. 165. Parasmilia fittoni Milne Edwards & Haime, 1848: Baron-Szabo, p. 165. Parasmilia punctata Milne Edwards & Haime, 1848: Baron-Szabo, p. 165. Parasmilia cincta Nielsen 1922: Löser et al., p. 487 (older synonyms cited therein). Parasmilia cylindrica Milne-Edwards & Haime 1851: Löser et al., p. 488 (older synonyms cited therein). Parasmilia cylindrica: Bernecker & Weidlich, p. 8.

Dimensions. d=16–18 mm; s=48; h=up to 70 mm. Description. Cylindrical, slightly bent; costosepta thin, arranged in 4 complete cycles in 6 systems; columella fasciculate or reduced. Remarks. The synonymy list given above is based on the idea that the specimens used to create the taxa P. punctata, P. cylindrica, P. danica, P. fittoni, and P. cincta represent different ontogenetical stages of the species P. elongata. The species P. fittoni, P. elongata, P. cylindrica, P. biseriata, and P. punctata are characterized by the skeletal dimensions of d=16–18 mm, s=48, and h=up to 70 mm. In the species P. cincta and P. biseriata the number of septa ranges from 30–32 in a corallite diameter of 8 mm. The specimen upon which the latter species was described seems to closely correspond to the juvenile stage of the species P. elongata. This idea is supported by the fact that both smaller and larger stages have been documented from the same localities: Upper Campanian of Denmark and Upper Danian of Denmark. Type location of species. Lower Maastrichtian of Belgium (Ciply). Distribution. Cretaceous of France and Germany, Turonian and Campanian of Great Britain, Lower Campanian of Turkmenistan, Upper Campanian-Maastrichtian of Denmark (Fakse, Stevns Klint), Lower Maastrichtian of Belgium (Ciply) and ?Ukraine, Maastrichtian of Kazakhstan and Russia, Danian of Austria, Upper Danian of Denmark.


BARON-SZABO

Genus Faksephyllia Floris, 1972 Type species. Caryophyllia faxoensis Beck, in Lyell, 1837, Middle Danian of Denmark (Fakse limestone).

Diagnosis. Colonial, dendroid-fasciculate or phaceloid. Gemmation intracalicinal, distomodaeal. Costosepta compact, laterally covered by small, rounded granules. Pali and columella absent, but trabecular extensions of septal axial ends can form a pseudo-columella. Endothecal dissepiments vesicular. Wall septothecal.

Faksephyllia faxoensis (Beck, 1835) Pl. 6, figs 6–8 parsv1820 v*1835 1922 1972 1987 v1990 (v)1996 v2005

Madreporites hyppurinus: Schlotheim, p. 356. Caryophyllia faxoensis: Beck, in Lyell, p. 249, fig. 4 (topotypes studied). Haplophyllia faxensis, Beck: Nielsen, p. 14, pl. 1, figs 1–7. Faksephyllia faxoensis gen. n. & Beck, in Lyell (1837) sp.: Floris, p. 73, pl. 4, figs 7–11, pl. 5, figs 1–5 (older synonyms cited therein). Faksephyllia faxoensis (Beck, in Lyell, 1837): Kuzmicheva, p. 138, pl. 24, figs 7–8. Faksephyllia faxoensis (Beck, 1835): Bernecker & Weidlich, p. 113, pl. 26, figs 3, 4, pl. 29, figs 5–8, pl. 30, figs 3, 4 (topotypes studied). Hexakoralle, Morphotyp 7: Tragelehn, p. 198, pl. 60, fig. 9. Faksephyllia faxoensis (Beck, 1835): Bernecker & Weidlich, p. 11ff, fig. 5B, fig. 6 (2), fig. 6B (topotypes studied).

Dimensions. d=3–4.5 mm, juveniles 1.5–2.5 mm; s=30–40, in juveniles around 24. Description. Finely branched, phaceloid colony; corallites circular to irregularly elliptical in outline; septa developed in 4 incomplete cycles in 6 systems in corallites that are not influenced by budding; septa thin, straight, oldest of which reach corallite centre where they might fuse; branching angle often between 30°–45°, can be up to 90°. Type locality of species. Middle Danian of Denmark (Fakse limestone). Distribution. Paleocene of Austria, Middle Danian of Denmark (Fakse and Greenland) and Sweden (Limhamn).

Genus Caryosmilia Wanner, 1902 Type species. Caryosmilia granosa Wanner, 1902, Danian of Egypt.

Diagnosis. Solitary, trochoid, fixed. Septa compact, nearly smooth marginally. Columella feebly developed. Paliform lobes opposite third cycle septa. Endothecal dissepiments deep in corallum. Remarks. Wanner (1902, p. 103) created the genus Caryosmilia without depicting a holotype of the type species. Because he assigned to this species specimens from two different localities which differ in age (Babel Jasmund corresponds to the Danian whereas the sediments in the desert region north of Dachel range from the Maastrichtian to the Paleocene), the exact type locality and age of the type species was never defined. Therefore, the specimen NRZ-PZ, Cn 47523 from Babel Jasmund, Danian of Egypt is here chosen as the lectotype of the type species (Caryosmilia granosa Wanner, 1902) of the genus Caryosmilia Wanner, 1902.



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Affinities. Similar to Parasmilia but columella feebly developed and paliform lobes opposite 3rd cycle septa.

Caryosmilia granosa Wanner, 1902 Pl. 7, figs 1a, b v*1902 1903 1930 non1955 1981 1996 2000 2002

Caryosmilia granosa n. sp.: Wanner, p. 103, pl. 14, figs 14–15. Caryosmilia granosa, Wanner, 1902: Fourtau, p. 345, pl. 5, figs 8–9. Caryosmilia abeli nov. spec.: Kühn, p. 546, pl. 1 (26), fig. 5. Caryosmilia granosa Wann.: Rossi Ronchetti, p. 119, pl. 12, figs 5a–c. Caryosmilia granosa Wanner: Barthel & Herrmann-Degen, p. 162. Asterosmilia decapali Hassan & Salama, 1969–70: Schuster, p. 82, fig. 44. Caryosmilia granosa, Wanner 1902: Löser, p. 18. Caryosmilia granosa Wanner, 1902: Baron-Szabo, p. 165, text-fig. 65.

Dimensions. d (min)=6–10 mm; d (max)=8–14 mm; d (min)/d (max)=around 0.75; s=48; h=10–20 mm. Description. Trochoid, elliptical in outline; septa arranged in 4 compete cycles in 6 systems; S1 and S2 nearly equal, remaining septa alternate in length and thickness. Remarks. In having finely granulated septa, a well-developed spongy columella, and no paliform structures in front of third cycle septa, the specimens described as Caryosmilia granosa by Rossi Ronchetti (1955, p. 119) rather correspond to the genus Parasmilia, most likely representing Parasmilia centralis. According to Kühn (1930) the species Caryosmilia abeli from the Danian of Austria is distinguished from the species C. granosa in having trabecular prolongations of septal axial ends that connect with the columella, and more strongly developed paliform lobes. Because these characteristics are interpreted as non-specific variations, the two species are considered to be synonymous. PLATE 7: Fig. 1 Caryosmilia granosa Wanner, 1902, holotype, NRM-PZ, Wanner coll., Cn 47523, Danian of Egypt. 1a, longitudinal view, scale bar: 4 mm; 1b, cross view, scale bar: 3.5 mm. Fig. 2 Flabellum conoideum Vaughan, 1900 (holotype of Flabellum texense Vaughan & Popenoe, 1935), NMNH, M371034, Paleocene of the USA (Texas), cross view, scale bar: 3.5 mm. Fig. 3 Desmophyllum excavatum (Hagenow, 1839), BSPG, most likely Bölsche coll., labeled as Coelosmilia sacheri Reuss, not cataloged, Senonian (?Upper Campanian) of Germany, scale bar: 4 mm. 3a, cross view; 3b, longitudinal view. Fig. 4 Desmophyllum excavatum (Hagenow, 1839), NRM-PZ, Wanner coll., Cn 47532, Danian of Egypt, scale bar: 1.5 mm. 4a, cross view; 4b, longitudinal view. Fig. 5 Smilotrochus cornucopiae (Duncan, 1869), cast of holotype in the collections of BMNH, Maastrichtian of England, scale bar: 3 mm. 5a, cross view; 5b, longitudinal view. Fig. 6 Smilotrochus milneri (Gregory, 1898), NRM-PZ, Wanner coll., Cn 47533, Danian of Egypt, cross view, scale bar: 3.5 mm. Fig. 7 Smilotrochus milneri (Gregory, 1898) (paratype of Smilotrochus jacobi Alloiteau, 1936), MNHN, in box with holotype Mo5213, Campanian-Maastrichtian of Madagascar, cross view, polished surface, scale bar: 4.5 mm. Fig. 8 Smilotrochus milneri (Gregory, 1898) (paratype of Smilotrochus jacobi Alloiteau, 1936), MNHN, in box with holotype Mo5213, Campanian-Maastrichtian of Madagascar, cross view, polished surface, scale bar: 4.5 mm. Fig. 9 Flabellum conoideum Vaughan, 1900 (syntype of Flabellum conoideum mathewsense Vaughan, 1900), NMNH, M157853, Paleocene of the USA (Texas), scale bar: 3.5 mm. 9a, cross view; 9b, longitudinal view. Fig. 10 Flabellum conoideum Vaughan, 1900 (syntype of Flabellum conoideum mathewsense Vaughan, 1900), NMNH, M157853, Paleocene of the USA (Texas), scale bar: 3.5 mm. 10a, cross view; 10b, longitudinal view.


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Type locality of species. Danian of Egypt. Distribution. Maastrichtian of Tunisia and Madagascar, Maastrichtian-Paleocene of Egypt, Danian of Austria. New Material. Danian of Egypt (Babel Jasmund), NRZ-PZ, sample nos.: Cn 47518 and 47523–24.

Genus Asterosmilia Duncan, 1867 (=Sphenotrochopsis Alloiteau & Tissier, 1958, Type species. Sphenotrochopsis alloiteaui Alloiteau & Tissier, 1958, Paleocene of France) Type species. Trochocyathus abnormalis Duncan, 1863, Miocene of the West-Indies (subsequent designation Vaughan, 1919).

Diagnosis. Solitary, trochoid-ceratoid, generally free. Costosepta compact, minutely and densely granulated laterally. Costae prominent or flat. Pali or paliform lobes next to last one or two cycles, usually opposite S3. Columella lamellar at surface, trabecular below. Wall septothecal to septoparathecal. Epithecal wall pellicular when present. Remarks. Alloiteau & Tissier (1958) described the genus Sphenotrochopsis from the Upper Danian of Belgium. According to the authors, the genus is characterized by a trabecular columella that appears lamellar, compact costosepta that are finely and densely granulated laterally, paliform structures that occur before oldest cycle septa (for the species Sphenotrochopsis alloiteaui that means before the cycles next to last one or two cycles), and a septothecal wall (parathecal only at very top of corallum). Synonymy of Sphenotrochopsis with Asterosmilia is therefore indicated.

Asterosmilia alloiteaui (Alloiteau & Tissier, 1958) Text-Fig. 12 (v)*1958 (v)1958 (v)1958 (v)1958 (v)1958

Sphenotrochopsis alloiteaui nov. sp.: Alloiteau & Tissier, p. 269, pl. 1, figs 1a–b’. Sphenotrochopsis chavani nov. sp.: Alloiteau & Tissier, p. 271, pl. 1, figs 2a–b’. Sphenotrochopsis straeleni nov. sp.: Alloiteau & Tissier, p. 272, pl. 1, figs 3a–b’. Kionotrochus briarti nov. sp.: Alloiteau & Tissier, p. 277, pl. 1, figs 7a–b’. Kionotrochus montensis nov. sp.: Alloiteau & Tissier, p. 280, pl. 1, figs 6a–b’.

Dimensions. d (min)=3.8–6.5 mm; d (max)=4.4–7.5 mm; d (min)/d (max)=0.75–0.87; s=36–64. Description. Trochoid-ceratoid corallum, slightly compressed in outline; costosepta are arranged in 6 systems, in corallite diameters of 4–5 mm around 40 septa are found, in a corallite diameter of around 6 mm 4 complete cycles of septa are present; Remarks. As pointed out by Cairns (1997), the specimens ascribed to Kionotrochus by Alloiteau & Tissier (1958) differ from this genus in having an epithecal wall, a spongy columella, and an apparently attached corallum. In showing 1) paliform structures before S3 (as seen in the type material illustrated on Pl. 1, figs 6a–7b’), 2) finely granulated, compact costosepta, 3) a trabecular columella that appears lamellar at surface, and 4) both septothecal-septoparathecal and epithecal walls, the taxa originally described as K. briarti and K. montensis correspond to the genus Asterosmilia. Regarding the species originally described as K. alloiteaui and K. chavani see Remarks section above for the genus Asterosmilia.


BARON-SZABO

Type locality of species. Upper Danian of Belgium. Distribution. Upper Danian of Belgium.

Text-Fig. 12 Asterosmilia alloiteaui (Alloiteau & Tissier, 1958). A, B, holotype of Asterosmilia alloiteaui (Alloiteau & Tissier, 1958) based on the illustrations in Alloiteau & Tissier (1958), figured as Sphenotrochus alloiteaui Alloiteau & Tissier, 1958, Upper Danian of Belgium, A, cross view of corallum; B, upper surface longitudinal view; C, D, holotype of Sphenotrochus chavani Alloiteau & Tissier, 1958 (=junior synonym of Asterosmilia alloiteaui [Alloiteau & Tissier, 1958]), based on the illustrations in Alloiteau & Tissier (1958), Upper Danian of Belgium, C, cross view of corallum; D, upper surface longitudinal view; E, F, holotype of Sphenotrochus straeleni Alloiteau & Tissier, 1958 (=junior synonym of Asterosmilia alloiteaui [Alloiteau & Tissier, 1958]), based on the illustrations in Alloiteau & Tissier (1958), Upper Danian of Belgium, E, cross view of corallum; F, upper surface longitudinal view; G, H, holotype of Kionotrochus briarti Alloiteau & Tissier, 1958 (=junior synonym of Asterosmilia alloiteaui [Alloiteau & Tissier, 1958]), based on the illustration in Alloiteau & Tissier (1958), Upper Danian of Belgium, G, cross view of corallum; H, upper surface longitudinal view; I, J, holotype of Kionotrochus montensis Alloiteau & Tissier, 1958 (=junior synonym of Asterosmilia alloiteaui [Alloiteau & Tissier, 1958]), based on the illustration in Alloiteau & Tissier (1958), Upper Danian of Belgium, I, cross view of corallum; J, upper surface longitudinal view, scale bar: 5 mm.

Genus Dasmosmilia Pourtalès, 1880 Type species. Dasmosmilia lymani Pourtalès, 1880, Recent, West Indies (off Grenada).

Diagnosis. Solitary, trochoid, fixed. Costosepta compact, margins smooth or slightly granular. Axial structures formed by trabecular extensions of inner ends of septa, resulting in false columella and false pali. Endothecal dissepiments thin, sparse. Epitheca rudimentary.

Dasmosmilia kochii (Pratz, 1910) Text-Fig. 13 *1910

Turbinolia kochii, Pratz; nov. sp.: Pratz, p. 299, pl. 23, figs 1–1b.



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v1933 2000 2002

Dasmosmilia reesidei n. sp.: Wells, p. 139, pl. 12, figs 8–9, pl. 14, figs 25–29. Dasmosmilia reesidei, Wells 1933: Löser, p. 27. Dasmosmilia reesidei Wells, 1933: Baron-Szabo, p. 166.

Text-Fig. 13 Dasmosmilia kochii (Pratz, 1910). A, type specimen as figured in Pratz (1910) as Turbinolia kochii Pratz, 1910, Upper Campanian of Serbia, cross view; B, NMNH, Coates coll., sample no.: J-71-16Aa, Upper Maastrichtian of Jamaica; C, NMNH, Coates coll., sample no. J-71-16AbC, Upper Maastrichtian of Jamaica; scale bar: 4 mm

Dimensions. d (min)=5–20 mm; d (max)=5–21 mm; d (min)/d (max)=0.83–1; s=up to 100; h=up to 23 mm. Description. Solitary, turbinate-trochoid; circular to slightly elliptical in outline; costosepta arranged in 5 complete cycles in adult specimens, alternating in thickness, unequal in length; in coralla of a diameter of around 5–7 mm, 48 septa corresponding to 4 complete cycles are developed; columella is irregularly spongytrabecular and fused with trabecular prolongations of inner ends of septa of the first two sets; wall parathecal to septoparathecal; endothecal dissepiments sparse, mainly occurring in the lower part of the corallum; relicts of an epithecal wall present. Remarks. The specimens from Jamaica (Figs 13B and C) and from Serbia (Fruska Gora) have smaller dimensions than the specimens of the type series of Dasmosmilia reesidei. However, according to their septa development and their morphological appearance it is suggested that they represent juvenile stages of the species D. kochii. Type locality of species. Upper Campanian of Serbia (Fruska Gora). Distribution. Upper Campanian of Serbia (Fruska Gora), Maastrichtian of the USA (see Wells 1933: Navarro Formation, Texas; see Sohl, 1987: Alabama, Mississippi, Texas), Upper Maastrichtian of Jamaica ( this paper). New Material. Upper Maastrichtian of Jamaica, NMNH, Coates coll., sample nos.: J-71-16Aa; J-7116Ab (=Shaw Castle, Maldon Formation).

Genus Smilotrochus Milne Edwards & Haime, 1851 (=Dungulia Oppenheim, 1930b, Type species. Coelosmilia milneri Gregory, 1898, Lower Eocene of Egypt [Dungul Wells]). Type species. Trochosmilia tuberosa Milne Edwards & Haime, 1850a (=Turbinolia compressa Morris, 1843), Albian of England (Devonshire); original designation by Milne Edwards & Haime, 1851b.


BARON-SZABO

Diagnosis. Solitary, trochoid or subturbinate, fixed. Costosepta compact, granulated laterally. Septa often fuse which can result in the appearance of pseudo-dendrophyllid arrangements or in the forming of a a pseudocolumella when septal axial ends fuse in the corallite centre. Columella absent. Septal axial ends often project and may form short or very elongate paliform structures. Wall parathecal or septothecal. Endothecal dissepiments developed, but deep in corallum. Remarks. Gregory (1898) described the solitary species Coelosmilia milneri which, according to Gregory (1898, p. 249), is distinguished from the genus Smilotrochus only by “less simple and less crowded septa”, as well as “broader interseptal loculi”. While it remains unclear what exactly Gregory referred to as “simple” or “less simple” septa, respectively, the character of density of septa (Gregory used the term “crowded septa”, resulting in “narrower interseptal loculi”) could be considered a specific but not a generic feature. Therefore, the species Coelosmilia milneri (type species of the genus Dungulia) is considered a species of Smilotrochus. Hence, because it is the type species of Dungulia, the latter represents a junior synonym of Smilotrochus. In creating the genus Dungulia, Oppenheim (1930b) used non-type material to prove and establish a relation to the dendrophylliid group of eupsammiids. Because in the original description of Coelosmilia milneri, Gregory (1898) solely compared his new species with other caryophylliid species, a relation only with caryophylliid forms can be assumed. However, the presence of Trochosmilia-like septa (=rather thin, often finely granulated septa) which have the tendency to fuse was already documented by Milne Edwards & Haime (1850a) and later included in the first generic description of Smilotrochus by Milne Edwards (1857, vol. II, p. 70). Especially in taxa of this genus which have flexuous septa, as already reported for the Maastrichtian form Smilotrochus hagenowi Milne Edwards (1857, vol. 2, p. 71), the fusion of their axial ends can result in a septal pattern that resembles the dendrophyllid type (compare specimens figured on Pl. 7, figs 5a and 6–8).

Smilotrochus cornucopiae (Duncan, 1869) Pl. 7, figs 5a, b v*1869 1914 1922 1987 2000 (v)2005

Coelosmilia cornucopiae, n. sp.: Duncan, vol. 2, p. 8, pl. 3, figs 6–10. Coelosmilia cornucopiae Duncan 1869: Felix, pars 7, p. 219. Smilotrochus faxoensis, Forchhammer and Steenstrup: Nielsen, p. 29, pl. 3, figs 1, 1a, pl. 4, figs 19–22. Smilotrochus ponderosus (Forchhammer and Steenstrup, 1850): Kuzmicheva, p. 132, pl. 22, fig. 8, pl. 23, fig. 1. Coelosmilia cornucopiae, Duncan 1869: Löser, p. 20. Smilotrochus faxoensis: Bernecker & Weidlich, p. 16ff, fig. 7A, figs 7 (1, 4, 5).

Dimensions. d (min)=8–21 mm; d (max)=8–25 mm; d (min)/d (max)=0.8–0.92; s=30–48; h=20–70 mm. Description. Solitary, trochoid, slightly elliptical in outline; costosepta arranged in 4 complete cycles in 6 irregular system in corallites that are 15 mm and larger in diameter; in more adult corallites of around 15 mm in diameter S1 and S2 are nearly equal and reach the axial region; in younger specimens 6 septa reach corallite centre; remaining septa irregularly alternate in length and thickness. Remarks. The specimens assigned to Smilotrochus ponderosus in Kuzmicheva (1987) are slightly larger and have more septa than the species S. cornucopiae. However, the ratio of the corallite diameter completely corresponds to Duncan’s species and has a number of septa that would expected to be found in a slightly larger corallum. Therefore, it is suggested that the larger specimens represent ontogenetically later stages of the species S. cornucopiae. The specimens described as Smilotrochus faxoensis from the Danian of Denmark and Sweden completely correspond to the juvenile to nearly adult stages observed in the type specimen of S. cornucopiae. Therefore, SCLERACTINIAN CORALS OF THE K/T-BOUNDARY


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they are intepreted to be synonymous. Type locality of species. Maastrichtian of England (Trimmingham, Norfolk) Distribution. Maastrichtian of England, Kazakhstan, and Ukraine, Danian of Denmark (Stevns Klint) and Sweden (Limhamn).

Smilotrochus galeriformis (Kner, 1848) Text-Fig. 14 v*1848 1852 1857 1880 ?1880 1914 1914 1925 1927 1927 1974 1987 1987 2002 2002 2002 2002 2002 2002

Turbinolia galeriformis: Kner, vol. 3, p. 34, pl. 5, fig. 8 (topotypes studied). Coelosmilia galeriformis Rss.: Reuss, p. 118, pl. 17, figs 1a–b. Smilotrochus Hagenowi: Milne Edwards, vol. 2, p. 71. Smilotrochus Jakhmari, Duncan: Duncan, p. 19, pl. 1, figs 14–17. Blagrovia simplex, Duncan: Duncan, p. 28, pl. 14, figs 11–13. Coelosmilia galeriformis Kner sp. 1850: Felix, pars 7, p. 220. Smilotrochus Hagenowi M. Edwards 1857: Felix, pars 7, p. 230. Smilotrochus Hagenowi M. Edw.: Umbgrove, p. 116, pl. 11, fig. 28. Smilotrochus galeriformis Kner: Siemiradzki, p. 314, pl. 1, fig. 4. Smilotrochus Nagorzanensis n. sp.: Siemiradzki, p. 316, pl. 1, fig. 8. Smilotrochus galeriformis (Kner, 1848): Kuzmicheva, p. 63, pl. 10, figs 1–2. Smilotrochus grandis Siemiradzki, 1927: Kuzmicheva, p. 130, pl. 22, fig. 5. Smilotrochus galeriformis (Kner, 1848): Kuzmicheva, p. 131, pl. 22, figs 6–7b. Smilotrochus galeriformis (Kner 1848): Löser et al., p. 605 (older synonyms cited therein). Smilotrochus hagenowi, Milne-Edwards 1857: Löser et al., p. 606 (older synonmys cited therein). Smilotrochus nagorzanensis, Siemiradzki 1927: Löser et al., p. 607 (older synonyms cited therein). Smilotrochus galeriformis (Kner, 1848): Baron-Szabo, p. 170. Smilotrochus hagenowi Milne Edwards, 1857: Baron-Szabo, p. 170. Smilotrochus nagorzanensis Siemiradzki, 1927: Baron-Szabo, p. 170.

Dimensions. d (min)=7.5–38 mm; d (max)=10–50 mm; d (min)/d (max)=0.71–0.96; s=96+s6; h=up to 80 mm.

Text-Fig. 14 Smilotrochus galeriformis (Kner, 1848). A, B, based on the illustration in Kuzmicheva (1987), Maastrichtian of Ukraine; A, longitudinal view of septal apparatus; B, longitudinal view of corallum, scale bar: 10 mm.

Description. Solitary, trochoid, elliptical-compressed (adult stages) or nearly cylindrical (juvenile stages)


BARON-SZABO

in outline; costosepta thin, straight or wavy, arranged in 4 complete cycles in 6 systems in a corallite of around 10 mm in diameter, 5 complete cycles present in a corallite of around 30 mm in diameter; in larger corallites the beginning of a sixth cycles is present; septa nearly equal in juvenile stages, alternate in length and thickness in more adult corallites. Remarks. The specific characteristics of this taxon given above are based on the description and illustrations in Reuss (1852) which represent topotypes of the species S. galeriformis. Type locality of species. Upper Campanian of Ukraine (Nagorzany, Lwowskaya) Distribution. Cretaceous of Poland, Upper Cretaceous of Sweden (Limhamn), Upper Santonian-Lower Campanian of Romania, Upper Campanian-Maastrichtian of Ukraine, Maastrichtian of Azerbaijan, Russia, and Kazakhstan, Upper Maastrichtian of The Netherlands, Danian of Pakistan.

Smilotrochus ponderosus (Nielsen, 1922) Fig. 15 *1922 1927 non1987 1987 2002 2002 2002

Coelosmilia ponderosa, (Forchhammer and Steenstrup): Nielsen, p. 223, pl. 4, figs 1–2. Smilotrochus grandis n. sp.: Siemiradzki, p. 315, pl. 1, fig. 1. Smilotrochus ponderosus (Forchhammer and Steenstrup, 1850): Kuzmicheva, p. 132, pl. 22, fig. 8, pl. 23, fig. 1. Smilotrochus excavatus (Hagenov, 1839): Kuzmicheva, p. 133, pl. 23, figs 2–4. Smilotrochus grandis, Siemiradzki 1927: Löser et al., p. 606 (older synonyms cited therein). Smilotrochus ponderosa (Nielsen 1922): Löser et al., p. 607 (older synonyms cited therein). Smilotrochus ponderosus (Nielsen, 1922): Baron-Szabo, p. 170.

Dimensions. d (min)=35–40 mm; d (max)=38–45 mm; d (min)/d (max)=0.88–0.92; s=56– ca. 70; h=46–100 mm. Description. Solitary, trochoid, slightly elliptical in outline; septa arranged in 4 complete cycles in 6 systems, regularly alternating; the beginning of a fifth cycle present in a corallite of 35–40 mm in diameter.

Text-Fig. 15 Smilotrochus ponderosus (Nielsen, 1922), based on the illustration in Nielsen (1922), Upper Campanian of Denmark (Amter Aalborg), longitudinal view, scale bar: 20 mm.

Type locality of species. Upper Campanian of Denmark (Amter Aalborg). SCLERACTINIAN CORALS OF THE K/T-BOUNDARY


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Distribution. Cenomanian of Ukraine, Lower Santonian of Poland, Upper Campanian-Maastrichtian of Denmark (Amter Aalborg), Maastrichtian of Kazakhstan, Russia, and Ukraine.

Smilotrochus milneri (Gregory, 1898) Pl. 7, figs 6–8 *1898 v1902 v1936 v1996 1996 v1999 2000 2002

Coelosmilia Milneri, n. sp.: Gregory, p. 248, pl. 8, figs 1–3. Coelosmilia Milneri Gregory: Wanner, p. 102, pl. 14, figs 19–24. Smilotrochus Jacobi (nov. sp.): Alloiteau, p. 12, pl. 6, figs 3–38. Smilotrochus jacobi Alloiteau, 1936: Baron-Szabo, in Baron-Szabo & Steuber, p. 20, pl. 8, figs 5–6. Dungulia milneri (Gregory, 1898): Schuster, Appendix list. Smilotrochus jacobi Alloiteau, 1936: Baron-Szabo, p. 451, pl. 4, fig. 2. Smilotrochus jacobi, Alloiteau 1936: Löser, p. 73. Smilotrochus jacobi Alloiteau, 1936: Baron-Szabo, p. 170.

Dimensions. d (min)=9–15 mm; d (max)=26–30 mm; d (min)/d (max)=0.35–0.73; s=40–96; h=up to 35 mm. Description. Solitary, trochoid; compressed in adult stages that have a maximum corallite diameter of 30 mm (ratio d (min)/d (max)=0.35), elliptical in younger ontogenetical stages; in corallite diameters of d (max) ranging between 15–18 mm the ratio d (min)/d (max) varies between 0.5 and 0.73; costosepta thin, wavy to flexuous, arranged in 5 complete cycles in a corallite of d (max) 26–30 mm; in corallites of d (max) 15–18 mm the number of septa is around 48, arranged in 4 complete or nearly complete irregular cycles in 6 systems; 12 (younger stages)–24 (late adult stage) septa equal, reaching the axial region. Remarks. See discussion above in the Remarks chapter below genus description of Smilotrochus. Type locality of species. Lower Eocene of Egypt (Dungul Wells). Distribution. Aptian of Greece, Upper Turonian of Austria (Gosau Group at Weissenbachalm), Campanian-Maastrichtian of Madagascar (Narinda), Paleocene-Lower Eocene of Egypt. New Material. Danian of Egypt (Babel Jasmund), NRM-PZ, sample nos.: Cn 47533–35. Subfamily Desmophyllinae Vaughan & Wells, 1943 (=Family Parasmiliidae Alloiteau, 1952a p.p.) Diagnosis. Solitary and colonial. Ahermatypic. Colony formation by intra- and extratentacular budding. Endothecal dissepiments deep and sparse. Coenosteum dense where developed. Columella rudimentary or absent. Pali and epitheca absent.

Genus Desmophyllum Ehrenberg, 1834 Type species. Desmophyllum dianthus Ehrenberg, 1834 (=Desmophyllum cristagalli Milne Edwards & Haime, 1848c), Recent, Gulf of Gascony.

Diagnosis. Solitary, trochoid to subturbinate, fixed. Costosepta compact, exsert, granular marginally. Costae strong near calice, rudimentary basally and covered by stereome in most species. Endotheca present or absent. Columella absent or very small. Pali absent. Desmophyllum excavatum (Hagenow, 1839)


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Pl. 7, figs 3a–4b *1839 1850a 1852 v1852 1852 1857 1857 1858–61 1858–61 1864 1866 v1866 v1866 v1867 v1867 1869 1881 1881 1885 v1902 1914 1914 1914 1914 1922 1955 v1969 1982 1987 non1987 1996 2000 2000 2000 2000 2002 2005

Turbinolia excavata nob.: Hagenow, p. 289. Coelosmilia laxa: Milne Edwards & Haime, p. 52, pl. 8, figs 4a–c. Turbinolia excavata var. cylindrata: Quenstedt, p. 655, pl. 59, fig. 1. Coelosmilia Sacheri Rss.: Reuss, p. 119, pl. 17, figs 2, 2a–b. Coelosmilia cupuliformis Rss.: Reuss, p. 119, pl. 17, figs 3–5. Coelosmilia laxa: Milne Edwards, vol. 2, p. 178. Coelosmilia ? excavata: Milne Edwards, vol. 2, p. 179. Coelosmilia laxa: de Fromentel, p. 102. Coelosmilia ? excavata: de Fromentel, p. 102. Coelosmilia ? excavata: de Fromentel, p. 287. Coelosmilia laxa M. Edw. u. Haime: Bölsche, p. 463. Coelosmilia cupuliformis Reuss: Bölsche, p. 463. Coelosmilia Sacheri Reuss: Bölsche, p. 464. Coelosmilia cupuliformis Reuss: Bölsche, p. 27. Coelosmilia Sacheri Reuss: Bölsche, p. 28. Trochosmilia (Coelosmilia) laxa: Duncan, vol. 2, p. 8, pl. 3, figs 11–17, pl. 4, figs 9–12. Turbinolia excavata var. cylindrata: Quenstedt, p. 839, pl. 176, fig. 29. Coelosmilia laxa: Quenstedt, p. 986, pl. 181, fig. 27. Turbinolia excavata: Quenstedt, p. 1011, pl. 82, fig. Coelosmilia libyca n. sp.: Wanner, p. 102, pl. 14, figs 16–18. Coelosmilia cupuliformis Reuss 1852: Felix, pars 7, p. 219. Coelosmilia laxa E. H. 1850: Felix, pars 7, p. 220 (older synonyms cited therein). Coelosmilia excavata Hagenow sp. 1839: Felix, pars 7, p. 221. Coelosmilia Sacheri Reuss 1852: Felix, pars 7, p. 221. Coelosmilia excavata, v. Hagenow sp.: Nielsen, p. 222, pl. 2, figs 21–28, pl. 3, figs 8–8b (older synonyms cited therein). Coelosmilia laxa M.-Edw. e Haime: Rossi Ronchetti, p. 103, pl. 11, fig. 1. Dungulia libyca (Wanner): Hassam & Salama, p. 88, pl. 1, figs 4a–c (topotypes studied). Desmophyllum laxum (Edwards et Haime, 1851): Kuzmicheva, p. 23, pl. 1, figs 2a–b. Desmophyllum laxum (Edwards et Haime, 1851): Kuzmicheva, p. 128, pl. 22, figs 2–3. Smilotrochus excavatus (Hagenov, 1839): Kuzmicheva, p. 133, pl. 23, figs 2–4. Dungulia libyca (Wanner, 1902): Schuster, Apendix list. Coelosmilia cupuliformis Reuss 1854: Löser, p. 20. Desmophyllum laxa (Milne-Edwards & Haime 1851): Löser, p. 28. Desmophyllum sacheri, Reuss 1854: Löser, p. 28. Parasmilia excavata (Hagenow 1839): Löser, p. 60. Desmophyllum laxa (Milne Edwards & Haime, 1850): Baron-Szabo, p. 168. Parasmilia excavata: Bernecker & Weidlich, 8ff.

Dimensions. d (min)=9–34 mm; d (max)=10–38 mm; d (min)/d (max)=0.80–1; s=24–96+s; h=13–100 mm. Description. Solitary, trochoid to subturbinate-subpatellate; circular in outline in juvenile stages, becoming slightly elliptical in later ontogenetical phases; costosepta thin, developed in 5 complete cycles in 6 systems in late adult stages; septa of the incomplete sixth cycle short, almost rudimentary; 3 complete cycles with the beginning of a fourth one developed in corallum of d=9–13 mm; 4 complete cycles with or without the beginning of a fifth one developed in a corallum of d (max) 15 mm. Type locality of species. Lower Maastrichtian of Germany (Rügen Island). Distribution. Senonian of Germany and Great Britain, Lower Maastrichtian of Germany (Rügen Island), SCLERACTINIAN CORALS OF THE K/T-BOUNDARY


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Maastrichtian of Denmark, (Fakse and Stevns Klint), Kazakhstan, Libya, Russia, Ukraine, and Turkmenistan, Paleocene of Egypt. New Material. Danian of Egypt (Babel Jasmund), NRM-PZ, sample no.: Cn 47532. Family Flabellidae Bourne, 1905 Diagnosis. Solitary, fixed or free; ahermatypic; neotenic. Wall epithecal, usually smooth, thickened internally by stereome. Septa compact, nonexsert, formed by one fan system of simple trabeculae, smooth marginally, smooth or granulated laterally. Pali, synapticulae, and dissepiments absent. Paliform lobes present in one genus. Columella usually rudimentary or absent, may be lamellar or papillose. Remarks. The diagnosis of this family is based on the one given by Cairns (1989) who provided detailed discussions regarding the Family Flabellidae.

Genus Flabellum Lesson, 1831 Type species. Flabellum parvoninum Lesson, 1831, Recent, Pacific.

Diagnosis. Solitary, cuneiform or compressed turbinate. Base tending to become free in adult stage. Transverse division lacking. Septa numerous (4–7 cycles), smooth laterally, crested marginally. Calicular edge smooth or jagged. Calicinal pit deep. Columella rudimentary: a simple fusion of lower, axial edges of major septa.

?Flabellum anderssoni (Felix, 1909) Text-Fig. 16

Text-Fig. 16 ?Flabellum anderssoni (Felix, 1909), syntype NRZ Cn85, Maastrichtian of Antarctic Peninsula, cross view, scale bar: 10 mm (from Filkorn 1994, see full reference in Reference list; copyright 1994 by the American Geophysical Union; reproduced by permission of American Geophysical Union). *1909

Parasmilia Anderssoni nov. sp.: Felix, p. 8, pl. 1, figs 7–7d.


BARON-SZABO

1914 1949 v1967 (v)1972 v1994 2000

Parasmilia Anderssoni Felix 1909: Felix, pars 7, p. 227. Parasmilia Anderssoni Felix: Furque & Camacho, p. 276, 293, and 295. Flabellum primitivum nov. spec.: Kühn & Traub, p. 14, pl. 1, figs 11, 12. Flabellum groenlandicum sp. n.: Floris, p. 81, pl. 5, figs 6A, B, pl. 6, figs 1–33, text-fig. 16. ?Flabellum anderssoni (Felix, 1909): Filkorn, p. 67, figs 24 (1–3) and 25 (1–3) (older synonyms cited therein). Flabellum anderssoni (Felix 1909): Löser, p. 36.

Dimensions. d (min)=1.5–15 mm; d (max)=6–21 mm; d (min)/d (max)=0.71–0.82, often ranging from 0.6–0.7 in juvenile stages; s=24–48; h=up to 23 mm. Description. Solitary, trochoid-subcylindrical; slightly elliptical in outline in more adult stages, compressed in juvenile coralla; costosepta straight, arranged in 3 to 4 cycles in 6 systems; 3 cycles are complete when corallite is around d (max) 15 mm; 12 septa reach the axial region; trabecular extensions of their inner ends fuse forming a weakly developed pseudo-columella. Remarks. Filkorn (1994, p. 67 ff.) re-evaluated the taxon Parasmilia anderssoni and tentatively transferred it to the genus Flabellum. However, the septal development also seems characteristic of juvenile stages of the genus Flabellosmilia (Baron-Szabo 2003). The species F. primitivum closely corresponds to juvenile stages of F. anderssoni; therefore, their synonymy is suggested. Type locality of species. Maastrichtian of Antarctic Peninsula (Seymour Island). Distribution. Maastrichtian (Seymour Island)-Danian (Tierra del Fuego) of Antarctica, Paleocene of Austria, Upper Danian of Denmark (Greenland).

Flabellum conoideum Vaughan, 1900 Pl. 7, figs 2, 9a–10b v*1900 v1900 v1935 v1935 v1935 (v)1977 (v)1977 ?1993

Flabellum conoideum sp. nov.: Vaughan, p. 56, pl. 3, figs 1–4. Flabellum conoideum var. mathewsense var. nov.: Vaughan, p. 58, pl. 3, figs5–6a. Flabellum conoideum Vaughan: Vaughan & Popenoe, p. 326, pl. 3, fig. 1. Flabellum conoideum var. Mathewsense Vaughan: Vaughan & Popenoe, p. 327, pl. 3, fig. 2. Flabellum texense Vaughan & Popenoe new species: Vaughan & Popenoe, p. 327, pl. 3, figs 3–5. Flabellum conoideum Vaughan: Toulmin, p. 145, pl. 1, figs 3, 4. Flabellum conoideum mathewsense Vaughan: Toulmin, p. 145, pl. 1, figs 5, 6. Caryophyllia conodeum ? Vaughan and Popenoe: Hansen et al., p. 257, Table 1.

Dimensions. d (min)=6.5–16 mm; d (max)=7.6–18 mm; d (min)/d (max)=0.75–0.9; s=48–72; h=up to 14 mm. Description. Slightly compressed corallum; septa developed in 4 cycles in 6 systems with the beginning of a fifth one; in corallite of around 7 mm in diameter 48 septa are present; in adult coralla S1, S2, and some of S3 dominant; the variety F. mathewsense has well-developed costae corresponding to S1 and S2, but grades directly into the typical form of F. conoideum. Type locality of species. Paleocene of the USA (Prairie Creek, Wilcox County, Alabama). Distribution. Paleocene of the USA (both members of Porters Creek Formation, Alabama; Kincaid and Wills Point Formations, Texas).

Flabellum cuneiforme Lonsdale, 1845 SCLERACTINIAN CORALS OF THE K/T-BOUNDARY


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Pl. 8, figs 1a, b v*1845 v1864 parsv1900 v1900

Flabellum (?) cuneiforme: Lonsdale, p. 512. Flabellum remondianum: Gabb, p. 207, pl. 26, fig. 199 (topotypes studied). Flabellum cuneiforme Lonsdale: Vaughan, p. 60, pl. 3, fig. 10, non fig. 12 (older synonyms cited therein) (topotypes studied). Flabellum remondianum Gabb: Vaughan, p. 67, pl. 4, figs 5, 6 (older synonyms cited therein) (topotypes studied).

Dimensions. d (min)=up to 12 mm; d (max)=up to 26.5 mm; d (min)/d (max)=0.4–0.6; h=up to 33 mm; Description. Flabellate corallum, narrow to compressed in outline, septa developed in 4 size orders in adult stages; generally, septa of first two size orders reach axial region. Costae rather smooth and straight. Remarks. In addition to accepting the variety Flabellum cuneiforme var. pachyphyllum, Vaughan (1900, p. 62ff.) established another four varieties of F. cuneiforme: pachyphyllum, acutiforme, fragile, and wallesi. Vaughan based these new variations on differences in their morphological appearance like smoothness of external surface, compressed or wide appearance in cross view, thickness of epitheca, presence or absence of lateral wings, transverse lines of growth, and the lack or development of prominent costae. The main problem regarding Vaughan’s investigation occurs due to the fact that Vaughan (1900, pl. III, fig. 12) assigned a specimen to Flabellum cuneiforme Lonsdale which significantly differed from the holotype of F. cuneiforme Lonsdale in exactly these external features (compare figure of type of Flabellum cuneiforme Lonsdale in Vaughan, 1900, pl. III, fig. 10). If these two specimens of Flabellum cuneiforme Lonsdale figured by Vaughan were considered the same species, they would span the range of variation which included at least three of his described variations (pachyphyllum, acutiforme, and wallesi) as documented by Vaughan (1900, p. 62ff., pl. III, figs 13, 15, 19, and 23). While it cannot be decided in the current study how much, if at all, these characteristics were controlled by (paleo-) environmental factors, it can be stated that the type material of F. cuneiforme Lonsdale and F. remondianum Gabb (as given the synonmy list above) correspond to each other by having 4 septal size orders, a rather compressed corallum of d (min)/d (max) around 0.5, and rather smooth and straight costae. Therefore, they are considered to be synonymous. PLATE 8 Fig. 1 Flabellum cuneiforme Lonsdale, 1845 (topotype of Flabellum remondianum Gabb, 1864), CAS, 69419, Paleocene of the USA (California), scale bar: 6 mm. 1a, cross view; 1b, longitudinal view. Fig. 2 ?Flabellum fresnoense Durham, 1943, holotype, CAS, 68349, Paleocene of the USA (California), scale bar: 2.5 mm. 2a, cross view; 2b, longitudinal view. Fig. 3 Alveolocyathus felixi (Filkorn, 1994), holotype, NMNH, 92996, Maastrichtian of Antarctic Peninsula, cross view, upper surface, scale bar: 1 mm. Fig. 4 Palocyathus seymourensis Filkorn, 1994, holotype, NMNH, 93050, Maastrichtian of Antarctic Peninsula, cross view, upper surface, scale bar: 1 mm. Fig. 5 Bothrophoria ornata Felix, 1909, topotype,NMNH, 92999, Maastrichtian of Antarctic Peninsula, scale bar: 2 mm. 5a, cross view; 5b, longitudinal view (photographs courtesy S. Cairns). Fig. 6 Platytrochus claibornensis Gregorio, 1890 (holotype of Platytrochus primaevus Vaughan & Popenoe, 1935), NMNH, M371031, Paleocene of the USA (Texas), scale bar: 1 mm. 6a, cross view; 6b, longitudinal view. Fig. 7 Actinhelia elegans (Goldfuss, 1826), topotype, MB, Schlotheim coll., MB.K.3688, Maastrichtian of The Netherlands. 7a, close-up, scale bar: 2.5 mm; 7b, upper surface of colony, scale bar: 5 mm. Fig. 8 Actinhelia elegans (Goldfuss, 1826), NMNH, Coates coll., 339b, Middle-Upper Maastrichtian of Jamaica, cross thin section, scale bar: 2 mm. Fig. 9 Actinacis haueri Reuss, 1854 (holotype of Montipora cretacea Umbgrove, 1925), RMNH, Umbgrove coll., 29072, Upper Maastrichtian of The Netherlands, upper surfase of colony in ‘steinkern’ preservation, scale bar: 1 mm. Fig. 10 Actinacis barretti Wells, 1934, NMNH, Berryhill coll., PR-7 series, Danian of Puerto Rico, oblique cross view of colony, polished surface, scale bar: 1 mm. Fig. 11 Actinacis martiniana d’Orbigny, 1850, NMNH, Coates coll., no. 517, Maastrichtian of Jamaica, cross thin section, scale bar: 2 mm. Fig. 12 Actinacis reussi Oppenheim, 1930, NMNH, Coates


BARON-SZABO

coll., no. 363, Middle-Upper Maastrichtian of Jamaica, cross thin section, scale bar: 4.5 mm.

Type locality of species. Eocene (Claibornian) of the USA (Eutaw Springs, Alabama).



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Distribution. Paleocene-Eocene (Claibornian) of the USA (Paleocene of California, Lodo Formation; Eocene of Alabama, South Carolina).

?Flabellum fresnoense Durham, 1943 Pl. 8, figs 2a, b v1943 2000

Flabellum fresnoense Durham, n. sp.: Durham, p. 197, pl. 32, figs 2–3. Flabellum fresnoense, Durham 1943: Löser, p. 36.

Dimensions. d=5.7 x 7.9 mm; d (min)/d (max)=0.5–0.72; s=16+s3; h (broken specimen)=7.4 mm. Description. Solitary, trochoid-subcylindrical; slightly flattened in outline; costosepta arranged in 2 cycles with the beginning of a third cycle in 8 systems; 16 septa reach the axial region. Remarks. The holotype of the species Flabellum fresnoense represents a small juvenile fragmented specimen which leaves unanswered questions regarding its septal development and expected habitus of the adult corallum. As shown in recent studies on the ontogenetical developments in solitary forms of the Upper Cretaceous, like e.g., Aulosmilia and Flabellosmilia (Baron-Szabo 2003), the septal arrangement can vary significantly during ontogeny in that changes from septal developments in cycles to arrangements in size orders often occur in these genera. In the present case the septal apparatus might not necessarily be arranged in cycles (here in 8 systems) but could very well represent a development in two with the beginning of a third size order. Moreover, the holotype of Flabellum fresnoense very closely resembles juvenile stages of Flabellosmilia (see e.g., Baron-Szabo 2003, pl. 17, fig. 1). While one significant feature of the latter genus is its diamond-shape (in cross view), in its juvenile stages however, it often shows rather flabellate to elliptical outlines. The diamond-shape often develops in later ontogenetical stages in that the corallum increasingly becomes more flabellate and at the same time the septa that are positioned in the central area of the corallum (meaning perpendicular to the distant edges of the corallum) become more dominant in length, widening the corallum in these places. In Flabellum fresnoense such a feature might be displayed. In addition, in having reduced costae, the form shows a feature that is characteristic of the genus Flabellosmilia. Type location of species. Paleocene of the USA (California, Fresno County, Lodo Formation, Cheney Well #1). Distribution. Paleocene of the USA (California, Fresno County, Lodo Formation, Cheney Well #1).

Family Turbinoliidae Milne Edwards & Haime, 1848 Diagnosis. Solitary, small and free, except for species having transverse division. Corallum conical, bowlshaped, or cylindrical. Costae originate independently or by trifurcation. surfaces granular, serrate, or smooth in texture. Intercostae present in some genera, and septa and costae alternate in others. Intercostal regions solid, pitted, or perforate. Epitheca absent. Septa usually exsert and usually hexamerally arranged in 3 to 4 cycles (28–48 septa); however, range of septal number is 10 to 72, including decahedral symmetry. Pali and paliform lobes in various arrangements or absent. Columella papillose, styliform, lamellar, fascicular, or absent. Endotheca absent.

Genus Turbinolia Lamarck, 1816


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Type species. Turbinolia sulcata Lamarck, 1816, Middle Eocene of France.

Diagnosis. Solitary, conical, circular in cross section and small (rarely exceeding 3.5 mm) in diameter. Costae independent in origin, usually well-developed, smooth ridges, and C1–2 sometimes thickened basally. Intercostae present or absent, depending on species; in type species present as alignment of low mounds, becoming low ridges only near calice. Series of circular pits up to 70 µm in diameter flank each costa, each bordered on distal and proximal edges by small thecal buttresses oriented perpendicular to the costa and often fused to the intercostae, if present. Thecal pits thus appear to form a double column, often in alternating arrangement. Septa exsert, hexamerally arranged in 2-4 cycles (12–48 septa). Pali and paliform lobes absent. Columella quite variable, including styliform, stellate, hexameral, and lamellar. Remarks. The diagnosis of this genus is based on the one given by Cairns (1989), who provided re-examinations and detailed discussions regarding the turbinoliids.

Turbinolia dickersoni Nomland, 1916 Text-Fig. 17 (v)*1916 (v)1944 (v)1944 v1945 v1945 (v)1960 1969 1992 1997 1997 1997 1997 1997

Turbinolia dickersoni, n. sp.: Nomland, p. 61, pl. 3, figs 5–8. Turbinolia midwayenesis Monsour, n. sp.: Monsour, p. 114, pl. 21, figs 4, 5. Turbinolia tenuis Monsour, n. sp.: Monsour, p. 114, pl. 21, figs 7, 8. Turbinolia barbadiana Wells, n. sp.: Wells, p. 13, pl. 3, figs 8–10. Turbinolia barbadiana var. crassicostata Wells, n. var.: Wells, p. 14, pl. 3, fig. 11. Turbinolia rosetta Howe, n.sp.: Howe, p. 1020, text-figs 1a, b. Turbinolia frescoenis n. sp.: Barta-Calmus, p. 818, pl. 1, figs 9, 9a. Turbinolia dickersoni Nomland, 1916: Squires & Demetrion, p. 24, figs 45–46. Turbinolia rosetta Howe, 1960: Cairns, p. 24. Turbinolia barbadensis Wells, 1945: Cairns, p. 24. Turbinolia barbadiana var. crassicostata Wells, 1945: Cairns, p. 24. Turbinolia dickersoni Nomland, 1916: Cairns, p. 24. Turbinolia frescoenis Barta-Calmus, 1969: Cairns, p. 24.

Dimensions. d=up to 3.75 mm; s=24; costae=24–48; h=2–10.5 mm. Description. Trochoid corallum, Corallum elongate, slender, conical, straight; costosepta developed in 3 complete cycles in 6 systems; S1 and S2 strongly fused to columella; S3 unite with S1; 24 costae correspond to septa, in larger specimens additional 24 costae not representing septa present. Remarks. In having 24 septa arranged in 3 cycles in 6 systems and a corallite diameter of up to 3.5 mm the species T. barbadiana corresponds very well with the taxon Turbinolia sulcata Lamarck from the Middle Eocene of France. However, no septa of the latter fuse with the columella or with each other as they do in T. dickersoni. Type locality of species. Lower Eocene of the USA (California). Distribution. Paleocene of Ivory Coast and the USA (Alabama, Mississippi), Eocene of Mexico and the USA (Alabama, Louisiana, Mississippi, and Texas), Lower Eocene of Barbados (Upper Scotland Formation) and the USA (California), Upper Eocene of Colombia, Oligocene of Peru (Mirador facies).



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Text-Fig. 17 Turbinolia dickersoni Nomland, 1916. A, B, as figured in Howe (1960, as holotype of Turbinolia rosetta Howe, 1960), Paleocene of the USA (Alabama), scale bar: 2.5 mm; A, longitudinal view; B, cross view.

Genus Bothrophoria Felix, 1909 Type species. Bothrophoria ornata Felix, 1909, Maastrichtian of Antarctic Peninsula.

Diagnosis. Solitary, conical-cuneiform, free, with rounded base. Calice elliptical in outline. Costosepta compact, finely granulated laterally. Costae serrated, pali before all but the last cycle, P3 and P4 lamellar and broad. Columella papillose to spongy.

Bothrophoria ornata Felix, 1909 Pl. 8, figs 5a, b (v)*1909 (v)1994 (v)1997 2000 v2002

Bothrophoria ornata nov. gen. nov. sp.: Felix, p. 9, pl. 1, figs 8–11. Bothrophoria ornata Felix, 1909: Filkorn, p. 42, figs 14 (1–4)–15 (1–4) (older synonyms cited therein). Bothrophoria ornata Felix, 1909: Cairns, p. 18, figs 1L, 4L. Bothrophoria ornata, Felix 1909: Löser, p. 15. Bothrophoria ornata Felix, 1909: Baron-Szabo, p. 174, pl. 122, figs 1–2.

Dimensions. d (min)=4.6–8 mm; d (max)=6–11 mm; d (min)/d (max)=0.67–0.89; s=48–62; h=5.5–10.5 mm. Description. Turbinate-subcuneiform corallum; costosepta arranged in 4 cycles in 6 systems; S1 and S2


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nearly equal; pali long, sometimes discontinuous; columella spongy, well-developed. Type locality of species. Maastrichtian of Antarctic Peninsula (Snow Hill Island). Distribution. Campanian-Maastrichtian of Antarctic Peninsula (Seymour Island and Snow Hill Island).

Genus Alveolocyathus Filkorn, 1994 Type species. Alveolocyathus nordenskjoeldi Filkorn, 1994, Maastrichtian of Antarctic Peninsula (López de Bertodano Formation, Seymour Island).

Diagnosis. Solitary, conical, free, circular in outline with pointed base. Costosepta compact, hexamerally arranged, Dentate laterally. Costae serrate. Higher cycle costae (C3–4) originate by trifurcation. Pali in front of S2, and lamellar. Columella papillose to rudimentary.

Alveolocyathus felixi (Filkorn, 1994) Pl. 8, fig. 3 v*1994 v1994 v1994 v1997 v1997 v1997 2000 2000 v2002 v2002

Conocyathus felixi n. sp.: Filkorn, p. 47, figs 16 (1–4). Alveolocyathus nordenskjoeldi n. gen. n. sp.: Filkorn, p. 61, figs 21 (1–4). Laminocyathus wellsi n. sp.: Filkorn, p. 64, figs 22 (1–4). Laminocyathus wellsi Filkorn, 1994: Cairns, p. 19, figs 2b, 5d. Alveolocyathus nordenskjoeldi: Cairns, p. 60–61, pl. 2, fig. 2c; Pl. 5, fig. 5e. Conocyathus felixi: Cairns, p. 61. Alveolocyathus nordenskjoeldi, Filkorn 1995: Löser, p. 7. Laminocyathus wellsi, Filkorn 1995: Löser, p. 45. Alveolocyathus nordenskjoeldi Filkorn, 1994: Baron-Szabo, p. 174, text-fig. 72. Laminocyathus wellsi Filkorn, 1994: Baron-Szabo, p. 175, text-fig. 73.

Dimensions. d=7.5–8.3 mm; s=48; h=7.5 mm Description. Conical with pointed base; circular in outline; costosepta arranged in 4 cycles in 6 systems. Remarks. Filkorn (1994, p. 47 and 61) described the forms Conocyathus felixi and Alveolocyathus nordenskjoeldi. Later, Cairns (1997, p. 20) united these two forms, considering Conocyathus felixi as the junior synonym. However, because the latter was established 14 pages before A. nordenskjoeldi, it has to be considered the senior synonym. The validity of A. nordenskjoeldi as the type species of the genus Alveolocyathus is not affected by its position as a junior synonym (ICZN 67.1.2). Type locality of species. Maastrichtian of Seymour Island (Antarctic Peninsula). Distribution. Maastrichtian of Seymour Island (Antarctic Peninsula).

Genus Wellsotrochus Squires, 1960 (=Kangiliacyathus Floris, 1972; Type species. Kangiliacyathus groenlandicus Floris, 1972, Lower Danian of Denmark [Greenland]); (=Levicyathus Filkorn, 1994, Type species. Levicyathus cairnsi Filkorn, 1994, Maastrichtian of Antarctic Peninsula [López de Bertodano Formation, Seymour Island]).



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Type species. Wellsia cyathiformis Squires, 1958, Campanian-Maastrichtian (Haumurian) of New Zealand.

Diagnosis. Solitary, discoidal, bowl-shaped to variably conical, ?free, with flat, rounded, or somewhat pointed base. Calice circular to slightly elliptical in outline. Costosepta compact, hexamerally arranged. Costae presumed granular. Pali and paliform lobes absent. Columella styliform to sublamellar. Wall imperforate. Remarks. In creating the genus Wellsotrochus (ex Wellsia) Squires (1958, p. 57) described its growth form simply as “bowl-shaped” but he included specimens of various growth types, e.g., patellate, turbinatetrochoid, and bowl-shaped, indicating a wide range of growth types as one characteristic of this genus. It should be noted, however, that in most specimens the early juvenile stage is closest to the turbinate form.

Wellsotrochus cyathiformis (Squires, 1958) Text-Fig. 18 v*1958 1962 1962 1972 v1994 v1997 1997 1997 2000 2000 2002 v2002

Wellsia cyathiformis n. sp.: Squires, p. 57, pl. 10, figs 16–20. Wellsotrochus cyathiformis (Squires): Squires, p. 145. Wellsotrochus conicus, n. sp.: Squires, p. 145, pl. 3, figs 6–7. Kangiliacyathus groenlandicus gen. n. & sp. n.: Floris, p. 67, pl. 4, figs 2A–3. Levicyathus cairnsi n. sp.: Filkorn, p. 60, figs 20 (1–4). Levicyathus cairnsi Filkorn, 1994: Cairns, p. 18, figs 2d and 5a. Wellsotrochus cyathiformis (Squires, 1958): Cairns, p. 21, fig. 2j. Wellsotrochus conicus Squires, 1958: Cairns, p. 22. Levicyathus cairnsi, Filkorn 1995: Löser, p. 48. Wellsotrochus cyathiformis, Squires 1958: Löser, p. 86. Wellsotrochus cyathiformis (Squires, 1958): Baron-Szabo, p. 175, text-fig. 74. Levicyathus cairnsi Filkorn, 1994: Baron-Szabo, p. 176, pl. 122, figs 5–6.

Dimensions. d (min)=3.5–8 mm; d (max)=5–9.5 mm; d (min)/d (max)=0.61–1; s=24–48+s5; h=3–6.3 mm. Description. Solitary, variably conical or bowl-shaped to discoidal; circular to elliptical in outline; costosepta arranged in 3 to 4 cycles in 6 systems; septa of a beginning fifth cycle in largest specimens present. Type locality of species. Campanian-Maastrichtian (Haumurian) of New Zealand. Distribution. Campanian-Maastrichtian of New Zealand, Maastrichtian of Antarctic Peninsula, Lower Danian of Denmark (Greenland).

Wellsotrochus discus Squires, 1962 Fig. 18 v*1962 1997

Wellsotrochus discus, n. sp.: Squires, p. 146, pl. 1, figs 6–8. Wellsotrochus discus Squires, 1958: Cairns, p. 22.

Dimensions. d=9.4–11.7 mm; s=48; h=up to 5 mm. Description. Solitary, bowl-shaped to discoidal; circular to elliptical in outline; costosepta arranged in 4 cycles in 6 systems, alternating in length and thickness. Type locality of species. Campanian-Maastrichtian (Haumurian) of New Zealand. Distribution. Campanian-Maastrichtian of New Zealand.


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Text-Fig. 18 A, Wellsotrochus discus Squires, 1962, holotype, Co 1303, based on the illustrations in Squires (1962), cross view, polished surface, Campanian-Maastrichtian of New Zealand, scale bar: 5 mm. B, Wellsotrochus discus Squires, 1962, paratype, Co 1304, based on the illustrations in Squires (1962), longitudinal view, Campanian-Maastrichtian of New Zealand, scale bar: 4 mm. C-F, Wellsotrochus cyathiformis (Squires, 1958). C, Lower Danian of Denmark (Greenland) as illustrated in Floris (1972, figured as type material of Kangiliacyathus groenlandicus Floris, 1972), scale bar: 2 mm. D, holotype of Wellsotrochus cyathiformis (Squires, 1958), AUG 9a, Campanian-Maastrichtian (Haumurian) of New Zealand, base of specimen, oblique, scale bar: 2.5 mm (photograph courtesy S. Cairns); E, F, Maastrichtian of Antarctic Peninsula (USNM 93038, holotype of Levicyathus cairnsi Filkorn, 1994) (photograph courtesy S. Cairns), scale bar: 2 mm; E, cross view, upper surface; F, longitudinal view, upper surface. SCLERACTINIAN CORALS OF THE K/T-BOUNDARY


95

Genus Palocyathus Filkorn, 1994 Type species. Palocyathus seymourensis Filkorn, 1994, Maastrichtian of Antarctic Peninsula (López de Bertodano Formation, Seymour Island).

Diagnosis. Solitary, conical-cuneiform, free, with pointed base and circular in outline. Costosepta compact, exsert, hexamerally arranged, finely granulated laterally. Costae narrow, beaded ridges, higher cycle costae (C3–4) originate by trifurcation near corallum base. Pali in front of S2 and S3, small. Columella papillose.

Palocyathus seymourensis Filkorn, 1994 Pl. 8, fig. 4 v*1994 v1997 2000 v2002

Palocyathus seymourensis n. sp.: Filkorn, p. 57, figs 19 (1–4). Palocyathus seymourensis Filkorn, 1994: Cairns, p. 18, figs 1k, 4k. Palocyathus seymourensis, Filkorn 1995: Löser, p. 58. Palocyathus seymourensis Filkorn, 1994: Baron-Szabo, p. 176, text-fig. 75.

Dimensions. d (min)=4.4–8.2 mm; d (max)=4.4–9.1 mm; d (min)/d (max)=0.85–1; s=48; h=3.2–8.5 mm. Description. Solitary, turbinate-cuneiform; intercostal regions greater than width of costae and exteriorly pitted; costosepta arranged in 4 complete cycles in 6 systems; pali in front of S3 usually fused to their common S2. Type location of species. Maastrichtian of Antarctic Peninsula (López de Bertodano Formation, Seymour Island). Distribution. Maastrichtian of Antarctic Peninsula. Genus Platytrochus Milne Edwards & Haime, 1848 (=Aldrichiella Vaughan, 1900, Type species. Aldrichiella elegans Vaughan, 1900, Upper Eocene [Jacksonian] of the USA); (=Dominicotrochus Vaughan, in Vaughan & Hoffmeister, 1925, Type species. Dominicotrochus dominicensis Vaughan, in Vaughan & Hoffmeister, 1925, Middle Miocene of the Dominican Republic, Martinique); (=Koilotrochus Tenison-Woods, 1878, Type species. Koilotrochus vacuus Tenison-Woods, 1878, Middle Miocene of Australia [Victoria]) Type species. Turbinolia stokesii Lea, 1833, Middle Eocene of the USA (Claibornian) (subsequent designation Milne Edwards & Haime, 1850a).

Diagnosis. Solitary, corallum laterally compressed, often cuneiform, with calice elliptical in cross section and rarely more than 10 mm in diameter. Alate thecal edge costae may be present or absent. Costae smooth granular, type species having coarsely granular costae. Intercostae and thecal pits absent. Higher-cycle costae originate by trifurcation. Septa exsert, hexamerally arranged in 3–4 cycles, and in some cases (e.g. P. stokesii) separated from their corresponding costae by notch at calicular edge. Pali and paliform lobes absent. Columella papillose, consisting of 2 or more rows of papillae.


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Platytrochus claibornensis de Gregorio, 1890 Pl. 8, Fig. 6 *1890 v1935 (v)1975 (v)1987 1997 1997

Platytrochus Claibornensis de Greg.: de Gregorio, p. 255, pl. 45, figs 21, 22. Platytrochus primaevus n. sp.: Vaughan & Popenoe, p. 336, pl. 3, figs 19, 20. Platytrochus claibornensis Gregorio, 1890: Kuzmicheva, p. 21, pl. 2, figs 3a, 3b. Platytrochus claibornensis Gregorio, 1890: Kuzmicheva, p. 140, pl. 24, figs 10a, 10b. Platytrochus primaevus Vaughan and Popenoe, 1935: Cairns, p. 29. Platytrochus claibornensis (de Gregorio, 1890): Cairns, p. 29.

Dimensions. d (min)=3–4 mm; d (max)=4.5–6 mm; d (min)/d (max)=0.5–0.67; s=36–44; h=up to 9 mm. Description. Conical corallum, compressed in outline; costosepta arranged in 4 incomplete cycles in 6 systems; S1, S2, and some of S3 extend to corallite centre. Type locality of species. Middle Eocene (Upper Claibornian) of the USA (Alabama). Distribution. Middle-Upper Paleocene of Ukraine, Paleocene (Midwayan, Texas)-Middle Eocene (Upper Claibornian, Alabama) of the USA.

Suborder Fungiina Verrill, 1865 Diagnosis. Solitary and colonial. Septa fenestrate, formed by simple or compound trabeculae united by simple or compound synapticulae, margins beaded or dentate.

Family Fungiidae Dana, 1846 (=?Funginellidae Alloiteau, 1952a; =Asteroseriidae Alloiteau, 1952a; ?Cyclastraeidae Alloiteau, 1952a) Diagnosis. Solitary and colonial, fixed or free, mostly hermatypic. Solitary corallites discoidal or elongateoval, aboral surface flat or concave. Colonies discoidal or elongate-oval, free or foliaceous and fixed. Colony formation by incomplete intratentacular polystomodaeal budding. Wall synapticulothecal, often secondarily septothecal or thickened. Septa numerous, fenestrate in early stages, perforate or solid, composed of a single fan system of compound trabeculae producing simple or compound marginal dentations, laterally united by stout compound synapticulae. Axis of trabecular divergence horizontal in ephebic stage. Costae continuous or broken into spinose projections. Columella trabecular, feeble. Dissepiments absent. Epitheca only in early anthoblast stage. Homeomorphic with Cunnolitidae.

Genus Cycloseris Milne Edwards & Haime, 1849 (=?Funginella d'Orbigny, 1850, Type species. Funginella neocomiensis d'Orbigny, 1850, Hauterivian of France); (=Asteroseris Fromentel, 1870, Type species. Asteroseris coronula Fromentel, 1870, Cenomanian of France); (=?Cyclastraea Alloiteau, 1952a, Type species. Cyclolites spinosa Fromentel, 1864, Cenomanian of France); (=?Cyclophyllopsis Alloiteau, 1952a; Type species. Cyclolites aptiensis Fromentel, 1870, Aptian of France); (=Pseudocycloseris Alloiteau, 1957, Type species. Pseudocycloseris uxacalcensis Alloiteau, 1957, Turonian of France). Type species. Fungia cyclolites Lamarck, 1801, Recent, Pacific Ocean, off Australia (subsequent designation Milne Edwards & Haime, 1849a).

Diagnosis. Solitary, discoidal to cupolate. Costosepta compact, subcompact in younger cycles, with acute SCLERACTINIAN CORALS OF THE K/T-BOUNDARY


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dentations. (?Pseudo-) Columella trabecular. Synapticulae present. Endothecal dissepiments absent. Wall synapticulothecal.

Cycloseris (?) wellsi Baron-Szabo, 2002 Text-Fig. 19 parsv1934 v*2002

Cyclolites jamaicaensis, new species: Wells, p. 84–85, non specimen 3 (paratype). Cycloseris (?) wellsi sp. nov.: Baron-Szabo, p. 98, pl. 65, fig. 2.

Text-Fig. 19 Cycloseris (?) wellsi Baron-Szabo, 2002, holotype USNM I-147658, Campanian-Maastrichtian of Jamaica, upper surface, scale bar: 10 mm.

Dimensions. d=32 x 34 mm; s/mm=6/2 ; h=12 mm. Description. Cupolate corallum; septa arranged in complete cycles in 6 systems; oldest septa more dominant; columella trabecular, net-like, fused with axial ends of septa. Type locality of species. Campanian-Maastrichtian of Jamaica. Distribution. Campanian-Maastrichtian of Jamaica.


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Family Fungiacyathidae Chevalier, 1987 Diagnosis. Corallum simple, cupolate, free. Septotheca horizontal to concave and usually quite fragile. Septa and costae in direct correspondence with one another, 48 or 96 of each. Costae usually thin and serrate but may be rounded and granular. Septa imperforate, composed of a single fan system of widely spaced compound trabeculae that often terminate in tall septal spines. Septal faces usually carinate. Adjacent septa united by T- or Y- shaped synapticular plates that originate from the thecal base. Pali sometimes present. Columella spongy to irregular trabecular. Dissepiments absent. Polyp completely invests corallum. Exclusively ahermatypic (see Chevalier & Beauvais 1987). Remarks. The diagnosis represents the emended version of this family given by Cairns (1989, p. 5), who provided detailed discussions regarding the Family Fungiacyathidae.

Genus Fungiacyathus Sars, 1872 Type species. Fungiacyathus fragilis Sars, 1872, Recent, northern Atlantic.

Diagnosis. Having the characters of the family.

Fungiacyathus deltoidophorus (Felix, 1909) Text-Fig. 20 ?1902 *1909 1909 1909 cf.v1945 1949 v1994 v1994 v1994 ?1996 2000 2000 2000 2002 2002 2002

?Brachycyathus daniensis n. sp.: Wanner, p. 98, pl. 13, figs 10–10b. Cycloseris deltoidophora nov. sp.: Felix, p. 4, pl. 1, figs 1–2. Cycloseris antarctica nov. sp.: Felix, p. 7, pl. 1, figs 3–4. Cycloseris larseni nov. sp.: Felix, p. 8, pl. 1, figs 5–6. Fungiacyathus sp.: Wells, p. 7, pl. 2, figs 7–8. Cycloseris antarctica Felix: Furque & Camacho, p. 276. Fungiacyathus deltoidophorus (Felix, 1909): Filkorn, p. 29, figs 6–9 (older synonyms cited therein) (topotypes studied). Fungiacyathus larseni (Felix, 1909): Filkorn, p. 34, figs 10–11. Fungiacyathus antarcticus (Felix, 1909): Filkorn, p. 37, fig. 12. Discocyathus daniensis (Wanner, 1902): Schuster, Appendix list. Cycloseris antarctica, Felix 1909: Löser, p. 27. Cycloseris deltoidophora, Felix 1909: Löser, p. 27. Cycloseris larseni, Felix 1909: Löser, p. 27. Fungiacyathus antarcticus (Felix, 1909): Baron-Szabo, p. 98. Fungiacyathus deltoidophorus (Felix, 1909): Baron-Szabo, p. 98. Fungiacyathus larseni (Felix, 1909): Baron-Szabo, p. 98.

Dimensions. d=4–26 mm; s=48–96. Description. Solitary, cupolate, free; base of corallum flat or convex; septa correspond with costae; columella weakly developed.



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Text-Fig. 20 Fungiacyathus deltoidophorus (Felix, 1909), topotype, USNM 93024, Maastrichtian of Antarctic Peninsula, upper surface, oblique, scale bar: 2 mm (from Filkorn 1994, see full reference in Reference list; copyright 1994 by the American Geophysical Union; reproduced by permission of American Geophysical Union).

Remarks. Based on corallite diameter and number of septa, Felix (1909; see also Filkorn 1994) distinguished 3 different species: Fungiacyathus antarcticus (Felix, 1909), F. deltoidophorus (Felix, 1909), and F. larseni (Felix, 1909): d=4–14 mm and 48 septa in F. deltoidophorus, d=up to 14 mm and 48 septa in F. larseni, and d=up to 26 mm and 48–96 septa in F. antarcticus. The large overlap in the dimensions of the skeletal elements of F. deltoidophorus and F. larseni was previously pointed out by Filkorn (1994, p. 33–34). In addition to the dimensions of skeletal elements, Felix also included characteristics like the development of the bottom (flat or convex) and the length of the costae and relative size difference of the finely granulated central region of the aboral surface. All of these characters are interpreted as environmentally induced, and are therefore non-specific variations. Moreover, the form F. antarcticus appears to correspond to a later ontogenetical stage of F. larseni and F. deltoidophorus. Therefore, the three species are considered synonymous. In having 48 septa in a corallite of around 8 mm in diameter, the specimen described from the Lower Eocene of Barbados by Wells (1945) fits well into the above proposed model and closely corresponds to the species F. deltoidophorus. However, because Wells’ specimen represents a fragment its assignment is provisional. From the Danian of Egypt Wanner (1902) described the species ?Brachycyathus daniensis. Because the


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genus Brachycyathus is generally considered a junior synonym of Discocyathus (for generic characteristics see chapter on Discocyathus in this paper), Wanner’s species was transferred to the latter genus (e.g., Schuster 1996). However, in lacking pali before S3 the species B. daniensis differs from the genus Discocyathus, whereas the development of septa (compact septa that seem to be granulated to carinate laterally), the lack of endothecal structures, and the presence of a trabecular columella, supports the idea that the taxon B. daniensis belongs to the genus Fungiacyathus. In addition, Wanner’s species is characterized by 48 septa in corallite diameters ranging between 8–16 mm, thus closely agreeing with the species F. deltoidophorus. However, because of the problematic preservation and the fact that Wanner’s type material is lost and could therefore not be studied, its synonymy with the latter species is provisional. Type location of species. Maastrichtian of Antarctic Peninsula. Distribution. Campanian-Maastrichtian of Antarctic Peninsula (Seymour Island and Snow Hill Island), ?Paleocene of Egypt, Middle Eocene of Barbados (Upper Scotland Formation).

Family Acrosmiliidae Alloiteau, 1952 (=Leptophylliidae Vaughan, 1905, p.p.) Diagnosis. Solitary. Wall synapticulothecal. Costosepta perforated, formed by compound trabeculae, perforations mainly restricted to the axial region, compact peripherally. Distal margin of costae and septa moniliform. Endotheca well-developed.

Genus Acrosmilia d’Orbigny, 1849 (=Turbinoseris Duncan, 1873, Type species. Turbinoseris eocaenica Duncan, 1873, Eocene [limestone of St. Bartholomew] of the West Indies) Type species. Turbinolia cernua Michelin, 1846, Coniacian of France (Aude).

Diagnosis. Solitary, turbinate to patellate. Costosepta compact peripherally, perforate inwardly and near upper margins, strongly beaded marginally, granular and pennular laterally. Columella feebly developed, parietal. Synapticulae abundant. Endothecal dissepiments thin, vesicular to subtabulate. Synapticulothecate. Septal microstructure consists of compound trabeculae.

Acrosmilia conica d'Orbigny, 1850 Text-Fig. 21 v*1850 1851b v1854 1857 v1858–61 vpars1867 1880 1880 1880

Acrosmilia conica: d'Orbigny, vol 2, p. 203. Trochosmilia ? conica (d'Orbigny): Milne Edwards & Haime, p. 47. Leptophyllia clavata: Reuss, p. 101, pl. 6, figs 3–6. Trochosmilia ? conica (d'Orbigny): Milne Edwards, vol. 2, p. 165. Acrosmilia conica: de Fromentel, p. 100. Leptophyllia conica: de Fromentel, p. 302, pl. 45, non figs 2 a–c. Turbinoseris epithecata, Duncan: Duncan, p. 49, pl. 7, figs 8–9. Turbinoseris Ranikoti, Duncan: Duncan, p. 49, pl. 7, figs 10–11. Turbinoseris Haimei, Duncan: Duncan, p. 50, pl. 7, figs 4–7.



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1880 v1903a 1914 1914 1930a 1941a ?1956 ?1978 1981 1982 1986 v1996 v1999 2000 2000 2000 2002 v2002 2002 v2004

Turbinoseris Indica, Duncan: Duncan, p. 50, pl. 7, figs 1–3. Leptophyllia clavata Reuss: Felix, p. 200. Leptophyllia clavata Reuss 1854: Felix, pars 7, p. 194. Leptophyllia conica d'Orbigny sp. 1850: Felix, pars 7, p. 194. Leptophyllia conica d'Orbigny: Oppenheim, p. 148–150, pl. 16, figs 6–9, pl. 20, figs 1–2. Haplaraea ? discrepans, n. sp.: Wells, p. 289, pl. 1, fig. 2. Acrosmilia clavata (Reuss): Wells, part F, p. F. 385, fig. 277.2. Acrosmilia conica d'Orbigny 1850: Turnšek, p. 82, 113, pl. 16, figs 1–7. Trochosmilia tuba ? Fromentel: Abed & El-Asaad, p. 275, pl. 1, figs 3a–b. Acrosmilia conica d'Orbigny 1850: Beauvais, vol. 2, p. 138–140, pl. 40, figs 2–3 (older synonym cited herein). Acrosmilia conica (d'Orbigny 1850): Tchéchmédjiéva, p. 68ff. Leptophyllia clavata Reuss, 1854: Eliášová, pl. 2, fig. 4, pl. 4, fig. 2. Acrosmilia conica d'Orbigny, 1850: Baron-Szabo, p. 459, pl. 1, fig. 3. Acrosmilia conica d'Orbigny, 1850: Löser, p. 5. Haplaraea discrepans Wells, 1941: Löser, p. 39. Leptophyllia clavata, Reuss 1854: Löser, p. 47. Acrosmilia clavata (Reuss, 1854): Baron-Szabo, p. 99. Acrosmilia conica d'Orbigny, 1850: Baron-Szabo, p. 99, pl. 66, figs 1, 3, 5. Haplaraea discrepans Wells, 1941: Baron-Szabo, p. 103. Acrosmilia conica d'Orbigny: Moosleitner, p. 177, pl. 75, figs 12, 13.

Dimensions. d (min)=22–30 mm; d (max)=25–33 mm; d (min)/d (max)=0.85–0.96; s=96–192, s/mm=9–11/5.

Text-Fig. 21 Acrosmilia conica d'Orbigny, 1850. A, Paleocene of India (type material of Turbinoseris ranikoti Duncan, 1880), as figured in Duncan (1880), upper surface, lateral view, scale bar: 14 mm; B, close-up of A (magnification estimated because no data were given by Duncan), scale bar: 6 mm.


BARON-SZABO

Description. Turbinate to subpatellate corallum, subcircular to elliptical in outline; septa subcompact (primary, secondary and tertiary septa) or porous (quaternary and younger septa), thin, have rounded or spiny granules and a few pennulae laterally, arranged in 5–6 complete cycles in 6 systems; up to 24 septa extend to the center of the calice, where trabecular prolongations of their inner ends may meet and fuse with the columella. Younger septa alternate in length. The columella is weakly developed, papillose. Synapticulae are irregularly disposed. Endotheca consists of thin, subtabulate dissepiments. The wall is synapticulothecal. In places an epithecal wall (mono- or multilamellar) is present. Type locality of species. Coniacian of France. Distribution. Upper Turonian-Campanian of Austria (Gosau Group), Lower Coniacian-Upper Santonian of southern France, ?Senonian of Georgia (in Caucasia), ?Santonian of Slovenia/northwestern Croatia, Campanian of Cuba, Upper Campanian of Bulgaria, Campanian-Lower Maastrichtian of central Saudi Arabia, Upper Maastrichtian of Jamaica (this paper), Paleocene of India. New Material. Upper Maastrichtian of Jamaica, NMNH, Coates coll., sample no. J-71-42-16L2 (=Rio Minho).

Family Actinacididae Vaughan & Wells, 1943 Diagnosis. Colonial; colony formation by intra- and extratentacular budding. Basal epitheca present. Corallites synapticulothecate. Septa relatively few, composed of one system of 3 to 5 nearly vertical simple trabeculae, with numerous pores, laterally united by simple synapticulae, the innermost trabeculae often differentiated as paliform lobes. Septocostae often scarcely distinguishable, even absent when the distal ends of the septa bifurcate and become lost in a coenenchyme composed of discontinuous vertical trabeculae united horizontally by simple synapticulae; coenenchyme spinose or vermiculate on the surface. Columella absent or composed of one or more trabecular pillars. Endotheca thin, subtabulate.

Genus Actinacis d’Orbigny, 1849 Type species. Actinacis martiniana d’Orbigny, 1850, Upper Santonian of France (Figuères).

Diagnosis. Colony plocoid. Colony formation by extracalicinal budding. Corallites are embedded in a reticulate, porous coenosteum. Costosepta have few, but large perforations. Anastomosis present. Septal flanks granular. Wall synapticulothecate, incomplete. Columella parietal or substyliform or formed by elongated segments. No pali. Synapticulae abundant. Endothecal dissepiments sparse. Skeletal microstructure consists of simple and compound trabeculae.

Actinacis barretti Wells, 1934 Pl. 8, fig. 10 v*1934 v1960 1974 1974

Actinacis barretti: Wells, p. 101, pl. 4, figs 1–2. Actinacis barretti Wells: Berryhill, et al., p. 151. Actinacis caribiensis: Frost & Langenheim, p. 217, pl. 71, figs 1–5. Actinacis sp. cf. A. barretti Wells: Frost & Langenheim, p. 219, pl. 72, figs 1–5.



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Porites anguillensis Vaughan: Frost & Langenheim, p. 221, pl. 75, figs 1–3. Goniopora copoyensis n. sp.: Frost & Langenheim, p. 238, pl. 81, figs 5–6. Actinacis alabamiensis: Bryan, p. 426ff. Actinacis barretti Wells, 1934: Budd et al., p. 593. Actinacis caribiensis Frost and Langenheim, 1974: Budd et al., p. 593. Actinacis alabamiensis: Stemann, in Bryan et al., p. 33, text-fig. 1A. Actinacis barretti Wells, 1934: Vecsei & Moussavian, p. 131, pl. 36, fig. 3.

Dimensions. d=0.8–1.3 mm; c-c=1.5–2.5 mm; s=16–22, in juvenile corallites the number may be 12, in oldest corallites the number may reach 24. Description. Colonial, submassive to ramose, plocoid; gemmation extracalicinal; costosepta subcompact, thin, granulated laterally, arranged in varying systems; columella papillose to substyliform; wall synapticulothecate, incomplete; pali absent; synapticulae abundant. Endothecal dissepiments sparse. Remarks. Re-examination of the type specimen of Actinacis barretti revealed that the dimensions of the skeletal elements considerably vary from those originally given by Wells (1934). He reported 24 septa arranged in 6 systems and a corallite diameter of not more than 1 mm. From the Middle Eocene of Mexico Frost & Langenheim (1974) described the species Actinacis caribiensis which is supposed to be characterized by a corallite diameter of 0.7–1.2 mm and 16 septa developed in 8 systems. However, in the holotype the number of septa ranges mainly from 14–20 (12 septa in juvenile corallites) and the septal arrangement is generally in 8 but also in 6 systems. Therefore, it is considered a junior synonym of Wells‘ species. Type locality of species. Middle Eocene of Jamaica (Yellow Limestone). Distribution. Paleocene of Italy (Maiella Platform) and the USA (Alabama, Salt Mountain Limestone), Danian of Puerto Rico, Middle Eocene of Jamaica (Yellow Limestone) and Mexico (San Juan Formation), Upper Oligocene of Mexico (La Quinta Formation).

Actinacis haueri Reuss, 1854 Pl. 8, fig. 9 v*1854 1858–61 1860 v1903a 1914 v1925 v1930a non1956 (v)1972 v1982 2000 2000 v2001 v2002 v2002 v2006

Actinacis haueri m.: Reuss, p. 128, pl. 8, figs 13–14 (topotypes studied). Actinacis haueri: de Fromentel, p. 250. Actinacis haueri: Milne Edwards, vol. 3, p. 170. Actinacis haueri Reuss: Felix, p. 176, text.fig. 2. Actinacis haueri Reuss 1854: Felix, pars 7, p. 240. Montipora cretacea spec. nov.: Umbgrove, p. 120, pl. 8, fig. 4. Actinacis haueri Reuss: Oppenheim, p. 6, pl. 1, figs 7–8 (topotypes studied). Actinacis haueri Reuss: Bendukidze, p. 119. ‘Coral’: Samuel et al., pl. 156, fig. 1. Actinacis haueri Reuss 1854: Beauvais, vol. 2, p. 268, pl. 48, fig. 3 (topotypes studied). Actinacis haueri, Reuss 1854: Löser, p. 5. Montipora cretacea, Umbgrove 1926: Löser, p. 53. Actinacis haueri Reuss, 1854: Baron-Szabo, p. 260, fig. 1 G. Actinacis haueri Reuss, 1854: Baron-Szabo, p. 100, pl. 69, fig. 3. Montipora cretacea Umbgrove, 1925: Leloux, p. 15, pl. 1, fig. 2. Actinacis haueri Reuss, 1854: Baron-Szabo et al., p. 19, fig. 5.1.


BARON-SZABO

Dimensions. d=0.6–1 mm; c-c=2–3 mm; s=22–24. Description. Ramose, plocoid colony; corallites circular; costosepta arranged in 2–3 cycles in 6 systems; columella made of several segments. Remarks. Because the holotype was lost, Beauvais (1982, vol. 2, p. 269) chose a neotype from the Santonian marls of the Zimmergraben (Gosau Group, Austria). Type locality of species. Santonian of Austria (Zimmergraben at Gosau Group). Distribution. Upper Turonian-Coniacian (Theresienstein reef) and Coniacian-Santonian (Gosau Group) of Austria, Lower Coniacian and Upper Santonian of southern France, Maastrichtian of Mexico (Cardenas Formation), Upper Maastrichtian of The Netherlands, Montian-Thanetian of Slovakia (Western Carpathians).

Actinacis martiniana d’Orbigny, 1850 Pl. 8; fig. 11 *1850 nonv1854 1881 1906 1914 1914 v1930a v1930a v1930a v1933 1956 1958 v1982 v1982 v1982 1987 v1988 1993 1995 nonv1997 non 1997 v1998 v1998 2000 2000 2000 v2000 v2001 nonv2002 v2002 non2003

Actinacis Martiniana d’Orb., 1849: d’Orbigny, vol. 2, p. 209. Actinacis Martiniana d’Orbigny: Reuss, p. 127, pl. 24, figs 12–15. Actinacis haueri Reuss: Quenstedt, vol. 6, p. 900, pl. 178, fig. 28. Actinacis cymatoclysta n. sp.: Felix, p. 43, pl. 3, figs 4–4a. Actinacis cymatoclysta Felix 1906: Felix, pars 7, p. 239. Actinacis Martiniana d’Orbigny 1849: Felix, pars 7, p. 240. Actinacis parvistella n. sp.: Oppenheim, p. 9, pl. 1, figs 3, 3a (topotypes studied). Actinacis multilamellata n. sp.: Oppenheim, p. 13, pl. 15, fig. 2 (topotypes studied). Actinacis multipartita: Oppenheim, p. 20, pl. 15, figs 4, 4a (topotypes studied). Actinacis valverdensis: Wells, p. 120, pl. 11, figs 1, 2. Actinacis haueri Reuss: Bendukidze, p. 119. Actinacis martiniana d’Orb.: Alloiteau, p. 177, pl. 29, figs 1–3. Actinacis parvistella Oppenheim 1930: Beauvais, vol. 2, p. 273, pl. 49, figs 1, 2, pl. 69, fig. 3 (topotypes studied). Actinacis multilamellata Oppenheim 1930: Beauvais, vol. 2, p. 274, pl. 49, figs 3, 4, pl. 69, fig. 2 (topotypes studied). Actinacis multipartita Oppenheim: Beauvais, vol. 2, p. 277, pl. 49, fig. 6, pl. 50, fig. 1 (older synonyms cited therein) (topotypes studied). Actinacis cognata Oppenheim, 1901: Kuzmicheva, p. 159, pl. 29, fig. 1. Actinacis cognata Oppenheim, 1901: Drobne, et al., p. 188, pl. 33, figs 2–4. Actinacis cognata Oppenheim, 1901: Alvarez Perez, p. 230, pl. 22, figs 1–3. Actinacis cognata Oppenheim, 1901: Bosellini & Russo, p. 218, pl. 1, figs 1–4, text-figs 2–3. Actinacis martiniana d’Orbigny, 1850: Baron-Szabo, p. 78, pl. 11, figs 2, 5–6. Actinacis martiniana d’Orbigny, 1850: Turnšek, p. 4, pl. 4, figs A–C. Actinacis cognata Oppenheim, 1901: Turnšek, in Turnšek & Drobne, p. 137, pl. 8, figs 3–4. Actinacis parvistella Oppenheim, 1930: Baron-Szabo, p. 147, pl. 11, fig. 1. Actinacis cymatoclysta, Felix 1906: Löser, p. 5. Actinacis martiniana, d’Orbigny, 1850: Löser, p. 5. Actinacis parvistella, Oppenheim 1930: Löser, p. 5. Actinacis parvistella Oppenheim, 1930: Baron-Szabo, p. 118, pl. 11, figs 1, 3. Actinacis parvistella Oppenheim, 1930: Baron-Szabo, p. 258, figs 1A, 2E. Actinacis martiniana d’Orbigny, 1850: Baron-Szabo, p. 100, pl. 67, figs 3–4. Actinacis parvistella Oppenheim, 1930: Baron-Szabo, p. 100, pl. 67, fig. 6, pl. 68, figs 1–2. Actinacis martiniana d’Orbigny 1850: Götz, p. 5ff., pl. 1, fig. 2.



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Actinacis parvistella Oppenheim, 1930: Baron-Szabo et al., p. 20, fig. 5.2.

Dimensions. d=1–1.2 mm, late adult stages up to 1.5 mm, juvenile around 0.8 mm; c-c=1.5–3 mm; s=18–22, up to 24 septa in late adult stages. Description. Encrusting-lamellar or irregular multicolumnar-ramose, plocoid colony; calices circular or slightly oval in outline, separated by a vermiculate coenosteum; costosepta, thin, straight, generally developed in 3 complete cycles in 6 systems or rarely in undeterminable systems. Remarks. Beauvais (1982, vol. 2, p. 267–268) stated that in the holotype of Actinacis martiniana the corallite diameter ranged from 0.5–1 mm. However, in the illustrations of the holotype presented by Beauvais (1982, pl. 48, figs 2a–b) the corallite diameter generally ranges from 1–1.2 mm, with juvenile corallites of around 0.8 mm and adult ones of up to 1.5 mm. The specimens assigned to Actinacis cognata Oppenheim, 1901, in the synonmy list above by Kuzmicheva (1987, p. 159), Drobne, et al. (1988, p. 188), Alvarez Perez (1993, p. 230), Bosellini & Russo (1995, p. 218), and Turnšek & Drobne (1998, p. 137) are characterized by corallite diameters ranging between 0.7 and up to around 1.3 mm, and septal numbers of up to 24, thus very closely agreeing with A. martiniana. In A. cognata the corallites are smaller than 1 mm in diameter and the number of septa is 12–16 (Oppenheim 1901, p. 182). Moreover, the latter might actually be a junior synonym of Actinacis digitata Fritsch, 1878, from the Eocene of Borneo, which is characterized by corallite diameters of 0.7 mm to up to nearly 1 mm and 15–20 septa. Type locality of species. Upper Santonian of southern France (Figuières). Distribution. Middle Albian of Texas, Upper Turonian-Santonian of Austria (Gosau Group), Coniacian and Upper Santonian of southern France (Corbières, Figuières), Senonian of Ukraine, Santonian of Georgia (in Caucasus), Campanian of northern Spain (Catalonia), Maastrichtian of Mexico (Cardenas Formation), Maastrichtian of Jamaica (new material), Middle-Upper Maastrichtian of the UAE/Oman border region, Paleocene of Slovenia (Adriatic Platform) and Austria (new material), Upper Paleocene-Lower Eocene of Java, Eocene of Italy, Spain, and Ukraine. New Material. Paleocene of Austria (Kambühel-Kalke), NMNH, Baron-Szabo coll., sample no.: NMNH 1068601; Maastrichtian of Jamaica, NMNH, Coates coll., sample nos.: 326a; 516b; 517 (=Ducketts Land Settlement); 570i; 570r; 570s (=probably Cambridge area).

Actinacis remesi Felix, 1903b Text-Fig. 22 v*1903b ?1906 1911 v1914 (v)1925 non1925 1930a v1957 ?1966 1976 1982 1987

Actinacis remeši n. sp.: Felix, p. 567, text-figs 1, 2 (topotypes studied). Litharaea distans n. sp.: Felix, p. 43, pl. 3, figs 3–3a. Actinacis remeši Felix: Trauth, p. 155, pl. 4, fig. 1, text-fig. 4. Actinacis remeši Felix: Felix, pars 7, p. 240 (topotypes studied). Actinacis remeši Felix: Vetters, p. 9, pl. 1, figs 5, 6. Actinacis remeši Felix: Kühn, p. 131. Actinacis remeši Felix: Oppenheim, p. 10, pl. XV, figs 3, 3a. Actinacis vignyensis nov. sp.: Alloiteau, pl. 7, fig. 4, pl. 18, fig. 9 (topotypes studied). Actinacis remeši Felix: Morycowa & Lefeld, p. 538, pl. 33, figs 1, 2. Actinacis remeši Felix 1903: Turnšek, in Turnšek & Buser, pp. 63, 83, pl. 20, figs 5, 6. Actinacis remeši Felix 1903: Beauvais, vol. 2, table 12. Actinacis vignyensis Alloiteau, 1957: Meyer, p. 25, pl. 4, figs 8–9.


BARON-SZABO

v1989 2000 2000 v2000 v2001 v2002 v2004

Actinacis remeši Felix 1903: Eliášová, p. 96, pl. 50, figs 2–3, pl. 56, fig. 3, pl. 59, figs 1–2, pl. 60, fig. 1. Actinacis remesi, Felix 1903: Löser, p. 5. Actinacis vignyensis, Alloiteau 1957: Löser, p. 5. Actinacis remesi Felix, 1903: Baron-Szabo, p. 119, pl. 10, figs 3, 6. Actinacis remesi Felix, 1903: Baron-Szabo, p. 260, fig. 1 F. Actinacis remesi Felix, 1903: Baron-Szabo, p. 100, pl. 69, figs 4, 6–7. Actinacis remesi Felix: Moosleitner, p. 180, pl. 78, fig. 10.

Text-Fig. 22 Actinacis remesi Felix, 1903, Maastrichtian of Austria based on the illustration in Vetters (1925), cross view of colony, scale bar: 4 mm.

Dimensions. d=1.8–2.5 mm, juveniles around 1.5 mm; c-c=2.5–4 mm; s=17–22; in juvenile corallite the number is 15. Description. Massive-lamellar and plocoid colony; corallites circular or elongated in outline; septal arrangement generally irregular, but in some calices a septal development in 6 systems is perceptible; pali or paliform lobes occur irregularly. Type locality of species. Upper Cenomanian-Lower Santonian of the Czech Republic. Distribution. ?Urgonian of Poland, Upper Cenomanian-Lower Santonian of the Czech Republic, TuroSCLERACTINIAN CORALS OF THE K/T-BOUNDARY


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nian-Senonian of Libya, Upper Turonian-Coniacian (Theresienstein reef), Santonian-Campanian (Gosau Group), and Maastrichtian (northern alpine flysch) of Austria, Middle-Upper Maastrichtian of UAE/Oman border region, resediment in Senonian breccia of Slovenia, Danian of France (Vigny).

Actinacis reussi Oppenheim, 1930 Pl. 8; fig. 12 v*1930a v1982 (v)1994 (v)1997 (v)1997 2000 2002 v2003 (v)2005

Actinacis reussi n. sp.: Oppenheim, p. 8, pl. 1, figs 6, 6a, pl. 10, figs 2, 2a, pl. 15, fig. 6 (topotypes studied). Actinacis reussi Oppenheim 1930: Beauvais, vol. 2, p. 271, pl. 48, figs 5, 6. (topotypes studied) Actinacis reussi Oppenheim 1930: Turnšek, p. 14, pl. 10, figs 1–3. Actinacis martiniana d’Orbigny, 1850: Turnšek, p. 4, pl. 4, figs A–C. Actinacis reussi Oppenheim 1930: Turnšek, p. 5, pl. 5, figs A–C. Actinacis reussi, Oppenheim 1930: Löser, p. 5. Actinacis reussi Oppenheim, 1930: Baron-Szabo, p. 100. Actinacis martiniana d’Orbigny 1850: Götz, p. 5ff., pl. 1, fig. 2. Actinacis cognata Oppenheim: Baceta et al., p. 128ff, fig. 8B.

Dimensions. d=1.1–1.8 mm, juveniles around 0.8 mm; c-c=1.8–3.5 (4.5) mm; s=24, in juveniles around 22, in late adult stages up to 28. Description. Massive-knobby, encrusting-lamellar or subdendroid, plocoid colony, with corallites that spiral around finger-like branches; circular or slightly oval calices scattered irregularly over the surface of colony; costosepta generally developed in 3 complete cycles. Remarks. According to Beauvais (1982, vol. 2, p. 271–272), two syntypes exist of the species Actinacis reussi. He chose the specimen No. 24262 of the Oppenheim collection as the lectotype and gave its dimensions which considerably vary from the ones given by Oppenheim in his original description. According to Beauvais (1982), the dimensions of skeletal elements of the lectotype are: d=0.8–1.7 mm, c-c=2–3 mm, and s=24. Type locality of species. Santonian of Austria (Gosau Group at Pass Gsch⎫tt-Graben). Distribution. Lower Coniacian of southern France, Upper Coniacian-Santonian of Austria (Gosau Group), Santonian-Campanian of Slovenia, Santonian and Upper Campanian of Spain, Middle-Upper Maastrichtian of Jamaica (new material), Danian of northern Spain. New Material. Middle-Upper Maastrichtian of Jamaica, NMNH, Coates coll., sample nos.: 299; 304a; 305a; 343; 344; 345; 363; 395b; 568f-II; 568g; 568h; (=Jerusalem Mountain Inlier); 326; 338; 487d; 487e; 487f; 511; 516c; 527h; 527i; 527j; 527k (=Ducketts Land Settlement); 380B (=Vaughnsfield); 533u (Shaw Castle, Maldon Formation); 546; 548m; J-71-34c (=Rio Minho); 570m; 570o (=probably Cambridge area).

Genus Actinhelia d'Orbigny, 1849 Type species. Astrea elegans Goldfuss, 1826, Maastrichtian of the Netherlands (Maastricht).

Diagnosis. Colonial, massive, cerioid or subplocoid. Gemmation extracalicinal and intracalicinal. Costosepta compact with rare pores, confluent or nonconfluent. Columella trabecular or absent. Endothecal and perithecal dissepiments thin, vesicular. Wall synapticulothecal and septoparathecal with pores.


BARON-SZABO

Actinhelia elegans (Goldfuss, 1826) Pl. 8 , figs 7a, b, 8 v1820 v*1826 1850 1881 1914 1925 1999 2000 v2002 v2003 2005 v2006

Milleporites cellulosus Linn.: Schlotheim, p. 363. Astrea elegans: Goldfuss, vol. 1, p. 69, pl. 23, fig. 6. Actinhelia elegans: d'Orbigny, p. 278. Astrea elegans Goldfuss sp. (minor): Quenstedt, p. 856, pl. 176, fig. 53. Actinohelia elegans Goldfuss sp. 1826: Felix, pars 7, p. 234. Actinohelia elegans Goldfuss sp.: Umbgrove, p. 112, pl. 11, fig. 29. Actinhelia elegans: Leloux, p. 193, fig. 2. Actinhelia elegans (Goldfuss, 1826): Löser, p. 7. Actinhelia elegans (Goldfuss, 1826): Baron-Szabo, pl. 69, fig. 4, pl. 70, fig. 1. Actinhelia elegans (Goldfuss, 1826): Schafhauser et al., p. 190. Actinhelia elegans (Goldfuss, 1826): Filkorn, p. 124, fig. 2i. Actinhelia elegans (Goldfuss, 1826): Baron-Szabo et al., p. 21, figs 5.9, 5.10.

Dimensions. d=1.5–2 mm; d (juvenile); around 1 mm; c-c=1.8–2.2 mm; s=16 + s (8s1 + 8s2+s) Description. Colonial, massive, cerioid or subplocoid; gemmation extracalicinal and intracalicinal; costosepta compact with rare pores, confluent or nonconfluent, arranged in 2 complete cycles in 8 systems; septal flanks finely granulated; anastomosis common; corallites directly united by a common wall or separated by a vermiculate coenosteum; columella trabecular or absent; endothecal and perithecal dissepiments thin, vesicular; wall synapticulothecal and septoparathecal with pores. Remarks. Because the holotype of A. elegans is preserved as a steinkern, fine skeletal structures are present only in some places. Moreover, corallites that are in advanced stages of intracalicinal budding are much more frequent than they are in the Jamaican material. However, because both the holotype of A. elegans and the Jamaican specimens are characterized by the above given features of the genus Actinhelia and, in addition, show skeletal elements of the same dimensions, they are considered the same. During a recent visit to the Natural History Museum in Paris material collected by Alloiteau from the Danian of France (Vigny) was studied by the author. Among these corals a specimen was found which closely corresponds to the taxon Actinhelia elegans. Type locality of species. Maastrichtian of The Netherlands (St. Pietersberg). Distribution. Maastrichtian of the Netherlands, Maastrichtian of Mexico (Ocozocuautla and Cardenas Formations), Middle-Upper Maastrichtian of Jamaica (new material), Danian of France (Vigny; unpublished data, Baron-Szabo, 2003). New Material. Middle-Upper Maastrichtian of Jamaica, NMNH, Coates coll., sample nos.: 339a; 339b; 416; 562g-II (=Jerusalem Mountain Inlier); 485p; 496c-I; 500a; 501; 527e-II; J-66-31-14-II (=Ducketts Land Settlement); 533t; 533v (Shaw Castle, Maldon Formation); 589c (=probably Cambridge railway area); J3502a3 (=Point Flamstead).

Genus Elephantaria Oppenheim, 1930 Type species. Elephantaria lindstroemi Oppenheim, 1930a, Santonian of Austria (Gosau Group).

Diagnosis. Colonial, submassive to lamellar, subthamnasterioid. Gemmation extracalicinal. Corallites embedded in a porous-reticulate coenosteum, connected by irregularly confluent septa. Costosepta reduced, subcomSCLERACTINIAN CORALS OF THE K/T-BOUNDARY


109

pact to porous, granulate laterally. Columellar trabeculae and synapticulae present. Endothecal dissepiments thin, vesicular. Subgenus. Bosnopsammia Oppenheim, 1909 (Type species. Bosnopsammia katzeri Oppenheim, 1909, Eocene of Bosnia); like Elephantaria but trabecular portions in corallite center form larger trabecular-papillose columella with a more or less distinct development of paliform lobes. Remarks. Based on a single specimen, Oppenheim (1909, p. 315) described the colonial coral Bosnopsammia. Oppenheim stated that it showed eupsammiid characteristics, closely resembling the genera Coenopsammia and Stereopsammia. However, following re-investigation of a large number of specimens collected from the type locality during the years after he had published the first description, Oppenheim (1912, p. 106) revised his original diagnosis of the genus, stating that in well preserved specimens the development of septa, as well as wall, and corallite integration rather indicated that it was a taxon closely related to the genus Actinacis. According to Oppenheim (1912), the genus Bosnopsammia is characterized by corallites embedded in a porous coenosteum with costosepta irregularly connecting them, and with frequently occurring nonconfluent septa, extracalicinal budding, costosepta having pores and granulated margins, and corallite centers filled with trabecular-papillose columella with a more or less distinct development of paliform lobes. Therefore, as a subgenus, the synonymy with the taxon Elephantaria is suggested.

Elephantaria lindstroemi Oppenheim, 1930a Pl. 9; Fig. 1 v*1930a ?1937 1956 v1998 v1999 2000 v2001 v2002

Elephantaria lindstroemi n. sp.: Oppenheim, p. 2, pl. 39, figs 6–7 (topotypes studied). Elephantaria lindstroemi Oppenheim: Kühn & Andrusov, p. 4–5. Elephantaria lindstroemi Oppenheim: Wells, p. F393, fig. 286,2. Goniopora hrpeljensis n. sp.: Turnšek, in Turnšek & Drobne, p. 138, pl. 9, figs 1–4. Elephantaria lindstroemi Oppenheim, 1930: Baron-Szabo, p. 457, pl. 6, fig. 4. Elephantaria lindstroemi, Oppenheim 1930: Löser, p. 33. Elephantaria lindstroemi Oppenheim, 1930: Baron-Szabo, p. 260, fig. 2F. Elephantaria lindstroemi Oppenheim, 1930: Baron-Szabo, pl. 70, fig. 3.

PLATE 9 Fig. 1 Elephantaria lindstroemi Oppenheim, 1930, NMNH, Coates coll., no. 545r, Maastrichtian of Jamaica, cross thin section, scale bar: 4 mm. Fig. 2 Astraraea media (Sowerby, 1832), topotype, NHMW, Reuss coll., 1864-XI-1320, Turonian-Campanian of Austria. 2a, close-up, scale bar: 3 mm; 2b, upper surface of colony, scale bar: 10 mm. Fig. 3 Astraraea media (Sowerby, 1832), NMNH, Coates coll., no. 568g-II, Middle-Upper Maastrichtian of Jamaica, cross thin section, scale bar: 3 mm. Fig. 4 Pleurocora arachnoides (Knorr & Walch, 1777) (syntypes of Pleurocora alternans Milne Edwards & Haime, 1848d), MNHN, Milne Edwards & Haime coll., Mo 1268, Campanian-Maastrichtian of Belgium. 4a, upper surface, cross view, scale bar: 3 mm; 4b, longitudinal view of colony, scale bar: 3 mm. Fig. 5 Pseudofavia grandiflora (Reuss, 1854), BMNH, AZ 406, Middle-Upper Maastrichtian of the UAE/Oman border region. 5a, thin section, longitudinal view, scale bar: 4 mm; 5b, thin section, cross view, scale bar: 4 mm. Fig. 6 Astraraea multiradiata (Reuss, 1854), BMNH, AZ 408, Middle-Upper Maastrichtian of the UAE/Oman border region, cross thin section, scale bar: 4 mm.


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111

Dimensions. c-c=3–8 mm, up to 10 mm in peripheral areas of colony; s=9–20, in largest corallites 24 septa possible; size of colony=3–10 cm in length, about 2–6 cm in width. Description. The corallum is irregularly massive-multicolumniform, with corallites that are embedded in a vermiculated coenosteum; septa short, thick, subcompact, and equal in thickness. About 6 septa reach the axial region, in front of which pali or paliform lobes might occur. Septal flanks have thick rounded granules and pennulae. The columella is papillose, weakly developed. Synapticulae are disposed throughout the colony. There is no wall between the corallites. The endotheca consists of thin tabulate or vesicular dissepiments. Microstructure is poorly preserved, but in places thick polyaxial trabeculae are present. Remarks. In having a subthamnasterioid corallum, with corallites that are both embedded in a porousreticulate coenosteum and are often hardly distinguishable from it, as well as having subcompact to porous septa with very short or reduced costae, the specimens described as Goniopora hrpeljensis in Turnšek & Drobne (1998) from the Thanetian of the Adriatic platform closely correspond to the genus Elephantaria. In Goniopora the corallites are arranged in the form of a cerioid colony and generally have distinct synapticulothecal-parathecal developments. Type locality of species. Turonian-Campanian of Austria (Gosau Group). Distribution. Turonian-Campanian of Austria (Gosau Group), ?Upper Santonian-Lower Campanian of Slovakia, Campanian-Maastrichtian of Jamaica (new material), Thanetian of Italy and Slovenia (Adriatic platform). New Material. Maastrichtian of Jamaica, NMNH, Coates coll., sample nos.: 314; 4482/2; 450a; 490f; 496L; (=Ducketts Land Settlement); 531 (=Vaughnsfield); 539b; J-71-12-11Ea (=Shaw Castle, Maldon Formation); 545r (=Rio Minho); 570a; 570k ; 591; (=probably Cambridge area); 590c (=probably Maldon Inlier); J-66-27 (=Catadupa); J-71-14c4 (=Sunderland); J-72-13A-2w; J-72-14 (=Marchmont); Campanian of Jamaica, NMNH, Coates coll., sample nos.: J-71-1A (=Green Island Inlier);

Family Haplaraeidae Vaughan & Wells, 1943 (=Felixaraeidae M. Beauvais, 1982; =Astraraeidae M. Beauvais, 1982) Diagnosis. Solitary and colonial. Colony formation by intra- and extratentacular budding. Corallite wall synapticulothecal, poorly defined or absent, costate, usually epithecate, porous. Septa exsert, composed of one fan system of compound trabeculae, with vertical axis of divergence, irregularly porous and thick. Dissepiments thin. Columella absent or parietal or feebly developed.

Subfamily Haplaraeinae Vaughan & Wells, 1943 Diagnosis. Solitary and colonial. Polyps large in size. Trabeculae thick. Synapticulae numerous. Columella parietal. Dissepiments thin, large. Remarks. Vaughan & Wells (1943) did not quantitatively define the size of polyps which they only referred to as 'large', but the nominatform of the subfamily (Haplaraea) was established on specimens having a calicular diameter ranging from 22–29 mm.


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Genus Astraraea Felix, 1900 (=Trechmannaria Wells, 1935, Type species. T. montanaroae Wells, 1935, Campanian of Jamaica); (=Valliseris Alloiteau, 1957, Type species. V. rennensis Alloiteau, 1957, Upper Cenomanian of France). Type species. Thamnastrea multiradiata Reuss, 1854, Upper Santonian of Austria.

Diagnosis. Colonial, thamnasterioid. Corallites can be arranged in short meandroi series. Gemmation intratentacular mono- to tristomodaeal. Corallites permanently monocentric. Radial elements biseptal with perforations arranged in vertical rows. Distal margins with rounded (moniliform) and subequal denticles. Lateral septal faces ornamented with rounded granulae. Synapticulae numerous. Columella parietal-papillose. Endothecal dissepiments sparse, thin. No wall. Remarks. Alloiteau (1957, p. 319) described the genus Valliseris from the Cenomanian of France and gave the following characteristics: “Colonial, corallites isolated or arranged in short meandroid series, corallite series separated by tectiform collines, septa biseptal with large pores, gemmation intracalicinal, no wall, corallites connected by septa, endothecal dissepiments thin, septal distal margins moniliform, granulated laterally, synapticulae numerous, columella parietal-papillose.” However, in the illustration of the holotype of the type species of Valliseris (see Alloiteau, 1957, pl. 16, fig. 2), corallites that are arranged in short series are generally separated by tholiform, and only rarely by tectiform, collines. Tholiform collines are also present in Astraraea when the corallites occur in small groups. In addition, in Valliseris, isolated corallites occur irregularly, which results in a thamnasterioid corallite integration. Therefore, the genus Valliseris Alloiteau is considered a junior synonym of Astraraea Felix. The genus Trechmannaria Wells, 1935, from the Campanian of Jamaica, is characterized by skeletal structures that are identical to the ones found in Valliseris and Astraraea: “Colonial, massive, thamnasterioidmeandroid, gemmation intracalicinal, corallite centres distinct, arranged in short series, separated by tholiform to tectiform collines, septa confluent, subcompact to perforated, granulated laterally, moniliform distally, synapticulae numerous, endothecal dissepiments sparse, columella spongy-papillose, no wall between the corallites.” In addition, re-examination of the type material of Trechmannaria Wells revealed that isolated corallites also occur. Therefore, the genus Trechmannaria is considered a junior synonym of Astraraea Felix.

Astraraea media (Sowerby, 1832) Pl. 9, figs 2a, b, 3 v*1832 1850 1851a v1854 1857 1858–61 1900 v1903a 1914 1930a ?1937a 1939a ?1945

Astrea media: Sowerby, in Sedgwick & Murchison, p. 417, pl. 37, fig. 5 9 (topotypes studied). Synastrea media (Sowerby): Milne Edwards & Haime, 3. ser., vol. 12, p. 150. Thamnastrea media: Milne Edwards & Haime, p. 109. Thamnastrea media Sowerby: Reuss, p. 119, pl. 19, figs 3–4. Thamnastrea media: Milne Edwards, vol. 2, p. 568. Synastrea media: de Fromentel, p. 220. Astraraea media (Sowerby): Felix, p. 3. Astraraea media (Sowerby): Felix, p. 187. Astraraea media Sowerby sp. 1832: Felix, pars 7, p. 205. Astraraea media (Sowerby): Oppenheim, p. 42, pl. 10, fig. 7, pl. 11, figs 5–6. Synastrea media (Sowerby): Bataller, p. 196. Astraraea senessei n. sp.: Alloiteau, p. 20, pl. 1, fig. 9. Synastrea media Sowerby sp. 1832: Bataller, p. 45.



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v1982 1982 v1998 2000 2002 v2002 2002 v2003 v2004

Astraraea media (Sowerby) 1832: Beauvais, vol. 2, p. 31, pl. 24, fig. 4. Astraraea senessei Alloiteau 1939: Beauvais, vol. 2, p. 34, pl. 25, fig. 2. Astraraea senessei Alloiteau, 1939: Baron-Szabo, p. 146, pl. 5, fig. 5. Astraraea senessei, Alloiteau 1939: Löser, p. 10. Astraraea media (Sowerby 1832): Löser et al., p. 83 (older synonyms cited therein). Astraraea media (Sowerby, 1832): Baron-Szabo, p. 104, pl. 73, fig. 1. Astraraea senessei Alloiteau, 1939: Baron-Szabo, p. 104. Astraraea media (Sowerby, 1832): Baron-Szabo, p. 137, pl. 21, figs 3, 6. Astraraea media (Sowerby): Moosleitner, p. 174 and 182, pl. 72, fig. 3, pl. 80, fig. 1.

Dimensions. c-c=4.5–7.5 mm; s=20–40; s/mm=5–7/2. Description. Colonial, thamnasterioid. Gemmation intratentacular mono- to tristomodaeal. Corallites permanently monocentric. Radial elements biseptal with perforations arranged in vertical rows. Distal margin with rounded and subequal denticles. Lateral septal faces ornamented with rounded granulae. Synapticulae numerous. columella parietal-papillose; endothecal dissepiments sparse, thin; no wall between the corallites. Type locality of species. Santonian-Campanian of Austria (Gosau Group). Distribution. ?Turonian of northern Spain, Turonian-Lower Coniacian and Upper Santonian of southern France, ?Upper Santonian and Campanian of northern Spain (Catalonia), Upper Turonian-Campanian of Austria (Gosau Group) and Croatia, Maastrichtian of France, Middle-Upper Maastrichtian of Jamaica (new material). New Material. Middle-Upper Maastrichtian of Jamaica, NMNH, Coates coll., sample nos.: 539b-II (=Jerusalem Mountain Inlier); 568g-II (=probably Cambridge area).

Astraraea multiradiata (Reuss, 1854) Pl. 9, fig. 6 v*1854 1858–61 1900 v1903a 1914 v1935 1952a v1982 v2000 2002 v2002 v2002

Thamnastraea multiradiata m.: Reuss, p. 118, pl. 7, fig. 1. Synastraea multiradiata: de Fromentel, p. 219. Astraraea multiradiata Reuss sp.: Felix, p. 38. Astraraea multiradiata Reuss sp.: Felix, p. 186, text-figs 10, 11. Astraraea multiradiata Reuss sp. 1854: Felix, pars 7, p. 205. Trechmannaria montanaroae, sp. n.: Wells, p. 190, pl. 11, figs 2, 3. Astraraea multiradiata (Reuss, 1854): Alloiteau, p. 608, pl. 2, fig. 9. Astraraea multiradiata (Reuss) 1854: Beauvais, vol. 2, p. 30, pl. 24, fig. 3, pl. 25, fig. 1 (older synonyms cited therein). Astraraea multiradiata (Reuss, 1854): Baron-Szabo, p. 116, pl. 8, fig. 5. Astraraea multiradiata (Reuss 1854): Löser et al., p. 84 (older synonmys cited therein). Astraraea multiradiata (Reuss, 1854): Baron-Szabo, p. 104, pl. 71, figs 1–3. Trechmannaria montanaroae Wells, 1935: Baron-Szabo, p. 107, pl. 75, figs 2, 4.

Dimensions. c-c=10–22 mm; s/mm=4–5/ 2. Description. Massive, thamnasterioid colony with calices, which are regularly disposed or arranged in small groups of up to 3 corallites; septa subequal to irregularly alternating; up to 30 septa extend to the columella; columella is spongy-papillose, well-developed. Type locality of species. Upper Santonian of Austria (Gosau Group). Distribution. Cenomanian of Azerbaijan, Coniacian of southern France (Corbières, Aude), Upper Turo-


BARON-SZABO

nian-Santonian of Austria (Gosau Group), Santonian of northern Spain, Lower Senonian of Curaçao, Campanian of Hungary and Jamaica, Middle-Upper Maastrichtian of the UAE/Oman border region.

Genus Pleurocora Milne Edwards & Haime, 1848 (=Psammiophora Fromentel, 1870, Type species. Psammiophora cenomana Fromentel, 1870, Cenomanian of France); (=Phyllohelia Alloiteau, 1952a, Type species. Pleurocora explanata Milne Edwards & Haime, 1848d, Maastrichtian of Belgium); (=Latohelia Löser, 1987, Type species. Synhelia reptans Počta, 1887, Cenomanian of the Czech Republic); (=? Barycora Alloiteau, 1952a, Type species. Barycora corbariensis Alloiteau, 1952a, Lower Coniacian of France). Type species. Lithodendron gemmans Michelin, 1846, Lower Coniacian of France (see Milne Edwards & Haime, 1848d).

Diagnosis. Colonial, subplocoid to subphacelo-dendroid. Corallites cylindrical, very short, with coenosteum between them formed by costae and exothecal dissepiments. Costosepta compact or subcompact, granulated. Pali opposite all but last cycle. Wall dense, synapticulothecal. Columella papillose. Endothecal dissepiments thin.

Pleurocora arachnoides (Knorr & Walch, 1777) Pl. 9, Fig. 4 v*1777 1778 1799 v1826 v1846 v1848d v1848d v1848d 1850 v1851a v1857 v1857 1857 1881 1903a 1903a 1906 1914 1914 1914 1914 1914 1914 v1925 1930 v1982

Astroites arachnoides: Knorr & Walch, vol. 3, p. 193, pl. 6e, fig. 6 (topotypes studied). Astroites arachnoides: Schröter, vol. 3, p. 461, pl. 9, fig. 3. Madrepores: Faujas-Saint-Fond, pl. 41, fig. 6. Astraea arachnoides: Goldfuss, vol. 1, p. 70, pl. 23, figs 9a–c (topotypes studied). Lithodendron gemmans N. Michelin, p. 305, pl. 72, fig. 6. Pleurocora gemmans: Milne Edwards & Haime, p. 310. Pleurocora alternans: Milne Edwards & Haime, p. 312. Pleurocora Koninckii: Milne Edwards & Haime, p. 312 (topotypes studied) Phyllocoenia arachnoides: d’Orbigny, vol. 2, p. 277. Astraea Riemsdycki: Milne Edwards & Haime, p. 98. Heliastraea Riemsdycki: Milne Edwards, vol. 2, p. 469. Pleurocora alternans: Milne Edwards, vol. 2, p. 603. Pleurocora Koninckii: Milne Edwards, vol. 2, p. 603. Astraea arachnoides Goldfuss: Quenstedt, p. 855, pl. 176, fig. 50. Haplohelia ornata nov. sp.: Felix, p. 323, pl. 17, figs 14–15. Oculina ogilviae nov. sp.: Felix, p. 323, pl. 23, figs 4, 5. Pleurocora Angelisi n. sp.: Felix, p. 47, pl. 3, figs 2–2b. Orbicella Riemsdycki E. H. sp. 1851: Felix, pars 7, p. 167. Pleurocora alternans E. H. 1849: Felix, pars 7, p. 171. Pleurocora Angelisi Felix 1906: Felix, pars. 7, p. 171. Pleurocora gemmans Michelin sp. 1847: Felix, pars 7, p. 171. Pleurocora Koninckii E. H. 1849: Felix, pars 7, p. 172. Haplohelia ornata Felix 1903.: Felix, pars 7, p. 233. Orbicella Riemsdijcki var. conica (nov. var.): Umbgrove, p. 103, pl. 8, fig. 3. Pleurocora cf. alternans Milne-Edwards & Haime: Kühn & Andrusov, p. 5–6. Haplohelia ornata Felix 1903: Beauvais, vol. 1, p. 131, pl. 11, figs 2a–b (topotypes studied).



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1987 (v)?1991 v1997 v1998 nonv1999 v1999 1999 2000 2000 2000 2000 2000 2000 2000 2000 2002 2002 2002 2002 v2002 2002 2002 2002 v2002 v2004

Haplohelia regularis, D’Orbigny: Meyer, pl. 5, figs 8–9. “Branching coral”: Bryan, p. 430, Fig. 7C. Pleurocora cf. alternans Milne-Edwards & Haime, 1849: Baron-Szabo, p. 77, pl. 10, figs 4, 6. Latohelia cicularia n. sp.: Baron-Szabo, p. 150, pl. 9, figs 1–4, pl. 10, fig. 2. Pleurocora cf. alternans Milne Edwards & Haime, 1849: Baron-Szabo, p. 455, pl. 3, fig. 2. Latohelia cf. cicularia Baron-Szabo, 1998: Baron-Szabo, p. 461, pl. 5, fig. 3. Montastrea arachnoides: Leloux, p. 193, fig. 2. Heliastraea riemsdycki (Milne-Edwards & Haime 1851): Löser, p. 40. Latohelia cicularia, Baron-Szabo 1998: Löser, p. 46. Montastraea arachnoides (Knorr & Walch 1777): Löser, p. 53. Orbicella riemsdijcki var. conica, Umbgrove 1926: Löser, p. 57. Pleurocora alternans, Milne-Edwards & Haime 1849: Löser, p. 66. Pleurocora angelisi, Felix 1906: Löser, p. 66. Pleurocora gemmans (Michelin 1847): Löser, p. 66. Pleurocora konincki, Milne-Edwards & Haime: Löser, p. 66. ‘solitary coral’: Bauer et al., pl. 4, fig. 9. Pleurocora alternans Milne Edwards & Haime, 1849: Baron-Szabo, p. 106. Pleurocora angelisi Felix, 1906: Baron-Szabo, p. 106. Pleurocora cicularia (Baron-Szabo, 1998): Baron-Szabo, p. 106. Pleurocora gemmans (Michelin, 1846): Baron-Szabo, p. 105, pl. 73, figs 2–3, 5, 8–9, pl. 74, figs 1–4, 6. Pleurocora ornata (Felix 1903): Baron-Szabo, p. 106. Haplohelia ogilviae (Felix 1903): Löser et al, p. 328 (older synonmys cited therein). Haplohelia ornata, Felix 1903: Löser et al, p. 328 (older synonmys cited therein). Montastrea arachnoides conica (Umbgrove, 1925): Leloux, p. 22. Pleurocora gemmans (Michelin): Moosleitner, p. 180, pl. 78, figs 6, 12, 13.

Dimensions. d=2–3.5 mm, juvenile around 1.5 mm; c-c: 3–5 mm (in submassive colonies); s=24–48. Description. Submassive, plocoid to subdendroid colony; costosepta arranged in 3 to 4 complete cycles, depending on ontogenetical stage of corallite; columella spongy-papillose, generally well-developed. Remarks. Re-examination of type and original material revealed that the species Pleurocora gemmans (Michelin), P. alternans Milne Edwards & Haime, P. koninckii Milne Edwards & Haime, P. ogilviae (Felix), P. ornata (Felix), and P. circularia (Baron-Szabo) have to be considered junior synonyms of P. arachnoides (Knorr & Walch) because all of them are characterized by a corallite diameter of 2–3.5 mm (juvenile around 1.5. mm), corallite distance of 3–5 mm, and septa numbering from 24–48 and developed in 6 systems. Milne Edwards & Haime (1851a, p. 98) described the species Astraea riemsdycki. Later, Milne Edwards (1857, p. 469) transferred this taxon to the genus Heliastraea and gave the dimensions d: 3–4 mm and s: 60–70. Re-examination of the specimen (=?holotype of) Heliastraea riemsdycki, collection number M-561 [L 27] in the Michelin/Milne Edwards & Haime collection, MNHN by the author of this paper revealed that this specimen has all characterics of a subplocoid representative of Pleurocora and skeletal elements (d: 2–3.5 mm, juvenile 1.5 mm, s: up to 48) that closely correspond to the species P. arachnoides. Type locality of species. Maastrichtian of The Netherlands (St. Pietersberg). Distribution. Uppermost Cenomanian of Egypt (Sinai), Turonian-Campanian of Austria (Gosau Group), Lower Coniacian and Santonian of France, Senonian of Ukraine (Delyatin, Iwano-Frankowskaya), Upper Santonian-Lower Campanian of Romania, Campanian of northern Spain (Torallola), Campanian-Maastrichtian of Belgium (Obourg, Mons), Maastrichtian of The Netherlands (St. Pietersberg), Danian of France (Vigny), ?Paleocene of USA (Alabama, Salt Mountain Limestone).


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Pleurocora explanata Milne Edwards & Haime, 1848 Text-Fig. 23 *1848d 1857 1914 1987 2000 2002

Pleurocora explanata: Milne Edwards & Haime, vol. 4, p. 311, pl. 7, fig. 10. Pleurocora explanata: Milne Edwards, vol. 2, p. 602. Pleurocora explanata E. H. 1849: Felix, pars 7, p. 171. Phyllohelia explanata (Edwards et Haime, 1848): Kuzmicheva, p. 110, pl. 17, figs 6a–b. Pleurocora explanata, Milne-Edwards & Haime 1848: Löser, p. 66. Pleurocora explanata Milne Edwards & Haime, 1848: Baron-Szabo, p. 106.

Dimensions. d=3.5–4 mm; s=48 + s5. Description. Submassive, plocoid to subdendroid colony; corallites circular in outline; costosepta thin, richly dentate laterally, arranged in 4–5 cycles.; pali ?large. Type locality of species. Campanian-Maastrichtian of Belgium (Obourg, Mons). Distribution. Campanian-Maastrichtian of Belgium (Obourg, Mons), Danian-Montian of Ukraine.

Text-Fig. 23 Pleurocora explanata Milne Edwards & Haime, 1848d, type specimen as illustrated in Milne Edwards & Haime (1848d), Campanian-Maastrichtian of Belgium, upper surface of colony, scale bar: 4 mm.

Pleurocora haueri Milne Edwards & Haime, 1848 Text-Fig. 24 v*1848d v1854 1857 v1903a 1909 1914 1914 1972 v1994 v1999 2000 2000 2002

Pleurocora Haueri: Milne Edwards & Haime, p. 312 (topotypes studied). Pleurocora Haueri M. Edw. et H.: Reuss, p. 112, pl. 6, figs 26–27 (topotypes studied). Pleurocora Haueri: Milne Edwards, vol. 2, p. 603. Pleurocora Haueri M. Edwards et J. Haime: Felix, p. 267 (topotypes studied). Oculina Nordenskjöldi nov. sp.:Felix, p. 11, pl. 1, figs 12–13. Pleurocora Haueri E. H. 1849: Felix, pars 7, p. 172. Oculina Nordenskjöldi Felix 1909: Felix, pars 7, p. 232. O. nordenskjoldi Felix, 1909: Floris, p. 37. ?Oculina nordenskjoeldi (Felix, 1909): Filkorn, p. 71, figs 26 (1–4) (older synonyms cited therein). Pleurocora cf. alternans Milne-Edwards & Haime, 1849: Baron-Szabo, p. 455, pl. 3, fig. 2. Pleurocora nordenskjoldi, Felix 1909: Löser, p. 56. Pleurocora haueri, Milne-Edwards & Haime 1849: Löser, p. 66. Pleurocora haueri Milne Edwards & Haime, 1848: Baron-Szabo, p. 106.



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Text-Fig. 24 Pleurocora haueri Milne Edwards & Haime, 1848d, Campanian-Maastrichtian of Antarctica (syntype of Oculina Nordenskjˆldi Felix, 1909, NRZ Cn94), cross view, oblique, polished surface, scale bar: 5 mm (from Filkorn 1994, see full reference in Reference list; copyright 1994 by the American Geophysical Union; reproduced by permission of American Geophysical Union).

Dimensions. d=5–7 mm, juveniles around 3.5 mm; s=32–48 Description. Subdendroid colony; costosepta developed in 4 incomplete or complete cycles in 6 systems; pali occur in front of oldest cycle, usually fuse with the columella; columella trabecular, irregularly papillose or made of twisted segments. Remarks. The systematic position of Oculina Nordenskjöldi Felix has been dicussed for nearly a century. In the original description even Felix himself expressed some doubts about the assignment of the syntypes to the genus Oculina. Filkorn (1994, figs 26 [1–2]) gave the first photographic illustration of the syntypes. He also expressed doubts about the grouping with Oculina. As previously pointed out by Filkorn, characteristics of the species include compact to subcompact septa, development of synapticulothecal (septothecal in places) wall and pali, as well as a trabecular columella which is irregularly papillose or fused, forming twisted segments.. Therefore, the assignment to the genus Pleurocora is suggested. Type locality of species. Turonian-Campanian of Austria (Gosau Group). Distribution. Turonian-Campanian of Austria (Gosau Group), Campanian-Maastrichtian of Antarctica (Seymour Island, Snow Hill Island).

Genus Pseudofavia Oppenheim, 1930 Type species. Parastraea grandiflora Reuss, 1854, Turonian-Campanian (probably Santonian) of Austria (Gosau Group).

Diagnosis. Colonial, massive, plocoid to subcerioid. Gemmation extracalicinal. Corallites separated by a vermiculate coenosteum. Costae short. Costosepta compact to subcompact, nonconfluent, granulated laterally.


BARON-SZABO

Anastomosis present. Columella trabecular. Synapticulae abundant. Endothecal dissepiments vesicular, sparse. Wall synapticulothecal to parathecal, incomplete.

Pseudofavia grandiflora (Reuss, 1854) Pl. 9, figs 5a, b v*1854 v1903a 1930a 1943 1976 v1982 (v)1997 2000 v2000 v2002

Parastraea grandiflora m: Reuss, p. 120, pl. 16, fig. 10. Parastraea grandiflora Reuss: Felix, p. 181, fig. 7. Pseudofavia grandiflora (Reuss): Oppenheim, p. 65, pl. XL, fig. 5. Pseudofavia grandiflora (Reuss): Vaughan & Wells, p. 133, pl. 16, fig. 10. Pseudofavia grandiflora (Reuss 1854): Turnšek, in Turnšek & Buser, p. 60, 81, pl. 18, figs 1–3. Pseudofavia grandiflora (Reuss) 1854: Beauvais, vol. 2, p. 16, pl. 23, fig. 3 (older synonyms cited therein). Pseudofavia grandiflora (Reuss 1854): Turnšek, p. 174, figs 174 A–E. Pseudofavia grandiflora (Reuss 1854): Löser, p. 70. Pseudofavia grandiflora (Reuss, 1854): Baron-Szabo, p. 116, pl. 8, fig. 4. Pseudofavia grandiflora (Reuss, 1854): Baron-Szabo, p. 107, pl. 76, figs 1–4.

Dimensions. d (max)=11–15 mm; d (min)=9–12 mm; c-c=9–11 mm; s=(44) 48–60; size of the colony: about 9 cm in diameter. Description. Massive, cerioid colony; corallites polygonal in outline, separated by a vermiculate coenosteum, which is up to 2 mm in thickness; septa arranged in 4 complete with the beginning of a fifth cycle in 6 systems, regularly alternating in length, but nearly equal in thickness; axial ends may meet and fuse with the columella, or dissociate into paliform structures; columella spongy-papillose or made of elongate segments; occurrence of synapticulae mainly restricted to both the peripheral areas of corallite and the peritheca. Remarks. The description of the type of Pseudofavia grandiflora (Reuss) by Beauvais (1982, vol. 2, p. 16–17) differs from the type material in that it does not reflect the whole spectrum of variation in the specimen. Beauvais gave 21.5–35.5 mm for the calicular diameter, which apparently was the size range of the largest corallites only (see syntype in Baron-Szabo 2002, pl. 76, fig. 1). In the type specimen corallites having a maximum diameter of 15–22 mm (11 mm in juveniles) are most common. Type locality of species. Turonian-Campanian of Austria (Gosau Group). Distribution. Turonian-Campanian of Austria (Gosau Group), Senonian breccia (resediment) of Slovenia, ?Senonian of Georgia (in Caucasia), Middle-Upper Maastrichtian of the UAE/Oman border region.

Genus Stiboriopsis Vaughan, 1899 Type species. Stiboriopsis jamaicaensis Vaughan, 1899, Campanian (-?Maastrichtian) of Jamaica.

Diagnosis. Colonial, massive, subthamnasterioid-subplocoid to meandroid. Gemmation by septal budding and fission. Corallites isolated or arranged in short meandroid series. Costosepta compact, with rare pores, confluent to nonconfluent, dentate to granulate laterally. Paliform structures can be present before principle septa. Columella trabecular or rudimentary. Endothecal dissepiments abundant, corallites united by their costae or separated by exothecal dissepiments. Remarks. The systematic position of Stiboriopsis Vaughan (1899) has been the subject of much discussion. Wells (1956) placed it in the family Haplaraeidae as a synonym of Trechmannaria Wells, 1935. AccordSCLERACTINIAN CORALS OF THE K/T-BOUNDARY


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ing to Alloiteau (1957), this genus belongs to the Faviidae. As pointed out by Baron-Szabo & González-León (1999, p. 485) the synonymy with Trechmannaria Wells seems questionable, because Stiboriopsis Vaughan is described as having rather compact structures, but Trechmannaria represents a form that, e.g., has corallite centres which are linked by porous lamellae. Moreover, all additional skeletal features of Trechmannaria Wells closely correspond to the ones of Astraraea Felix (see Remarks of Astraraea Felix). Stiboriopsis Vaughan resembles montlivaltiids and faviids in having thick, straight septa and similar microstructure, but is distinguished from them by: (1) having large but rare septal pores (as shown in the illustration in Vaughan 1899, pl. 38, fig. 4), (2) variable ornamentation on the inner ends of septa and (3) trabecular structures in the centre of the corallite (true columella or trabecular prolongations of the inner ends of septa). The diagnosis given above slightly differs from the original by Vaughan (1899, p. 237) and is based on the features seen in the figure of the type specimen in Vaughan (1899, pl. 38, fig. 4).

Text-Fig. 25 Stiboriopsis jamaicaensis Vaughan, 1899. A, Maastrichtian of Madagascar (holotype of Andemantastraea ambatryensis Alloiteau 1951a as figured in Alloiteau 1951a), cross view of colony, scale bar: 4 mm ; B, type material as figured in Vaughan (1899), Campanian (-?Maastrichtian) and Eocene of Jamaica (Blue Mountain series), cross view of colony, scale bar: 4 mm.

Stiboriopsis jamaicaensis Vaughan, 1899 Text-Fig. 25 *1899 v1951a 1999 2000

Stiboriopsis jamaicaensis, sp. nov.: Vaughan, p. 238, pl. 38, figs 2–4. Andemantastraea ambatryensis nov. sp.: Alloiteau, p. 48, pl. 8, figs 3–4. Stiboriopsis jamaicaensis Vaughan, 1899: Baron-Szabo & Gonzalez-Leon, p. 485. Andemantastraea ambatryensis, Alloiteau 1951: Löser, p. 8.


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Stiboriopsis jamaicaensis, Vaughan 1899: Löser, p. 74. Stiboriopsis jamaicaensis Vaughan, 1899: Baron-Szabo, p. 107, text-fig. 37.

Dimensions. d (max., isolated corallites): 7–9 mm; d (series, wall-wall): 3.5–7 mm; s (isolated corallites): 24+s4; s/mm: 20/10. Description. Massive, subthamnasterioid colony, subplocoid when corallites isolated, or corallites arranged in meandroid series; calicinal series generally short, consisting of 3–4 corallites; costosepta arranged in 3 complete cycles, with the beginning of a fourth cycle; corallites united by their costae or separated by up to 2 mm wide intercalicinal spaces formed by exothecal dissepiments. Type locality of species. Campanian (-?Maastrichtian) and Eocene of Jamaica (Blue Mountain series). Distribution. Campanian (-?Maastrichtian) and Eocene of Jamaica (Blue Mountain series), Maastrichtian of Madagascar (Betioky).

Subfamily Meandrophylliinae Roniewicz, 1976 Diagnosis. Polyps small. Gemmation intracalicinal. Costosepta confluent, except the collines. Columella trabecular, simple or papillose, feebly developed. Dissepiments tabulate. Trabeculae thin. Remarks. Roniewicz (1976) did not give any specific information as to the size of polyps she referred to as ‘small’, but the dimensions of the corallites of the nominatform of the subfamily (Meandrophyllia) are 2–4 mm in diameter.

Genus Meandrophyllia d’Orbigny, 1849 (=Meandraraea Étallon, 1859, Type species. M. gresslyi Étallon, 1859, Jurassic of France); (=Uxacalcaraea Alloiteau, 1957, Type species. U. heberti Alloiteau, 1957, Turonian of France); (=Michelinaraea Alloiteau, 1957, Type species. Meandrina ataciana Michelin, 1846, Upper Santonian of France). Type species. Meandrina lotharinga Michelin, 1843, Upper Jurassic of France (Meuse).

Diagnosis. Colonial, massive to lamellar, meandroid. Gemmation intracalicinal. Corallites isolated or arranged in short, wavy calicinal series. Series separated by tholiform to tectiform collines. Costosepta confluent to subconfluent, perforated. Columella spongy-papillose. Synapticulae present. Wall synapticulothecal, incomplete.

Meandrophyllia oceani (de Fromentel, 1877) Pl. 10, figs 1a, b v*1877 1914 v1936 1937a v1937a 1982 ?1995 ?1997

Latimaeandra Oceani: de Fromentel, p. 450, pl. 107, fig. 4. Latimaeandra Oceani de Fromentel: Felix, pars 7, p. 176. Meandraraea Sampelayoi Bataller: Bataller, p. 42, figs 14–15. Maeandraraea massilienis d’Orbingy sp.: Bataller, p. 209, text-fig. on p. 210. Meandraraea Sampelayoi Bataller sp. 1936: Bataller, p. 211, text-fig. on p. 211. Meandrarea oceani (de Fromentel) 1877: Beauvais, vol. 2, p. 221, pl. 43, figs 5–6. Meandrarea oceani (de Fromentel, 1887): Tchéchmédjiéva, p. 74, pl. 19, fig. 4. Dimorphomeandra viaderi n. sp.: Reig Oriol, p. 26, pl. 4, figs 4–5.



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Meandrarea oceani (de Fromentel, 1873): Löser, p. 47. Meandrophyllia oceani (Fromentel, 1873): Baron-Szabo, p. 110. Meandrophyllia oceani (Fromentel, 1873): Schafhauser et al., p. 190, tab.1. Meandrophyllia oceani (Fromentel, 1873): Baron-Szabo et al., p. 22, fig. 4.5.

Dimensions. d (max)=3–4 mm; c-c (same series)=2.5–5.5 mm; s/mm: 7–9/2. Description. Colonial, massive to lamellar, meandroid; gemmation intracalicinal; corallites isolated or arranged in short series or calicinal clusters, consisting of up to 3 corallites; series separated by generally tholiform but occasionally tectiform collines; costosepta confluent to subconfluent, nearly equal in thickness, perforated, granular and ?pennular laterally; columella spongy-papillose; well-developed; synapticulae present; wall synapticulothecal, incomplete. Remarks. In having intracalicinal-marginal budding, very well separated corallites, a very reduced corallite wall and a very reduced columella, the material described from the Upper Campanian of Bulgaria in Tchéchmédjiéva (1995) rather corresponds to the genus Lophomeandra Beauvais, 1982, and closely resembles the species Lophomeandra polygonata Beauvais, 1982. According to the description and illustration provided by Bataller (1937a, p. 209, text-fig. on p. 210), the material assigned to Meandraraea massiliensis d’Orbigny seems to closely correspond to the genus Meandrophyllia and belongs to the species Meandrophyllia oceani (de Fromentel, 1877) (see Baron-Szabo et al. 2006). Type locality of species. Coniacian of France (Corbi⎝res). Distribution. Coniacian-Santonian of France, Upper Santonian of Austria (Gosau Group), Upper Campanian of ?Bulgaria, Campanian-Maastrichtian of northern Spain, Maastrichtian of Mexico (Cardenas Formation), Campanian and Middle-Upper Maastrichtian of Jamaica (new material). New Material. Middle-Upper Maastrichtian of Jamaica, NMNH, Coates coll., sample no.: J-71-2a (=Marchmont); Campanian of Jamaica, NMNH, Coates coll., sample nos.: J-71-3a; J-71-3b (=Green Island).

Meandrophyllia textilis (Goldfuss, 1826) Pl. 10, figs 6a, b v*1826 1850b 1850 1851a 1857 1914 1925 1999 2002 2002

Astrea textilis nobis: Goldfuss, vol. 1, p. 68, pl. 23, figs 3a–b. Synastrea textilis: Milne Edwards & Haime, vol. 4, 3. ser., p. 156. Synastrea textilis: d‘Orbigny, vol. 2, p. 277. Thamnastrea textilis: Milne Edwards & Haime, p. 114. Thamnastraea textilis: Milne Edwards, vol. 2, p. 566. Thamnastraea textilis Goldfuss sp. 1826: Felix, pars 7, p. 199. Thamnastraea (c.f. Dimorphastraea) textilis Goldfuss spec.: Umbgrove, p. 110. Fungiastrea ? textilis: Leloux, p. 193, fig. 2. Fungiastrea textilis (Goldfuss 1826): Löser et al., p. 317 (older synonyms cited therein). Fungiastrea textilis (Goldfuss, 1826): Baron-Szabo, p. 147.

Dimensions. d (max)=1.8–2 mm; c-c (same and adjacent series)=2–3.5 mm; s/mm=8–10/2. Description. Meandroid colony; corallites arranged in very short meandroid series, generally consisting of 2 or 3 corallite centres; series separated by tholiform collines; septa subequal. Remarks. The taxonomic position of Astrea textilis Goldfuss has been discussed for nearly 2 centuries. Because the holotype of this species is preserved as a mold (=negative imprint, see Pl. 10, fig. 6a), it falsely resembles forms that have compact (typical of Thamnastrea) or compact to subcompact septa (typical of


BARON-SZABO

Fungiastrea), no wall between the corallites (typical of Thamnastrea, Fungiastrea, and Synastrea), and very numerous synapticulae (typical of Synastrea). However, as can be seen in the “positive image” of the holotype presented here (Pl. 10, fig. 6b), the specimen formerly described as Astrea textilis represents a meandroid colony that has rather perforated and costate septa, and corallites that are isolated or arranged in short meandroid series which are separated by an incomplete synapticulothecal wall. Therefore, the species Astrea textilis is here transferred to the genus Meandrophyllia d’Orbigny. Type locality of species. Upper Maastrichtian of The Netherlands (St. Pietersberg). Distribution. Upper Maastrichtian of The Netherlands.

Meandrophyllia velamentosa (Goldfuss, 1826) Pl. 10, figs 4a–c v*1826 v1826 1850 1850 1851a 1851a 1857 1857 1914 1914 v1925 v1925 v1999 v1999 2002 2002 2002 2002

Astrea velamentosa nobis: Goldfuss, vol. 1, p. 68, pl. 23, figs 4a–b. Astrea gyrosa nobis: Goldfuss, vol. 1, p. 68, pl. 23, fig. 5. Synastrea filamentosa: d‘Orbigny, vol. 2, p. 277. Synastrea gyrosa: d‘Orbigny, vol. 2, p. 277. Thamnastrea velamentosa: Milne Edwards & Haime, p. 113. Parastraea ? gyrosa: Milne Edwards & Haime, p. 116. Favia ? gyrosa: Milne Edwards, vol. 2, p. 441. Thamnastraea velamentosa: Milne Edwards, vol. 2, p. 563. Favia ? gyrosa Goldfuss sp. 1826: Felix, pars 7, p. 173. Thamnastraea velamentosa Goldfuss sp. 1826: Felix, pars 7, p. 199. Thamnstraea velamentosa Goldfuss sp.: Umbgrove, p. 110. Thamnstraea gyrosa Goldfuss sp., gen. nov. nom: Umbgrove, p. 112, pl. 11, fig. 18. Fungiastrea ? Velamentosa: Leloux, p. 193, fig. 2. Meandrophyllia gyrosa: Leloux, p. 193, fig. 2. Fungiastrea velamentosa (Goldfuss 1826): Löser et al., p. 317 (older synonyms cited therein). Meandrophyllia gyrosa (Goldfuss 1826): Löser et al., p. 401 (older synonyms cited therein). Meandrophyllia gyrosa (Goldfuss, 1826): Baron-Szabo, p. 110. Fungiastrea velamentosa (Goldfuss, 1826): Baron-Szabo, p. 147.

Dimensions. d (max)=1.5 mm; d (min)=0.8–1.2 mm; c-c (same and adjacent series)=1.2–2.4 mm; s/ mm=10–14/2. Description. Meandroid colony; corallites generally isolated, sometimes arranged in short meandroid series, consisting of up to 4 corallite centres; series separated by tholiform or tectiform collines; septa developed in 2 irregularly occurring size orders. Remarks. The taxonomic history of Astrea velamentosa Goldfuss very closely resembles the one of Astrea textilis Goldfuss in that the holotype of this species is preserved as a mold (=negative imprint, see Pl. 10, fig. 4a), it therefore falsely resembles forms that have compact (typical of Thamnastrea) or compact to subcompact septa (typical of Fungiastrea), no wall between corallites (typical of Thamnastrea, Fungiastrea, and Synastrea), and very numerous synapticulae (typical of Synastrea). However, as can be seen in the “positive image” of the holotype presented here (Pl. 10, figs 4b–c), the specimen formerly described as Astrea velamentosa represents a meandroid colony that has rather perforated and costate septa and corallites that are isolated or arranged in short meandroid series which are separated by an incomplete synapticulothecal wall. Therefore, the species Astrea velamentosa is here transferred to the genus Meandrophyllia d’Orbigny.



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Type locality of species. Upper Maastrichtian of The Netherlands (St. Pietersberg). Distribution. Upper Maastrichtian of The Netherlands.


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PLATE 10 Fig. 1 Meandrophyllia oceani (Fromentel, 1873), NMNH, Coates coll., no. J-71-2a, Middle-Upper Maastrichtian of Jamaica. 1a, cross thin section, scale bar: 1.5 mm; 1b, thin section, longitudinal view, scale bar: 2 mm. Fig. 2 Cyathoseris catadupensis (Vaughan, 1899), NMNH, Coates coll., no. 520, Maastrichtian of Jamaica, cross thin section, scale bar: 3 mm. Fig. 3 Cyathoseris catadupensis (Vaughan, 1899), holotype, MCZ, 114216, Campanian-Eocene of Jamaica, upper surface of colony, scale bar: 2 mm. Fig. 4 Meandrophyllia velamentosa (Goldfuss, 1826), holotype, IPB, Goldfuss coll., no. 229, Upper Maastrichtian of The Netherlands. 4a, cross view of colony in ‘steinkern’ preservation, scale bar: 3 mm; 4b, “positive sketch” (presented for the first time), scale bar: 3 mm; 4c, close-up of Fig. 4b, scale bar: 2 mm. Fig. 5 Brachymeandra leptophylla (Reuss, 1854), BMNH, AZ 476, Middle-Upper Maastrichtian of the UAE/Oman region, cross thin section of colony, scale bar: 5 mm. Fig. 6 Meandrophyllia textilis (Goldfuss, 1826), holotype, IPB, Goldfuss coll., no. 228, Upper Maastrichtian of The Netherlands. 6a, cross view of colony in ‘steinkern’ preservation, scale bar: 3 mm; 6b, “positive sketch” (presented for the first time), scale bar: 3 mm.

Genus Brachymeandra Alloiteau, 1957 (=Brachycoenia Beauvais, 1982, Type species. Adelastraea leptophylla Reuss, 1854, Upper Santonian of Austria). Type species. Brachymeandra delphinensis Alloiteau, 1957, Turonian of France.

Diagnosis. Colonial, massive, thamnasterioid, plocoid to submeandroid superficially. Gemmation intracalicinal. Costosepta subcompact or porous, subconfluent or confluent. Marginally beaded, finely granulated laterally. Columella parietal-papillose. Paliform structures present. Synapticulae abundant. Endothecal dissepiments thin, subtabulate. Perithecal wall can be present. Generally no wall between the calices. Remarks. Based on the species Adelastraea leptophylla Reuss, 1854, M. Beauvais (1982, vol. 2: 47) created the genus Brachycoenia, which differs only slightly from the genus Brachymeandra Alloiteau. Beauvais mentions the existence of paliform structures in Brachymeandra as the main difference between the two genera. However, later investigations of the type material of Brachycoenia (Baron-Szabo 2002, 2003) revealed the occasional existence of paliform structures. Therefore, the separation of these taxa does not seem justified.

Brachymeandra leptophylla (Reuss, 1854) Pl. 10, fig. 5 v*1854 1857 1858–61 v1903a 1914 ?1956 v1982 1986 ?1987 ?1987 1995 v1999 2000 v2000

Adelastraea leptophylla m.: Reuss, p. 115, pl. 12, figs 3, 4 (topotypes studied). Confusastraea leptophylla: Milne Edwards, vol. 2, p. 484. Confusastraea leptophylla: de Fromentel, p. 210. Confusastraea leptophylla M. Edwards (Reuss sp.): Felix, p. 293 (topotypes studied). Confusastraea leptophylla Reuss sp. 1854: Felix, pars 7, p. 169. Confusastraea leptophyllia (Reuss): Bendukidze, p. 85, pl. 9, figs 2–2a. Brachycoenia leptophylla (Reuss) 1854: Beauvais, vol. 2, p. 48, pl. 26, fig. 7, pl. 27, fig. 1. Brachycoenia leptophylla (Reuss, 1854): Tchéchmédjiéva, p. 68ff. Reussicoenia sp. (R. vignyensis n. sp.): Meyer, pl. 5, fig. 3. Thalamocoenia sp. (R. vignyensis n. sp.): Meyer, pl. 5, fig. 3. Brachycoenia leptophylla (Reuss, 1854): Tchéchmédjiéva, p. 60, pl. 12, figs 11, 12.. Brachycoenia leptophyllia (Reuss, 1854): Baron-Szabo, p. 455, pl. 1, fig. 6, pl. 4, fig. 3. Brachycoenia leptophylla (Reuss 1854): Löser, p. 15. Brachycoenia leptophylla (Reuss, 1854): Baron-Szabo, p. 119, pl. 11, fig. 2.



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Brachymeandra leptophylla (Reuss, 1854): Baron-Szabo, p. 111, pl. 78, figs1–3. Brachymeandra leptophylla (Reuss): Moosleitner, p. 182, pl. 80, fig. 2.

Dimensions. c-c=5–10 mm; s=36–60, in late budding stages the number of septa may be larger; s/mm=6–7/2; size of the colony=7 cm in diameter. Description. The massive and thamnasterioid colony has slightly protuberant calices, which appear to be subplocoid. Increase in corallites is due to intracalicinal gemmation. Costosepta are subcompact or porous, confluent or subconfluent, nearly equal in thickness, and have pennulae and spiniform or coarse, rounded granules laterally. Their inner ends have the tendency to fuse. About 20 septa reach the centre of the calice. The columella is spongy-papillose, well-developed. Paliform structures can be frequently observed. Synapticulae are irregularly disposed. There is no wall between the corallites. Endotheca is formed by numerous vesicular or subtabulate dissepiments. Septal microstructure is made of thick monaxial and polyaxial trabeculae. Type locality of species. Upper Santonian of Austria (Gosau Group). Distribution. ?Senonian of Georgia (in Caucasia), Santonian-Campanian of Austria (Gosau Group), Upper Campanian of Bulgaria, Middle-Upper Maastrichtian of the UAE/Oman region, ?Danian of France (Vigny).

Family Agariciidae Gray, 1847 (=Lamellofungiidae Alloiteau, 1957; =?Andemantastraeidae Alloiteau, 1952a) Diagnosis. Solitary and colonial; hermatypic. Colony formation mainly by intratentacular budding. Wall absent or synapticulothecal, and usually becoming solid. Septa formed by one fan system of simple trabeculae, rarely porous, margins beaded, directly confluent between centres, united by some compound synapticulae. Endothecal dissepiments mostly absent. Columella trabecular or absent.

Genus Cyathoseris Milne Edwards & Haime, 1849a Type species. Pavonia infundibuliformis Blainville, 1830, Eocene of France (Auvert) (see Milne Edwards & Haime, 1849a).

Diagnosis. Colonial, crateriform, thamnasterioid. Gemmation intracalicinal-circumoral followed by intratentacular-marginal budding with terminal forking. Septa compact to subcompact, confluent. Septal margins beaded. Columella papillose, feebly developed. Synapticulae present. Endothecal dissepiments thin, few in number. Synapticulothecal wall incomplete or absent.

Cyathoseris catadupensis (Vaughan, 1899) Pl. 10, figs 2, 3 v*1899 v1952a v1957 (v)1996 2000 2000

Mesomorpha catadupensis, sp. nov.: Vaughan, p. 246, pl. 41, figs 1–2, 4. Pseudoseris senessei: Alloiteau, p. 663. Pseudoseris senessei Alloiteau: Alloiteau, p. 299, pl. 7, fig. 6, pl. 9, fig. 13. Cyathoseris sp.: Schuster, p. 72, pl. 14, figs 2–4. Mesomorpha catadupensis, Vaughan 1899: Löser, p. 51. Pseudoseris senessei, Alloiteau 1957: Löser, p. 70.


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2002 v2002 (v)2005

Mesomorpha catadupensis Vaughan, 1899: Baron-Szabo, p. 65. Cyathoseris senessei (Alloiteau, 1952a): Baron-Szabo, p. 117, pl. 82, figs 4–5. Mesomorpha catadupensis Vaughan, 1899: Filkorn, p. 124, fig. 2j.

Dimensions. c-c (series)=3–5 mm; c-c (between series)=3–6 mm; s/mm=5–7/2. Description. Massive-lamellar and thamnasterioid colony; corallites grouped in concentric series in juvenile colonies, becoming meandroid in later stages; isolated calices occur frequently; gemmation intracalicinal; septa confluent, straight, subcompact, porous in places, nearly equal in thickness; lateral surfaces covered with rounded granules, varying in size and shape, or long spiny ornamentations; about 10 septa reach the calicinal center, where their inner ends may terminate into claviform thickenings or form prolongations, uniting with neighbouring septa; columella rudimentary or made of a few papillae; synapticulae present throughout the colony; wall synapticulothecal, incomplete; endothecal dissepiments thin, vesicular, rare. Remarks. In having corallites arranged in concentric series, a small papillose columella, compact septa with beaded margins, and a synapticulothecal wall which is incomplete, the material described by Vaughan (1899) as Mesomorpha catadupensis seems to rather correspond to the genus Cyathoseris. Type locality of species. Campanian-Eocene of Jamaica (Catadupa beds). Distribution. Upper Santonian of France, Campanian-Eocene of Jamaica (Catadupa beds and new material), Paleocene of Egypt. New Material. Maastrichtian of Jamaica, NMNH, Coates coll., sample nos.: 379 (=Jerusalem Mountain Inlier); 495a; 520 (=Ducketts Land Settlement).

Cyathoseris formosa d’Achiardi, 1875 Pl. 11, fig. 1 v*1875 1901 1902 1912 1925 v1992 1997 v2003 v2006

Cyathoseris ? formosa, m.: d’Achiardi, p. 75, pl. 14, fig. 4 (type probably lost, but topotypes studied). Cyathoseris cf. formosa d‘Ach.: Oppenheim, p. 205. Cyathoseris formosa: Osasco, p. 116. Cyathoseris formosa d‘Ach.: Oppenheim, p. 116. Cyathoseris formosa d’Achiardi 1875: Felix, pars 28, p. 123 (older synonyms cited therein). Cyathoseris ? eocenica Stemann n. sp.: Budd et al., p. 576, figs. 3.9–3.10. Cyathoseris ? eocenica Stemann, 1992: Bryan et al., p. 34, text-fig. 1C. Cyathoseris formosa d’Achiardi 1875: Schafhauser et al., p. 190. Cyathoseris formosa d’Achiardi, 1875: Baron-Szabo et al., p. 26, fig. 5.7.

Dimensions. d (primary corallite)=20–30 mm; d (secondary ones)=3–9 mm; c-c (series)=5–13 mm; c-c (between series)=5–24 mm; s/mm= 4–7/2; s (secondary corallites)=up to 36. Description. Massive-lamellar, thamnasterioid colony; gemmation circumoral; corallites (secondary) grouped around the main (primary) corallite in meandroid series; septa confluent, straight; thick and compact to subcompact septa alternate with thin and often porous ones; ornamented by delicate, spiniform and very elongated spiny granules laterally; up to 20 septa reach the calicinal center; columella spongy papillose; synapticulae numerous, disposed over the entire colony; wall synapticulothecal, incomplete or absent; endothecal dissepiments sparse, thin, vesicular. Type locality of species. Eocene of Italy (Friaul). Distribution. Maastrichtian of Mexico (Cardenas Formation), Paleocene of the USA (Alabama, Salt



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Mountain), Eocene of Bosnia and Italy, Upper Eocene of Panama, Lower Oligocene of Slovenia and Italy.

Genus Pavona Lamarck, 1801 (=Pseudastraea Reuss, 1864, Type species. Pseudastraea columnaris Reuss, 1864, Oligocene of Slovenia [Gornji Grad]) (=Hydnophorabacia d’Achiardi, 1875, Type species. Hydnophorabacia variabilis d’Achiardi, 1875, Eocene of Italy [Friul]) (= Reussastraea d'Achiardi, 1875, Type species.Reussastraea granulosa d'Achiardi, 1875, Eocene of Italy [Friul]). Type species. Madrepora cistata Ellis & Solander, 1786, Recent, Red Sea.

Diagnosis. Colonial, variably thamnasterioid-submeandroid, hydnophoroid, submassive to foliaceous. Gemmation intracalicinal, often resulting in calicinal series separated by generally tectiform collines. Costosepta compact to subcompact, confluent or nonconfluent between corallites of adjacent series, confluent between corallites of same series. Corallite wall absent in same corallite series, synapticulothecal between series, often becoming solid, well-developed. Columella trabecular.

Pavona bronni (Haime, 1850) Text-Fig. 26 v*1850 v1856 1873 1880 1914 1921 1925 v1973 v1980 non1995 2002

Hydnophora Bronni: Haime, in Bellardi, p. 229 (topotypes studied). Monticularia venusta: Catullo, p. 76, pl. 17, figs 2a–b (topotypes studied). Thamnastrea Brevipes: Stoliczka, p. 42, pl. 9, figs 2–3. Reussastraea grandis, Duncan: Duncan, p. 45, pl. 10, figs 1–4. Thamnastraea brevipes Stoliczka 1873: Felix, pars 7, p. 195. Mycetoseris minuta n. f.: Prever, p. 72, pl. 10, figs 5–6. Pavonia grandis Duncan sp. 1880: Felix, pars 28, p. 129. Hydnophora venusta (Catullo) 1856: Barta-Calmus, p. 303, pl. 20, figs 1–6.. Pavona bronni (Haime, 1850): Pfister, p. 58, pl. 2, figs 1–5, pl. 3, figs 5–8 (older synonyms cited therein). Fungiastrea ? brevipes (Stoliczka 1873): Löser & Raeder, p. 48. Fungiastrea brevipes (Stoliczka, 1873): Baron-Szabo, p. 147.

Dimensions. d=5–8 mm, in areas of intense budding the distance is around 3 mm; s=24 to up to around 30, around 18 in juvenile corallites; s/mm=5–8/5. Description. Thamnasterioid colony, corallites irregularly disposed over surface; corallites circumscribed, appear cerioid; costosepta, generally 6–12 of which reach corallite centre, are equal or irregularly alternate in length and thickness; columella often formed by only one papilla, appearing sublamellar or irregularly styliform. Remarks. For over a century the taxonomic position of Thamnastrea brevipes from the Maastrichtian of India as originally proposed by Stoliczka was accepted. In 1995, Löser & Raeder tentatively grouped this taxon with the genus Fungiastrea. However, the apparent presence of costae as well as thick and very coarsely granulated septa of the agariciid type exclude the species T. brevipes from both genera. In addition to these features, in showing compact to subcompact septa that are confluent or nonconfluent, and corallites that are in thamnasterioid to submeandroid arrangement, the Indian material closely corresponds to the genus Pavona.


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Text-Fig. 26 Pavona bronni (Haime, 1850). A, B, topotype, MPUR 3477, Oligocene of Italy; A, upper surface, scale bar: 4 mm; B, close-up, scale bar: 7 mm; C, sketch of upper surface of colony as figured in Duncan (1880), as type material of Reussastraea grandis, Upper Paleocene of Pakistan, scale bar: 8 mm; D, E, illustrations of upper surface view of colony as figured in Stoliczka (1873, as holotype of Thamnastraea brevipes Stoliczka, 1873), Maastrichtian of India; D, scale bar: 14 mm; E, scale bar: 8 mm.

Pfister (1980, p. 58–60) studied around 150 specimens of Pavona bronni (Haime, 1850) including type material and material collections from the Oligocene of Italy. She came to the conclusion that the species P. bronni was characterized by an extremely wide range of variation regarding the dimensions of its skeletal structures such as collines, polyp integration (cerioid, hydnophoroid, meandroid, sometimes circumorally arranged), septal thickness, corallite diameter, number of septa in each corallite, septal density in meandroid/ hydnophoroid series, frequency regarding the occurrence of individual corallites vs. corallites arranged in meandroid series, and a very variably developed columella (spongy, papillose, sublamellar, consisting of only one papilla, etc), the latter of which, in general, is rudimentary at the upper surface. Based on this material, Pfister could show that in many specimens several sets of characteristics co-occurred, each of which had been used to define individual species. Therefore, she taxonomically united these forms with Pavona bronni but grouped them in 5 morphological categories, which form a continuous series with Pavona bronni forma a (=Pavona minuta [Prever, 1921]) on one end and Pavona bronni forma e (=Monitcularia venusta Catullo, 1856) on the other. The specimens presented here (Fig. 26) closely correspond to the morphology and range of variation of Pavona bronni: the specimens in Fig. 26 A and B (= Pavona bronni [Haime, 1850]) and the specimen in Fig. 26 C (= Reussastraea grandis Duncan, 1880) correspond to Pavona bronni forma b, whereas the specimen in Fig. 26 D and E (= Thamnastrea brevipes Stoliczka, 1873) corresponds to Pavona bronni forma a. Therefore, these species are considered synonymous. Type locality of species. Oligocene of Italy (Castelgomberto). Distribution. Maastrichtian of India (Arrialoor Group), Upper Paleocene of Pakistan, Eocene of northern Spain (Catalonia), Oligocene of Italy.



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Genus Morphastrea d’Orbigny, 1850 Type species. Agaricia ludoviciana Michelin, 1845, Cenomanian of France (Mans).

Diagnosis. Colonial, lamellar, thamnasterioid-submeandroid. Gemmation extracalicinal and marginal. Septa confluent to subconfluent, coarsely granulated laterally, pores present. Synapticulae abundant. Columella styliform. No wall between the corallites. Trabeculae thick.

Morphastrea escharoides (Goldfuss, 1826) Pl. 11, figs 3, 4 v*1826 1850 1851a 1857 1884 1914 v1925 v1935 1999 2000 v2002

Astrea escharoides nobis: Goldfuss, vol. 1, p. 68, pl. 23, fig. 2. Morphastrea escharoides: d’Orbigny, vol. 2, p. 277. Dimorphastea escharoides: Milne Edwards & Haime, p. 108. Dimorphastea escharoides: Milne Edwards, vol. 2, p. 586. Thamnastraea escharoides: de Fromentel, p. 592. Dimorphastea escharoides Goldfuss sp. 1826: Felix, pars 7, p. 200. Dimorphastraea escharoides Goldfuss spec.: Umbgrove, p. 111. Meandrarea clarendonensis, sp. n.: Wells, p. 191, pl. 11, fig. 1. Synastrea escharoides: Leloux, p. 193, fig. 2. Synastrea escharoides (Goldfuss 1826): Löser, p. 77. Morphastrea escharoides (Goldfuss, 1826): Baron-Szabo, p. 118–119, text-fig. 42.

Dimensions. c-c=4–8 mm; s/mm=15–20/5. Description. Meandro-thamnasterioid colony; corallites irregularly distributed or arranged in meandroid series; septa compact with some pores, confluent to subconfluent, subequal; up to 12 septa reach calicinal center; columella irregularly trabecular or styliform, deep in corallites. Type locality of species. Upper Maastrichtian of The Netherlands. Distribution. Campanian-Maastrichtian of Jamaica, Upper Maastrichtian of The Netherlands.

PLATE 11 Fig. 1 Cyathoseris formosa d'Achiardi, 1875, UNAM, IGM 8721, Maastrichtian of Mexico, cross thin section, scale bar: 8 mm. Fig. 2 Anisoria vidali (Mallada, 1892), topotype (holotype of Anisoria batalleri Reig Oriol, 1987), MGSB, Reig Oriol coll., 44251, Maastrichtian of Spain, upper surface of colony, scale bar: 15 mm. Fig. 3 Morphastrea escharoides (Goldfuss, 1826), holotype, IPB, Goldfuss, coll., 227, Upper Maastrichtian of The Netherlands, upper surface of colony in ‘steinkern’ preservation, scale bar: 9 mm. Fig. 4 Morphastrea escharoides (Goldfuss, 1826) (holotype of Meandrarea clarendonensis Wells, 1935), BMNH, R.28929, Campanian-Maastrichtian of Jamaica, upper surface of colony, scale bar: 9 mm. Fig. 5 Trochoseris aperta Duncan, 1864, NMNH, Coates coll., no. 569c, Maastrichtian of Jamaica. 5a, close-up, scale bar: 3 mm; 5b, cross thin section, scale bar: 6 mm. Fig. 6 Trochoseris aperta Duncan, 1864, holotype, BMNH R.39647, Eocene of Pakistan, upper surface of corallum, scale bar: 10 mm.


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Genus Heterogyra Reuss, 1868 (=Felixastraea Oppenheim, 1930a, Type species. Cyathoseris zitteli Felix, 1903a, Santonian of Austria [Gosau Group]). Type species. Heterogyra lobata Reuss, 1868, Oligocene of Italy.

Diagnosis. Colonial, massive to submassive or explanate, thamnasterioid-meandroid. Gemmation intracalicinal (polystomodaeal budding with terminal forking). Calicinal series free laterally near summits or separated by tectiform to tholiform collines, discontinuous. Ambulacrae present in places. Corallites arranged in calicinal series or calicinal centres with lamellar linkages. Costosepta compact, can be subcompact in younger septa, subconfluent to confluent, laterally finely granulated. Columella parietal-papillose. Synapticulae abundant. Endothecal dissepiments thin, subtabulate. Wall synapticulothecal, often becoming solid.

Heterogyra murchisoni (d’Archiac & Haime, 1853) Text-Fig. 27 v*1853 1880 ?1880 1902 v1927 1996

Pachyseris Murchisoni n. sp.: d’Archiac & Haime, p. 194, pl. 12, figs 9a, 9b. Pachyseris Murchisoni, d’Archiac & Haime: Duncan, p. 46, pl. 14, figs 3, 4. Cyathoseris orientalis, Duncan: Duncan, p. 47, pl. 6, figs 7–10. Oroseris undata n. sp.: Wanner, p. 104, pl. 14, fig. 13. Pachyseris Murchisoni J. Haime sp. 1851: Felix, pars 35, p. 368 (older synonmys cited therein). Comoseris undata (Wanner, 1902): Schuster, Appendix list.

Dimensions. d=2.5–6.5 mm; c-c (series)=5–10 mm, in areas of intense budding the distance might be smaller; s/mm=5–8/2. Description. Thamnasterioid-meandroid colony; corallites often arranged in calicinal series that are up to 30 mm long; individual corallites irregularly scattered over colony; septa arranged in 2–3 size orders, in areas of intense budding septa of a fourth size order may be present.

Text-Fig. 27 Heterogyra murchisoni (d’Archiac & Haime, 1853). A, holotype BMNH R.29143, Eocene of Pakistan, upper surface of colony; B, sketch of upper surface of colony as provided by Duncan (1880, figured as type material of Cyathoseris orientalis), Paleocene of Pakistan; C, sketch of upper surface of colony in Wanner (1902, figured as Oroseris undata Wanner, 1902), Upper Danian of Egypt, (size calculated based on the description by Wanner 1902, according to which 32 septa are found in1 cm); scale bar: 10 mm.

Remarks. Wanner (1902) stated that in the specimen he described as Oroseris undata there was no colu-


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mella. However, in the illustration provided by Wanner subpapillose columellar structures seem to be present, thus, in addition to all other characters that are present regarding corallite integration, septal and wall developments, as well as axial structures of the corallite series, it closely agrees with the genus Heterogyra. Because the collines present in the specimen that Wanner (1902) described as Oroseris undata, are mainly steep and tectiform, this specimen seems to differ from the type specimen of Heterogyra murchisoni (d’Archiac & Haime, 1853) and the material decribed by Duncan (1880). Both of the latter have calicinal series that are rather flat and separated by mainly tholiform collines. But as shown in Figure 27, both types of collines are present in all of the specimens. Therefore, they are considered to be within the range of variation of the same species. Type locality of species. Eocene of Pakistan (probably near Hyderabad). Distribution. Paleocene-Eocene of Pakistan, Upper Danian of Egypt, Eocene of Bosnia, France, and Spain, Miocene of Borneo and Pakistan.

Genus Anisoria Vidal, 1917 (=Plesiodiploria Alloiteau, 1960. Type species. Plesiodiploria scutifera: Alloiteau, 1960, Maastrichtian of Spain). Type species. Meandrina vidali Mallada, 1892, Maastrichtian of Spain.

Diagnosis. Colonial, meandroid. Gemmation intracalicinal. Calicinal centres indistinct, arranged in long, wavy series, separated by ambulacrae. Septa compact, granulated. Columella absent but trabecular extensions of septal axial ends may form pseudo-columella. Synapticulae present. Wall parathecal-septoparathecal. Endothecal dissepiments numerous, occurring near wall. Remarks. According to Alloiteau (1957) the microstructure of Anisoria is analogous to the kind in Trochosmilia corniculum.

Anisoria vidali (Mallada, 1892) Pl. 11, fig. 2 v*1892 v1917 v1937a 1957 v1960 v1987 v1987 2000 v2002

Meandrina Vidali: Mallada, p. 160 (topotype studied). Anisoria vidali: Vidal, p. 5, pl. 1, figs 1–4, pl. 2, fig. 6 (topotype studied). Anisoria Vidali Mallada sp. 1917: Bataller, p. 166, 2 text-figs on p. 166 (topotype studied). Anisoria vidali (Mallada, 1892): Alloiteau, p. 283. Plesiodiploria scutifera nov. sp.: Alloiteau, p. 94, pl. 1, figs 4–7. Anisoria vidali (Mallada in Vidal 1917): Reig Oriol, p. 5, pl. 1, figs 1–5, pl. 2, figs 2–3. Anisoria batalleri n. sp.: Reig Oriol, p. 6, pl. 1, figs 6, 7, pl. 2, fig.. 1. Anisoria vidali, Vidal, 1917: Löser, p. 8. Anisoria vidali (Mallada, 1892): Baron-Szabo, p. 120, text-fig. 44

Dimensions. c-c (wall-wall)=2–2.5 mm, in late budding stage up to 3 mm; ambulacrae: up to around 8 mm; s/ mm=14–16/5. Description. Meandroid corallum; calicinal series separated by ambulacrae, the latter of which are generally between 1–4 mm wide (estimated from the illustrations in Vidal 1917, and Bataller 1937a); septa compact, nonconfluent, finely granulated laterally, arranged in 2 size orders; S3 irregularly occur.



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Remarks. Because in Mallada (1892) the type material of Anisoria vidali (Mallada, 1892) was mentioned without description or illustration, the information regarding skeletal elements of the holotype were taken from Vidal (1917) and Bataller (1937a), both authors of which provided proper documentation of the type material. The skeletal structures of the holotype of Anisoria batalleri Reig Oriol, 1987, from the Maastrichtian of northern Spain closely correspond to all of the features of Anisoria vidali. Therefore, the two species are interpreted to be synonymous. Distribution. Campanian-Maastrichtian of Spain.

Genus Valloria Vidal, 1874 Type species. Valloria egozcuei Vidal, 1874, Maastrichtian of Spain.

Diagnosis. Colonial, massive, meandroid. Gemmation intracalicinal. Calicinal series separated by ambulacrae. Costosepta confluent, compact with perforated axial ends. Synapticulae numerous, mainly peripherally. Endothecal dissepiments sparse or absent. Columella substyliform to sublamellar. No wall between calicinal series. Affinities. Septa as in Heterogyra, but corallum meandroid, columella substyliform to sublamellar, and wall absent.

Text-Fig. 28 Valloria egozcuei Vidal, 1874, sketch based on the illustration of the holotype in Vidal (1874), Maastrichtian of Spain, upper surface of colony,scale bar: 8 mm.

Valloria egozcuei Vidal, 1874 Text-Fig. 28 *1874 1892

Valloria Egozcuei, n. sp.: Vidal, p. 39, pl. 7, figs 44a–b. Valloria Egozcuei Vidal: Mallada, p. 160.


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1937a 1957 2002 2002

Valloria Egozcuei Vidal 1874: Bataller, p. 164, text-fig. on p. 164. Valloria Egozcuei Vidal (1874): Alloiteau, p. 325. Valloria egozcuei Vidal 1874: Löser et al., p. 711 (older synonyms cited therein). Valloria egozcuei Vidal, 1874: Baron-Szabo, p. 121, text-fig. 45.

Dimenisons. d (wall-wall)=1 mm; c-c=1.5–5 mm; s/mm=33/5. Description. Massive, meandroid colony; calicinal series long and straight or very wavy, separated by ambulacrae which are around 1–3 mm; costosepta arranged in 2 alternating size orders; Type locality of species. Maastrichtian of Spain (La Posa, Isona, Lléida). Distribution. Maastrichtian of Spain.

Genus Trochoseris Milne Edwards & Haime, 1849 (=Elliptoseris Duncan, 1880, Type species. Trochoseris aperta Duncan, 1864, Eocene of Pakistan). Type species. Anthophyllum distortum Michelin, 1844, Eocene of France (Auvert) (see Milne Edwards & Haime, 1849a).

Diagnosis. Solitary, turbinate or trochoid, fixed. Septa are subcompact to porous, beaded marginally. Columella papillose. Synapticulae abundant. Endothecal dissepiments thin, sparse or absent. Wall synapticulothecal.

Trochoseris aperta Duncan, 1864 Pl. 11; figs 5a–6 v*1864 v1880 1880 1880 v1880 v1899 v1899 v1919 v1919 1925 1925 1925 v1934 v1941a ?1974 1981 1987 1987 ?1991 1992 v1992 2000 v2002

Trochoseris aperta, nobis: Duncan, p. 303, pl. 19, fig. 5. Elliptoseris aperta, Duncan: Duncan, p. 48, pl. 8, figs 3–6 (topotypes studied). Turbinoseris elegans, Duncan: Duncan, p. 51, pl. 16, figs 3, 4. Trochoseris difformis, Reuss: Duncan, p. 47, pl. 11, figs 9, 10. Trochoseris aperta, nobis: Duncan, p. 107, pl. 27, figs 9, 10. Trochosmilia hilli, sp. nov.: Vaughan, p. 233, pl. 36, figs 1–4. Trochoseris catadupensis, sp. nov.: Vaughan, p. 242, pl. 39, figs 5, 6 Topotypes studied). Trochoseris catadupensis Vaughan: Vaughan, p. 426. Trochoseris meinzeri, new species: Vaughan, p. 426, pl. 106, figs 2, 2a, 2b. Trochoseris catadupensis Vaughan 1899: Felix, pars 28, p. 120. Trochoseris meinzeri Vaughan 1919: Felix, pars 28, p. 121. Elliptoseris aperta Duncan 1880 Felix, pars 28, p. 147. Trochoseris catadupensis Vaughan: Wells, p. 78, pl. 2, figs 9, 10. Trochoseris catadupensis Vaughan: Wells, p. 288, pl. 1, fig. 1. Trochoseris (?) sp. cf. T. meinzeri Vaughan: Frost & Langenheim, p. 197, pl. 61, fig. 4. Trochosmilia oldhami Duncan: Abed & El-Asa‘ad, p. 275, pl. 1, figs 2a–b. Trochoseris catadupensis Vaughan: Kuzmicheva, p. 60. Fungiacyathus bodrakensis Kuzmicheva, sp. nov.: Kuzmicheva, p. 96, pl. 11, figs 1, 2. ?Trochoseris sp.: Stemann, in Bryan et al., p. 33. Trochoseris catadupensis Vaughan, 1899: Budd et al., p. 593. Trochoseris meinzeri Vaughan, 1919: Budd et al., p. 593. Trochoseris catadupensis Vaughan 1899: Löser, p. 82. Trochoseris catadupensis Vaughan: Mitchell, p. 6 ff., table 1 (topotypes studied).



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v2002 v2003 2005 2005 v2006

Trochoseris catadupensis Vaughan, 1899: Baron-Szabo, p. 122, pl. 84, figs 3, 5. Trochoseris catadupensis Vaughan, 1899: Schafhauser et al., p. 190, tab. 1. Trochosmilia hilli Vaughan, 1899: Filkorn, p. 121, fig. 2d. Trochoseris catadupensis Vaughan, 1899: Filkorn et al., p. 123, fig. 2h. Trochoseris aperta Duncan, 1864: Baron-Szabo et al., p. 25, fig. 5.4.

Dimensions. d (max)=6–74 mm; d (min)=7–70 mm; s (adult)=300–500; s/mm=12–14 (16 areas of intense budding)/5; d (min)/d (max) (adult)=0.70–0.89; d (min)/d (max) (juvenile)=0.53–0.68. Description. Solitary, trochoid or patellate, expanding rapidly from the base of attachment; calice broad and shallow in adult stages, slightly elongated in outline; calice deep and concave in juvenile stages, elliptical in outline; calicular pit circular or elongate; costosepta long, straight or wavy, crowded, compact, subcompact to porous in younger cycles, and developed in 6 to 7 cycles in 6 systems in adult stages; in places an irregular development following the Pourtalès present; lateral septal surfaces covered by delicate, subcarinate granulations, spiniform and rounded granules, and very elongated spiny prolongations; twelve to 40 septa extend to calicinal center, joining spongy-papillose columella; synapticulae numerous, disposed over the entire corallite; endothecal dissepiments sparse, thin, vesicular; wall synapticulothecal; epicostate lamellae are present; microstructure is composed of thick monaxial trabeculae, giving off secondary ones, and ?polyaxial trabeculae. Remarks. Compared to the specimen that is figured in Plate 11, figs 5a–b, the image presented on Plate 11, fig. 6 represents more juvenile corallite which is characterized by straighter septa and a deeper corallum. In later ontogentical stages, the central part of the corallum becomes rather broad and flat with wavy, winglike peripheral areas and flexuous septa. In the more juvenile corallites, the beginning of wing-like developments in the peripheral areas can be seen in the central upper and left part of the specimen in Plate 11, fig. 6. Type locality of species. Eocene of Pakistan (Sind, Kurachee). Distribution. Campanian of Cuba, Campanian-Lower Maastrichtian of central Saudi Arabia, CampanianMaastrichtian (new material) and Eocene of Jamaica, Maastrichtian of Mexico (Ocozocuautla and Cardenas Formations), ?Paleocene of the USA (Alabama, Salt Mountain Limestone), Danian of Ukraine, PaleoceneEocene of Pakistan, Eocene of Cuba and Panama, ?Upper Eocene of Mexico (Ixtaclum shale). New Material. Maastrichtian of Jamaica, NMNH, Coates coll., sample nos.: 315a; 415a; 427a (=Jerusalem Mountain Inlier);497p; J-71-129A (=Ducketts Land Settlement); 533w; 533x; 533y (=Shaw Castle, Maldon Formation); 565; 569a; 569c; 569d; 590b; 593a; (=probably Cambridge railway area); J-71-7a; (=Glenbrook); J-71-13A-2v; (=Vaughnsfield); J3444h (=Welcome Hall).

Genus Vaughanoseris Wells, 1934 Type species. Vaughanoseris catadupensis Wells, 1934, Maastrichtian of Jamaica.

Diagnosis. Solitary, discoid-patellate, slightly elliptical in outline. Free, fixed in early stages. Costosepta compact, granulate. Columella spongy-papillose. Synapticulae, endothecal dissepiments, and epitheca present.

Vaughanoseris catadupensis Wells, 1934 Pl. 12, figs 1a–c v*1934

Vaughanoseris Catadupensis, new species: Wells, p. 81, pl. 3, figs 11–13, pl. 5, fig. 3.


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v1956 1984 2000 2002 v2002

Vaughanoseris catadupensis Wells 1934: Wells, p. F381, figs 272.1a–c. Vaughanoseris cf. V. catadupensis (Vaughan) Wells, 1934: Pal. et al., p. 58, pl. 1, 5–6, pl. 3, fig. 26. Vaughanoseris catadupensis, Wells 1934: Löser, p. 86. Vaughanoseris catadupensis Wells: Mitchell, p. 6ff., table 1. Vaughanoseris catadupensis Wells, 1934: Baron-Szabo, p. 123, pl. 85, figs 1, 4–5.

Dimensions. d=30 x 32 mm; s=192; h=15 mm. Description. Discoid-patellate corallum; costosepta straight, arranged in 6 complete cycles in 6 systems; S1 and S2 nearly equal; ?paliform structures before S3 and S4 irregularly present. Type locality of species. Maastrichtian of Jamaica (Catadupa Midway, Cambridge). Distribution. Campanian-Maastrichtian of Jamaica and India (Ladakh).

Genus Ogilviastraea Oppenheim, 1930 Type species. Placohelia bigemmis Felix, 1903a, Santonian of Austria (Gosau Group)

Diagnosis. Colonial, dendroid. Gemmation intracalicinal (bigemmation). Calices are subcircular or elliptical. Perithecal dissepiments abundant, vesicular. Costosepta compact, arranged radially or bilaterally, finely granulated laterally. Paliform structures and synapticulae present. Columella trabecular, thin and lamellar or variably spongy-papillose. Endothecal dissepiments few in number. Wall synapticulothecal-septothecal, forming a stereozone.

Ogilviastraea bigemmis (Felix, 1903) Pl. 12, fig. 2 v*1903a 1930a ?1952b v1982 ?1984 ?1987 2000 2000 2000 v2001 v2002 2002

Placohelia bigemmis nov. sp.: Felix, p. 325, pl. 22, figs 12, 13. Ogilviastraea bigemmis (Felix): Oppenheim, p. 357, pl. 34, figs 8–12. Diblasus sp.: Alloiteau, p. 12, pl. 1, figs 13–15. Ogilviastraea bigemmis (Felix): Beauvais, 1982: vol. 2, p. 113, pl. 37, fig. 4. Ogilviastraea stoliczkae sp. nov.: Pal et al., p. 59, pl. 2, fig. 19. Euphyllia turgainensis Kuzmicheva, sp. nov.: Kuzmicheva, p.142, pl. 25, figs 2a–b. Euphyllia turgainensis, Kuzmicheva 1987: Löser, p. 35. Ogilviastraea bigemmis, Felix 1903: Löser, p. 56. Ogilviastraea stoliczkae, Pal et al., 1984: Löser, p. 56. Ogilviastraea cf. bigemmis (Felix, 1903): Baron-Szabo, p. 265, fig. 3A. Ogilviastraea bigemmis (Felix, 1903): Baron-Szabo, p 84, pl. 85, figs 2–3, pl. 86, fig. 1. Ogilviastraea stoliczkae Pal et al., 1984: Baron-Szabo, p. 84.

Dimensions. d (max)=up to 15 mm; d (min)=up to 9 mm; s=5–7/2. Description. Dendroid colony with slightly elongated calices becoming elliptical due to bigemmation; calices subcircular or elliptical in outline; perithecal dissepiments abundant, vesicular; costosepta compact, covered with spiniform and rounded granules laterally; septa arranged in up to 5 irregularly developed size orders, radially or bilaterally; paliform structures and synapticulae present; columella trabecular, forming a thin lamella or variably spongy-papillose; endothecal dissepiments thin, subtabulate, few in number; wall syn-



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apticulothecal-septothecal, forming a stereozone. Type locality of species. Santonian of Austria (Gosau Group). Distribution. Turonian-Santonian of Austria, ?Campanian-?Maastrichtian of India (Ladakh), ?Lower Maastrichtian of Kazakhstan, ?Maastrichtian of Senegal, Middle-Upper Maastrichtian of Jamaica (new material). New Material. Middle-Upper Maastrichtian of Jamaica, NMNH, Coates coll., sample nos.: J-66-13-55a (=Jerusalem Mountain Inlier); 595c (=probably Cambridge area).

Family Agathiphylliidae Vaughan & Wells, 1943 Diagnosis. Solitary and colonial. Colony formation by extratentacular budding. Corallite wall synapticulothecal near calice, septothecal or parathecal below. Septocostae more or less confluent between calices. Septa of lower cycles composed of two fan systems of mostly compound trabeculae; higher cycles with one fan system, frequently perforate along inner ends; upper margins with acute transverse dentations. Inner fan system usually projecting as paliform lobes. Columella composed of twisted trabecular processes, well developed. Endothecal and exothecal dissepiments abundant, vesicular.

Genus Agathiphyllia Reuss, 1864 Type species. Agathiphyllia explanata Reuss, 1864, Oligocene of Italy (designation by Vaughan, 1919).

Diagnosis. Colonial, submassive, plocoid, Gemmation extracalicinal. Costosepta compact, nonconfluent; costae strongly developed. Columella trabecular; strong paliform lobes present. Wall synapticulothecal, septothecal due to secondary thickening. Endothecal dissepiments vesicular.

Agathiphyllia bosniaca (Oppenheim, 1912) Pl. 12, fig. 5 parsv1900 *1912 1925 (v)1996 v1997

Columastraea bicoronata, sp. nov.: Gregory, p. 32, pl. 2, Fig. 7 non Figs 8, 9. Helistraea bosniaca n. sp.: Oppenheim, p. 119, pl. 16, figs 1–1c. Orbicella bosniaca Oppenheim 1912: Felix, pars 28, p. 63. Hexakoralle, Morphotyp 17: Tragelehn, p. 198, pl. 62, fig. 7. Agathiphyllia cf. blaviensis Chevalier, 1954: Vecsei & Moussavian, p. 128, pl. 35, fig. 1.

PLATE 12 Fig. 1 Vaughanoseris catadupensis Wells, 1934, holotype, NMNH, I74485, Maastrichtian of Jamaica, scale bars: 7 mm. 1a, cross view; 1b, base of corallum; 1c, lateral view. Fig. 2 Ogilviastraea bigemmis (Felix, 1903), NMNH, Coates coll., no. 595c, Middle-Upper Maastrichtian of Jamaica, cross thin section, scale bar: 2 mm. Fig. 3 Goniopora imperatoris Vaughan, 1919, BMNH, AZ 415, cross thin section, Middle-Upper Maastrichtian of the UAE/Oman border region, scale bar: 2 mm. Fig. 4 Goniopora imperatoris Vaughan, 1919, NMNH, Coates coll., no. 570c, cross thin section, Maastrichtian of Jamaica, scale bar: 2 mm. Fig. 5 Agathiphyllia bosniaca (Oppenheim, 1912), BMNH, R.5033 (syntype of Columnastraea bicoronata Gregory, 1900), Upper Paleocene of Somalia, upper surface, scale bar: 3 mm. Fig. 6 Goniopora websteri (Bowerbank, 1840) (figured as Litharaeaopsis subepithecata [Oppenheim, 1912], in Drobne et al. 1988), SAZU, Dv 24-982c, cross thin section, Paleocene of Slovenia, scale bar: 5 mm (photograph courtesy D. Turnšek). Fig. 7


BARON-SZABO

Goniopora elegans (Leymerie, 1846), BMNH, AZ 417, Middle-Upper Maastrichtian of the UAE/Oman border region, cross thin section, scale bar: 4 mm.



139

Dimensions. d (max.)=3–5.5 mm, juvenile corallites may be around 2.5 mm; c-c=3.5–6.5 mm; s=24+s4, in juvenile corallites the number usually around 20. Description. Colonial, massive, corallites circular to slightly elliptical in outline; costosepta compact, developed in 3 complete cycles with the beginning of the fourth cycle in 6 systems; S1 reach corallite centre; columella spongy-papillose; generally 12 paliform lobes present. Remarks. The type collection of Columastraea bicoronata Gregory, 1900, consists of specimens belonging to different taxonomic groups. The specimen BMNH R.5035 belongs to the genus Stephanaxophyllia and was chosen to be the lectotype of the species bicoronata (Baron-Szabo 2006, p. 61). The second syntype BMNH R.5033 belongs to the genus Agathiphyllia and corresponds to Oppenheim’s species A. bosniaca. Type locality of species. Eocene of Bosnia. Distribution. Paleocene of Austria and Italy, Upper Paleocene of Somalia, Eocene of Bosnia.

Genus Pattalophyllia D’Achiardi, 1867 (=Leptaxis Reuss, 1867 non Lowe, 1852, Type species. Leptaxis elliptica Reuss, 1867, Oligocene of Italy [Monte Grumi]); (=Stephanosmilia Reuss, 1874 non de Fromentel, 1862 [=Cricocyathus Quenstedt, 1881], Type species. Cyathophyllia annulata Reuss, 1868, Oligocene of Italy [Vicentin]); (=Petrophylliella Felix, 1925. Type species. Caryophyllia grumi Catullo, 1852, Oligocene of Italy [Monte Grumi]). (=Vivesastraea Alloiteau & Tissier, 1958, Type species. Vivesastraea villattae Alloiteau & Tissier, 1958, Paleocene of France); (=Pattalophylliopsis Vaudois-Miéja, 1964, Type species. Pattalophylliopsis epitheca Vaudois-Miéja, 1964, Eocene of France [Cluses]); (=Chevalieriphyllia Russo, 1979, Type species. Pattalophyllia costata d’Achiardi, 1881, Eocene of Italy) Type species. Turbinolia subinflata Catullo, 1856, Eocene of Italy.

Diagnosis. Solitary, variably conical (e.g. turbinate, trochoid). Temporary colonial status due to lateral gemmation may occur. Septa compact-subcompact (older cycles) to perforated (younger cycles), strongly granulated laterally. Pali irregularly before all septal cycles, generally very elongate. Columella trabecular, variably and well-developed (e.g., spongy, papillose, net-like, lamellar). Endothecal dissepiments thin, vesicular, numerous. Wall synapticulothecal and septothecal. Peripheral stereozone present. Epithecal wall present or reduced. Remarks. Alloiteau & Tissier (1958) created the genus Vivesastraea and presented several figures of the type material, one of which does not correspond to the photographs of the holotype nor to the diagnosis of the genus. According to the authors, their text-fig. 3 is an illustration of the holotype of the type species. This textfigure shows a specimen with 24 septa arranged according to the Pourtalés plan, a lamellar columella, as well as the absence of a stereozone and paliform lobes. Even though the text-figure corresponds exactly to the description of the type species, Vivesastraea villattae, it completely differs from the diagnosis the authors gave for the genus Vivesastraea. However, both the description and text-figure 3 match in every detail the ones presented for a specimen they assigned to Asterosmilia prolifer, illustrated on plate 2, fig. 4’b’. According to Alloiteau & Tissier (1958), the specimen illustrated on plate 2, figures 3 a–b and 3 a’–b’ represents the holotype of the type species of the genus Vivesastraea. These figures very well correspond to the generic diagnosis. Even though they differ from the species description, they are accepted here as representing the holotype. Because a description of the type species has never been provided, the illustrations of the type as pre-


BARON-SZABO

sented in Alloiteau & Tissier (1958, pl. 2, figs 3 a–b and 3 a’–b’) can be used to give the following specific diagnosis: Solitary, turbinate; corallite diameter 14 mm, height 10 mm; costosepta compact (older cycles) to subcompact (younger cycles), coarsely granulated and carinated laterally, around 100 septa present, which are arranged in (?) 5 complete cycles in (?) 6 systems, first 3 cycles reach the corallite centre; paliform lobes elongate, occurring irregularly before septa of all cycles; wall synapticulothecal and septothecal; peripheral stereozone present; columella trabecular; synapticulae abundant; endothecal dissepiments sparse. Based on the species Pattalophyllia costata d’Achiardi, 1881, Russo (1979, p. 56) created the genus Chevalieriphyllia, indicating that it differed from Pattalophyllia only by lacking pali. Although no pali can be seen in the figure of the specimen called a “syntype” of Chevalieriphyllia costata see Russo (1979, pl. 5, fig. 5), pali appear to be irregularly present in the figure of the specimen that Russo identified as the “lectotype” of Chevalieriphyllia costata in Russo (1979, pl. 5, figs 2a and 2b). Therefore, the genus Chevalieriphyllia Russo, 1979, is considered a junior synonmy of Pattalophyllia. In the original description of Stephanosmilia Reuss, 1874 (see Reuss 1874, p. 28), it is stated that the type specimens (1874/3/54 #4 and #5, originals of the figures in Reuss, 1874, pl. 43, figs 4 and 5) lacked endothecal dissepiments (…‘Mangel an Endothecallamellen‘…). However, re-investigation of the type material revealed that endothecal dissepiments occur deep in the corallum and are very abundant. Thus, since the absence of this feature represented the only difference to Pattalophyllia, the genus Cricocyathus Quenstedt, 1881 (ex Stephanosmilia Reuss, 1874 non Fromentel, 1862) closely corresponds to the genus Pattalophyllia D’Achiardi, 1867, in all skeletal structures.

Pattalophyllia grumi (Catullo, 1852) Text-Fig. 29 v1852 v1856 v1868 v1874 v1874 1881 1885 1885 v1901 1925 1941 (v)1958 v1960 v1973 v1973 1974 1979

Caryophyllia grumi: Catullo, p. 11, pl. 1, fig. 2 (topotypes studied). Caryophyllia grumi Catullo: Catullo, p. 45, pl. 6, fig. 2 (topotypes studied). Cyathophyllia annulata: Reuss, pl. 1, fig. 10. Trochocyathus peziza: Reuss, p. 19, pl. 54, fig. 5. Stephanosmilia annulata (Reuss): Reuss, p. 28, pl. 43, figs 3–5, pl. 46, figs 5–6. Pattalophyllia costata: d’Achiardi, p. 241. Circophyllia annulata (Reuss): Felix, p. 394–397. Petrophyllia grumi (Catullo): Felix, p. 398–400. Pattalophyllia cyclolitoides Michelin sp.: Oppenheim, p. 60, pl. 2, figs 1–7b. Petrophylliella grumi Catullo sp. 1847: Felix, pars 28, p. 59. Circophyllia truncata (Brongniart): Solè Sabaris, p. 150. Vivesastraea villattae nov. sp.: Alloiteau & Tissier, p. 261, pl. 2, figs 3a–b’. Cricocyathus tarraconensis nov. sp.: Alloiteau, p. 104, pl. 2, fig. 9. Petrophylliella grumi (Catullo): Barta-Calmus, p. 246, pl. 11, figs 1–2 (topotypes studied). Cricocyathus annulatus (Reuss): Barta-Calmus, p. 422–424, pl. 49, figs 22–23. Syzygophyllia (?) sp.: Frost & Langenheim, p. 289, pl. 111, fig. 1. Russo

pars1979

Chevalieriphyllia costata (d’Achiardi): Russo, p. 56-58, pl. 5, figs 2a-3b non fig. 5, pl. 6, figs 1a2.

v1980 1992

Cricocyathus grumi (Catullo): Pfister, p. 79, pl. 14, fig. 6 (topotypes studied). Pattalophyllia costata d’Achiardi, 1881: Darga, p. 80.



141

1993 (v)1996 (v)1996 2002 (v)2002

Cricocyathus grumi (Catullo) 1852: Alvarez Perez, p. 180, pl. 11, figs 5–8 (older synonyms cited therein). Solitäre Hexakoralle: Tragelehn, p. 199, pl. 63, fig. 8. Pattalophyllia costata d’Achiardi, 1881: Schuster, p. 77, pl. 17, figs 2a–c. Cricocyathus tarraconensis Alloiteau 1958: Löser et al., p. 173. Cricocyathus annulatus (Reuss, 1868): Schuster, p. 27, pl. 9, figs 7, 8.

Text-Fig. 29 Pattalophyllia grumi (Catullo, 1852), as figured in Alloiteau & Tissier (1958, as holotype of Vivesastraea villattae Alloiteau & Tissier, 1958), Paleocene of France. A, cross view of corallum; B, longitudinal view; scale bar: 7 mm.

Dimensions. d (min)=9–21 mm; d (max)=11.6–45 mm; s=up to 192+s7; s/mm=5–7/2.; in a corallite of around 10 mm in diameter around 80 septa occur; there are about 120 septa in a corallite of around 20 mm. Description. Solitary, turbinate, ceratoid, or patellate. Costosepta thick and compact in older cycles, distinctly thinner, subcompact to porous in younger cycles. Thirty to about 50 septa reach the axial region. Septal flanks strongly beaded. Anastomosis frequently present. Pali very elongate, irregularly in front of all septal cycles. Columella well-developed, spongy or net-like. Synapticulae present, more frequent in the vicinity of the wall. Endothecal dissepiments very abundant. Wall synapticulothecal to synapticulo-parathecal, septoparathecal in places. Epithecal wall can be present. Microstructure is of the agariciid type. Type locality of species. Oligocene of Italy (Monte Grumi). Distribution. Campanian of northern Spain (Torallola), Paleocene of Austria, Paleocene-Eocene of France, Lower Eocene of Egypt, Eocene of Spain, Middle Eocene of Mexico (San Juan Formation), Upper Eocene of Germany (Bavaria), Eocene-Oligocene of Italy, Upper Oligocene of Iran (Abadeh).

Family Poritidae Gray, 1842 Diagnosis. Colonial; colony formation by extratentacular budding. Corallites (except Napopora and Synaraea) closely united without coenenchyme, bounded by one or more synapticular rings. Septa (except Alveopora) composed of 3 to 8 nearly vertical trabeculae, loosely united, with more or less regular pores. Innermost trabeculae of certain septa differentiated as pali. Septa ento- and ectocoelic. One columellar trabecula usually present.


BARON-SZABO

Genus Goniopora Blainville, 1830 Type species. Goniopora pedunculata Quoy and Gaimard, in Blainville, 1830, Recent, New Guinea. Diagnosis. Colonial. massive, columniform or ramose, rarely incrusting. Gemmation extracalicinal. Corallites united closely or separated by a reticulate coenosteum. Septa subcompact to porous, arranged bilaterally. Pali present. Columella spongy or made of twisted segments. Synapticulae present. Endothecal dissepiments thin, few in number. Wall parathecal or synapticulothecal, incomplete.

Goniopora elegans (Leymerie, 1846) Pl. 12, fig. 7 *1846 v1868 1883 1884 1880 v1900 1912 1915 1925 1941b 1941 1949 1975 1987 nonv1988 cf.1996 1996 nonv1998 v2000 v2001 v2002

Porites elegans: Leymerie, p. 358, pl. 13. Dictyaraea elegans (Leymerie): Reuss, p. 163, pl. 15, figs 6, 7. Litharaea sp.: Pratz, p. 223. Goniaraea elegans (Leymerie): Felix, p. 424, pl. 3, figs 1, 2. Porites superposita, Duncan: Duncan, p. 57, pl. 14, figs 5, 6. Litharaea parkinsoni, sp. nov.: Gregory, 37, pl. 2, figs 13, 14. Goniaraea elegans (Leymerie): Oppenheim, p. 98, pl. 12 (3), figs 2, 3, text-fig. 2. Goniaraea elegans (Leymerie): Dainelli, p. 216. Goniaraea elegans Leymerie sp. 1846: Felix, pars 28, p. 285 (older synonyms cited therein). Goniopora nomlandi, n. sp.: Wells, p. 11 (311), pl. 1, fig. 6. Goniaraea elegans (Leymerie): Solè Sabaris, 398, pl. 8, fig. 52. Goniopora Ameliana Defr. sp. 1826: Alloiteau, p. 11, pl. 8, fig. 2. Goniaraea elegans (Leymerie): Kuzmicheva, p. 26, pl. 3, fig. 6. Goniopora elegans (Leymerie): Kuzmicheva, p. 158, pl. 28, fig. 7. Goniopora elegans (Leymerie, 1846): Drobne et al., p. 188, pl. 34, figs 1–3. Favites tenuiseptata (Duncan): Metwally, p. 387, figs 4g–h. Goniopora elegans (Leymerie): Schuster, p. 73, pl. 15, fig. 4. Goniopora elegans (Leymerie, 1846): Turnšek, in Turnšek & Drobne, p. 138, pl. 8, fig. 5. Goniopora elegans (Leymerie, 1846): Baron-Szabo, p. 118, pl. 9, fig. 2. Goniopora elegans (Leymerie, 1846): Baron-Szabo, p. 262, fig. 2 C. Goniopora elegans (Leymerie, 1846): Baron-Szabo, p. 128, pl. 89, fig. 5.

Dimensions. d=2.5–4.5 mm; c-c=3.5–6 mm; s=20–26. Description. Massive, cerioid; calices rounded or irregularly polygonal in outline, separated by a reticulated coenosteum; septa arranged in 2–3 cycles, about 10 of which reach centre of the calice; another set of up to 10 septa reaches about half the length of the oldest ones, sometimes fusing with septa of the preceding cycle; columella irregular spongy-papillose or formed by twisted segments. Remarks. The specimens described and figured as Favites tenuiseptata from the Upper Maastrichtian of the Oman Mountains in Metwally (1996) clearly show all characteristics of the genus Goniopora as described above, but seem to have a greater number of septa (around 24–28) and a slightly larger corallite diameter (3–5.5 mm). Type locality of species. Eocene of France (Corbières). Distribution. Upper Turonian-Coniacian of Austria (Theresienstein reef), Middle-Upper Maastrichtian of the UAE/Oman border region, Lower Paleocene of Ukraine, Upper Danian of Pakistan, Eocene of Egypt, France, Bosnia, Slovak. Carpathians, Somalia, Peru, and Spain, Eocene-Lower Oligocene of Italy.



143

Goniopora websteri (Bowerbank, 1840) Pl. 12; fig. 6 v*1840 v1850a 1880 1880 1912 1914 1925 1975 1981 1987 v1988 v1998 v1998 2000 2000 2002

Astrea websteri: Bowerbank, p. 23, figs A, B (topotypes studied). Litharaea websteri (Bowerbank): Milne Edwards & Haime, p. 38, pl. 6, figs 1a–c (topotypes studied). Litharaea epithecata, Duncan: Duncan, p. 23, pl. 24, figs 1–9. Litharaea epithecata, nobis, var. hemisphaerica: Duncan, p. 24, pl. 2, fig. 10. Litharaea subepithecata n. sp.: Oppenheim, p. 103, pl. 11, figs 7, 8, pl. 12, figs 12, 12a. Goniopora epithecata Duncan sp. 1880: Felix, pars 7, p. 241. Litharaea websteri Bowerbank sp. 1840: Felix, pars 28, p. 285 (older synonyms cited therein). Goniopora websteri (Bowerbank, 1840): Kuzmicheva, p. 26, pl. 3, fig. 5. Goniopora epithecata Duncan: Abed & El-Asa‘ad, p. 287, pl. 3, figs 4a–d. Goniopora websteri (Bowerbank, 1840): Kuzmicheva, p. 157, pl. 28, fig. 6. Litharaeaopsis subepithecata (Oppenheim, 1912): Drobne et al., p. 189, pl. 35, figs 1–4. Litharaea subepithecata Oppenheim, 1912: Turnšek, in Turnšek & Drobne, p. 138, pl. 10, figs 1, 2. Litharaea websteri (Bowerbank, 1840): Turnšek, in Turnšek & Drobne, p. 139, pl. 10, figs 3, 4. Goniopora epithecata (Duncan 1880): Löser, p. 38. Goniopora epithecata var. hemisphaerica (Duncan 1880): Löser, p. 38. Goniopora epithecata (Duncan 1880): Baron-Szabo, p. 128.

Dimensions. d=5–6.5 mm; juveniles around 4 mm; c-c=5–6 mm; s=24. Description. Knobby-sublamellar corallum; corallites irregularly polygonal in outline; septa arranged in 3 complete cycles in 6 systems; S1 and S2 nearly equal, reaching the axial region; columella irregular spongypapillose or formed by twisted segments. Type locality of species. Middle Eocene of England (Bracklesham). Distribution. Campanian of central Saudi Arabia, Paleocene of India, Slovenia, and Ukraine, Eocene of Bosnia and England.]

Goniopora imperatoris Vaughan, 1919 Pl. 12; figs 3, 4 v1873 v*1919 1929 1929 1929 1929 1974 1979 v1986 v1988 v1989 1992 v1994 1997 v1998

Actinacis rollei, Reuss: Duncan, p. 561. Goniopora imperatoris: Vaughan, p. 493, pl. 142, figs 3, 3a. Goniopora casacadensis Vaughan, 1919: Coryell & Ohlsen, p. 222, pl. 40, fig. 5. Goniopora imperatoris Vaughan, 1919: Coryell & Ohlsen, p. 224, pl. 41, fig. 2. Goniopora portoricensis Vaughan, 1919: Coryell & Ohlsen, p. 225, pl. 41, fig. 5. Goniopora regularis (Duncan), 1863: Coryell & Ohlsen, p. 226, pl. 42, figs 1–1a. Goniopora regularis (Duncan): Frost & Langenheim, p. 234, pl. 80, figs 4–7. Goniopora regularis: Frost & Weiss, p. 1113ff, fig. 8. Goniopora imperatoris Vaughan: Foster, p. 85, pl. 35, figs 3–7, pl. 36, figs 1–7, pl. 37, figs 1–4, text-figs 2–3, 6, 8, 11, 13, 14 (older synonyms cited therein). Goniopora elegans (Leymerie, 1846): Drobne et al., p. 188, pl. 34, figs 1–3. Goniopora imperatoris: Budd et al., p. 267ff. Goniopora sp., cf. G. vaughani Nomland, 1916: Squires & Demetrion, p. 19, figs 33–35. Goniopora imperatoris: Budd et al., pp. 952, 957, 965, 974. Goniopora antiqua: Reig Oriol, p. 27, pl. 5, fig. 7. Goniopora elegans (Leymerie, 1846): Turnšek, in Turnšek & Drobne, p. 138, pl. 8, fig. 5.


BARON-SZABO

2002 v2000 v2002

Goniopora antiqua, Reig Oriol 1997: Löser et al, p. 322. Goniopora imperatoris Vaughan, 1919: Baron-Szabo, p. 118, pl. 9, fig. 3. Goniopora imperatoris Vaughan, 1919: Baron-Szabo, p. 127, pl. 89, figs 2–3.

Dimensions. d=1.8–2.8 mm, juveniles are as small as 1.4 mm; c-c=1.8–4 mm; s=20–29. Description. Ramose or massive, with calices that are rounded or polygonal in outline; septa subcompact, thin, equal in thickness, and have delicate granulations laterally; septal arrangement is bilaterally; one set, consisting of about 6 septa, extends to center of corallite, where the septa might dissociate to form pali or trabecular prolongations, which join with the columella; about 6 septa, forming the second set, reach three quarters the length of the oldest ones; youngest septa are nearly equal with the septa of the preceding set; pali or paliform lobes irregularly occur axial to septa of every set; columella spongy or made of thin, twisted segments; wall synapticulothecal, incomplete. Type locality of species. Middle Miocene of Panama (La Boca Formation). Distribution. Upper Santonian-Lower Campanian of northern Spain, Middle-Upper Maastrichtian of the UAE/Oman border region, Maastrichtian of Jamaica (new material), Danian of Italy and Slovenia (Dolenja vas), Middle Eocene of St. Bartholomew and Mexico (Bateque Formation), Upper Oligocene of Puerto Rico, Mexico (La Quinta Formation), and Antigua (Antigua Formation), Lower Miocene of Anguilla (Anguilla Formation) and ?Georgia (Chattahoochee Formation), Lower-Middle Miocene of Florida (Tampa Formation, Panama (Emperador Limestone), and Anguilla (Anguilla Formation), Middle Miocene of Panama (La Boca Formation) and Puerto Rico (Ponce Formation), Neogene of the Dominican Republic, Middle-Upper Pliocene of Jamaica (Bowden Formation) and Florida (Pinecrest Sandstone). New Material. Maastrichtian of Jamaica, NMNH, Coates coll., sample nos.: 304b (=Jerusalem Mountain Inlier); 354a; 482; 490e; 506; 507; (=Ducketts Land Settlement); 570c; 570h; 570L (=probably Cambridge area); J-71-14d4 (=Road Sunderland Black); J-71-34p; J-71-40 (=Rio Minho).

Goniopora microscopica (Duncan, 1863) Pl. 13, figs 1, 2 v*1863 1867 1880 v1900 v1919 1986

Alveopora microscopica: Duncan, p. 426, pl. 14, fig. 5. Alveopora microscopica Duncan: Duncan, p. 25. Isastraea punctata, Duncan: Duncan, p. 44, pl. 18, figs 10–13. Litharaea Colae, sp. nov.: Gregory, p. 37, pl. 2, figs 12a, b. Goniopora regularis var. microscopica (Duncan): Vaughan, p. 492. Goniopora microscopica (Duncan): Foster, p. 86.

Dimensions. d=1.2–1.7 mm, juvenile around 0.9 mm; c-c=2.2–3.2 mm; s=24–26. Description. Massive, columniform or ramose, subcerioid colony; calices rounded or irregularly polygonal in outline; septa thin, nearly equal, arranged bilaterally; about 10 septa reach the center of the calice; a set of 7 to 10 septa reaches about half the length of S1, sometimes fusing with septa of the preceding cycle; youngest septa distinctly shorter; columella irregular spongy-papillose or formed by twisted segments; wall parathecal or synapticulothecal, incomplete; endothecal dissepiments very thin, vesicular. Type locality of species. Upper Oligocene of Antigua (Antigua Formation). Distribution. Middle-Upper Maastrichtian of Jamaica (this paper), Paleocene of Pakistan (Jhirk, Sind), Upper Paleocene of Somalia, Upper Oligocene (Antigua Formation) of Antigua.



145

New Material. Middle-Upper Maastrichtian of Jamaica, NMNH, Coates coll., sample nos.: 350; 359a; 370b; 387a; 393a (=Jerusalem Mountain Inlier).

Goniopora reussiana (Duncan, 1865) Pl. 13, Fig. 3 v*1865 1868 1899 v1906 1929 1934 v1934 v1934 1941a v1945 1974 1974 1992 2002 2002 v2002

Porites Reussiana, spec. nov.: Duncan, in Duncan & Wall, p. 8, pl. 1, fig. 2 (topotypes studied). Porites Reussiana, Dunc.: Duncan, p. 25. Porites reussiana, Duncan, Vaughan, p. 246. Goniopora jamaica1: Bernard, p. 159. Goniopora decaturensis Vaughan, 1919: Coryell & Ohlsen, p. 223, pl. 40, fig. 7. Goniopora taberi, n. sp.: Wells, p. 155 (11), pl. 2, figs 5–7. Goniopora reussiana (Duncan): Wells, p. 90, pl. 4, fig. 18, pl. 5, figs 4–5. Goniopora trechmanni, new species: Wells, p. 91, pl. 3, figs 16–17. Goniopora reussiana (Duncan), 1865: Wells, p. 293 (11). Goniopora hedbergi Wells, n. sp.: Wells, p. 7, pl. 2, figs 5–6 Goniopora cascadensis Vaughan: Frost & Langenheim, p. 232, pl. 79, figs 3–7. Goniopora taberi Wells: Frost & Langenheim, p. 236, pl. 81, figs 1–4. Goniopora taberi Wells, 1934: Budd et al., p. 572 and 593, table 1. Goniopora reussiana (Duncan 1865): Löser, et al. p. 323. Goniopora reussiana (Duncan, 1865): Baron-Szabo, p. 128. Goniopora trechmanni Wells, 1934: Baron-Szabo, p. 128, pl. 89, fig. 4.

Dimensions. d=2–3 mm, juveniles as small as 1.8 mm; c-c=2.5–4 mm; s=18–24, s (juveniles)=12. Description. Ramose-multicolumnar to submassive, cerioid to subplocoid; septa generally 24 in number, arranged in 3 cycles in 6 systems, alternating in length and thickness; S1 extend to corallite center; pali before S1; columella generally small and weakly developed. Remarks. In the specimen figured as Goniopora reussiana (Duncan, 1865) (which is the holotype of Goniopora trechmanni Wells, 1934) on Plate, Fig. 3, the corallites are generally not closely united but rather are separated by a porous coenosteum. In this feature the specimen shows great resemblance to the genus Actinacis d’Orbigny, however, the septal arrangement is bilateral, pali are present, and in areas in which corallites are more densely spaced, cerioid polyp integration is found. Therefore, the specimen is grouped with the genus Goniopora. Type locality of species. Campanian of Jamaica. Distribution. Campanian-Maastrichtian of Cuba, Campanian-Maastrichtian and Eocene of Jamaica, Eocene of Cuba and Barbados (Upper Scotland Formation), Middle Eocene (San Juan Formation) and Upper Oligocene (La Quinta Formation) of Mexico, Upper Oligocene of Puerto Rico.

Family Micrabaciidae Vaughan, 1905 Diagnosis. Simple, free, cupoloid, nonepithecate. Costae alternating in position with septa, beaded or smooth, fixed to septa by regularly spaced synapticular bars. Septa ecto- and entocoelic, composed of one fan system of simple trabeculae, more or less perforate, margins strongly dentate with transverse beads or spiniform teeth. Dissepiments absent. Synapticulae simple, mostly in costal region. Columella composed of trabecular processes, loose, spongy or compact, often elongate.


BARON-SZABO

Genus Micrabacia Milne Edwards & Haime, 1849 Type species. Fungia coronula Goldfuss, 1827, Cenomanian of Germany (see Milne Edwards & Haime, 1849a).

Diagnosis. Solitary, cupolate, free, small, up to 15 mm in diameter. Septa perforate. Septal margins nearly vertical peripherally, finely granulated laterally. Wall synapticulothecal.

Micrabacia radiata (Goldfuss, 1827) Pl. 13, figs 5a–6b v*1827 v1840 1841 1850 1861 v1864 1867 pars1914 v1916 1916 ?1916 1916 v1916 v1916 1928 1928 1928 1928 v1933 1933 1933 v1933 1933 v1933 1933 ?1933 1933 non(v)1975 non1987 1996 2000 2000 2000 2000 2000 2000 2000 2000

Fungia radiata nobis: Goldfuss, p. 47 and 49, pl. 14, fig. 1, pl. 14, fig. 8. Fungia clathrata nob.: Hagenow, p. 648, pl. 9, figs 3a–c. Fungia radiata: Roemer, p. 25. Cyclolites radiata ?: d’Orbigny, vol. 2, p. 275. Micrabacia coronula Meek & Hayden, p. 430, nom. nud. (non Goldfuss). Micrabacia americana Meek: Meek, p. 1, pl. 28, figs 1a–d. Micrabacia senoniensis n. sp.: Bölsche, p. 36, pl. 3, fig. 1. Micrabacia astreata Goldfuss sp. 1826: Felix, pars 7, p. 184. Micrabacia rotatilis sp. nov.: Stephenson, p. 753, pl. 49, figs 1–4. Micrabacia americana Meek and Hayden: Stephenson, p. 118, pl. 20, figs 4–5. Micrabacia americana var. multicostata n. var.Stephenson, p. 119, pl. 20, fig. 6. Micrabacia rotatilis, var. georgiana Stephenson, p. 120, pl. 21, figs 5–8. Micrabacia mineolensis n. sp.: Stephenson, p. 122, pl. 23, figs 6–8. Micrabacia mississippiensis n. sp.: Stephenson, p. 123, pl. 23, figs 9–11. Micrabacia americana Stephenson: Whitney, p. 226. Micrabacia mississippiensis Stephenson: Whitney, p. 226. Micrabacia mineolensis Stephenson: Whitney, p. 226. Micrabacia mineolensis Stephenson: Adkins, p. 77. Micrabacia navarroensis n. sp.: Wells, p. 159 (241), pl. 15, figs 16–17. Micrabacia rotatilis, var. georgiana Stephenson: Wells, p. 161 (243). Micrabacia arkensasensis n. sp.: Wells, p. 163 (245), pl. 15, figs 21–22. Micrabacia stephensoni n. sp.: Wells, p. 164 (246), pl. 15, figs 18–20. Micrabacia mississippiensis Stephenson: Wells, p. 165 (247). Micrabacia taylorensis n. sp.: Wells, p. 166 (248), pl. 15, figs 1–7. Micrabacia americana Meek: Wells, p. 168 (250), pl. 16. Micrabacia americana multicostata Stephenson: Wells, p. 169 (251), pl. 13, fig. 9. Micrabacia mineolensis Stephenson: Wells, p. 170 (252), pl. 15, figs 8–13. Micrabacia rotatilis Stephenson, 1916: Kuzmicheva, p. 25, pl. 3, figs 3a–c. Micrabacia rotatilis Stephenson, 1916: Kuzmicheva, p. 156, pl. 28, figs 4–5. Micrabacia rotatilis Stephenson, 1916: Schuster, listed in appendix Micrabacia americana, Meek & Hayden 1876: Löser, p. 51. Micrabacia americana var. multicostata Stephenson 1916: Löser, p. 51. Micrabacia arkensasensis, Wells 1933: Löser, p. 51. Cyclabacia clathrata, Hagenow 1840: Löser, p. 24. Micrabacia mineolensis, Stephenson: Löser, p. 51. Micrabacia mississippiensis, Stephenson 1916: Löser, p. 51. Micrabacia navarroensis, Wells 1933: Löser, p. 51. Fungia radiata, Goldfuss 1826: Löser, p. 37.



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2000 2000 2000 2000 2002 2002 2002 2002 v2002 2002 2002 v2002 2002 2002 2002 v2002 v2002

Micrabacia rotatilis, Stephenson 1916: Löser, p. 51. Micrabacia rotatilis var. georgiana, Stephenson 1916: Löser, p. 51. Micrabacia senoniensis, Bölsche 1866: Löser, p. 51. Micrabacia taylorensis, Wells 1933: Löser, p. 52. Micrabacia americana Meek and Hayden, 1876: Baron-Szabo, p. 130. Micrabacia americana multicostata Stephenson, 1916: Baron-Szabo, p. 130. Micrabacia arkensasensis Wells, 1933: Baron-Szabo, p. 130. Micrabacia rotatilis Stephenson, 1916: Baron-Szabo, p. 130. Micrabacia hilgardi Stephenson, 1917: Baron-Szabo, p. 131, pl. 92, fig. 6. Micrabacia mississippiensis Stephenson, 1917: Baron-Szabo, p. 131. Micrabacia navarroensis Wells, 1933: Baron-Szabo, p. 131. Micrabacia radiata (Goldfuss, 1826): Baron-Szabo, p. 131, pl. 92, fig. 5. Micrabacia rotatilis Stephenson, 1916: Baron-Szabo, p. 130. Micrabacia rotatilis georgiana Stephenson, 1916: Baron-Szabo, p. 131. Micrabacia senoniensis Bölsche, 1866: Baron-Szabo, p. 131. Micrabacia stephensoni Wells, 1933: Baron-Szabo, p. 131, pl. pl. 92, fig. 2. Micrabacia taylorensis Wells, 1933: Baron-Szabo, p. 131, pl. 92, fig. 3.

Dimensions. d (adult)=6–9 mm; s (adult)=96; h=up to 4 mm. Description. Cupolate-discoid, free, slightly convex to flat; septa arranged in 5 complete cycles in 6 systems in adult specimens; septa nearly equal in thickness. PLATE 13 Fig. 1 Goniopora microscopica (Duncan, 1863), NMNH, Coates coll., no. 370b, cross thin section, Middle-Upper Maastrichtian of Jamaica, scale bar: 4 mm. Fig. 2 Goniopora microscopica (Duncan, 1863) (holotype of Litharaea colae Gregory, 1900), BMNH, R. 5043, upper surface, Upper Paleocene of Somalia, scale bar: 3.5 mm. Fig. 3 Goniopora reussiana (Duncan, 1865) (holotype of Goniopora trechmanni Wells, 1934), NMNH, I74482, upper surface of colony, Maastrichtian of Jamaica, scale bar: 4 mm. Fig. 4 Micrabacia marylandica Stephenson, 1916, holotype, NMNH, I203612, Maastrichtian of the USA (Maryland), scale bar: 2 mm. 4a, cross view; 4b, base of corallum. Fig. 5 Micrabacia radiata (Goldfuss, 1826), syntype, IPB, Goldfuss coll. 166, Campanian of Germany, scale bar: 2.5 mm. 5a, cross view; 5b, base of corallum. Fig. 6 Micrabacia radiata (Goldfuss, 1826) (holotype of Micrabacia mississippiensis Stephenson, 1916), NMNH, I32008, Maastrichtian of the USA (Mississippi), scale bar: 2.5 mm. 6a, cross view; 6b, base of corallum. Fig. 7 Stephanophyllia lanquinei Alloiteau, 1936, holotype, MNHN, Mo5214, upper surface, Campanian-Maastrichtian of Madagascar, scale bar: 3 mm. Fig. 8 Stephanophyllia lanquinei Alloiteau, 1936, UANLCE MAAS 217, Lower Maastrichtian of Mexico. 8a, upper surface, scale bar: 4 mm; 8b, base of corallum, scale bar: 4 mm. Fig. 9 Stephanophyllia cribraria (Stephenson, 1916), holotype, USNM, I31996, Campanian of the USA (North Carolina). 9a, base of corallum, scale bar: 2.5 mm; 9b, cross view, upper surface, scale bar: 2.5 mm. Fig. 10 Siderastrea adkinsi (Wells, 1934), NMNH, Berryhill coll., PR-7 series, Danian of Puerto Rico. 10a, longitudinal view, polished surface, scale bar: 4 mm; 10b, cross view, polished surface, scale bar: 4 mm. Fig. 11 Siderastrea adkinsi (Wells, 1934), holotype, NMNH, I74483, upper surface, Maastrichtian of Jamaica, scale bar: 3 mm.


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149

Remarks. Wells (1933, p. 154ff. [235ff.]) grouped the species he assigned to Micrabacia into two major categories which he called “Regularis” and “Irregularis” and he differentiated them on the basis on the following criteria: 1) ratio of corallum height to corallum diameter; 2) ratio of length of costae C5 to corallum diameter: and 3) ornamentation of costae. The “Regularis” group consists of only three taxa (M. marylandica Stephenson, M. hilgardi Stephenson, and M. hilgardi occidentalis Wells). The “Irregularis” group includes M. arkensasensis Wells, M. taylorensis Wells, M. mississippiensis Stephenson, M. mineolensis Stephenson, M. rotatilis Stephenson, M. rotatilis var. georgiana Stephenson, M. navarroensis Wells, M. cribraria Stephenson, M. stephensoni Wells, M. americana Meek and Hayden, and M. americana multicostata Stephenson. Wells further subdivided the “Irregularis” group into three subgroups: the “Mineolensis” group; the “Rotatilis” group, and the “Americana” or “Cribraria” group (in the “Synopsis of the species of Micrabacia”, Wells [1933, p. 237] called it the “Americana” group, but in the text dealing with the species descriptions, Wells [1933, p. 244] called it the “Cribraria” group). Wells calculated the mean value of the ratios of the height/ diameter and costae C5/diameter for each taxon, and he indicated that these mean values, as well as the costae ornamentation, could be used to separate taxa within these groups and subgroups. There are major problems with this approach besides the fact that the “Cribraria” subgroup is based on a taxon that belongs to the genus Stephanophyllia (see Remarks following description of Stephanophyllia cribraria below). The first problem is that Wells, in many cases, had only a small number of specimens from which he derived the mean ratios (he gives individual measurements for corallum height and diameter, but not for length of costae C5). Consequently, if only one or two specimens are not included in a data set, the mean ratio changes significantly. For example, specimens assigned to M. stephensoni have “height/diameter” ratios ranging from 0.37 to 0.54, and Wells reported a mean ratio of 0.44 (Note: for these same specimens the author of the current report calculated a different mean value of 0.46). Regardless, if the two specimens with the highest ratios had not been included in the calculation, the average ratio would have decreased to 0.43, and if the single specimen having the lowest ratio had not been included, the mean value would have increased to 0.48. Because the mean ratio values given by Wells are similar in many groups (for seven taxa the group mean height/diameter ratios range from 0.40 to 0.46), and highly subject to change depending on the number of specimens available, they can not be considered representative of the typical characteristics of the taxa. A similar problem occurs with taxa for which Wells had only one or a few specimens (e.g., M. mississippiensis, known only from the holotype and with a height/diameter ratio of 0.33, or M. hilgardi occidentalis, with only four specimens having a mean ratio of 0.51). In the first case, it can not be determined how typical the holotype is in terms of the shape of the corallum. In the second case, the mean ratio falls with the range of ratios for individual specimens of both the M. hilgardi and M. stephensoni. Furthermore, the lowest height/ diameter ratios of individual specimens of M. stephensoni are identical to those of specimens of M. americana, M. marylandica, and M. navarroensis. Although Wells does not give the individual measurements of the length of costae C5, the mean costae C5/diameter ratios are so similar for some taxa (0.14-0.19 for five taxa, and 0.21-0.25 for four others), that it is highly likely that, as in the case of the height/diameter ratios, individual ratio values for one taxon would be identical to the mean value for another taxon. Furthermore, Wells (1933, p. 236) himself pointed out that costae length “is hard to determine unless the specimens are mature”. This introduces another potential source of error in the calculation. Even when the both types of ratios are considered together, there are problems with separating some taxa, as in the case of M. rotatilis and M. rotatilis var. georgiana. Both taxa have the same mean costae C5/diameter ratio of 0.30, and their mean height/diameter” ratios differ by only 0.01 (0.42 in M. rotatilis var. georgiana and 0.43 in M. rotatilis). On an individual specimen basis, all the specimens of M. rotatilis var. georgiana


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described by Wells have “height/diameter”ratios that are identical to several specimens in the M. rotatilis group, and they, therefore, cannot be distinguished using Wells’ model, especially when, as in the mentioned example, the same or a comparable type of ornamentation of costae occurs in these specimens. Regarding the last point, Wells (1933, p. 236) himself, mentions that “The ornamentation of the costae is an important specific character, but account must be taken of destruction of granulations or beads due to wear”. In conclusion, it appears unlikely that Wells’ models could be used to reliably assign individual specimens to any of the above mentioned nominal taxa. Furthermore, based on the re-examination of type material, it was determined that the only taxonomic characters that can be used to differentiate species of Micrabacia are septal development (equal vs. alternating in thickness) and number of septa relative to corallite diameter. Using these criteria, forms with five complete cycles in six systems developed in corallites of 6-9 mm diameter, and with septa that are equal or nearly equal in thickness, are considered to represent the same taxon, and all the species listed in the synonymy above do have these characteristics. Type locality of species. Campanian of Germany (Aachen). Distribution. Campanian (Aachen)-Lower Maastrichtian (Rügen Island) of Germany, Maastrichtian of the USA (Arkansas, Colorado, Georgia, Mississippi, Montana, New Mexico, South Dakota, Texas), Paleocene-Eocene of Egypt.

Micrabacia marylandica Stephenson, 1916 Pl. 13, figs 4a, b v*1916 v1917 v1917 1926 1928 1933 1933 2000 2000 2000 2002 2002 2002

Micrabacia marylandica sp. nov.: Stephenson, p. 755, pl. 48, figs 1–4. Micrabacia hilgardi n. sp.: Stephenson, p. 120, pl. 22, figs 1–6. Micrabacia marylandica Stephenson: Stephenson, p. 121, pl. 22, figs 7–10. Micrabacia hilgardi Stephenson: Wade, p. 26, pl. 1, figs 3–4, 6–8. Micrabacia hilgardi Stephenson: Whitney, p. 226. Micrabacia hilgardi Stephenson: Wells, p. 156 (238). Micrabacia hilgardi occidentalis n. var.: Wells, p. 158 (240), pl. 15, figs 14–15. Micrabacia hilgardi, Stephenson 1916: Löser, p. 51. Micrabacia hilgardi occidentalis Wells 1933: Löser, p. 51. Micrabacia marylandica, Stephenson 1916: Löser, p. 51. Micrabacia hilgardi Stephenson, 1917: Baron-Szabo, p. 131. Micrabacia hilgardi occidentalis Wells, 1933: Baron-Szabo, p. 131. Micrabacia marylandica Stephenson, 1916: Baron-Szabo, p. 131.

Dimensions. d=2–7 mm; s=up to 96; h=up to 3 mm. Description. Cupolate, free, corallum flat, disc-like; septa arranged in 5 complete cycles in 6 systems; S1 and some of S2 slightly more dominant, join axially forming V-shapes similar as in the genus Deltocyathus; remaining septa regularly alternating in length but subequal in thickness with thickened axial ends. Remarks. Wells (1933, p. 154ff. [235ff.]) revised 14 species of Micrabacia, upon which he created a model that had several significant problems due to which the distinguishing of species is impossible (see discussion in Remarks of Micrabacia radiata [Goldfuss]). Therefore, based on the re-examination of type material, forms that have 1) 5 complete cycles in 6 systems developed in corallites ranging between 5–7 mm in diameter, and 2) septa that are unequal in thickness in that S1 and some of S2 are slightly more dominant and 3) have fusing axial ends that form V-shapes similar to the development in the genus Deltocyathus; and 4)



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remaining septa that regularly alternate in length but are subequal in thickness with thickened axial ends, are considered synonymous. Type locality of species. Maastrichtian of the USA (E. costata zone, Monmouth Formation, Maryland). Distribution. Maastrichtian of the USA (Alabama, Mississippi, Tennessee, Texas).

Genus Stephanophyllia Michelin, 1841 Type species. Fungia elegans Bronn, 1837, Miocene of Italy (see Milne Edwards & Haime, 1849a).

Diagnosis. Solitary, cupolate, free. Generally larger than Micrabacia. Septa perforate. Septal margins forming peripheral shelf, dentate laterally. Wall synapticulothecal.

Stephanophyllia cribraria (Stephenson, 1916) Pl. 13, figs 9a, b v*1916 v1923 v1926 v1928 v1933 (v)1938 1967 2002 2002

Micrabacia cribaria, n. sp.: Stephenson, p. 117, pl. 20, figs 1–3. Micrabacia cribraria Stephenson: Stephenson, p. 66, pl. 9, figs 15–17. Micrabacia cribraria Stephenson: Wade, p. 27, pl. 1, figs 9–10. Micrabacia cribaria Stephenson: Whitney, p. 226. Micrabacia cribraria Stephenson: Wells, p. 162 (244). Stephanophyllia regularis n. sp.: Traub, p. 38, pl. 1, fig. 6. Stephanophyllia regularis Traub, 1938: Kühn & Traub, p. 7. Micrabacia cribraria, Stephenson 1916: Löser et al, p. 415 (older synonyms cited therein). Micrabacia cribraria Stephenson, 1917: Baron-Szabo, p. 130.

Dimensions. d=5.5 mm; s=96. Description. Flat to cupolate, free; S1 equal or thinner than S2. Remarks. Re-examination of the type material revealed that it has perforate septa forming a marginal shelf. Therefore, the species M. cribraria belongs to the genus Stephanophyllia. Because no adequately preserved material from the Maastrichtian-Paleocene time period was available for study, the holotype of M. cribraria from the Campanian of the USA is shown in Pl. 13, figs 9a–b. Type locality of species. Campanian of the USA (Black Creek Formation, Neuse River, North Carolina). Distribution. Middle-Upper Turonian of northern Germany, Upper Santonian of Mexico, Santonian (Matawan, New Jersey; Bullock County, Georgia), Campanian (Black Creek Formation, Haddonfield, Lorilland, Crawfords Corner, North Carolina; Bullock County, Georgia), and Maastrichtian (Bullock and Lee Counties, Georgia; Union Springs, Alabama; McNairy County, Tennessee) of the USA, Paleocene of Austria.

Stephanophyllia indica Duncan, 1880 Text-Fig. 30 *1880 1970 1996

Stephanophyllia indica, Duncan: Duncan, p. 56, pl. 8, figs 7–10. Stephanophyllia intermedia sp. nov.: Hassan & Salama, p. 1, figs 1a–c. Stephanophyllia intermedia Hassan & Salama, 1970: Schuster, Appendix list.


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Dimensions. d=15–18 mm; s=48 (?+s5); h=4–7 mm. Description. Cupolate, free; septa arranged in 4 complete cycles in 6 systems; S1 and S2 slightly more dominant, join axially forming V-shapes similar to those in the genus Deltocyathus; remaining septa equal in thickness and regularly alternating in length. Type locality of species. Upper Paleocene of Pakistan (Jhirk, Sind). Distribution. Lower Danian of Egypt, Upper Paleocene of Pakistan.

Text-Fig. 30 Stephanophyllia indica Duncan, 1880, as figured in Duncan (1880), Upper Paleocene of Pakistan. A, cross view of corallum, scale bar: 4 mm; B, lateral view, close-up, scale bar: 4 mm; C, longitudinal view of corallum, scale bar: 9 mm.

Stephanophyllia lanquinei Alloiteau, 1936 Pl. 13, figs 7–8b v*1936 1958 (v)1975 1987 2000

Stephanophyllia Lanquinei (nov. sp.): Alloiteau, p. 15, pl. 6, figs 29–32. Stephanophyllia Lanquinei All.: Alloiteau, p. 205. Micrabacia rotatilis Stephenson, 1916: Kuzmicheva, p. 25, pl. 3, figs 3a–c. Micrabacia rotatilis Stephenson, 1916: Kuzmicheva, p. 156, pl. 28, figs 4–5. Stephanophyllia lanquinei, Alloiteau 1936: Löser, p. 74.

Dimensions. d=8–11 mm; s=96; h=up to 4 mm. Description. Cupolate, free; septa arranged in 5 complete cycles in 6 systems; S1 and S2 slightly more dominant axially, join axially forming V-shapes similar to those in the genus Deltocyathus; remaining septa equal in thickness and regularly alternating in length. Type locality of species. Campanian-Maastrichtian of Madagascar. Distribution. Campanian-Maastrichtian of Madagascar, Lower Maastrichtian of Mexico (Cerralvo, this paper), Danian of Ukraine. New Material. Lower Maastrichtian of Mexico, UANLCE MAAS 217. Family Siderastreidae Vaughan & Wells, 1943 Diagnosis. Colonial, rarely solitary; hermatypic. Colony formation by intra- and extratentacular budding. Synapticulothecate. Septa composed of one fan system of small, simple or compound trabeculae. Septa strongly granulated laterally, more or less porous, margins beaded or dentate, laterally united by simple synapticulae. Columella composed of one or more papillary trabeculae. Endothecal dissepiments present.



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Genus Siderastrea Blainville, 1830 Type species. Madrepora radians Pallas, 1766, Recent, West Indies.

Diagnosis. Colonial, massive, ramose or incrusting, cerioid. Gemmation extracalicinal. Septa compact to subcompact. Columella trabecular, papillose. Endothecal dissepiments vesicular. Wall synapticulothecal, formed by several synapticular rings, well-developed.

Siderastrea adkinsi (Wells, 1934) Pl. 13, figs 10a–11 v*1934 v1960 2000 2002 v2002 v2003 v2004 v2006

Synastrea adkinsi: Wells p. 87, pl. 3, figs 14, 15. Synastrea cf. adkinsi Wells: Berryhill, et al., p. 151. Synastrea adkinsi, Wells 1934: Löser, p. 77. Synastrea adkinsi Wells : Mitchell, p. 6 ff., table 1. Siderastrea cf. scottica Wells: Baron-Szabo, pl. 92, fig. 4. Siderastrea adkinsi (Wells, 1934): Schafhauser et al., p. 190, tab. 1. Siderastrea adkinsi (Wells, 1934): Baron-Szabo et al., p. R79. Siderastrea adkinsi (Wells, 1934): Baron-Szabo et al., p. 16, fig. 4.7.

Dimensions. d=4.5–6.5 (8) mm; juvenile corallites may be smaller (2–3.5 mm); c-c=5–10 mm, in areas of intense budding distance is around 3.5 mm; s=48, up to about 70; in juvenile corallites the number of septa may be 20–30; s/mm= 7–10/2; colony size: massive or encrusting= 5–25 cm in diameter; columnar or branching= up to 15 cm in length, with branches that are 5 cm in diameter. Description. Massive, branching, domal, or encrusting, cerioid; calices diversely polygonal in outline; septa thin, subcompact or porous, subconfluent or nonconfluent, sometimes directly uniting with the neighboring polyp, arranged in 4 complete cycles in 6 systems, laterally covered with rounded granules, pennulae and ?menianae; up to about 20 septa reach corallite center; columella spongy-papillose; wall synapticulothecal, incomplete; endothecal dissepiments thin, vesicular. Remarks. In the original description of the type material Wells (1934, p. 87) gave a distance between corallite centers of 5 to 6.5 mm. He also reported that the septa were exclusively confluent and that the corallite walls and dissepiments were absent. However, re-examination of the type material showed that it differs from the original description by: 1) having a corallite wall, formed by synapticulae, 2) endothecal dissepiments, 3) mainly non- and subconfluent septa, and 4) a corallite distance of 4.5–8 mm (3.5 juvenile). In regard to the first three itens, the assignment to the genus Siderastrea is suggested. Because the specimen of S. adkinsi figured on Plate 13, fig. 11 is an upper surface view, its appearance slightly differs from the images of 10a and 10b, the latter of which show polished surfaces of a colony. Type locality of species. Maastrichtian of Jamaica (Catadupa Midway). Distribution. Campanian-Maastrichtian (new material) of Jamaica, Maastrichtian of Mexico (Cardenas Formation), Danian of Argentina and Puerto Rico. New Material. Maastrichtian of Jamaica, NMNH, Coates, coll., sample nos.: 311; 312; 325; 454; 454a; 454b; 454c; 454d; 457a; 458a; 459-I; 465-I; 466; 466a; 466b; 468a; 468b; 468c; 468e; 468g; 469; 469a; 469b; 469c; 469d; 469e; 469f-III; 470f; 470g; 474-II-2; 492d-II; 521 (=Ducketts Land Settlement); 380a; J-71-13p3; J-71-13r3; J-71-13s3; (=Vaughnsfield); 548a; J-71-42-17n (=Rio Minho); 570g-I; JAG 18.5a; JAG 18.5b


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(=probably Cambridge railway area); J-71-17f-I; J-71-17i-I; J-71-18c (=Shaw Castle, Maldon Formation); J72-2h (=Marchmont Inlier); J3502d (=Point Flamstead).

Siderastrea conferta (Duncan, 1863) Pl. 14, fig. 1 v*1863 v1867 v1900 v1919 1925 1925 1974 1974 1989 1994

Isastrea conferta, spec. nov.: Duncan, p. 422, pl. 14, fig. 2 (topotypes studied). Isastrea conferta, Dunc.: Duncan, p. 25 (topotypes studied). Siderastrea hexagonalis sp. nov.: Vaughan, p. 155, pl. 18, figs 1–4. Siderastrea conferta (Duncan): Vaughan, p. 451, pl. 117, fig. 3, pl. 120, figs 1–4, pl. 121, figs 1–2a. Siderastrea conferta Duncan sp. 1863: Felix, pars 28, p. 131. Siderastrea hexagonalis Vaughan 1919: Felix, pars 28, p. 132. Siderastrea (Siderastrea) conferta (Duncan): Frost & Langenheim, p. 206, pl. 66, figs 1–6. Siderastrea (Siderastrea) hexagonalis Vaughan: Frost & Langenheim, p. 210, pl. 68, figs 1–3. Siderastrea conferta: Budd et al., pp. 267ff. Siderastrea conferta: Budd et al., pp. 957, 965, 974.

Dimensions. d (max)=3.5–7.5 mm, juveniles as small as 2.5 mm; c-c=3–6 mm; s=50–82. Description. Massive, cerioid; costosepta arranged in 4 complete and a beginning fifth cycle in 6 systems; up to 12 septa reach corallite center; columella formed by a small number of papillae or spongy. Remarks. The forms listed in synonymy above are all characterized by corallite diameter ranging between 3.5–7.5 mm, septa in more than 4 complete cycles in 6 systems and reaching nearly 5 complete cycles in latest adult stages, and a corallite wall generally formed by 1 to 2 synapticular rings. Type locality of species. Lower-Middle Miocene of Anguilla (Anguilla Formation). Distribution. Paleocene of the USA (Midwayan, Alabama), Lower Eocene (Lecheria limestone) and Upper Oligocene of Mexico (Rancho Berlin Sandstone and La Quinta Formation), and Puerto Rico, LowerMiddle Miocene of Anguilla (Anguilla Formation).

Siderastrea vancouverensis Vaughan, 1923 Pl. 14, fig. 2 v*1923 1927 v2003 v2006

Siderastrea vancouverensis, sp. n.: Vaughan, p. 175, pl. 40, figs 1–2. Siderastrea vancouverensis Vaughan 1923: Felix, pars 35, p. 373 Siderastrea vancouverensis Vaughan, 1923: Schafhauser et al., p. 190, tab. 1. Siderastrea vancouverensis Vaughan, 1923: Baron-Szabo et al., p. 15, fig. 4.8.

Dimensions. d (max)=7–11 mm, juvenile corallites may be 3–5.5 mm; d (min)=6–8 mm; c-c=4–10 mm; s=48– up to around 70.



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PLATE 14 Fig. 1 Siderastrea conferta (Duncan, 1863) (holotype of Siderastrea hexagonalis Vaughan 1919), NMNH, M158317, upper surface, Paleocene of the USA (Alabama), scale bar: 5 mm. Fig. 2 Siderastrea vancouverensis Vaughan, 1923, UNAM, IGM 8728, cross thin section, Maastrichtian of Mexico, scale bar: 2 mm. Fig. 3 Siderofungia morloti (Reuss, 1864), NMNH, Coates coll., no. J-66-31-42, Middle-Upper Maastrichtian of Jamaica. 3a, cross thin section, scale bar: 3 mm; 3b, thin section, longitudinal view, scale bar: 2 mm. Fig. 4 Synastrea agaricites (Goldfuss, 1826), NMNH, Coates coll., no. J3461c, cross thin section, Middle-Upper Maastrichtian of Jamaica, scale bar: 4 mm. Fig. 5 Pironastrea discoides d'Archiardi, 1875, SAZU, Pa-11 (11001b), thin section, cross view, oblique, Thanetian of Italy, scale bar: 1 mm. Fig. 6 Pironastrea discoides d'Archiardi, 1875, syntype, corresponding to pl. 18, fig. 2 in d’Achiardi, 1875), Paleontological Institute, Pisa d’Achiardi coll., not cataloged, upper surface, Eocene of Italy, scale bar: 3 mm. Fig. 7 Hindeastraea garloica (Tchéchmédjiéva, 1975), BSP, Stinnesbeck coll., no. CH-1, cross thin section, Campanian-Maastrichtian of Argentina, scale bar: 2 mm. Fig. 8 Hindeastraea discoidea White, 1888, holotype, NMNH, I19166 (-001), upper surface, Maastrichtian of the USA, scale bar: 3 mm. Fig. 9 Hindeastraea discoidea White, 1888, paratype, NMNH, I19166 (-002), upper surface, Maastrichtian of the USA, scale bar: 3 mm. Fig. 10 Hindeastraea discoidea White, 1888, paratype, NMNH, I19166 (-003), upper surface, Maastrichtian of the USA, scale bar: 3 mm.

Description. Cerioid colony; corallites polygonal; septa compact to subcompact, nonconfluent to confluent, arranged in 4 complete cycles with the beginning of the fifth cycle in 6 systems, granulated and pennulated laterally; columella trabecular, papillose; endothecal dissepiments vesicular; wall synapticulothecal, incomplete. Type locality of species. Oligocene or Miocene of Vancouver Island. Distribution. Maastrichtian of Mexico (Cardenas Formation), Oligocene or Miocene of Vancouver Island.

Genus Siderofungia Reis, 1889 Type species. Columnastrea bella Reuss, 1869, Oligocene of Italy (Crosara).

Diagnosis. Colonial, massive, thamnasterioid. Gemmation extracalicinal. Corallites have a slight polygonal appearance. Septa confluent, subcompact to porous, granulated laterally. Synapticulae present. Columella trabecular. Corallite walls indistinct. Endothecal dissepiments thin.

Siderofungia morloti (Reuss, 1864) Pl. 14, fig. 3a, b v*1864 v1869 1880 v1889 1909 1949 v1973 1975 1987

Astraea Morloti m.: Reuss, p. 22, pl. 6, figs 1a, 1b. Columnastrea bella: Reuss, p. 243, pl. 21, figs 5a–b. Astraea Morloti, Reuss: Duncan, p. 44, pl. 7, figs 15, 16. Siderofungia bella (Reuss): Reis, p. 110, pl. 4, fig. 19. Siderofungia bella (Reuss): Felix, p. 125. Siderofungia andrusovi: Alloiteau, p. 9, pl. 1, figs 2a–b, pl. 6, fig. 2. Siderofungia bella (Reuss) 1869: Barta-Calmus, p. 512, pl. 55, figs 1, 2, 7 (older synonyms cited therein). Siderastraea zelinskae Kuzmicheva, sp. nov.: Kuzmicheva, p. 24, pl. 3, fig. 2. Siderofungia zelinskae (Kuzmicheva, 1975): Kuzmicheva, p. 88, pl. 6, figs 1–2, pl. 7, fig. 1.



157

Dimensions. d (adult)=2.8–5 mm; d (juvenile)=2–2.5 mm; s (adult)=20–36; s (juvenile)=12–18; c-c=2.2–6 mm. Description. Colonial, massive, thamnasterioid; gemmation extracalicinal; corallites slightly polygonal in outline; septa confluent, nearly equal in thickness, subcompact to porous, with spiniform granules laterally; up to 12 septa extend to the axial region, their inner ends dissociate forming paliform structures; synapticulae present, rare; columella trabecular, papillose to substyliform; endothecal dissepiments thin, abundant. Type locality of species. Oligocene of Slovenia. Distribution. Middle-Upper Maastrichtian of Jamaica (new material), Upper Paleocene of Pakistan, Eocene of Slovakia and Ukraine, Upper Eocene of Spain, Oligocene of Germany, Italy, and Slovenia, Upper Oligocene of Hungary. New Material. Middle-Upper Maastrichtian of Jamaica, NMNH, Coates coll., sample no.: J-66-31-42 (=Ducketts Land Settlement).

Genus Sideroseris Wells, 1945 (=Rhabdophylliopsis Alloiteau & Tissier, 1958, Type species. Rhabdophyllia tenuis Alloiteau & Tissier, 1958 [non Reuss, 1868] =Rhabdophylliopsis alloiteaui Barta-Calmus, 1973, Danian of France). Type species. Sideroseris durhami Wells, 1945, Eocene of Barbados.

Diagnosis. Solitary, subcylindrical to trochoid (corallite diameter to around 10 mm) and colonial, arranged in phaceloid clumps. Solitary stage probably with a corallite height to 10 mm. Costosepta thin, compact, strongly granulated laterally. Columella papillose, made of a small number of trabecular rods. Synapticulae mainly peripherally. Dissepiments absent. Wall synapticulothecal, perforate. Remarks. Wells (1945) created the genus Sideroseris using a single corallum that was conical in shape. Later, in Middle Eocene sediments from Mexico, Frost & Langenheim (1974) discovered phaceloid clumps produced by polyps that were identical with Sideroseris in every skeletal structure except for the type of polyp integration. The same observation was made by Baron-Szabo et al. (2004) studying material form Danian sediments of Argentina. In addition, numerous individual polyps were found together with the phaceloid clumps. Moreover, specimens that were in the early stages of producing additional polyps by extracalicinal budding occurred frequently (Baron-Szabo, unpublished data). Generally, early budding stages were found in corallites that were at least around 10 mm in height. While the presence or absence of colonial development in scleractinians represents a very important generic feature, there is a small number of scleractinian genera that seem to be able to occur in both forms: These taxa exist as solitary forms up to a certain ontogenetical stage after which they develop into a colonial form. This feature has been observed for, e.g., Trachyphyllia Milne Edwards & Haime (Baron-Szabo 2006) and also applies to additional forms like Syzygophyllia Reuss (BaronSzabo unpublished data, Sanders & Baron-Szabo in prep.), whereas in phaceloid forms like Cladocora Ehrenberg, Calamophylliopsis Alloiteau, Dermosmilia Koby, Cladophyllia Milne Edwards & Haime, Carolastraea Eliášová, and many others, such “solitary” feature has never been observed, indicating that in genera of the latter type the first gemmation takes place at a very early ontogenetical stage without regard to the budding frequency at later stages. For example, in several species of Calamophylliopsis (compare Turnšek 1972), Cladophyllia (e.g., Baron-Szabo 2002, 2006), Carolastraea (e.g., Eliášová 1976) and others, long corallites are present that rarely branch (e.g. in the holotype of Cladophyllia dichotoma [Goldfuss 1826] branches are often up to 6 cm without budding), however, no forms have been reported that could be interpreted as solitary forms


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of any of these taxa. On the contrary, pieces of them were always identified as fragments of a colony. Sideroseris durhami Wells, 1945 Text-Fig. 31

Text-Fig. 31 Sideroseris durhami Wells, 1945. A, B, as figured in Alloiteau & Tissier (1958, as holotype of Rhabdophyllia tenuis Alloiteau & Tissier, 1958), Danian of France. A, upper surface of colony, scale bar: 8 mm; B, longitudinal view of colony, scale bar: 45 mm; C, D, holotype of Sideroseris durhami Wells, 1945, USNM 68331, Eocene of Barbados. C, cross view of corallite, scale bar: 4.5 mm; D, longitudinal view of corallum, scale bar: 6 mm. v*1945 (v)1958 v1973 1974 1991 v2004

Sideroseris durhami Wells, n.sp.: Wells, p. 6, pl. 2, figs 2–4. Rhabdophylliopsis tenuis Reuss sp.: Alloiteau & Tissier, p. 252, pl. 4, figs 1 a–b. Rhabdophylliopsis alloiteaui n. nov.: Barta-Calmus, p. 255. Sideroseris durhami Wells: Frost & Langenheim, p. 212, pl. 69, figs 1–9. Sideroseris new. sp.: Stemann, in Bryan et al., p. 33. Sideroseris durhami Wells, 1945: Baron-Szabo, et al., p. 79R.

Dimensions. d=4–9 mm, late adult stages 15 mm; s=55–up to around 70, in latest adult stages up to around 100 septa may occur. Description. Solitary, subcylindrical, or colonial, phaceloid with short branches; costosepta thin, closely set, arranged in 4 complete with the beginning of a fifth one in 6 systems, alternating in length; S1 reach corallite centre, fusing with papillose columella; epithecal wall irregularly present. Type locality of species. Eocene of Barbados. Distribution. Paleocene of the USA (Alabama, Salt Mountain Limestone), Danian of Argentina and France, Eocene of Barbados (Upper Scotland Formation), Middle Eocene of Mexico (San Juan Formation).

Genus Pironastrea d'Archiardi, 1875 (=Koilomorpha Alloiteau, 1952a, Type species. Meandrina aurasiaca Michelin, 1841, Turonian of France). Type species. Pironastrea discoides d'Archiardi, 1875, Eocene of Italy.



159

Diagnosis. Colonial, corallum explanate, subthamnasterioid to submeandroid; Gemmation intracalicinal-circumoral; corallites arranged in concentric rings, separated by tholiform collines; septa compact to subcompact; columella trabecular, papillose; endothecal dissepiments vesicular; wall synapticulothecal.

Pironastrea discoides d'Archiardi, 1875 Pl. 14, figs 5, 6 v*1875 1909 1925 v1992 v1998

Pironastrea discoides, m.: d'Archiardi, p. 197, pl. 18, figs 2a–3d. Pironastrea discoides d'Arch.: Oppenheim, p. 320. Pironastrea discoides d'Archiardi 1875: Felix, pars 28, p. 130 (older synonmys cited therein). Pironastrea sp.: Budd et al, p. 578, fig. 4.2. Pironastrea discoides d'Archiardi, 1875: Turnšek, in Turnšek & Drobne, p. 137, pl. 8, figs 1a–2.

Dimensions. c-c (same and adjacent series)=1.5–3 mm, in areas of intense budding corallite distance is smaller or indistinct; s=12–24; s/mm=25–35/5. Description. Mound-shaped or explanate; corallites monocentric or arranged in wavy series of distinct or indistinct corallite centers, separated by rather flat tholiform collines; septa subequal; columella spongy-trabecular or formed by 1 or 2 twisted segments. Type locality of species. Middle Eocene of Italy (Friul). Distribution. Thanetian-Eocene of Italy, Middle Eocene of Bosnia.

Genus Hindeastraea White, 1888 (=Plesiosiderastraea Tchéchmédjiéva, 1975. Type species. P. garloica Tchéchmédjiéva, 1975, Upper Campanian of Bulgaria). Type species. Hindeastraea discoidea White, 1888 Maastrichtian of Texas (Navarro Formation, USA).

Diagnosis. Colonial, massive, cerioid. Septa compact, with rare pores peripherally, numerous spiniform granules laterally. Anastomosis present. Columella spongy-papillose. Synapticulae present. Wall septothecal-synapticulothecal. Septothecal wall formed by inflexion and thickening of peripheral ends of septa. Endothecal dissepiments vesicular to subhorizontal. Remarks. Examniation of excellent photographic material provided by Jacob Leloux (Leiden) of the type material of Plesiosiderastraea garloica Tchéchmédjiéva and re-investigation of the syntypes of Hindeastraea discoidea White revealed that they completely correspond in all skeletal elements. Due to the small size of the syntypes of Hindeastraea discoidea White, 1888, the presence of sparse synapticulae and peripherally occurring septal pores have been overlooked for a long time. Moreover, in the type specimen of Plesiosiderastraea garloica Tchéchmédjiéva endothecal developmentare not as richly developed as stated by Tchéchmédjiéva (1975, p. 35), thus closely resembling the endotheca in Hindeastraea. Therefore, the two genera are considered to be synonymous.

Hindeastraea discoidea White, 1888 Pl. 14, figs 8–10


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?1880 v*1888 1898 1914 v1929 1951 2000 2000 v2002 v2002

Litharea grandis, Duncan: Duncan, p. 57, pl. 11, figs 11–13. Hindeastraea discoidea, sp. nov: White, p. 363, figs 1–5. Isastraea dicoidea (White): Frech, p. 322. Isastraea discoidea Ch. White sp. 1888: Felix, pars 7, p. 174. Hindeastraea collinensis, new species: Hoffmeister, p. 2, pl. 1, figs 1–2, pl. 2, figs 2–4. Hindeastraea discoidea White: Perkins, p. 5–10, pl. 1, figs 1–10, pl. 2, figs 1–9. Hindeastraea collinensis, Hoffmeister 1929: Löser, p. 41. Hindeastraea discoidea White 1888: Löser, p. 41. Hindeastraea discoidea White, 1888: Baron-Szabo, p. 61, pl. 44, figs 1–3. Hindeastraea collinensis Hoffmeister, 1929: Baron-Szabo, p. 61, pl. 43, fig. 6.

Dimensions. d (max)=5–8.5 mm, juvenile corallites are around 4 mm, corallites in late adult stages are around 10 mm; s=24–30. Description. Cerioid, submassive colony; corallites irregularly polygonal in outline; septa arranged in 3 complete cycles in 6 systems, alternating in length and thickness; septa of a beginning fourth cycle generally present; spongy-papillose columella well-developed; endothecal dissepiments vesicular, generally deep in corallum. Remarks. In having compact septa with rare pores, synapticulothecal and septothecal walls, the latter of which seems to be formed by inflexion and thickening of peripheral ends of septa, and in having a spongypapillose columella, the material described by Duncan (1880, pl. 11, figs 11–13) as Litharea grandis from the Danian of Pakistan seems to very closely correspond to the genus Hindeastraea White, 1888. Regarding the dimensions of skeletal elements, the description given by Duncan differs from the illustrations of the type material (e.g., Duncan gave a septal number of 24, the illustration shows corallites with up 30 septa as well as corallites with only around 18 septa. Furthermore, Duncan reported a maximum corallite diameter of generally 10 mm (to 12.5 mm), whereas the illustration shows a corallite diameter [d (max)] of generally 6.5–8.5 mm). Because that the type material was not available for study, the grouping with Hindeastraea discoidea is provisional. Type locality of species. Maastrichtian of the USA (Navarro Formation, Texas). Distribution. Campanian-Maastrichtian of the USA (Taylor marl and Navarro Formation, Texas), ?Danian of Pakistan.

Hindeastraea garloica (Tchéchmédjiéva, 1975) Pl. 14, fig. 7 (v)*1975 1986 (v)1988 1995

Plesiosiderastraea garloica gen. n., sp. n.: Tchéchmédjiéva, p. 34, pl. 1, figs 1–2. Plesiosiderastraea garloica Tchéchmédjiéva, 1975: Tchechmédjiéva, p. 67ff. Plesiosiderastraea tzankovi sp. n.: Tchéchmédjiéva, p. 239, pl. 1, figs 5, 6. Plesiosiderastraea tzankovi Tchéchmédjiéva, 1988: Tchechmédjiéva, p. 71, pl. 18, figs 6, 7.

2000

Plesiosiderastraea garloica, Cheshmedzhieva 1975: L⎞ser, p. 66.

2000 2002 2002

Plesiosiderastraea tzankovi, Cheshmedzhieva 1988: L⎞ser, p. 66. Plesiosiderastraea garloica Tchéchmédjiéva, 1975: Baron-Szabo, p. 133, text-fig. 49. Plesiosiderastraea tzankovi Tchéchmédjiéva,1988: Baron-Szabo, p. 133.

Dimensions. d (max)=3–6 mm, juvenile corallites are slightly smaller (around 2.7 mm); c-c=3–4 mm; s=24–28, in juvenile corallites 16–22. Description. Cerioid, massive colony; corallites irregularly polygonal in outline; septa arranged in 3 comSCLERACTINIAN CORALS OF THE K/T-BOUNDARY


161

plete cycles in 6 systems, alternating in length and thickness; in some corallites the beginning of a fourth cycle is present; spongy-papillose columella well-developed; endothecal dissepiments vesicular to subtabulate, deep in corallum or abundant throughout the corallite; wall generally thick. Type locality of species. Upper Campanian of Bulgaria. Distribution. Upper Campanian of Bulgaria, Campanian-Maastrichtian of Argentina (this paper). New Material. Campanian-Maastrichtian of Argentina, BSP, Stinnesbeck coll., sample no.: CH-1.

Suborder Microsolenina Morycowa & Roniewicz, 1995 Diagnosis. Radial elements costoseptal or biseptal in type, densely arranged, formed by compound trabeculae and structurally fenestrate. Trabeculae with longitudinal striation. Septal faces ornamented with pennulae which can fuse into menianes, or dissociate into lateral axes. Pennular edges ornamented. Synapticulae trabecular in origin, rod-like. Columella parietal or monotrabecular, if developed. Dissepiments vesicular, isometric or flat and wide, depending on family.

Family Synastreaidae Alloiteau, 1952 Diagnosis. Colonial. Costosepta exsert. Septal perforations near axial region and near upper margins. Septa strongly beaded marginally. Synapticulothecate. Endothecal dissepiments well-developed. Columella trabecular.

Genus Synastrea Milne Edwards & Haime, 1848 Type species. Astrea agaricites Goldfuss, 1826, Senonian of Austria (Nußbach) (see Milne Edwards & Haime, 1848b).

Diagnosis. Colonial, massive, thamnasterioid. Corallites isolated or arranged in very short meandroid series. Gemmation intracalicinal. Septa confluent, perforated, marginally moniliform, granulated laterally. Columella subpapillose, rudimentary. Synapticulae abundant. Endothecal dissepiments thin, sparse.

Synastrea agaricites (Goldfuss, 1829) Pl. 14; fig. 4 v*1829 1830 1834 1849b 1850 1851a v1854 1857 nonv1881 v1903a 1930a

Astrea agaricites nobis: Goldfuss, p. 66, pl. 23, fig. 9. Siderastrea agaricites (Goldfuss): Blainville, p. 336. Siderastrea agaricites (Goldfuss): Blainville, p. 370. Synastrea agaricites: Milne Edwards & Haime, p. 148. Synastrea agaricites: d'Orbigny, p. 206. Thamnastrea agaricites: Milne Edwards & Haime, p. 109. Thamnastrea agaricites M. Edw. et H.: Reuss, p. 118, pl. 19, figs 1, 2. Thamnastrea agaricites: Milne Edwards, p. 556. Astrea agaricites Goldfuss: Quenstedt, p. 889, pl. 178, fig. 4. Thamnastrea agaricites Goldfuss sp. 1826: Felix, p. 202. Synastrea agaricites Goldfuss sp.: Oppenheim, p. 156, pl. 22, figs 5, 5b, pl. 28, fig. 4, pl. 48, fig. 15.


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parsv1930a 1937a 1944 1945 1952a 1956 non1980 v1982 v1982 1986 2000 v2001 v2002 2003 v2004

Synastrea procera Reuss sp.: Oppenheim, p. 159, pl. 28, figs 5, 5a non 6 [topotypes studied]. Synastrea agaricites Goldfuss sp. 1826: Bataller, p. 191, text-fig. on p. 191 Older synonyms cited therein). Synastrea agaricites (Goldfuss): Klinghardt, p. 191, pl. 2, figs 7, 8. Synastrea agaricites Goldfuss sp. 1826: Bataller, p. 41. Synastrea agaricites (Goldfuss, 1826): Alloiteau, p. 665, pl. 4, fig. 3. Thamnastrea agaricites (Goldfuss): Bendukidze, p. 95, pl. 7, fig. 3. Synastrea agaricites (Goldfuss): Vidal, p. 60, pl. 11, fig. 6. Synastrea agaricites (Goldfuss) 1826: Beauvais, vol. 2, p. 122, pl. 37, fig. 3. Synastrea excelsa nov. sp.: Beauvais, vol. 2, p. 128, pl. 39, fig. 2 [topotypes studied]. Synastrea agaricites (Goldfuss, 1826): Tchéchmédjiéva, p. 67ff. Synastrea agaricites (Goldfuss 1826): Löser, p. 77. Synastrea agaricites (Goldfuss, 1826): Baron-Szabo, p. 266, fig. 3B. Synastrea agaricites (Goldfuss, 1826): Baron-Szabo, p. 140, pl. 100, figs 1–2, 4. Synastrea sp. cf. S. agaricites Goldfuss, 1826): Filkorn, p. 1. Synastrea agaricites (Goldfuss): Moosleitner, p. 184, pl. 82, fig. 4.

Dimensions. c-c=4–8.5 (10) mm; s=24–34; s/mm=5–6/2; size of the colony=10–20 cm in diameter. Description. The corallum is massive and thamnasterioid, with corallites that are regularly disposed or arranged in short series. Gemmation is due to intracalicinal budding. Septa are subcompact, confluent, nearly equal in thickness, and have thick rounded granules, pennulae and meniane laterally. Ten to 20 septa reach the center of the calice. Their inner ends might fuse with columella or dissociate to form pali or paliform lobes. Fusion of septal ends occur frequently. Synapticulae are scattered throughout the colony. The columella is spongy-papillose, well-developed. Endotheca consists of numerous subtabulate or vesicular dissepiments. There is no wall between the corallites. Type locality of species. Santonian of Austria (Gosau Group at Abtenau). Distribution. Upper Turonian-Campanian of Austria, Lower Coniacian and Upper Santonian of southern France (Corbière, Provence), Santonian and Maastrichtian of northern Spain (Catalonia), Senonian of Georgia (in Caucasia), Upper Campanian of Bulgaria, Maastrichtian of Turkey and Mexico (Ocozocuautla Formations, Middle-Upper Maastrichtian of Jamaica (this paper). New Material. Middle-Upper Maastrichtian of Jamaica, NMNH, Coates coll., sample nos.: 376; 412 (=Jerusalem Mountain Inlier); J3461c (=Welcome Hall).

Synastrea garumnica (Vidal, 1921) Pl. 15, fig. 1 *1921 1937a 1987 (v)1996 2000 2002

Thamnastraea garumnica: Vidal, p. 6, pl. 8, figs 11–12. Synastraea garumnica Vidal sp. 1921: Bataller, p. 195, 3 text-figs on p. 197 (older synonmys cited therein). Synastraea sp. (S. danica n. sp.): Meyer, p. 25, pl. 5, fig. 1. Dimorphastrea irradians Reuss, 1868: Schuster, p. 76, pl. 16, fig. Synastrea garumnica, Vidal 1921: Löser, p. 77. Synastrea garumnica (Vidal, 1921): Baron-Szabo, p. 140.

Dimensions. c-c=3–6 mm, in early budding stages the calicinal distance is smaller; s=20–36; s/mm=6–9/2. Description. Colonial, massive, thamnasterioid; gemmation intracalicinal; corallites irregularly disposed or arranged in short series; septa subcompact, confluent, nearly equal in thickness, thick rounded granules, SCLERACTINIAN CORALS OF THE K/T-BOUNDARY


163

pennulae and meniane laterally; about ten septa reach the center of the calice; inner septal ends can fuse with columella or dissociate to form pali or paliform lobes; anastomosis frequently occurring; synapticulae irregularly distributed; columella is spongy-papillose, endotheca dissepiments subtabulate or vesicular, abundant; no wall between the corallites. Type locality of species. Maastrichtian of northern Spain. Distribution. Maastrichtian of northern Spain, Maastrichtian of Jamaica (new material), Danian of France (Vigny) and Egypt. New Material. Maastrichtian of Jamaica, NMNH, Coats coll., sample nos.: 533m (=Shaw Castle, Maldon Formation); 585c (=probably Cambridge area); J3461e (=Welcome Hall)

Synastrea geometrica (Goldfuss, 1826) Pl. 15, figs 2a, b v*1826 1850b 1850 1851a v1854 1857 1857 1858–61 v1881 ?1899 v1903a non1909 1914 1914 v1925 ?1930a ?1937b (v)1951a 1982 1999 2000 2000

Astrea geometrica Goldfuss, p. 67, pl. 22, fig. 11. Synastrea ? geometrica: Milne Edwards & Haime, vol. 4, p. 156. Synastrea geometrica: d’Orbigny, vol. 2, p. 278. Thamnastrea geometrica: Milne Edwards & Haime, p. 114. Latomaeandra astraeoides m.: Reuss, p. 106, pl. 21, figs 7–8 (topotypes studied). Thamnastrea astraeoides: Milne Edwards, vol. 2, p. 570. Thamnastrea geometrica: Milne Edwards, vol. 2, p. 571. Synastraea astraeoides: de Fromentel, p. 220. Astrea agaricites Goldfuss: Quenstedt, p. 889, pl. 178, fig. 4. Latimeandra astraeoides Reuss: Söhle, p. 38. Latimaeandraraea astraeoides Felix (Reuss sp.): Felix, p. 216 (topotypes studied). Latimaeandraraea astraeoides Reuss: Prever, p. 102, pl. 9, fig. 5. Latimaeandraraea astraeoides Reuss sp. 1854: Felix, pars 6, p. 199, pars 7, p. 202. Thamnastrea geometrica Goldfuss sp. 1826: Felix, pars 7, p. 197. Thamnastraea geometrica Goldfuss sp.: Umbgrove, p. 109. Maeandraraea astraeoides Reuss: Oppenheim, p. 200. Maeandraraea astraeoides Reuss sp. 1854: Bataller, p. 307. Synastrea subflabellata nov. sp. Alloiteau: Alloiteau, p. 47, pl. 8, figs 1–2. Brachyseris astraeoides (Reuss) 1854: Beauvais, vol. 2, p. 239, pl. 45, figs 1a–c. Synastrea geometrica: Leloux, p. 193. Synastrea geometrica (Goldfuss 1826): Löser, p. 77 Synastrea subflabellata, Alloiteau 1951: Löser, p. 77.

PLATE 15 Fig. 1 Synastrea garumnica (Vidal, 1921), NMNH, Coates coll., no. 585c, thin section, Maastrichtian of Jamaica,scale bar: 4 mm. Fig. 2 Synastrea geometrica (Goldfuss 1826), holotype, IPB, Goldfuss coll., no. 225b, Upper Maastrichtian of The Netherlands, scale bar: 6 mm. 2a, upper surface in ‘steinkern’ preservation; 2b, ‘positive sketch (presented for the first time). Fig. 3 Cunnolites cancellata (Goldfuss, 1826), holotype, IPB, Goldfuss coll., no. 170, upper surface, Upper Maastrichtian of The Netherlands, scale bar: 3 mm; Fig. 4 Cunnolites tenuiradiata (Fromentel, 1863) (holotype of Cyclolites thomasi Wells, 1935), BMNH, R.30227, upper surface, ?Upper Cretaceous-?Eocene (Yellow Limestone) of Jamaica, scale bar: 8 mm. Fig. 5 Polyphylloseris microkothos n. sp., holotype, NMNH, Coates coll., no. J-66-19, MiddleUpper Maastrichtian. 5a, cross thin section, scale bar: 4 mm; 5b, thin section, longitudinal view, scale bar: 2 mm; 5c, close-up of Fig. 5a, scale bar: 2 mm.


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Dimensions. d=6–12 mm; c-c=7–14 mm; s=32–48; s/mm=4–6/2. Description. Thamnasterioid colony; costosepta confluent to rarely subconfluent, irregularly perforated, nearly equal, arranged in 3–4 cycles in 6 systems; columella papillose; synapticulae scattered throughout the colony; endothecal dissepiments thin, vesicular. Remarks. Because the type material of Latomaeandra astraeoides Reuss was lost, Beauvais (1982) chose a neotype. In the description of the neotype he gave the size of corallite diameters as 5–8 mm and distance of calicinal centers as 4–9 mm. However, in addition to the close resemblance to the genus Synastrea (thamnasterioid colony with isolated corallies or corallies arranged in very short meandroid series, gemmation intracalicinal, confluent and perforated septa, marginally moniliform, granulated laterally, columella parietal, synapticulae abundant, endothecal dissepiments thin), in the neotype the corallite diameter and distance of corallite centers range from 6–12 mm, thus closely agreeing with the dimensions given in the original description by Reuss (1854, p. 106). Therefore, it is considered a junior synonym of Synastrea geometrica. The corallites in the specimen described and illustrated by Prever (1909) have a corallite diameter of 4–6 mm and therefore differ from the present species. Type locality of species. Upper Maastrichtian of The Netherlands. Distribution. Santonian-Maastrichtian of Austria (Gosau Group), ?Lower Maastrichtian of Spain, Upper Maastrichtian of Madagascar and The Netherlands.

Family Microsolenidae Koby, 1889 Diagnosis. Simple and colonial. Colony formation by intra- and extratentacular budding. Corallite walls absent or slightly delineated by synapticular rings. Epitheca complete but very thin. Septa thin, regularly fenestrate. Synapticulae abundant, regularly distributed. Pennular edges directed upwards, ornamented with rounded, equal dentations. Endothecal and exothecal dissepiments thin, if developed. Columella parietal or styliform. Remarks. The presence of regularly confluent septocostae constitute the principal distinction between this family and the more highly developed actinacidids.

Genus Polyphylloseris de Fromentel, 1857 Type species. Polyphyllastrea convexa d'Orbigny, 1850, Hauterivian of France.

Diagnosis. Colonial, thamnasterioid to subplocoid. Gemmation intracalicinal. Corallites elevated, appearing mammelonate. Septa confluent, regularly perforated, pennular laterally. Columella trabecular. Synapticulae abundant. Endothecal dissepiments vesicular, numerous.

Polyphylloseris microkothos n. sp. Pl. 15, figs 5a–c Derivatio nominis. from Greek: micros=small, kothos=cup, referring to the small diameter of the corallite. Holotype. J-66-19. Locus typicus. Jerusalem Mountain Inlier, Jamaica. Stratum typicum. Titanosarcolites-limestone, Middle-Upper Maastrichtian.


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Dimensions. d=2–4 mm; c-c=4–8.5 mm; s=16–30. Diagnosis. Colonial, thamnasterioid to subplocoid; gemmation intracalicinal; corallites elevated, appearing mammelonate; costosepta confluent, nearly equal in thickness, regularly perforated, arranged in irregular systems, pennular laterally; columella spongy-trabecular or formed by twisted segments; synapticulae abundant; endothecal dissepiments vesicular to subtabulate, numerous. Comparison. The new species differs from all known Cretaceous forms by the smaller corallite diameter and the smaller number of septa. In e.g. P. simondsi Wells, 1932, from the Albian of Texas, the corallite diameter ranges from 3–5 mm and the number of septa varies from 40 to 50; in P. conophora (Felix, 1891) from the Barremian of Mexico the number of septa can exceed 60; in P. convexa (d'Orbigny, 1850) from the Hauterivian of France the number of septa is generally 76 and the diameter of the corallites ranges from 7–8 mm. Distribution. Titanosarcolites-limestone at Jerusalem Mountain Inlier, Jamaica, Middle-Upper Maastrichtian.

Family Cunnolitidae Alloiteau, 1952 (=ex Cyclolitidae d'Orbigny, 1851) Diagnosis. Solitary and colonial, subdiscoid, patellate, or cupolate. Hermatypic. Colony formation by circumoral or intratentacular budding. Corallite wall synapticulothecal. Epitheca present or absent. Costosepta as in Synastreidae. Septal perforations at inner ends and near upper margins. Septa strongly beaded marginally, perforations generally filled secondarily, axis of trabecular divergence inclined outward. Endothecal dissepiments and columella weak or absent. Homeomorphic with Fungiidae. Remarks. The validity of the genus Cunnolites has been discussed for about 250 years. Originally, the name Cunnolites was created 'pre-Linnean' by Barrère (1746), and was 'post-Linnean' renamed by Guettard, 1774, as Porpites ellipticus. Lamarck (1801) re-assigned the same specimen again and placed it as the third species in his newly created genus Cyclolites (=Cyclolites ellipticus Lamarck, 1801). Later investigations by Milne Edwards & Haime (1850) revealed that the two species placed before Cyclolites ellipticus, which are Cyclolites numismalis and Cyclolites hemisphaerica, belonged to Palaeozoic forms and had to be excluded from the synonymy of Cyclolites. At the same time they chose Cyclolites ellipticus Lamarck to be the type species of the genus. For priority reasons Alloiteau (1952a, 1957) did not accept the revision by Milne Edwards & Haime (1850). He argued that if the author of a new genus presented more than one species but did not designate a type species then the first species listed had to be acknowledged as the type species. And because in the case of Cyclolites the first species represents the type species of the Palaeozoic taxon Palaeocyclus Milne Edwards & Haime, he rejected the validity of the genus Cyclolites and created the taxon Cunnolites. This revision caused confusion in the literature in that some authors started using the new name established by Alloiteau, whereas others kept to the traditional name by Lamarck. The problematic situation was even more complicated by the creation of new genera named Plesiocunnolites Alloiteau, 1957, Plesiocunnolitopsis M. Beauvais, 1964, and Paracunnolites M. Beauvais, 1964. Some authors have accepted these as valid genera, others have ranked them in the subgenus level of Cyclolites or Cunnolites, and still others have put them in synonymy of Cyclolites or Cunnolites. This case can only be resolved by the Commission of Zoological Nomenclature. However, in order to avoid further confusion, the name Cunnolites, based on the type species Cunnolites barrerei Alloiteau, 1957, is tentatively accepted with all the consequences for higher taxonomic levels (e.g. giving the family name of Cunnolitidae Alloiteau, 1952a, priority over Cyclolitidae d'Orbigny, 1851). SCLERACTINIAN CORALS OF THE K/T-BOUNDARY


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According to Vaughan & Wells (1943), Wells (1956), and others, the Family Cunnolitidae Alloiteau, 1952a (=ex Cyclolitidae d'Orbigny, 1851) was characterized by solitary forms that were “free in ephebic stage”. However, as has been documented by, e.g., Sanders & Baron-Szabo (2007), forms of Cunnolites can very well stay attached throughout their whole life. Therefore, the characteristic “free in ephebic stage” was removed from the diagnosis of the family.

Genus Cunnolites Alloiteau, 1957 Type species. Cunnolites barrerei Alloiteau, 1957, Campanian of France.

Diagnosis. Solitary, cunnolitid (cupolate), free, circular or elliptical in outline. Base flat to concave. Calicular pit circular or elongate. Septa perforate (younger septa) to subcompact. Columella absent or feebly developed, trabecular. Synapticulae abundant. Endothecal dissepiments thin, few in number. Epitheca present or absent. Remarks. During the last century, numerous revisions have been carried out on the genus Cunnolites, with species differentiated mainly based on the ratios of their skeletal dimensions such as corallite diameter/ height of corallum, corallite diameter/size of calicular pit, etc. Recent studies on the ontogeny of Cunnolites, carried out by Baron-Szabo (2003) strongly suggest that the specific characteristics are restricted to the density of septa in peripheral areas of the corallum vs. septal density around calicinal pit, as well as the ratio of minimum/maximum corallite diameter (Table 5). Géczy (1954) carried out investigations on juvenile and adult stages in specimens from the Campanian of Hungary which he assigned to “Cyclotites”. However, because in his examination, forms of “Cyclotites” and specimens which are clearly colonial and most likely belong to the genus Aspidastraea Kühn (e.g, specimens figured on plate II, figs 2 and 14, specimen figured plate III, fig. 27) were mixed, the data were not used in present study.

Cunnolites cancellata (Goldfuss, 1826) Pl. 15, fig. 3 v*1826 1828 1830 1834 1846 1851b 1863 non1870 1873 1873 ?1880 1880 1880 1897 1903b 1910

Fungia cancellata Goldfuss, p. 48, pl. 14, fig. 5. Cyclolites cancellata: Morren, p. 50. Cyclolites cancellata, Goldf.: Blainville, vol. 60, p. 301. Cyclolites cancellata, Goldf.: Blainville, p. 338. Fungia filamentosa, sp. nov.: Forbes, p. 163, pl. 19, fig. 11. Cycloseris filamentosa: Milne Edwards & Haime, p. 127. Cycloseris minima: de Fromentel, p. 363, pl. 64, figs 3–3c. Cyclolites filamentosa: de Fromentel, p. 341, pl. 69, fig. 3. Cyclolites filamentosa, (Forbes): Stoliczka, p. 49, pl. 10, figs 10–12. Cyclolites orbiculus, Stoliczka: Stoliczka, p. 49, pl. 11, figs 2–2c. Cyclolites altavillensis, Defrance: Duncan, p. 54, pl. 16, figs 11. Cyclolites anomala, Duncan: Duncan, p. 54, pl. 17, figs 13–14. Cyclolites striata, Duncan: Duncan, p. 55, pl. 17, figs 7–8. Cyclolites Medlicotti : Noetling, p. 10, pl. 1, figs 11–12. Cyclolites cancellata Goldfuss: Felix, p. 50. Cyclolites aff. elliptica, Lamarck: Pratz, p. 310, pl. 24, fig. 6.


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pars1910 1914 1914 1914 1914 v1925 1933 1942 1981 1992 1996 2000 2000 2000 2002 2002 2002 2002 v2004

Cyclolites cereviviana Pratz, nov. sp.: Pratz, p. 311, pl. 24, fig. 5 non figs 3–4. Cyclolites cancellata Goldfuss sp. 1826: Felix, pars 7, p. 186 (older synonyms cited therein). Cyclolites filamentosa Forbes sp. 1846: Felix, pars 7, p. 188. Cyclolites Medlicotti Nötling 1897: Felix, pars 7, p. 190. Cyclolites orbiculus Stoliczka sp. 1873: Felix, pars 7, p. 191. Cyclolites cancellata, Goldf. sp.: Umbgrove, p. 108. Cyclolites medlicotti Noetling: Kühn, p. 178. Cyclolites regularis, Leym.: Maccagno, p. 792, pl. 1, figs 3–3b. Cyclolites medlicotti Noetling: Abed & El-Asa’ad, p. 283, pl. 2, figs 2a–d. Cunnolites filamentosa (Forbes 1846): Turnšek, p. 167, fig. 2. Cunnolites (Paracunnolites) minima (de Fromentel): Metwally, p. 383, figs 3d, e. Cunnolites medlicotti (Noetling 1897): Löser, p. 23. Cyclolites cancellata (Goldfuss 1826): Löser, p. 26. Cyclolites orbiculus, Stoliczka 1873: Löser, p. 26. Cyclolites minima de Fromentel 1863: Löser, et al. p. 199 (older synonyms cited therein). Cunnolites cancellata (Goldfuss, 1826): Baron-Szabo, p. 143. Cunnolites orbiculus (Stoliczka, 1873): Baron-Szabo, p. 143. Cunnolites medlicotti (Noetling, 1897): Baron-Szabo, p. 143. Cunnolites cancellata (Goldfuss): Moosleitner, p. 175, pl. 73, fig. 2.

Dimensions. d (min)/d (max)=0.9–1; s/mm=17–25/5 in peripheral areas, 22–25/5 around calicinal pit. Description. Solitary, cunnolitid corallum circular or nearly circular in outline; septa thin, densely packed, nearly equal. Remarks. The specimens examined, as listed in synonymy, are characterized by a corallite that is always circular or nearly circular in outline and numerous, rather equal septa that occur in nearly the same density throught the corallum. Type locality of species. Upper Maastrichtian of The Netherlands (St. Pietersberg). Distribution. Senonian of Portugal, Santonian-Campanian of Spain, Upper Campanian of Serbia (Fruska Gora), Campanian-Lower Maastrichtian of central Saudi Arabia, Campanian-Maastrichtian of Croatia, Maastrichtian of India (Veraghoor, Arrialoor group), Iran, Somalia, and Pakistan (Mari Hills), Upper MaastrichtianDanian of The Netherlands, Paleocene of India.

Cunnolites tenuiradiata (de Fromentel, 1863) Pl. 15, fig. 4 *1863 1870 1880 1880 ?pars1910 1914 v1930a v1935 1937b 1939 1945 non1980

Cyclolites tenuiradiata: de Fromentel, pl. 54. Cyclolites tenuiradiata: de Fromentel, p. 344. Cyclolites superba, Duncan: Duncan, p. 54, pl. 17, figs 11–12. Cyclolites Haimei, Duncan: Duncan, p. 54, pl. 17, figs 9–10. Cyclolites cereviviana Pratz; nov. sp.: Pratz, p. 311, pl. 24, figs 2–4 non fig. 5. Cyclolites tenuiradiata de Fromentel 1870: Felix, pars 7, p. 192. Cyclolites reussi var. portentosa n. sp.: Oppenheim, p. 118, pl. 17, fig. 4, pl. 18, fig. 9 (topotypes studied). Cyclolites thomasi, sp. n.: Wells, p. 190, pl. 12, figs 1–3. Cyclolites tenuiradiata de Fromentel 1870: Bataller, p. 306. Cyclolites tenuiradiatus de Fromentel nov. var.: Ciry, p. 246, 258, 294, and 300. Cyclolites tenuiradiata de Fromentel 1870: Bataller, p. 40. “Cyclolites” tenuiradiata de Fromentel 1864: Vidal, p. 70, pl. 7, figs 4–6.



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Cunnolites portentosus (Oppenheim) 1930: Beauvais, vol. 2, p. 194 (topotypes studied). Cunnolites (Cunnolites) tenuiradiata (de Fromentel): Al Asa’ad, p. 635, pl. 1, figs 7a–b. Cunnolites undulatus fossaenobilis Beauvais, 1982: Tchechmédjiéva, p. 72, pl. 18, figs 6, 7. Cunnolites subcircularis sulcatus Beauvais, 1982: Tchechmédjiéva, p. 73, pl. 18, fig. 12, pl. 19, fig. 1. Cunnolites (Cunnolites) tenuiradiata (de Fromentel): Metwally, p. 382, figs 2f–h. Cunnolites reussi var. portentosa (Oppenheim 1930): Löser, p. 22. Cunnolites tenuiradiata (de Fromentel 1863): Löser, p. 22. Cyclolites thomasi Wells 1935: Löser, p. 26. Cunnolites sp.: Baron-Szabo, p. 123, pl. 5, figs 2, 4. Cunnolites portentosus (Oppenheim, 1930): Baron-Szabo, p. 143, pl. 102, fig. 6. Cunnolites tenuiradiata (Fromentel, 1863): Baron-Szabo, p. 143.

Dimensions. d (min)/d (max)=0.67–1, varying during ontogeny but mainly ranging between 0.7 and 0.85; s/ mm: 15–20 (22)/5 in peripheral areas, 12–14/5 around calicinal pit. Description. Cunnolitid, circular or elliptical in outline; septa straight, thick, subequal or alternately arranged in 5 to 7 cycles; density of septa higher in peripheral areas. Remarks. The specimens examined, as listed in synonymy above, are characterized by a corallite that varies extremely in outline throughout ontogeny (measurements were taken from type material and illustrations of type material) and has a septal density that is lower around the calicinal pit than it is in the peripheral areas. Type locality of species. Cretaceous of France (Coustouges, Pyrénées-Oriental). Distribution. Cretaceous of France, Santonian of Austria (Gosau Group), Upper Campanian of Bulgaria and ?Serbia (Fruska Gora), Upper Campanian-Maastrichtian of the UAE/Oman border region, Maastrichtian of France (Royen), Santonian-?Maastrichtian of Spain, ?Upper Cretaceous-?Eocene (Yellow Limestone) of Jamaica (Spring Mtn., Parish of St. James), Paleocene of India.

Cunnolites scutellum (Reuss, 1854) Pl. 16, fig. 4 v1854 v*1854 1857 1870 ?1873 ?1873 1880 1903b 1903b 1914 pars1914 pars1914 v1927 v1930a v1930a 1933 1939

Cyclolites elliptica Lamarck: Reuss, p. 123, pl. 22, fig. 7, pl. 23, figs 1–2 (topotypes studied). Cyclolites scutellum m.: Reuss, p. 125, pl. 22, figs 1–3 (topotypes studied). Cyclolites elliptica Lamarck: Pictet, p. 423, pl. 106, fig. 4. Cyclolites filamentosa: de Fromentel, p. 341, pl. 69, fig. 3. (Forbes sp.) Cyclolites conoidea, Stoliczka: Stoliczka, p. 48, pl. 10, figs 4–5. Cyclolites faecata, Stoliczka: Stoliczka, p. 48, pl. 10, figs 6–9. Cyclolites crenulata, Duncan: Duncan, p. 52, pl. 17, figs 3–4 Cyclolites hemisphaerica Michelin (non Lamarck): Felix, p. 47. Cyclolites Choffati n. sp.: Felix, p. 48, pl. 3, figs 2–3. Cyclolites Choffati Felix 1903: Felix, pars 7, p. 186. Cyclolites elliptica Guettard sp. 1770: Felix, pars 7, p. 187. Cyclolites hemisphaerica Michelin 1847: Felix, pars 7, p. 189. Cyclolites felixi, n. sp.: Böhm, p. 191, pl. 11, figs 2a–b. Cyclolites ellipticus LK var. subcirlularis: Oppenheim, p. 79, pl. 3, fig. 4–4a (topotypes studied). Cyclolites profundus n. sp.: Oppenheim, p. 133, pl. 6, fig. 2 (topotypes studied). Cyclolites scutellum Reuss: Kühn, p. 178. Cyclolites ellipticus Lamarck: Ciry, p. 246.


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1939 v1957 (v)1975 v1982 v1982 1986 ?1992 (v)1995 1995 (v)1997 (v)1997 1994 2000 2000 ?2000 2000 2001 2002 2002 2002 2002 ?2002 ?2002 v2002 2002 2002

Cyclolites ellipticus Lamarck var. subcirlularis Oppenheim: Ciry, p. 246. Plesiocunnolites subcircularis nov. gen., nov. sp.: Alloiteau, p. 345, pl. 9, fig. 1, pl. 10, figs 2a–b, pl. 12, figs 7–8, pl. 18, fig. 10. Micrabacia disca Wu, sp. nov.: Wu, p. 105, pl. 8, figs 9–13. Cunnolites subcircularis (Alloiteau) 1957: Beauvais, vol. 2, p. 179, pl. 40, fig. 6, pl. 41, fig. 5 (topotypes studied). Cunnolites sororius profundus (Oppenheim) nov. s. sp.: Beauvais, vol. 2, p. 189 (topotypes studied). Cunnolites sororius profundus (Oppenheim, 1930): Tchechmédjiéva, p. 61ff. Cunnolites conoidea (Stoliczka, 1873): Turnšek, p. 167, fig. 2. Cunnolites (Paracunnolites) scutellum (Reuss, 1854): Abdel-Gawad & Gameil, p. 24, pl. 7, figs 3, 4. Cunnolites undulatus planus Beauvais, 1982: Tchechmédjiéva, p. 71, pl. 18, figs 4, 5. Cunnolites (Paracunnolites) scutellum (Reuss, 1854): Turnšek, p. 55, pl. 55, figs A–O. Cunnolites (Plesiocunnolites) faecata (Stoliczka 1873): Turnšek, p. 59, pl. 59, figs A–H. Micrabacia disca Wu: Lioa & Xia, p. 192, pl. 41, figs 4–7. Cunnolites profundus, Oppenheim 1930: Löser, p. 23. Cyclolites choffati, Felix 1903: Löser, p. 25. Cyclolites conoidea, Stoliczka 1873: Löser, p. 25. Plesiocunnolites subcircularis, Alloiteau 1957: Löser, p. 661. Micrabacia disca Wu: Löser & Liao, p. 666. Cyclolites scutellum Reuss 1854: Löser et al., p. 209 (older synonyms cited therein). Micrabacia disca Wu 1975: Löser et al., p. 415. Micrabacia disca Wu 1975: Baron-Szabo, p. 130. Cunnolites choffati (Felix, 1903): Baron-Szabo, p. 143. Cunnolites conoidea (Stoliczka, 1873): Baron-Szabo, p. 143. Cunnolites faecata (Stoliczka, 1873): Baron-Szabo, p. 143. Cunnolites felixi (Böhm, 1927): Baron-Szabo, p. 143, pl. 103, fig. 3. Cunnolites profundus (Oppenheim, 1930): Baron-Szabo, p. 143. Cunnolites scutellum (Reuss, 1854): Baron-Szabo, p. 143.

Dimensions. d (min)/d (max)=0.8–1; s/mm: 14–20/5 in peripheral areas, 15–22/5 around calicinal pit. Description. Solitary, cunnolitid corallum circular or slightly elliptical in outline; septa densely packed, very irregularly developed; septa density slightly higher around calicinal pit Remarks. In having a corallite that is nearly circular or circular throughout ontogeny (measurements were taken from type material and illustrations of type material), the specimens examined, as listed in the synonymy, closely correspond to C. cancellata, but they are distinguished from C. cancellata by having a lower septal density throughout the corallum. In the size and development of the corallum the specimens of Cyclolites conoidea described by Stoliczka (1873, p. 48) correspond well with C. scutellum. However, because in the original description no information is given about the number or density of septa, its synonymy with the latter is doubtful. Type locality of species. Upper Turonian-Campanian of Austria (Gosau Group). Distribution. Upper Cretaceous of Romania, Upper Turonian-Campanian of Austria (Gosau Group), Cenomanian of Ukraine, Senonian of Turkey, Santonian-Campanian of Portugal (Coimbra), Senonian of Spain, Upper Santonian-Lower Campanian of France, Santonian-Campanian of Greece and Slovenia, Upper Campanian of Bulgaria, Campanian-Maastrichtian of Jamaica (this paper), Serbia, ?Croatia, and Tibet, Maastrichtian of Iran, ?Maastrichtian (Valudayur, Arrialoor group)-Paleocene of India.



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Text-Fig. 32 Cunnolites giganteus (d’Orbigny, 1850), as figured in Quenstedt (1881, as Fungia robusta Quenstedt, 1881), Senonian of Germany. A, cross view, part of corallum; B, longitudinal view, part of corallum; scale bar: 8 mm.

Cunnolites giganteus (d’Orbigny, 1850) Text-Fig. 32 *1850 1851b v1854 1860 1863 1867 1881 1914 v1930a 1933 1992 1992 1992 1995 1995 1995 v1997 2002 2002 2002 2002 2002 2002 2002

Cyclolites gigantea: d’Orbigny, vol. 2, p. 201. Cyclolites Ligeriensis: Milne Edwards & Haime, p. 126. Cyclolites macrostoma m.: Reuss, p. 122, pl. 22, figs 8–10, pl. Cyclolites gigantea: Milne Edwards, vol. 3, p. 46. Cyclolites Ligeriensis: de Fromentel, p. 360, pls. 58, figs 2–2b. Cyclolites gigantea d’Orbigny: de Fromentel, p. 332, pls. 51–52. Fungia robusta: Quenstedt, p. 869, pl. 177, figs 23 d, o, s, y. Cyclolites gigantea d’Orbigny 1850: Felix, pars7, p. 189. Cyclolites Quenstedti n. sp.: Oppenheim, p. 127, pl. 6, figs 3a, b (topotypes studied). Cyclolites robustus Quenstedt: Kühn, p. 177. Cunnolites gigantea (d’Orbigny, 1850): Turnšek, p. 167, fig. 2. Cunnolites ligeriensis (Milne Edwards & Haime 1851): Turnšek, p. 167, fig. 2. Cunnolites quenstedti (Oppenheim 1930): Turnšek, p. 167, fig. 2. Cunnolites giganteus (d’Orbigny, 1850): Tchechmédjiéva, p. 70, pl. 17, fig. 3. Cunnolites depressus (Reuss, 1854): Tchechmédjiéva, p. 70, pl. 17, figs 4, 5. Cunnolites undulatus muthmannsdorfensis Beauvais, 1982: Tchechmédjiéva, p. 72, pl. 18, figs 9–12. Cunnolites (Plesiocunnolitopsis) robusta (Quenstedt) 1880: Turnšek, p. 63, pl. 63, figs A–F. Cyclolites gigantea d’Orbigny 1850: Löser et al., p. 194 (older synonyms cited therein). Cyclolites ligeriensis Milne Edwards & Haime 1850: Löser et al., p. 197 (older synonyms cited therein). Cyclolites quenstedti Oppenheim 1930: Löser et al., p. 206 (older synonyms cited therein). Plesiocunnolitopsis robusta (Quenstedt 1880): Löser et al., p. 549 (older synonyms cited therein). Cunnolites gigantea (d’Orbigny, 1850): Baron-Szabo, p. 143. Cunnolites ligeriensis (Milne Edwards & Haime 1851): Baron-Szabo, p. 143. Cunnolites macrostoma (Reuss, 1854): Baron-Szabo, p. 143.


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Cunnolites quenstedti (Oppenheim, 1930): Baron-Szabo, p. 143. Cunnolites robusta (Quenstedt, 1881): Baron-Szabo, p. 143. Cunnolites macrostoma (Reuss): Moosleitner, p. 175, pl. 73, figs 7, 8, 10, 11, 14.

Dimensions. d (min)/d (max)=0.73–0.97; s/mm: 8–15/5 in peripheral areas, 8–15/5 around calicinal pit. Description. Solitary, free, nearly circular to oval in outline; septal density equal throughout the corallum. Remarks. In having a corallite that varies substantially in outline throughout ontogeny (measurements were taken from type material and illustrations of type material), the specimens examined, as listed in the synonymy, closely correspond to C. tenuiradiata, but they are distinguished from C. tenuiradiata by having septal densities that are both equal and significantly lower throughout the corallum. Type locality of species. Lower Santonian of France (Le Beausset, Var). Distribution. Cretaceous of France, Upper Cretaceous of Romania and Spain, Lower Turonian and Upper Campanian of Bulgaria, Coniacian-Santonian of Austria, Santonian-Campanian of Slovenia, Senonian of Germany, Campanian of Hungary, Campanian-Maastrichtian of Serbia, Maastrichtian of Iran (Neyriz),

Cunnolites polymorphus (Goldfuss, 1826) Text-Fig. 33

Text-Fig. 33 Cunnolites polymorphus (Goldfuss, 1826), as figured in Duncan (1880), as type material of Cyclolites vicaryi Duncan, 1880, Paleocene of India. A, lateral view of corallum, upper surface, scale bar: 7 mm; B, cross view, close-up, scale bar: 1 mm. v*1826 v1826 v1826 1834 1847 1850 v1854 1851b 1851 1858–61 1860 1860

Fungia polymorpha nobis: Goldfuss, vol. 1, p. 48, pl. 14, figs 6 a–c, g–m. Fungia undulata nobis: Goldfuss, vol. 1, p. 49, pl. 14, fig. 7. Fungia discoidea nobis: Goldfuss, vol. 1, p. 50, pl. 14, fig. 7. Cyclolites discoidea, Goldf.: Blainville, p. 335. Cyclolites hemisphaerica n.: Michelin, p. 282, pl. 64, figs 2a–b. Funginella discoidea: d‘Orbigny, vol. 2, p. 202. Cyclolites placenta m.: Reuss, p. 125, pl. 17, figs 4–6. Cyclolites discoidea: Milne Edwards & Haime, vol. 2, p. 202. Cunnolites semigloboso Michelin: Leymerie, p. 199, pl. 8, fig. 14. Cyclolites placenta: de Fromentel, p. 124. Cyclolites placentula: Milne Edwards, vol. 3, p. 39. Cyclolites discoidea: Milne Edwards, vol. 3, p. 40.



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1860 1863 1863 1867 1870 1880 1880 1880 1881 1881 1881 1881 1881 1881 1881 1881 1897 v1903a v1903a v1903a v1903a 1910 1913 1914 1914 1914 v1930a v1930a v1930a v1930a v1930a v1930a 1939a 1941 1942 1942 ?1945 ?1945 ?1945 ?1945 1957 1957 1957 1964 1964 1964 1964 1970 1978 1978

Cyclolites polymorpha: Milne Edwards, vol. 3, p. 44. Cyclolites polymorpha: de Fromentel, p. 336, pl. 59, pl. 60, fig. 1. Cyclolites discoidea: de Fromentel, p. 351, pl. 65, pl. 66, fig. 1, pl. 69, fig. 2. Cyclolites placenta Reuss: de Fromentel, p. 333. Cyclolites reussi: de Fromentel, p. 338, pl. 55, fig. 3. Cyclolites Ranikoti, Duncan: Duncan, p. 52, pl. 17, figs 5–6. Cyclolites vicaryi, Haime: Duncan, p. 53, pl. 17, figs 1–2. Cyclolites regularis: Leymerie, p. 774. Fungia elliptica Lamarck: Quenstedt, p. 861, pl. 177, fig. 1. Fungia undulata rotunda: Quenstedt, p. 862, pl. 177, figs 2–3. Fungia dispar: Quenstedt, p. 862, pl. 177, fig. 4 (non fig. 24). Fungia polymorpha Goldfuss: Quenstedt, p. 863, pl. 177, fig. 7. Fungia discoidea: Quenstedt, p. 865, pl. 177, fig. 14. Fungia patellata: Quenstedt, p. 871, pl. 177, fig. 27. Fungia sellata: Quenstedt, p. 871, pl. 177, fig. 25. Fungia sororiae: Quenstedt, p. 874, pl. 177, fig. 30. Cyclolites regularis Leymerie: Nötling, p. 11, pl. 1, figs 13–14. Cyclolites placenta Reuss: Felix, p. 190. Cyclolites discoidea Blainville: Felix, p. 190. Cyclolites hemisphaerica Michelin: Felix, p. 192, pl. 18, fig. 2. Cyclolites polymorpha (Goldfuss): Felix, p. 198. Cyclolites polymorpha, Goldfuss; sp: Pratz, p. 310. Cyclolites krumbecki: Di Stefani, p. 264, pl. 23, fig. 6. Cyclolites placenta Reuss: Felix, pars 7, p. 191. Cyclolites polymorpha (Goldfuss): Felix, pars 7, p. 191. Cyclolites regularis Leymerie: Felix, pars 7, p. 192. Cyclolites placenta Reuss: Oppenheim, p. 89 (topotypes studied). Cyclolites weissermeli n. sp.: Oppenheim, p. 99, pl. 5, fig. 11 (topotypes studied). Cyclolites michelini nom. mut.: Oppenheim, p. 100, pl. 40, fig. 2, pl. 46, fig. 13 (topotypes studied). Cyclolites pulchellus n. sp.: Oppenheim, p. 105, pl. 51, Fig. 8, pl. 39, figs 9–10a (topotypes studied). Cyclolites undulatus var. plana: Oppenheim, p. 106, pl. 12, figs 2–2a (topotypes studied). Cyclolites sororiae (Quenstedt): Oppenheim, p. 121, pl. 3, fig. 5 (topotypes studied). Cyclolites senessei n. sp.: Alloiteau, p. 11, pl. 1, figs 12, 12a. Cyclolites reussi de Fromentel: Alloiteau, p. 71, pl. 30, figs 3–4. Cyclolites krumbecki De Stefani: Marini, p. 78, pl. 4, figs 1–3, pl. 5, figs 1–12. Cyclolites krumbecki var. orfellensis: Marini, p. 80, pl. 5, figs 13–18. Cyclolites placenta Reuss 1854: Bataller, p. 39. Cyclolites polymorpha Goldfuss sp. 1826: Bataller, p. 39. Cyclolites regularis Leymerie 1881: Bataller, p. 40. Cyclolites Reussi Fromentel 1863: Bataller, p. 40. Plesiocunnolites goldfussi: Alloiteau, p. 349. Cunnolites plani-alpici: Alloiteau, p. 349, pl. 9, fig. 4, pl. 11, figs 4, 5, 9. Cunnolites meringonensis: Alloiteau, pl. 3, figs 9–10, 16–17. Plesiocunnolites dispar (Quenstedt): Beauvais, p. 535, pl. 15, figs 1a–c. Plesiocunnolites sellatus (Quenstedt): Beauvais, p. 537, pl. 15, figs 4a–d. Cunnolites sororius (Quenstedt): Beauvais, p. 538, pl. 16, figs 2a–d. Plesiocunnolites reussi (Fromentel): Beauvais, p. 543, pl. 16, figs 5a–c. Cunnolites plani-alpici Alloiteau: Tschechmedjeva, p. 37, pl. 1, figs 3–4. Cunnolites (Cunnolites) reussi (Fromentel): Turnšek, p. 117, pl. 20, figs 1–6. Cunnolites (Cunnolites) sellata (Quenstedt): Turnšek, p. 117, pl. 20, figs 7–8.


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Cunnolites (Plesiocunnolites) dispar (Quenstedt): Turnšek, p. 121, pl. 27, figs 1–4. Cyclolites regularis Leymerie: Abed & El-Asaad, p. 285, pl. 2, figs 3a–b. Cunnolites placenta (Reuss) 1854: Beauvais, vol. 2, p. 163, pl. 40, fig. 7, pl. 41, fig. 1. Cunnolites senessei (Alloiteau) 1939: Beauvais, vol. 2, p. 165 (topotypes studied). Cunnolites sellatus (Quenstedt) 1881: Beauvais, vol. 2, p. 169 (topotypes studied). Cunnolites sellatus nefgrabenensis nov. s. sp.: Beauvais, vol. 2, p. 171, pl. 40, fig. 8 (topotypes studied). Cunnolites reussi (de Fromentel) 1870: Beauvais, vol. 2, p. 184, pl. 41, fig. 6 (topotypes studied). Cunnolites plani-alpici Alloiteau 1957: Beauvais, vol. 2, p. 189, pl. 41, figs 7–8 (topotypes studied). Cunnolites undulatus plana (Oppenheim) 1930: Beauvais, vol. 2, p. 198. Cunnolites undulatus (Goldfuss) 1826: Beauvais, vol. 2, p. 198 (older synonyms cited therein) (topotypes studied). Paracunnolites (?) discoideus (Goldfuss) 1826: Beauvais, vol. 2, p. 207, pl. 42, fig. 6 (topotypes studied). Cunnolites reussi (de Fromentel, 1870): Tchechmédjiéva, p. 63ff. Cunnolites sororius (Quenstedt, 1880): Tchechmédjiéva, p. 64ff. Cunnolites polymorphus (Goldfuss, 1826): Turnšek, p. 167, fig. 2. Cunnolites sororius (Quenstedt 1880): Turnšek, p. 167, fig. 2. Cunnolites ellipticus (Guettard, 1770): Turnšek, p. 167, fig. 2. Cunnolites (Cunnolites) reussi (Fromentel): Turnšek, p. 53, pl. 53, figs A–H. Cunnolites (Cunnolites) sellata (Quenstedt): Turnšek, p. 54, pl. 54, figs A–C. Cunnolites (Plesiocunnolites) dispar (Quenstedt) 1880: Turnšek, p. 58, pl. 58, figs A–D. Cunnolites (Plesiocunnolites) undulata (Goldfuss) 1826: Turnšek, p. 62, pl. 62, figs A–D. Cunnolites sp.: Baron-Szabo, p. 152, pl. 4, Fig. 1, pl. 8, Fig. 4, Tab. 4. Cunnolites regularis (Leymerie 1878): Löser & Liao, p. 666–667. Cyclolites discoidea (Goldfuss 1826): Löser et al., p. 189 (older synonyms cited therein). Cyclolites polymorpha (Goldfuss 1826): Löser et al., p. 204 (older synonyms cited therein). Cyclolites regularis Leymerie 1878: Löser et al., p. 206 (older synonyms cited therein). Cyclolites reussi (de Fromentel 1863): Löser et al., p. 207 (older synonyms cited therein). Cyclolites undulata undulata (Goldfuss 1826): Löser et al., p. 214 (older synonyms cited therein). Cunnolites dispar (Quenstedt, 1881): Baron-Szabo, p. 143. Cunnolites ellipticus (Guettard, 1770): Baron-Szabo, p. 143. Cunnolites plani-alpici Alloiteau 1957: Baron-Szabo, p. 143. Cunnolites michelini (Oppenheim, 1930): Baron-Szabo, p. 143. Cunnolites meringonensis Alloiteau 1957: Baron-Szabo, p. 143. Cunnolites placenta (Reuss, 1854): Baron-Szabo, p. 143. Cunnolites polymorpha (Goldfuss, 1826): Baron-Szabo, p. 143, pl. 102, fig. 6, pl. 103, fig. 2. Cunnolites pulchellus (Oppenheim, 1930): Baron-Szabo, p. 143. Cunnolites regularis (Leymerie, 1878): Baron-Szabo, p. 143. Cunnolites senessei Alloiteau, 1939: Baron-Szabo, p. 143. Cunnolites sellatus (Quenstedt, 1881): Baron-Szabo, p. 143. Cunnolites sellatus nefgrabenensis Beauvais, 1982: Baron-Szabo, p.143. Cunnolites sororius (Quenstedt, 1881): Baron-Szabo, p. 143. Cunnolites undulatus (Goldfuss, 1826): Baron-Szabo, p. 143, pl. 102, fig. 2. Cunnolites undulatus plana Beauvais, 1982: Baron-Szabo, p.143. Cunnolites weissermeli (Oppenheim, 1930): Baron-Szabo, p. 143. Cunnolites polymorpha (Goldfuss): Moosleitner, p. 175, pl. 73, figs 3, 4, 6, 12, 13.

Dimensions. d (min)/d (max)=0.7–0.98; s/mm=17–20/5 in peripheral areas, up to 25/5 in peripheral areas in growing stage, 17–20/5 around calicinal pit. Description. Solitary, free, nearly circular to oval in outline; septal density nearly equal throughout the



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corallum. Remarks. In having a corallite that varies substantially in outline throughout ontogeny (measurements were taken from type material and illustrations of type material), the specimens examined, as listed in the synonymy, show close affinities to C. giganteus and C. tenuiradiata, but they are distinguished from C. giganteus by having septal densities that are significantly larger, and differ from C. tenuiradiata by having septal densities that are both equal and larger throughout the corallum. Type locality of species. Cretaceous of France (Dauphiné, Haute Alpes). Distribution. Cretaceous of France and Spain, Albian of England (Norfolk, Hunstanton), Lower Cenomanian of Tibet, Upper Cretaceous of Romania, Italy, and Turkey, Lower Senonian of Portugal, SantonianCampanian of Slovenia, Romania, and Austria (Gosau Group), Campanian of Germany and Hungary, Upper Campanian of Serbia (Fruska Gora) and Bulgaria, Campanian-Maastrichtian of Serbia and Tibet (Yadong), Campanian-Lower Maastrichtian of central Saudi Arabia, Maastrichtian of Austria, Pakistan, and Libya, Paleocene of India.

Cunnolites angiostoma (Kühn, 1933) Pl. 16, Fig. 1 v*1933 1942 2002 2002

Cyclolites angiostoma nov. spec.: Kühn, p. 179, pl. 17, figs 5–6. Cyclolites elliptica Lamarck (Guettard sp.): Maccagno, p. 793, pl. 1, figs 5–5b. Cyclolites angiostoma, Kühn 1933: Löser et al., p. 185. Cunnolites angiostoma (Kühn, 1933): Baron-Szabo, p. 143.

Dimensions. d (min)/d (max)=0.85–0.89; d (min)=25–28 mm; d (max)=28–33 mm; s/mm=20–25/5 in peripheral areas, 25/5 around calicinal pit. Description. Solitary, free, somewhat elliptical in outline; septa thin, equal, densely packed; septal density nearly equal throughout the corallum but slightly larger around the calicinal pit; calicinal pit elongate. Type locality of species. Maastrichtian of Iran (Neyriz, Esfahan). Distribution. Maastrichtian of Iran and Somalia.

Genus Aspidastraea Kühn, 1933 (=Paradimorphastraea M. Beauvais, 1982, Type species. Dimorphastrea waehneri Felix, 1903a, Santonian of Austria [Gosau Group]). Type species. Aspidastraea orientalis Kühn, 1933, Senonian of Iran. Plate 16 Fig. 1 Cunnolites angiostoma (Kühn, 1933), holotype, BMNH, R. 5259, upper surface, Maastrichtian of Iran, scale bar: 5 mm. Fig. 2 Aspidastraea orientalis Kühn, 1933, holotype, BMNH, R. 30276, upper surface, Senonian of Iran, scale bar: 3 mm. Fig. 3 Aspidastraea clathrata (Goldfuss, 1826), holotype, IPB, Goldfuss coll., no. 226, upper surface, ‘steinkern preservation’, Upper Maastrichtian of The Netherlands, scale bar: 4 mm. Fig. 4 Cunnolites scutellum (Reuss, 1854) (holotype of Cyclolites felixi Böhm, 1927), BMNH, R. 25918, upper surface, Senonian of Turkey, scale bar: 4 mm. Fig. 5 Aspidastraea semhae (Kossmat, 1907), BSPG, 1975-I-129, Upper Maastrichtian of Iran (photographs courtesy Bayerische Staatssammlung Munich). 5a, upper surface of colony, scale bar: 8 mm; 5b, close-up, scale bar: 4 mm.


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Diagnosis. Colony forming a cupolate corallum. Gemmation circumoral. Septa porous or subcompact, and covered with numerous granules and pennulae laterally. Synapticulae abundant. Columella feebly developed, trabecular, or absent. No wall between the corallites. Endothecal dissepiments sparse. Aspidastraea clathrata (Goldfuss, 1826) Pl. 16, fig. 3 v*1826 1850 1881 v1925 1999 2000 2002

Astrea clathrata nobis: Goldfuss, vol. 1, p. 67, pl. 23, fig. 1. Synastrea clathrata: d’Orbigny, vol. 2, p. 278. Astraea clathrata Goldfuss=Quenstedt, p. 860. Thamnstraea clathrata Goldfuss sp.: Umbgrove, p. 110. Synastrea clathrata: Leloux, p. 193, fig. 2. Synastrea clathrata (Goldfuss 1826): Löser, p. 77. Synastrea clathrata (Goldfuss, 1826): Baron-Szabo, p. 140.

Dimensions. c-c=6–30 mm; s (secondary corallites)=around 12; s (main corallite)= around 60; s/mm=18–20/ 10. Description. Cast of cupulate corallum; central corallite about 10 mm in diameter, around which secondary corallites are arranged in irregular concentric series; secondary corallites 2–3 mm in diameter; septa porous to subcompact, confluent; synapticulae very abundant. Due to the problematic preservation of the specimen, exact dimensions of skeletal elements cannot be determined. Remarks. The systematic position of Astrea clathrata Goldfuss has been discussed for more than a century. D’Orbigny (1850) assumed that it was a thamnasterioid colony with subcompact septa typical of Synastrea. Quenstedt (1881, p. 860) stated that this specimen was more like a solitary coral of the Fungia-type because originally the septa were porous. The preservation of the holotype of Astrea clathrata Goldfuss, as a steinkern cast, made it possible to determine that the original skeleton was indeed very porous. Moreover, it very much resembles a solitary form. However, as shown in Fig. 3, Pl. 16, the central corallite appears surrounded by several very small, secondary corallites, that are in a very early ontogenetical stage (see BaronSzabo 2003, pl. 25, figs 2–4), thus completely agreeing with the genus Aspidastraea. Therefore, the assignment to the latter group is indicated. Type locality of species. Upper Maastrichtian of The Netherlands. Distribution. Upper Maastrichtian of The Netherlands.

Aspidastraea orientalis Kühn, 1933 Pl. 16, fig. 2 ?1880 v*1933 1943 1952a 1956 1987 v2000 2000 v2002

Thamnastraea Balli, Duncan: Duncan, p. 55, pl. 19, figs 1–3. Aspidastraea orientalis nov. spec.: Kühn, p. 179; pl. 17, fig. 7. Aspidastraea orientalis Kühn: Vaughan &Wells, p. 135. Aspidastraea orientalis Kühn, 1933: Alloiteau, p. 668. Aspidastraea orientalis Kühn: Wells, p. F387, fig. 280,3. Aspidastraea orientalis Kühn 1933: Kuzmicheva, p. 87, pl. V, fig. 5. Aspidastraea orientalis Kühn, 1933: Baron-Szabo, p. 124, pl. 5, fig. 6, pl. 10, figs 1, 4. Aspidastraea orientalis, Kühn 1933: Löser, p. 9. Aspidastraea orientalis Kühn, 1933: Baron-Szabo, p. 144, pl. 103, figs 4–5, pl. 104, figs 2–3.


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Aspidastraea orientalis Kühn, 1933: Baron-Szabo, p. 144, pl. 25, figs 1–5.

Dimensions. c-c (adjacent series)=3–5.5 mm, up to 7 mm in peripheral areas; c-c (same series)= 2.5–5 mm, around 1.5 mm in areas of intense budding; s(secondary corallites)=16–28, s (main corallites)= up to 60; s/ mm=16–20/5; h=5–25 mm; colony= 18–53 mm in diameter. Description. Corallum is in the form of a free colony, convex above, circular or slightly elliptical in outline, with corallites that are produced by circumoral budding; base flat or insignificantly concave, covered by a concentrically wrinkled epitheca; generally, one slightly larger central calice is present; rarely, two main calices may occur, or the size of the main calice might similar to the non-dominant corallites; septa straight or wavy, moderate and subequal in thickness, porous or subcompact, and covered with numerous granules and pennulae laterally; about 10 septa reach corallite centre, in main corallite up to 30 septa extend to the axial region; central part of corallites can be free or filled with a trabecular columella; thin endothecal dissepiments present; synapticulae very abundant; no wall between the corallites. Type locality of species. Senonian of Iran (Esfahan Basin). Distribution. Turonian of Armenia, Santonian of Austria (Gosau Group at Hochmoos), Senonian of Iran, Upper Maastrichtian of the UAE/Oman border region, ?Paleocene of India.

Aspidastraea semhae (Kossmat, 1907) Pl. 16, figs 5a, b *1907 1918 1922 1930 1931 v1933 1981 1981 2000 v2000 v2002

Aspidiscus semhae: Kossmat, p. 54, pl. 5, figs 9a–c. Aspidiscus semhae Kossmat: Fossa-Mancini, p. 145, pl. 14 [1], figs 10a–b. Aspidiscus semhae Kossmat: Hoppe, p. 72. Aspidiscus semhae Kossmat: Renz, p. 10. Aspidiscus semhae Kossmat: Renz, p. 5. Aspidastraea semhae (Kossmat): Kühn, p. 180 (topotypes studied). Cyclolites arabicus sp. nov.: Abed & El-Asa’ad, p. 279, pl. 3, figs 3a–b. Cyclolites hemispaerica Lamarck: Abed & El-Asa’ad, p. 282, pl. 2, figs 4a–c. Aspidastraea semhae (Kossmat 1907): Löser, p. 9. Aspidastraea semhae (Kossmat, 1907): Baron-Szabo, p. 124, pl. 12, figs 2, 4–5. Aspidastraea semhae (Kossmat, 1907): Baron-Szabo, p. 144, pl. 104, figs 1, 4.

Dimensions. c-c (same series)=2–8 mm, up to 10 mm in peripheral areas; c-c (adjacent series)=6–14 mm; s/ mm=9–12/5, in areas of intense budding up to 16/5. Description. Corallum is in the form of a free colony, convex above and elliptical in outline; base flat or slightly concave, sometimes covered by a smooth epitheca; corallites arranged in circumoral rows; generally, no central calice is present; in juvenile forms slightly dominant central corallites may be present; septa thick, straight or wavy, equal in thickness, and porous or subcompact, laterally covered with numerous pennulae and granules that vary in size and shape; 12 to about 24 septa reach corallite centre; axial part of corallites can be free or filled with a trabecular columella; endotheca consists of thin dissepiments; synapticulae very abundant; no wall between the corallites; microstructural features not preserved. Type locality of species. Cenomanian of the Arabian Gulf (Sokrota Island). Distribution. Upper Albian-Cenomanian of the Arabian Gulf (Sokrota Island), Campanian-Lower Maastrichtian of central Saudi Arabia, Upper Maastrichtian of Iran, Middle-Upper Maastrichtian of the UAE/Oman border region, Upper Maastrichtian of The Netherlands.



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Genus Paracycloseris Wells, 1934 Type species. Paracycloseris elizabethae Wells, 1934, Campanian of Jamaica.

Diagnosis. Solitary, patellate, circular in outline. Septa compact to subcompact in older cycles, porous in younger cycles. Septal arrangement irregularly developed according to the Pourtalès plan in younger cycles. Septal flanks covered with various types of granules and pennulae. Synapticulae abundant. Endothecal dissepiments vesicular, few in number. Wall synapticulothecal, irregularly developed. Epithecal wall generally present.

Paracycloseris nariensis (Duncan, 1880) Pl. 17; figs 1a–9 *1880 1925 1927 v1934 parsv1936 v1941a 1987 2000 2002 v2002 2002 2005

Trochocyathus Nariensis, Duncan: Duncan, p. 71, pl. 9, figs 1–13, pl. 16, figs 9, 10. Trochocyathus Nariensis Duncan 1880: Felix, pars 28, p. 201. Trochocyathus Nariensis Duncan 1880: Felix, pars 35, p. 431 (older synonyms cited therein). Paracycloseris elizabethae: Wells, p. 86, pl. 3, figs 5–10, pl. 5, figs 1–2. Leptocyathus hexacristatus (nov. sp.): Alloiteau, p. 11, pl. 6, figs 15, 16 and 19, non figs 13, 14, 17, and 18. Paracycloseris elizabethae Wells: Wells, p. 291 (9), pl. 2, fig. 1. Paracycloseris elizabethae Wells: Kuzmicheva, p. 60. Leptocyathus hexacristatus, Alloiteau 1936: Löser, p. 47. Paracycloseris elizabethae Wells: Mitchell, p. 6ff., table 1. Paracycloseris elizabethae Wells, 1934: Baron-Szabo, pl. 105, figs 1–5. Paracycloseris elizabethae Wells 1934: Löser et al., p. 479 (older synonyms cited therein). Paracycloseris elizabethae Wells, 1934: Filkorn et al, p. 125, fig. 2k.

PLATE 17 Fig. 1 Paracycloseris nariensis (Duncan, 1880) (holotype of Paracycloseris elizabethae Wells, 1934), NMNH, I-74489, “forma typicum”, adult specimen, Maastrichtian of Jamaica, scale bars: 5 mm. 1a, upper surface; 1b, base of corallum; 1c, lateral view of corallum. Fig. 2 Paracycloseris nariensis (Duncan, 1880) “forma typicum”, adult specimen, Maastrichtian of Jamaica. 2a, cross thin section, scale bar: 4 mm; 2b, close-up, scale bar: 1.5 mm. Fig. 3 Paracycloseris nariensis (Duncan, 1880), as figured in Duncan (1880), “forma robustum”, cross view, Eocene of Pakistan, scale bar: 8 mm. Fig. 4 Paracycloseris nariensis (Duncan, 1880), as figured in Duncan (1880), “forma robustum”, longitudinal view, Eocene of Pakistan,scale bar: 8 mm. Fig. 5 Paracycloseris nariensis (Duncan, 1880) (syntype of Leptocyathus hexacristatus Alloiteau, 1936), MNHN, Mo5212 (M-5), “forma typicum”, cross thin section, Maastrichtian of Madagascar, scale bar: 3 mm. Fig. 6 Paracycloseris nariensis (Duncan, 1880) (syntype of Leptocyathus hexacristatus Alloiteau, 1936), MNHN, Mo5212, “forma typicum”, upper surface, Maastrichtian of Madagascar, scale bar: 3 mm. Fig. 7 Paracycloseris nariensis (Duncan, 1880), NMNH, Coates coll., no. 557d, “mixed type of forma typicum and forma robustum”, longitudinal thin section, juvenile specimen, Maastrichtian of Jamaica, scale bar: 4 mm. Fig. 8 Paracycloseris nariensis (Duncan, 1880), NMNH, Coates coll., no. 557e, “mixed type of forma typicum and forma robustum”, cross thin section, intermediate stage between juvenile and adult, Maastrichtian of Jamaica, scale bar: 4 mm. Fig. 9 Paracycloseris nariensis (Duncan, 1880), NMNH, Coates coll., no. 557c, “mixed type of forma typicum and forma robustum”, cross thin section, juvenile specimen, Maastrichtian of Jamaica, scale bar: 4 mm.


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Dimensions. d (min)=9–31 mm; d (max)=10.5–35 mm; s=up to 200; d (min)/d (max)=0.85–1. Description. Corallum solitary, flat, slightly convex or conical on the base, concave above, circular in outline, with circular central pit; lower surface often covered by wrinkled epithecal wall. Septa long, straight, and arranged in 5–6 complete cycles, alternating in length; S1 and S2 generally subcompact and often distincly thicker than younger septa, reach axial region where trabecular prolongations of their inner ends dissociate to form paliform structures; columella well-developed, papillose; S3 subcompact or porous; younger septa porous. Remarks. According to Wells (1941a), three growth-forms can be distinguished in Paracycloseris: forma typicum, forma robustum, and forma turbinatum. These forms are shown on: Plate 17, figs 1–2b and 5–6: forma typicum, Plate 17, figs 3–4: forma robustum, Plate 17, figs 7–9: mixed types of forma typicum and forma turbinatum. Type locality of species. Eocene of Pakistan (Raduk, Nari Series). Distribution. Upper Cretaceous of Cuba and Romania, Campanian-Maastrichtian (new material) of Jamaica, Maastrichtian of Madagascar and Mexico (Ocozocuautla Formation [see Filkorn et al, 2005] and Cardenas Formation [see Wells, 1941a, p. 292]), Eocene of Pakistan, Miocene of Armenia. New Material. Maastrichtian of Jamaica, NMNH, Coates coll., sample nos.: 323; 336; 341-II; 369; 375a; 375c; 378; 391a; 391b; 415; 422; 423a; 427 (=Jerusalem Mountain Inlier); 527; J-71-128Ba; J-71-128Bb (=Ducketts Land Settlement); 535 (=Shaw Castle, Maldon Formation); 557a; 557b; 557c; 557d; 557e; 557f; J71-34a; J-71-34b-II; J-71-34c; J-71-34d; J-71-34e; J-71-34f; J-71-34g; J-71-34h; J-71-34i; J-71-34j; J-7134k; J-71-34L; J-71-34m; J-71-34n; J-71-42-16a; J-71-42-16b; J-71-42-16c; J-71-42-16d; J-71-42-16e; J-7142-16f; J-71-42-16g; J-71-42-16h; J-71-42-16i; J-71-42-16j; J-71-42-16k; J-71-42-16L; J-71-42-16m; J-7142-16n; J-71-42-16o; J-71-42-16p; J-71-42-16q; J-71-42-16r; J-71-42-16s; J-71-42-16a2; J-71-42-16b2; J-7142-16c2; J-71-42-16d2; J-71-42-16e2; J-71-42-16f2; J-71-42-16g2; J-71-42-16h2; J-71-42-16i2; J-71-4216j2; J-71-42-16k2; J-71-45a; J-71-47 (=Rio Minho); 585a-I (=probably Cambridge area); J-72-2a; (=Marchmont Inlier); J3527 (=Point Flamstead); J3669 (=Road Sunderland).

Family Latomeandridae Alloiteau, 1952 Diagnosis. Colonial. Gemmation extracalicinal. Septa costate or biseptal, made of simple or compound trabeculae. Sepral perforations abundant and large. Axial ends of septa not dissociated, anastomosis frequent. Endothecal dissepiments and synapticulae sparse. Columella parietal, weak. Wall synapticulothecal, complete or incomplete. Remarks. The nominatform of this family, Latomeandra Milne Edwards & Haime, 1849b, was subject to a major revision by Alloiteau (1957, p. 238 and 312 ff). He concluded that all Cretaceous forms assigned to this genus belonged to different groups and transferred them into the genera Koilomorpha Alloiteau, Meandraraea Étallon, Clausastrea d'Orbigny, Vallimeandra Alloiteau, and others. According to Alloiteau, the genus Latomeandra Milne Edwards & Haime is restricted to the Jurassic.

Genus Dimorphastrea d’Orbigny, 1850 Type species. Dimorphastrea grandiflora d’Orbigny, 1850, Barremian of France (Haute-Marne).


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Diagnosis. Colonial, massive, thamnasterioid. Gemmation circumoral. Septa subcompact with increasing perforations toward the axial end of the septa. Septal margins granular, septal flanks granular and pennular. Columella papillose. Synapticulae abundant. No wall between the corallites. Endothecal dissepiments thin, subhorizontal.

Dimorphastrea solida Umbgrove, 1925 Pl. 18, fig. 1 v*1925 v1930a ?1945 1982 1999 2000 v2002 2002

Dimorphastrea solida spec. nov.: Umbgrove, p. 115, pl. 9, fig. 7. Dimorphastrea ogilviae n. sp.: Oppenheim, p. 196, pl. 9, fig. 8, pl. 13, figs 2–2a, pl. 22, fig. 6 (topotypes studied). Dimorphastrea solida Umbgrove 1925: Bataller, p. 43. Dimorphastrea aff. sulcosa Reuss 1854: Beauvais, vol. 2, p. 87. Dimorphastrea solida: Leloux, p. 193. Dimorphastrea solida, Umbgrove 1925: Löser, p. 29. Dimorphastrea solida Umbgrove, 1925: Leloux, p. 14, pl. 1, fig. 1. Dimorphastrea solida Umbgrove, 1925: Baron-Szabo, p. 146.

Dimensions. c-c (same series)=6–10 mm; c-c (adjacent series)=10–13 mm; s/mm=8–10/10. Description. Cast of a thamnasterioid colony; corallites arranged in circumoral series; septa confluent, subcompact, straight, arranged in 3 irregularly occurring size orders. Type locality of species. Upper Maastrichtian of The Netherlands. Distribution. ?Santonian of Spain, Upper Santonian of Austria (Gosau Group at Neffgraben), Upper Maastrichtian-Danian of The Netherlands.

Genus Fungiastrea Alloiteau, 1952 Type species. Fungiastrea laganum Alloiteau, 1952, Upper Turonian of France (Vaucluse) (see Alloiteau, 1952a).

Diagnosis. Colonial, massive, thamnasterioid to submeandroid. Gemmation intracalicinal, extracalicinal in places. Calicinal centres distinct. Septa compact to subcompact, confluent, moderately granulated laterally. Columella spongy. Endothecal dissepiments thin, vesicular to subtabulate. Synapticulae rare. No wall between corallites.

Fungiastrea crespoi (Felix, 1891) Pl. 18, fig. 3 *1891 ?1995 v1996 v1997 v1997 v1999 v2000

Thamnastrea Crespoi nov. sp.: Felix, p. 146, pl. 22, fig. 5. Fungiastrea ? brevipes (Stoliczka 1873): Löser & Raeder, p. 48. Fungiastraea crespoi (Felix, 1891): Baron-Szabo, in Baron-Szabo & Steuber, p. 27, pl. 16, figs 2, 3. Fungiastraea crespoi (Felix, 1891): Sanders & Baron-Szabo, p. 74, pl. 21, fig. 8 (non fig. 7). Fungiastrea crespoi (Felix, 1891): Baron-Szabo, p. 87, pl. 16, fig. 3. Fungiastrea crespoi (Felix, 1891): Baron-Szabo & González-Léon, p.490, figs 5a, 6i. Fungastrea crespoi (Felix, 1891): Baron-Szabo, p. 121, pl. 11, fig. 4.



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Fungastrea crespoi (Felix, 1891): Baron-Szabo, p. 265, fig. 2 G. Fungiastrea crespoi (Felix 1891): Löser et al., p. 311 (older synonmys cited therein). Fungastrea crespoi (Felix, 1891): Baron-Szabo, p. 147, pl. 107, figs 2, 4. Fungastrea crespoi (Felix, 1891): Baron-Szabo et al., p. 201, pl. 38, fig. 1.

Dimensions. c-c=4–8 mm; s/mm=10–15/5. Description. The corallum is massive-lamellar and thamnasterioid. Septa are long, wavy, confluent, nearly equal in thickness, irregularly perforated, with rounded or spiniform granules and pennulae laterally. The columella is spongy-papillose. A small number of synapticulae are irregularly distributed. The endotheca is made of thin tabulate and slightly arched dissepiments. Type locality of species. Barremian of Mexico (Tehuacan, San Antonia de las Salinas). Distribution. Barremian of Mexico (Tehuacan), Barremian-Aptian of Trinidad and Tobago, Upper Barremian of Bulgaria, Lower Aptian of Austria, Germany, and Greece, Upper Aptian-Upper Albian of Iran, Aptian and Middle Albian of Mexico (Sonora), Cenomanian of northwestern Germany, Upper TuronianConiacian of Austria, Middle-Upper Maastrichtian of the UAE/Oman border region.

Fungiastraea flexuosa (Goldfuss, 1826) Pl. 18; figs 4a–5 v*1826 v1854 1857 1857 v1881 v1903a 1904 1911 1914 1914 v1925 v1930a ?1937b 1971 v1982 v1993

Astrea flexuosa nobis: Goldfuss, vol. 1, p. 67, pl. 12, figs 10a–b. Thamnastraea exigua m.: Reuss, p. 119, pl. 18, figs 5–6. Thamnastraea exigua: Milne Edwards, vol 2, p. 556. Thamnastrea flexuosa: Milne Edwards, vol. 2, p. 574. Astrea flexuosa: Quenstedt, p. 857, pl. 176, fig. 54. Thamnastraea exigua Reuss 1854: Felix, p. 209–210, text-fig. 15. Astraraea cf. flexuosa Felix (Goldfuss sp.): Felix, p. 383, pl. 3, fig. 7. Thamnastraea exigua Reuss: Trauth, p. 69, pl. 3, fig. 3. Thamnastraea exigua Reuss 1854: Felix, pars 7, p. 197. Astraraea flexuosa Goldfuss sp. 1826: Felix, pars 7, p. 204. Astraraea flexuosa Goldf. spec.: Umbgrove, p. 112. Synastraea exigua (Reuss): Oppenheim, p. 166, pl. 31, figs 12, 12a. Synastraea exigua Reuss sp. 1854: Bataller, p. 307. Fungiastraea aff. exigua (Reuss): Morycowa, p. 111–112, pl. 28, fig. 2. Dimorphomeandra exigua (Reuss) 1854: Beauvais, vol. 2, p. 55–56, pl. 26, fig. 1 (older synonyms cited therein). Fungiastraea cf. exigua (Reuss, 1854): Baron-Szabo, p. 162, pl. 5, figs 3a–b.

PLATE 18 Fig. 1 Dimorphastrea solida Umbgrove, 1925, syntype, RMNH, 29061, upper surface, ‘steinkern preservation’, Upper Maastrichtian of The Netherlands, scale bar: 12 mm. Fig. 2 Ellipsocoenia conferta (Umbgrove, 1925), holotype, RMNH, 29039, Maastrichtian of The Netherlands, scale bar: 8 mm. 2a, upper surface, ‘steinkern’ preservation; 2b, ‘positive sketch’ (presented for the first time). Fig. 3 Fungiastrea crespoi (Felix, 1891), BMNH, AZ 642, upper surface, MiddleUpper Maastrichtian of the UAE/Oman border region, scale bar: 6 mm. Fig. 4 Fungiastrea flexuosa (Goldfuss, 1826), holotype, IPB, Goldfuss coll., no. 224, Maastrichtian of The Netherlands, scale bar: 6 mm. 4a, upper surface, ‘steinkern’ preservation; 4b, ‘positive sketch’ (presented for the first time). Fig. 5 Fungiastrea flexuosa (Goldfuss, 1826), NMNH, Coates coll., no. 442, cross thin section, Maastrichtian of Jamaica, scale bar: 6 mm.


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Fungiastraea exigua (Reuss, 1854): Baron-Szabo, p. 87, pl. 16, figs 5–6. Fungiastraea exigua (Reuss, 1854): Baron-Szabo, p. 149, pl. 4, fig. 5. Fungiastraea exigua (Reuss, 1854): Baron-Szabo, p. 461, pl. 2, fig. 2, pl. 7, fig. 5. Fungiastraea exigua (Reuss 1854): Löser et al, p. 312. Astraraea flexuosa (Goldfuss, 1826): Baron-Szabo, p. 104, pl. 70, fig. 4. Fungiastraea exigua (Reuss, 1854): Baron-Szabo, p147. Fungiastraea exigua (Reuss, 1854): Baron-Szabo, p. 140, pl. 20, figs 4, 6.

Dimensions. c-c=3–5 mm, in peripheral areas of colony up to 6 mm; juvenile corallites as small as 2.5 mm; s=24–36, in corallite in budding stage the number of septa may be larger; s/mm=15–20/5. Description. Massive or lamellar, thamnasterioid corallum, with corallites that are regularly distributed over the colony. Corallites irregularly distributed or arranged in submeandroid series; septa confluent, occasionally subconfluent, subcompact, thin, nearly equal, straight or wavy; anastomosis is present; about 20 septa extend to corallite centre; well-developed spongy-papillose columella and paliform lobes fill axial region of corallite. Remarks. Re-examination of the holotype of Astrea flexuosa revealed that it differed from Astraraea in several ways: intracalicinal and extracalicinal budding, the presence of distinct corallite centres, and in the occurrence of endothecal dissepiments rather than synapticulae. It differs from the genus Thamnastrea by having corallites arranged in submeandroud series, compact and subcompact septa, and by lacking paliform teeth on the 1st order septa. Type locality of species. Maastrichtian of The Netherlands (St. Pietersberg). Distribution. Lower Aptian of Romania and the German/Austrian border region („Helvetikum“), AptianAlbian (Vasco-Cantabria), Lower Coniacian and Upper Santonian of France, Upper Turonian-Santonian of Austria (Gosau Group), Upper Santonian-Lower Campanian of Portugal (Azinhaga do Pinhal-de-Loura), Campanian and ?Maastrichtian of northern Spain (Catalonia), Maastrichtian of Jamaica (this paper) and The Netherlands. New Material. Maastrichtian of Jamaica, NMNH, Coates coll., sample nos.: 373b, 562 (=Jerusalem Mountain Inlier); 442; 442i; 442j (=Catadupa); 485; 485n (Ducketts Land Settlement); 586g (=probably Cambridge area); J3456c (=probably Welcome Hall); J3502a; J3502b3 (=Point Flamstead); J-71-41-16b4 (=Rio Minho).

Genus Ellipsocoenia d’Orbigny, 1850 (=Faviomorpha Reig Oriol, 1990, Type species. Favia bauzai Mallada, 1885, Eocene of Spain). Type species. Ellipsocoenia regularis d’Orbigny, 1850; Hauterivian of France (Fontenoy).

Diagnosis. Colonial, massive, plocoid. Gemmation intracalicinal. Calices circular or elliptical in outline, monocentric. Costosepta subcompact with large pores, nonconfluent. Columella trabecular, spongy-papillose. Wall synapticulothecal. Endothecal and exothecal dissepiments numerous.

Ellipsocoenia conferta (Umbgrove, 1925) Pl. 18, figs 2a, b v*1925

Orbicella conferta spec. nov.: Umbgrove, p. 105, pl. 8, fig. 6.


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v1925 2000 2000 2002 v2002

Favia Maastrichtensis spec. nov.: Umbgrove, p. 107, pl. 11, fig. 19. Favia maastrichtensis, Umbgrove 1926: Löser, p. 35. Orbicella conferta, Umbgrove 1926: Löser, p. 56. Favia conferta Umbgrove, 1925: Leloux, p. 13. Favia maastrichtensis Umbgrove, 1925: Leloux, p. 14, pl. 1, fig. 4.

Dimensions. d (max, lumen)= 3.5–8 mm; d (min, lumen)= 3–6 mm; c-c=6–10 mm; s=60, up to around 80. Description. Plocoid colony; corallites elliptical in outline; costosepta compact to subcompact, nonconfluent to subconfluent, arranged in 4 complete cycles with the beginning of a fifth cycle in 6 systems; anastomosis of S3 to S2 and S4 to S3 common; columella spongy-papillose, trabecular-lamellar in some corallites. Remarks. In having both subcompact septa and a synapticulothecal wall, as well as showing intracalicinal budding, the holotype originally described as Orbicella conferta closely corresponds to the genus Ellipsocoenia d’Orbigny but differs from the genus Orbicella Dana (=junior synonym of Montastrea Blainville according to Vaughan & Wells 1943, Wells 1956, and others), which is characterized by compact septa, extracalicinal budding, and a corallite wall that is septothecal and septoparathecal. The specimen orginally described as Favia maastrichtensis differs from the genus Favia Oken in having a synapticulothecal wall (parathecal and septothecal in Favia) and in showing intracalicinal budding with polyps that remain monocentric until comlpete separation of new polyps. In Favia up to three corallite centres remain connected, forming a small meandroid series. In addition, in Favia the intracalicinal budding takes place by septal division (sensu Morycowa and Roniewicz 1990). In the type material of the species E. conferta the corallites appear to be generally smaller than in the form E. maastrichtensis, which is apparently due to intense budding, in E. conferta. However, the range of corallite sizes overlap, therefore, in E. conferta the dimensions of the largest corallites correspond to the general corallite diameter in E. maastrichtensis, and the smallest ones in this form have the same diameter as the generally occurring corallites in E. conferta. Therefore, they are considered synonymous. Type locality of species. Maastrichtian of The Netherlands (Limburg). Distribution. Maastrichtian of The Netherlands.

Genus Baryphyllia Fromentel, 1857 (=Polyaraea Fritsch, 1878, Type species. Polyaraea gemmans Fritsch, 1878, Eocene of Borneo). Type species. Barysmilia gregaria d'Orbigny, 1850, Hauterivian of France.

Diagnosis. Colonial, massive, subplocoid. Gemmation intracalicinal. Corallites monocentric, projecting, closely united by costae and exothecal dissepiments. Costosepta fenestrate, granulated laterally, beaded marginally. Columella parietal. Synapticulae present, forming a stereozone.

Baryphyllia maxima (Umbgrove, 1925) Pl. 19, fig. 1 v*1925 non1987 2000

Orbicella maxima spec. nov.: Umbgrove, p. 104, pl. 10, fig. 14. Brachyphyllia maxima (Umbgrove, 1925): Meyer, p. 25, pl. 4, fig. 7. Montastrea maxima (Umbgrove, 1926): Löser, p. 53



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Montastrea maxima (Umbgrove, 1925): Leloux, p. 13.

Dimensions. d (min)=up to 14 mm; d (max)=up to 18 mm; s=96. Description. Massive, subplocoid colony; corallites projecting; costosepta irregularly porous, subcompact in older cycles, developed in 5 complete cycles in 6 systems; columella parietal, well-developed. Remarks. The material illustrated by Meyer (1987) as Brachyphyllia maxima (Umbgrove) shows all the characteristics of the genus Brachyphyllia. However, the type material of the species O. maxima corresponds to the genus Baryphyllia. Type locality of species. Maastrichtian of The Netherlands (St. Pietersberg) Distribution. Maastrichtian of The Netherlands.

Genus Gyrodendron Quenstedt, 1880 Type species. Gyrodendron lobatum Quenstedt, 1880 (=Chorisastrea dubia Becker, 1876), Upper Jurassic (Kimmeridgian) of Germany.

Diagnosis. Colonial, branching, fasciculate, formed by clusters of polycentric corallites. Gemmation intracalicinal. Costosepta subcompact to porous, granulated laterally. Columella trabecular, well-developed. Endothecal dissepiments thin, vesicular to subtabulate. Synapticulae present. Wall synapticulothecal.

Gyrodendron boltonae (Wells, 1934) Pl. 19, figs 2–4b v*1934 1995 v1998 2000 2000 2002 v2002 2002

Diplaraea (?) boltonae: Wells, p. 83, pl. 2, figs 1–2. Diplaraea posae n. sp.: Reig Oriol, p. 36, pl. 6, figs 4–5. Gyrodendron leptonema n. sp.: Baron-Szabo, p. 151, pl. 11, figs 2, 4, pl. 12, figs 5, 6. Diplaraea boltonae, Wells 1934: Löser, p. 30. Diplaraea posae, Reig Oriol 1995: Löser, p. 30. Gyrodendendron boltonae (Wells, 1934): Baron-Szabo, p. 152. Gyrodendendron leptonema Baron-Szabo, 1998: Baron-Szabo, p. 152, pl. 111, figs 2, 3. Gyrodendendron posae (Reig Oriol, 1995): Baron-Szabo, p. 152.

PLATE 19

Fig. 1 Baryphyllia maxima (Umbgrove, 1925), holotype, RMNH, 29030, upper surface, Maastrichtian of The Netherlands, scale bar: 4 mm. Fig. 2 Gyrodendendron boltonae (Wells, 1934), NMNH, Coates coll., no. 570b, cross thin section, Maastrichtian of Jamaica, scale bar: 5 mm. Fig. 3 Gyrodendendron boltonae (Wells, 1934), NMNH, Coates coll., no. J71-34o, cross thin section of adult corallite, Maastrichtian of Jamaica, scale bar: 5 mm. Fig. 4 Gyrodendendron boltonae (Wells, 1934), holotype, NMNH, I74479, Campanian of Jamaica. 4a, upper surface, scale bar: 4 mm; 4b, longitudinal view, scale bar: 6 mm. Fig. 5 Loboseris abbreviata (Reuss, 1854), NMNH, Coates coll., no. 595a, Middle-Upper Maastrichtian of Jamaica. 5a, close-up, scale bar: 3 mm; 5b, cross thin section, scale bar: 10 mm. Fig. 6 Loboseris abbreviata (Reuss, 1854), holotype, GBA, 1854/07/ 0050, upper surface, Santonian of Austria, scale bar: 5 mm.


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Dimensions. d (series)=4–21 mm; length of series=17–35 mm; c-c (in series)=5–14 mm; s=48, up to around 200 in latest adult stages; s/mm=6–10/2. Description. Flabello-meandroid (in juvenile stages) or phaceloid-fasciculate (in adult stages); corallites monocentric, grouped in short meandroid series; some corallites remain subpolygonal in outline within the series, giving the colony a subcerioid appearance; costosepta subcompact or perforate, frequently uniting, arranged in 3–4 size orders alternating in length and thickness; septal flanks covered with pennules and granules, varying in size and shape; columella parietal, irregularly developed; endotheca dissepiments numerous, thin, vesicular. The wall is synapticulothecal to septothecal. Remarks. The species G. boltonae and G. leptonema represent juvenile stages of colonies in which there are also corallites or corallite series present that correspond to the larger form described as posae. Therefore, their synonymy is suggested. Type locality of species. Campanian of Jamaica (Catadupa). Distribution. Campanian-Maastrichtian (this paper) of Jamaica, Campanian-Maastrichtian of Spain (Torallola, Pobla de Segur). New Material. Maastrichtian of Jamaica, NMNH, Coates coll., sample nos.: 379b (=Jerusalem Mountain Inlier); 431e; J-66-27 (=Catadupa railway); 515; 515c (=Ducketts Land Settlement); 550f; J-71-34o (=Rio Minho); 568e; 570b; 570j; 581; 584; 586f (=probably Cambridge railway area); J3444b; J3444c; J3444d; J3444f; (=Welcome Hall); J3502b2; J3502c (=Point Flamstead); J-71-11 (=Shaw Castle, Maldon Formation).

Genus Loboseris M. Beauvais, 1982 Type species. Mussa abbreviata Reuss, 1854, Santonian of Austria (Gosau Group).

Diagnosis. Colonial, massive, meandroid. Gemmation intracalicinal-terminal. Calices monocentric or in short wavy series. Peritheca consists of thin vesicular and tabular dissepiments. Costosepta perforate, nonconfluent, marginally granular. Columella parietal-spongy. Synapticulae and endothecal dissepiments abundant. Wall synapticulothecal. No wall between calices of the same series.

Loboseris abbreviata (Reuss, 1854) Pl. 19, figs 5a–6 v*1854 1857 1858–61 v1903a 1914 v1982 2000 v2002

Mussa abbreviata: Reuss, p. 104, pl. 4, figs 4–6. Favia ? abbreviata: Milne Edwards, vol. 2, p. 440. Thecosmilia abbreviata (Reuss): de Fromentel, p. 172. Thecosmilia abbreviata (Reuss): Felix, p. 243. Thecosmilia abbreviata Reuss sp. 1854: Felix, pars 7, p. 161. Loboseris abbreviata (Reuss) 1854: Beauvais, vol. II, p. 261–262, pl. 47, fig. 3. Loboseris abbreviata (Reuss 1854): Löser, p. 49. Loboseris abbreviata (Reuss, 1854): Baron-Szabo, pl. 113, figs 1–4.

Dimensions. d (max)=18–25 mm; d (min)=15–17 mm; s/mm=19–25/10. Description. Colonial, massive, meandroid; gemmation intracalicinal-terminal. calices monocentric or in short wavy series; peritheca consists of thin vesicular and tabular dissepiments; costosepta perforate, noncon-


BARON-SZABO

fluent, marginally granular; columella parietal-spongy; synapticulae and endothecal dissepiments abundant; wall synapticulothecal; no wall between calices of the same series. Type locality of species. Santonian of Austria (Gosau Group). Distribution. Santonian of Austria (Gosau Group), Middle-Upper Maastrichtian of Jamaica. New Material. Middle-Upper Maastrichtian of Jamaica, NMNH, Coates coll., sample no.: 595 (=probably Cambridge area); J-71-12 (=Shaw Castle, Maldon Formation); ?540 (has d ranging between 7 and 10 mm but same septal density as abbreviata =?juvenile form of abbreviata ) (=Shaw Castle, Maldon Formation). Family Brachiphylliidae Alloiteau, 1952 Diagnosis. Solitary and colonial. Gemmation extracalicinal. Septa nonconfluent. Anastomosis absent. Septal perforations regular. Axial ends of septa not dissociated. Endothecal dissepiments sparse. Synapticulae abundant. Wall synapticulothecal, incomplete. Columella well developed or rudimentary.

Genus Brachyphyllia Reuss, 1854 Type species. Brachyphyllia dormizieri Reuss, 1854, Santonian of Austria (Gosau Group); subsequent designation by Reuss (1864).

Diagnosis. Subplocoid to fasciculate colony. Gemmation extracalicinal and extracalicinal-marginal. Costosepta porous to subcompact, compact in oldest cycles, with strongly beaded margins. Columella well-developed, spongy. Endothecal dissepiments sparse. Synapticulae abundant. Wall parathecal-synapticulothecal.

Brachyphyllia depressa Reuss, 1854 Pl. 20, fig. 1 v*1854 1857 1858–61 1886 v1903a 1914 v1930a 1943 1956 v1982 1987 (v)1995 2000 v2002

Brachyphyllia depressa m.: Reuss, p. 103, pl. 2, figs 8–10 (topotypes studied). Brachyphyllia depressa: Milne Edwards, vol. 2, p. 480. Brachyphyllia depressa: de Fromentel, p. 209. Brachyphyllia depressa: de Fromentel, p. 578, pl. 168, figs 1–1b. Brachyphyllia depressa Reuss: Felix, p. 259. Brachyphyllia depressa Reuss 1854: Felix, pars 7, p. 169. Brachyphyllia depressa Reuss: Oppenheim, p. 339 (topotypes studied). Brachyphyllia depressa Reuss: Vaughan & Wells, pl. 11, figs 1–1a. Brachyphyllia. depressa Reuss: Wells, p. F. 381, figs 271.2a and 271.2b. Brachyphyllia depressa Reuss 1854: Beauvais, vol. 2, p. 225, pl. 43, figs 8a–b (topotypes studied). Brachyphyllia maxima (Umbgrove, 1925): Meyer, p. 25, pl. 4, fig. 7. Neocoeniopsis thraciensis sp. n.: Tchéchmédjiéva, p. 65, pl. 15, fig. 1. Brachyphyllia depressa, Reuss 1854: Löser, p. 16. Brachyphyllia depressa Reuss, 1854: Baron-Szabo, p. 153, pl. 115, fig. 1.



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PLATE 20 Fig. 1 Brachyphyllia depressa Reuss, 1854, topotype, NHMW, Reuss coll. no. 5, upper surface, Turonian-Campanian of Austria, scale bar: 8 mm. Fig. 2 Dermosmiliopsis orbignyi Alloiteau, 1952, UNAM, IGM 8724, Maastrichtian of Mexico. 2a, cross view, thin section, scale bar: 9 mm; 2b, close-up, scale bar: 4 mm. Fig. 3 Dermosmiliopsis tenuicosta (Reuss, 1854), NMNH, Coates coll., no. 535, Middle-Upper Maastrichtian of Jamaica. 3a, close-up, scale bar: 3 mm; 3b, cross view, thin section, scale bar: 4 mm. Fig. 4 Brachyphyllia felixi Baron-Szabo, 2000, NMNH, Coates coll., no. 545q, cross view, polished surface, Middle-Upper Maastrichtian of Jamaica, scale bar: 3 mm. Fig. 5 Brachyphyllia felixi Baron-Szabo, 2000, holotype, BMNH, AZ 2527, cross view, upper surface, Middle-Upper Maastrichtian of the UAE/ Oman border region, scale bar: 5 mm.

Dimensions. d=22–30 mm, juvenile corallites around 10 mm; s=40 (juvenile) up to ca. 100; s/mm=12–14/10. Description. Subplocoid to fasciculate, subcircular to elliptical in outline; costosepta arranged in 4 to 5 cycles in 6 systems in adult corallites, S1–S3 nearly equal in length, slightly alternating in thickness. Remarks. Neocoeniopsis thraciensis Tchéchmédjiéva, 1995, from the Maastrichtian of Bulgaria, differs from the genus Neocoeniopsis in forming a rather fasciculate colony by marginal budding and having abundant synapticulae that occur throughout the corallum (see plate 15, fig. 1 in Tchéchmédjiéva 1995). In Neocoeniopsis the corallite integration is plocoid, and synapticulae are sparse, mainly restricted to the vicinity of corallite wall. In addition, the septal apparatus of Neocoeniopsis thraciensis as shown on plate 15, fig. 1. in Tchéchmédjiéva (1995) closely corresponds to the brachyphylliid type and clearly differs from the kind developed in the pachyphylliids, to the latter of which, also in the opinion of the author Tchéchmédjiéva, Neocoeniopsis belongs. Because material stratigraphically corresponding to the Maastrichtian-Paleocene time period was not available, the Upper Cretaceous type material of the species B. depressa is illustrated on Fig. 1 on Pl. 20. Type locality of species. Turonian-Campanian of Austria (Gosau Group). Distribution. Turonian-Campanian of Austria (Gosau Group), Maastrichtian of Bulgaria, Danian of France (Vigny).

Brachyphyllia felixi Baron-Szabo, 2000 Pl. 20, figs 4, 5 non1848 non1854 v1903a non1903a 1905 pars1914 non1914 non1914 1930a non1978 *2000 v2002

Pleurocora haueri: Milne Edwards & Haime, vol. 11, p. 312. Pleurocora haueri M. Edw. et Haime: Reuss, p. 112, pl. 6, figs 26–27 (topotypes studied). Brachyphyllia haueri (Reuss): Felix, p. 261, pl. 20, fig. 15, text-fig. 29 (topotype studied). Pleurocora haueri M. Edw. e H.: Felix, p. 261. Brachyphyllia haueri (Reuss): Angelis d’Ossat, p. 200. Brachyphyllia haueri Reuss sp. 1854: Felix, pars 5, p. 32. Brachyphyllia haueri Reuss sp. 1854: Felix, pars 7, p. 170. Pleurocora haueri M. E. et H.: Felix, pars 7, p. 172. Brachyphyllia haueri Felix: Oppenheim, p. 343, pl. 46, figs 1–1b (topotypes studied). Pleurocora haueri Milne-Edwards & Haime 1848: Turnšek, in Turnšek & Polšak, p. 155, 173, pl. 11, figs 1–3. Brachyphyllia felixix sp. nov.: Baron-Szabo, p. 121, pl. 11, fig. 5. Brachyphyllia felixi Baron-Szabo, 2000: Baron-Szabo, p. 153, pl. 115, figs 3, 4.



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Dimensions. d=4–8 mm, in late adult stages up to 10 mm; s=60, up to about 100. Description. Plocoid colony with elevated corallites that are circular in outline. Gemmation is due to extracalicinal budding. Costosepta are subcompact or compact in older cycles, straight, thin, and arranged in 4 to 5 cycles in 6 systems, regularly alternating in length. Septa of the first 3 cycles are nearly equal in thickness. Septa of remaining cycles are distinctly thinner. About 12 septa reach the axial region, sometimes fusing with neighbouring septa or with the spongy-papillose columella. Lateral surfaces of septa are covered by spiny or rounded granules and pennulae. Synapticulae are distributed throughout the colony. The wall is synapticulothecal. Endotheca consists of delicate vesicular dissepiments. In early ontogenetic stages the corallite may be covered by an epithecal wall. Type locality of species. Middle-Upper Maastrichtian of the UAE/Oman border region (Jebel Huwayyah, Al Ain). Distribution. Aptian of northern Spain (Catalonia), Santonian-Campanian of Austria (Gosau Group), Middle-Upper Maastrichtian of the UAE/Oman border region and Jamaica (this paper). New Material. Middle-Upper Maastrichtian of Jamaica, NMNH, Coates coll., sample no.: 545q (=Rio Minho). Genus Dermosmiliopsis Alloiteau, 1952 Type species. Dermosmiliopsis orbignyi Alloiteau, 1952a, Upper Santonian of France (Aude).

Diagnosis. Phaceloid to subdendroid colony. Gemmation extracalicinal. Corallites monocentric. Septa porous, with strongly beaded margins. Columella spongy-papillose. Endothecal dissepiments sparse. Synapticulae abundant. Wall synapticulothecal.

Dermosmiliopsis orbignyi Alloiteau, 1952 Pl. 20, figs 2a, b *1952a 1957 1978 pars1982 2000 2002 v2006 v2006

Dermosmiliopsis Orbignyi ALL. 1950: Alloiteau, p. 671. nom. nud. Dermosmiliopsis Orbignyi nov. gen. nov. sp.: Alloiteau, pl. 4, figs 6a–b. Dermosmiliopsis orbignyi Alloiteau 1957: Turnšek & Polšak, p. 159, 176, pl. 15, figs 1–7. Barysmilia tenuicosta (Reuss) 1854: Beauvais, vol. 1, p. 182, pl. 14, fig. 9a non fig. 9b. Dermosmiliopsis orbignyi, Alloiteau 1952: Löser, p. 28. Dermosmiliopsis orbignyi Alloiteau, 1952: Baron-Szabo, p. 154. Dermosmiliopsis orbignyi Alloiteau, 1952: Schafhauser et al., p. 190, tab. 1. Dermosmiliopsis orbignyi Alloiteau, 1952: Baron-Szabo et al., p. 24, fig. 5.8.

Dimensions. d=10–17 mm, 6.5–9 mm in juvenile corallites; s=96, up to around 130, in juvenile corallites s=48–70. Description. Phaceloid-subdendroid colony; gemmation extracalicinal (marginal); septa subcompact to porous, with strongly beaded margins, arranged in 5 complete cycles with the beginning sixth cycle in 6 systems in adult corallites; columella spongy-papillose; endothecal dissepiments sparse, subtabulate, thin; synapticulae abundant; wall synapticulothecal. Type locality of species. Upper Santonian of France (Aude). Distribution. Upper Santonian of France, Santonian-Campanian of Croatia, Maastrichtian of Mexico (Cardenas Formation).


BARON-SZABO

Dermosmiliopsis tenuicosta (Reuss, 1854) Pl. 20, figs 3a, b v*1854 non1903a non 1961 1930a non 1937 non 1945 1978 pars1982 2002 v2002

Rhabdophyllia tenuicosta m.: Reuss, p. 105, pl. 6, figs 18 and 21 (topotypes studied). Stenosmilia tenuicosta (Reuss): Felix, p. 303, pl. 22, fig. 11, text-fig. 53. Stenosmilia tenuicosta (Reuss): Suraru, p. 655, pl. 1, fig. 3. Stenosmilia (?) tenuicosta (Reuss): Oppenheim, p. 434, pl. 42, figs 8–9. Stenosmilia tenuicostata Reuss sp. 1854: Bataller, p. 94. Stenosmilia tenuicostata Reuss sp. 1854: Bataller, p. 20. Dermosmiliopsis tenuicosta (Reuss 1854): Turnšek, in Turnšek & Polšak, p. 30 (158) and 48 (176), pl. 14, figs 1–5. Barysmilia tenuicosta (Reuss) 1854: Beauvais, vol. 1, p. 182, pl. 14, fig. 9b non fig. 9a. Dermosmiliopsis tenuicostata (Reuss 1854): Löser et al., p. 228 (older synonyms cited therein). Dermosmiliopsis tenuicosta (Reuss, 1854): Baron-Szabo, p. 154, pl. 116, figs 1–2.

Dimensions. d=4–8 mm; s=60–100. Description. Phaceloid to subdendroid colony; gemmation extracalicinal; corallites monocentric; septa subcompact to porous, with strongly beaded margins, arranged in 4–5 cycles in 6 systems; columella spongypapillose; endothecal dissepiments sparse, subtabulate, thin; synapticulae abundant; wall synapticulothecal. Type locality of species. Upper Turonian-Campanian of Austria (Gosau Group). Distribution. Upper Turonian-Campanian of Austria (Gosau Group), Santonian-Campanian of Slovenia, Croatia, and Spain, Middle-Upper Maastrichtian of Jamaica. New Material. Middle-Upper Maastrichtian of Jamaica, NMNH, Coates coll., sample nos.: 347; 353; 411 (=Jerusalem Mountain Inlier); 538 (=around two thousand pieces of fragments) (=Shaw Castle, Maldon Formation); J-71-13a3 (=Vaughnsfield).

Suborder Stylinina Alloiteau, 1952 Diagnosis. Colonial. Gemmation predominantly extracalicinal, rarely intracalicinal. Polyps generally circular, wall parathecal or septothecal (latter often thickened by stereozone). Radial elements (septa or costosepta) always compact and consisting of simple trabeculae, arranged in a single series or in a divergent system. Distal septal margins have very delicate denticles, sometimes subobsolete or made of granulae. Lateral septal surfaces ornamented with granulae, which can be very delicate and are aligned parallel to the distal margin. Auriculae present in some groups (e.g., nominatform Stylina), generally before 3rd cycle septa or younger. Endotheca present, generally abundant and vesicular, sometimes tabulate. Peritheca nearly always present.

Family Stylinidae d'Orbigny, 1851 Diagnosis. Colonial; colony formation by intra- and extratentacular budding. Corallites walls septothecal, often thickened by stereome. Endotheca tabular when present, rarely vesicular. Septa (so far as is known) composed of a single fan system of simple trabeculae, with smooth upper margins and smooth or finely granulated lateral surfaces. Columella styliform, lamellar or absent.



195

Genus Stylina Lamarck, 1816 (=Pseudocoenia d'Orbigny, 1850, Type species. Pseudocoenia bernardiana d'Orbigny, 1850, Jurassic of France). Type species. Stylina echinulata Lamarck, 1816, Sequanian (Jurassic) of France (Verdun).

Diagnosis. Colonial, massive, plocoid. Gemmation extracalicinal. Costosepta compact, laterally finely granulated. Columella styliform. Wall septothecal. Endothecal dissepiments tabular. Peritheca vesicular. Remarks. D'Orbigny (1850) established Pseudocoenia as a genus like 'Stylina lacking a columella' but without designating a type specimen for the type species. Later, Wells (1936) designated a lectotype which, however, turned out to have all characteristics of the genus Stylina Lamarck. Despite the fact that the lectotype differs from the generic concept of Pseudocoenia, the lectotype remains the valid type specimen of the type species of Pseudocoenia. Therefore, Pseudocoenia d'Orbigny, 1850, represents a junior synonym of Stylina Lamarck.

Stylina reussi Duncan, 1880 Text-Fig. 34 *1880 1925

Stylina Reussi, Duncan: Duncan, p. 30, pl. 10, figs 11–14. Stylina Reussi Duncan 1880: Felix, pars 28, p. 43.

Dimensions. d=2–2.5 mm, juvenile around 1.5 mm; c-c=2–3.5 mm; s=24.

Text-Fig. 34 Stylina reussi Duncan, 1880, as figured in Duncan (1880), Upper Paleocene of Pakistan, cross view, scale bar: 2 mm.

Description. Plocoid colony with densely packed circular to slightly elliptical corallites; costosepta arranged in 3 complete cycles in 6 systems, regularly alternating in length and thickness; S1 reach corallite centre and fuse with columella; in corallites of around 1.5 mm in diameter 12 septa are present; number of septa equals number of costae.


BARON-SZABO

Type locality of species. Upper Paleocene of Pakistan (Jhirk, Sind). Distribution. Upper Paleocene of Pakistan (Jhirk, Sind).

Genus STYLOSMILIA Milne Edwards & Haime, 1848a (=Pseudostylina Alloiteau, 1939b. Type species. Pseudostylina kovenkoi Alloiteau, 1939b, Upper Jurassic of Turkey). Type species. Stylosmilia michelini Milne Edwards & Haime, 1848a, Upper Jurassic of Germany (Raidersdorf).

Diagnosis. Colonial, phaceloid. Gemmation extracalicinal. Costosepta compact, covered by dense granular coenosteum. Columella styliform, strongly developed. Endothecal dissepiments subtabulate.

Stylosmilia kangpaensis Wu, 1975 Text-Fig. 35 (v)*1975 (v)1994 2000 2001 2002

Stylosmilia kangpaensis Wu, p. 102, pl. 8, figs 4–8. Stylosmilia kangpaensis Wu: Liao & Xia, p. 132, pl. 30, figs 4–7. Stylosmilia kangpaensis Wu 1975: Löser, p. 77. Stylosmilia kangpaensis Wu 1975: Löser & Liao, p. 666. Stylosmilia kangpaensis Wu, 1975: Baron-Szabo, p. 180.

Dimensions. d=4–5 mm, juvenile 2.5–3 mm; s=28–54. Description. Phaceloid-subdendroid corallum; corallites circular or slightly elongated in outline; costosepta compact, thin, finely granulated laterally, arranged in 3–4 cycles in 6 systems; S1 and S2 reach the center of the calice, sometimes fusing with styliform-sublamellar columella; endothecal dissepiments thin, subtabulate; wall septoparathecal. Type locality of species. Campanian-Maastrichtian of Tibet. Distribution. Campanian-Maastrichtian of Tibet.

Text-Fig. 35 Stylosmilia kangpaensis Wu, 1975, as illustrated in Wu (1975), Campanian-Maastrichtian of Tibet, scale bar: 5 mm.



197

Genus Tubicora Latham, 1929 (=Sagittastraea Alloiteau & Tissier, 1958, Type species. Sagittastraea pivetaui Alloiteau & Tissier, 1958, Paleocene of France). Type species. Tubicora somaliensis Latham, 1929, Eocene of Somalia.

Diagnosis. Colonial, phaceloid to subreptoid. Gemmation extracalicinal. Corallites cylindrical, connected by cross-tubuli. Costosepta compact. Columella styliform to substyliform. Endotheca scarcely developed. Wall septothecal and septoparathecal.

Tubicora aylmeri (Gregory, 1900) Pl. 21, fig. 1 v*1900 (v)1958 1993

Calamophyllia aylmeri, sp. nov.: Gregory, p. 31, pl. 1, fig. 3. Sagittastraea pivetaui nov. sp.: Alloiteau & Tissier, p. 247, pl. 2, figs 1 a–b. Tubicora aylmeri Gregory, 1900: Carbone, et al., p. 227, figs 11 f–g.

Dimensions. d=3–4.5 mm, juvenile around 2 mm; s=24+s4 . Description. Phaceloid corallum; corallites circular in outline connected by cross-tubuli which are 1.5–5 mm in diameter; costosepta generally straight; S1 and some of S2 reach center of calice; remaining septa alternate in length and thickness; columella styliform to substyliform; wall septoparathecal in upper part of corallite, septothecal deeper. Type locality of species. Upper Paleocene-Lower Eocene of Somalia. Distribution. Paleocene of France, Upper Paleocene-Lower Eocene of Somalia.

Family Agatheliidae L. & M. Beauvais, 1975 (=Ficariastraeidae M. Beauvais, 1982, =Hemiporitidae Alloiteau, 1952a p.p.) Diagnosis. Colonial, plocoid. Gemmation due to extracalicinal budding. Costosepta compact, arranged radially in 6 systems, beaded marginally. Columella styliform, or slightly spongy or lamellar. Endotheca sparsely developed. Septotheca formed by well-developed and abortive septa. Peritheca present, granulated. Microstructure of perithecal lamellae is identical with that in the Heterocoeniidae. PLATE 21 Fig. 1 Tubicora aylmeri (Gregory, 1900), holotype, BMNH, R. 5032. cross view of colony, polished surface, Upper Paleocene-Lower Eocene of Somalia, scale bar: 5 mm. Fig. 2 Euphyllia calyculata (Catullo, 1847) (holotype of Plocophyllia karstica Turnšek & Drobne, 1998), SAZU, DvW-23B (981a), Montian of Slovenia. 2a, cross thin section of corallite, scale bar: 4 mm. 2b, cross thin section of colony, scale bar: 4 mm. Fig. 3 Agathelia asperella Reuss, 1854, BMNH, AZ 895, cross thin section, Middle-Upper Maastrichtian of the UAE/Oman border region, scale bar: 4 mm. Fig. 4 Multicolumnastraea cyathiformis (Duncan, 1865), NMNH, Coates coll., no. 534d, cross thin section, Maastrichtian of Jamaica, scale bar: 2 mm. Fig. 5 Euphyllia calyculata (Catullo, 1847), topotype, GBA, 1874/3/56, Oligocene of Italy. 5a, upper surface, cross view, scale bar: 4 mm; 5b, longitudinal view, scale bar: 7 mm. Fig. 6 Cladophyllia cf. furcifera Roemer, 1888, NMNH, Coates coll., no. J-71-34r, cross thin section of corallite, Maastrichtian of Jamaica, scale bar: 2.5 mm. Fig. 7 Cladophyllia stewartae Wells, 1944, BMNH, AZ 575, Middle-Upper Maastrichtian of the UAE/Oman border region. 7a, close-up of corallite in budding stage, scale bar: 2 mm; 7b, close-up of corallite showing ‘septal division’,scale bar: 2 mm; 7c, cross thin section of colony, scale bar: 4 mm


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199

Genus Agathelia Reuss, 1854 Type species. Agathelia asperella Reuss, 1854, Upper Santonian of Austria (Neffgraben, Gosau Group).

Diagnosis. Colonial, massive, plocoid. Gemmation extracalicinal. Costosepta compact, radially or bilaterally arranged in 6 systems. Stunted septa present. Distal septal margin covered with small subequal denticles. Septal flanks ornamented with spiniform granulae aligned in a row that lies perpendicular to the distal margin. Columella feebly developed, parietal-spongy to lamellar. Endothecal dissepiments vesicular to subtabulate. Exothecal dissepiments vesicular, abundant. Septothecal wall with stunted septa. Wall covered by concentric perithecal lamellae. Perithecal sheets with granulate surface and separated by vesicular dissepiments. Microstructure of perithecal lamellae is identical with that in Heterocoeniidae.

Agathelia asperella Reuss, 1854 Pl. 21, fig. 3 v*1854 v1903a 1926 1937a v1975 1987 v1982 v1992 1994 v1997 2002 v2000 v2002 2003 v2003 v2004

Agathelia asperella m: Reuss, p. 82, pl. 9, figs 10–12 (topotypes and neotype studied). Agathelia asperella Reuss 1854: Felix, p. 262, text-figs 30, 32. Agathelia urgonica n. sp.: Dietrich, p. 75, pl. 5, fig. 1, pl. 8, fig. 2. Agathelia asperella Reuss 1854: Bataller, p. 141. Agathelia asperella Reuss: Beauvais & Beauvais, p. 567, text-fig. 1, fig. 2, text-fig. 2, figs 1, 2. Agathelia asperella Reuss, 1854: Kuzmicheva, p. 81, pl. II, fig. 1. Agathelia asperella Reuss 1854: Beauvais, vol 1, p. 44, pl. 61, fig. 7, pl. 62, figs 1, 2. Agathelia asperella Reuss, 1854: Eliàšovà, p. 405, pl. 6, fig. 1. Agathelia asperella Reuss: Liao & Xia, p. 69, pl. 5, text-figs 43a–c. Agathelia asperella Reuss, 1854: Baron-Szabo, p. 35, pl. 1, figs 1, 3, 5 (older synonyms cited therein). Agathelia asperella, Reuss 1854: Löser et al, p. 67. Agathelia asperella Reuss, 1854: Baron-Szabo, p. 108, pl. 4, fig. 4. Agathelia asperella Reuss, 1854: Baron-Szabo, p. 187, pl. 129, figs 3–5, pl. 130, figs 1–5. Agathelia asperella Reuss 1854: Götz, p. 5ff., pl. 1, fig. 3. Agathelia asperella Reuss, 1854: Baron-Szabo, p. 118, pl. 3, figs 2, 5, 6, pl. 4, figs 2, 7, 8. Agathelia asperella Reuss: Moosleitner, p. 174, 179, and 185, pl. 72, fig. 2, pl. 77, figs 2, 3, pl. 83, figs 1, 3.

Dimensions. d=3.5–6 mm; d (juvenile)=1.5–2.5 mm; dl=2.5–4.5 mm; c-c=3–5.5 mm, in areas of intense budding distance is 2.5 mm; s=24–48 +s5; s (juvenile)=16–22. Description. Massive, plocoid colony; calices often projecting, subcircular or elliptical in outline; costosepta arranged in 3 to 4 cycles in 6 systems; in some calices the beginning of a fifth cycle is present; S1 and S2 nearly equal in length and thickness, slightly alternating, extending to corallite centre; inner ends of S3 or S4 cuneiform or stretch out forming auriculae; columella lamellar to irregularly trabecular; wall is septothecal, in places parathecal. Remarks. Some authors have merged the species Stylina turbinata Trautschold, 1886, from the Hauterivian of Ukraine, which was transferred to the genus Agathelia Reuss by Bendukidze (1961), with the species Agathelia asperella Reuss. While the form Stylina turbinata Trautschold shows close resemblance to the genus Agathelia Reuss, its specific characteristics clearly differ from A. asperella Reuss. According to the original description and illustration by Trautschold (1886, p. 124, pl. III, figs 1a–b), the form A. turbinata is characterized by a corallite diameter ranging from 5–6 mm and 24 septa, which seem to be arranged in 12 sys-


BARON-SZABO

tems. In addition, in some corallites the number of septa appears to be distinctly smaller (ranging around 12). In A. asperella the septa are arranged in 6 systems and even in the most juvenile corallites the number of septa is larger than 12. Moreover, in corallites that are 5–6 mm in diameter the number of septa is often twice as large as in A. turbinata. Therefore, the latter species is excluded from the synonymy of A. asperella Reuss. Type locality of species. Santonian of Austria (Gosau Group at Neffgraben). Distribution. Berriasian-Valanginian of Tibet, Valanginian-Hauterivian of Crimea, Lower Aptian of Tanzania, Albian-Cenomanian of Tibet, Upper Cretaceous of Slovakia, Upper Cenomanian-Lower Turonian of the Czech Republic, (?Upper Turonian-) Lower Coniacian-Campanian of Austria (Gosau Group), Santonian of Armenia, Upper Santonian of Germany, Campanian-Maastrichtian of Spain, Middle-Upper Maastrichtian of the UAE/Oman border region.

Genus Multicolumnastraea Vaughan, 1899 Type species. Heliastraea cyathiformis Duncan, 1865, Campanian of Jamaica (Trout Hall) (see Duncan & Wall 1865).

Diagnosis. Colonial, massive, columnar, lamellar, plocoid. Gemmation extracalicinal. Costosepta compact. Septal margins regularly dentate. Costae rarely extend over the coenosteum. Columella formed by 3 or 4 pillars. Pali in front of first and second septal cycles. Wall septothecal. Endotheca well-developed, consisting of fine and densely packed dissepiments. Perithecal wall vesicular. Auriculae present before septal cycles younger than S2.

Multicolumnastraea cyathiformis (Duncan, 1865) Pl. 21, fig. 4 1865 v*1865 1867 1867 v1899 1919 1934 1956 v1960 ?1997 1994 v1998 2000 2000 2002 v2002 v2003 (v)2005 v2006

Heliastraea exsculpta: Duncan, in Duncan & Wall, p. 7, 8, 11. Heliastraea cyathiformis: Duncan, in Duncan & Wall, p. 7, 8, Pl. 1, figs 1a, b. Heliastraea exsculpta Duncan: Duncan, vol. 4, p. 24. Heliastraea cyathiformis Duncan: Duncan, vol. 4, p. 24. Multicolumnastraea cyathiformis (Duncan): Vaughan, p. 236, pl. 37, fig. 5, pl. 38, fig. 1. Multicolumnastraea cyathiformis (Duncan): Vaughan, p. 194. Multicolumnastraea cyathiformis (Duncan): Wells, p. 90. Multicolumnastraea cyathiformis (Duncan): Wells, p. F 406, fig. 302.3. Multicolumnastraea cyathiformis Duncan: Berryhill, et al., p. 151. Promadracis ramosus n. sp.: Reig Oriol, p. 12, pl. 1, fig. 7. Multicolumnastraea cyathiformis (Duncan): Liao & Xia, p. 177, pl. 53, figs 5–8. Multicolumnastraea cyathiformis (Duncan): Baron-Szabo, p. 131, pl. 6, fig. 4. Multicolumnastraea cyathiformis (Duncan & Wall 1865): Löser, p. 55. Promadracis ramosus, Reig Oriol 1997: Löser, p. 69. Multicolumnastraea cyathiformis (Duncan ): Mitchell, p. 6 ff., table 1. Multicolumnastraea cyathiformis (Duncan, 1865): Baron-Szabo, p. 189, pl. 131, figs 1–6. Multicolumnastraea cyathiformis (Duncan, 1865): Baron-Szabo, p. 119, pl. 3, fig. 7. Multicolumnastraea cyathiformis (Duncan, 1865): Filkorn et al., p. 118, fig. 2b. Multicolumnastraea cyathiformis (Duncan, 1865): Baron-Szabo et al., p. 12, fig. 4.3.

Dimensions. dl (adult)=1.5–2 mm; d (adult)=1.8–2.5 (3) mm; d (juvenile)=up to about 1 mm; c-c=2–3.5 mm, SCLERACTINIAN CORALS OF THE K/T-BOUNDARY


201

in regions of intensive budding the corallite distance may be smaller; s=24; ramose colonies= up to 15 cm in length and 5 cm in diameter; massive colonies=up to 8 cm in diameter; columniform colonies= up to 30 cm in length with branches that are 0.8–3 cm in diameter. Description. Colonial, columniform, ramose, or irregularly massive, plocoid; calices circular in outline; gemmation extracalicinal; costosepta nonconfluent or rarely subconfluent, compact, arranged in three cycles in 6 systems; septal anastomose frequently; costae strongly granulated and united by numerous subtabulate dissepiments; lateral surfaces of septa ornamented with spiniform granules or vertical carinae; paliform lobes sparse; S1 and S2 nearly equal, extending to the columella; S3 alternate in length with them; intercorallite areas crossed by costae when corallites close together; when more distant, costae merge into a porous, reticulated coenenchyme; columella made of isolated or joined papillae; wall septoparathecal to septothecal, rarely synapticulothecal; endotheca dissepiments thin, tabulate or vesicular; exothecal dissepiments tabulate or arched, well-developed. Type locality of species. Campanian of Jamaica (Trout Hall). Distribution. Santonian of Austria (Gosau Group), Campanian-Maastrichtian (new material) and Eocene of Jamaica, Campanian-Maastrichtian of Tibet; Campanian-?Maastrichtian of northern Spain (Catalonia), Maastrichtian of Mexico (Ocozocuautla and Cardenas Formations), Danian of Puerto Rico. New Material. Maastrichtian of Jamaica, NMNH, Coates, coll., sample nos.: 317a; 324; 327; 447; 460; 465-II; 469f-I; 485o; 485q; 485r; 485s; 490d; 491a-I; 491b; 516h; 493a; 497n; 497o; 497r; 497s; 497t; 497u; 497v; 497w; 497x; 497z; 498; 500d; 500e; 500f; 503; 506b; 508; 515e; J-71-20j; J-71-20k; J-71-20L; J-7120m; J-71-20n; J-71-20o; J-71-20p; J-71-20q; J-71-20r; J-71-20s; J-71-20t; J-71-20u; J-71-20v; J-71-20w; J71-20x; J-71-20y; J-71-20z (=Ducketts Land Settlement); 370c; 381; 387b; 403; 431d-II (=Jerusalem Mountain Inlier); 440m (=Catadupa); 532d; J-71-13v3; J-71-13w3; J-71-13x3; J-71-13y3; J-71-13z3; J-71-13a4; J71-13b4; J-71-13c4; J-71-13d4 (=Vaughnsfield); 533i; 533k; 533L; 533q; 533s; 533t-II; 534b (=Shaw Castle, Maldon Formation); 569c; 569g; 569h; 569i; 569j; 569k; 569L; 570i; 570t; 570u; 585b; 585e; 589a; 589c; 591e; Jag 18.9f; Jag 18.9g; Jag 18.9h; Jag 18.9i; Jag 18.9j; Jag 18.9k; Jag 18.9L; Jag 18.9m; Jag 18.9n (=probably Cambridge railway area); 578; -71-34q; J-71-103b; J-71Ab; J-71Ac; J-71Ad; J-71Ae; J-71Af (=Rio Minho).

Family Cladophylliidae Morycowa & Roniewicz, 1990 Diagnosis. Colonial, phaceloid. Radial elements of septal type. Trabeculae branched, arranged in series. Diameter of main trabeculae from 50 to 90 µm. Thin secondary trabeculae expressed on septal surfaces in the form of sharp granulae. Inner septal edge ornamented with auriculae. Septotheca formed by well-developed and abortive septa. Columella essential. Intratentacular budding through symmetrical division by forming septal wall. Remarks. The authors of the family Cladophylliidae Morycowa and Roniewicz, 1990, consider the phaceloid growth form in Cladophyllia and Apocladophyllia a pseudocolonial development. However, because the individual polyps most likely are of the same genetical composition (as a result of intratentacular budding) and because the polyps throughout their entire growth remain a colony physically (branches develop from one first polyp and stay together like branches in a tree), the more traditional interpretation of phaceloid integration of the polyps as a colonial form is followed here.


BARON-SZABO

Genus Cladophyllia Milne Edwards & Haime, 1851b Type species. Lithodendron dichotomum Goldfuss, 1826, Upper Jurassic of Germany (Giengen) (see Milne Edwards & Haime, 1851b); subsequent designation by Wells, 1933.

Diagnosis. Colonial, phaceloid. Corallites laterally free, temporarily fused with walls. Gemmation by septal division with succeeding dichotomic forking of corallites. Symmetry radial or radiobilateral. Corallites subcircular in cross section. Calicular edge sharp. Septa nonexsert. Septal faces with small and sharply pointed granulae. Inner edge with regular, auricular denticles. Interseptal anastomosis present. Columella essential, trabecular. Endotheca composed of tabulate dissepiments and incomplete ring of large peripheral dissepiments. Septotheca thick with transversely wrinkled surface. Trabeculae between 30 and 80 µm in diameter.

Cladophyllia cf. furcifera Roemer, 1888 Pl. 21, fig. 6 v*1888 1909 1914 1928 1933 1990 v1993 2002 2002

Cladophyllia furcifera n. sp.: Roemer, p. 8, pl. 1 (31), figs 4a, 4b (topotypes studied). Cladophyllia furcifera Roemer: Grabau & Shimer, vol I, p. 100, text-fig. 162. Cladophyllia furcifera Roemer 1888: Felix, pars 5, p. 41. Cladophyllia furcifera Roemer 1888: Adkins, p. 24. Cladophyllia furcifera Roemer 1888: Wells, p. 172, pl. 8, figs 5–8. Cladophyllia furcifera Roemer, 1888: Morycowa & Roniewicz, p. 168. Cladophyllia furcifera Roemer, 1888: Baron-Szabo, p. 160, pl. 4, fig. 3, text-fig. 6. Cladophyllia furcifera, Roemer 1888: Löser et al., p. 144 (older synonmys cited therein). Cladophyllia furcifera Roemer, 1888: Baron-Szabo, p. 181.

Dimensions. d=4–7 mm, juvenile around 2.5 mm; s=32–48 (+s), juvenile 24. Description. Phaceloid corallum; corallites circular in outline; costosepta compact, straight; S1 and some of S2 reach center of calice, where their inner ends may fuse with trabecular or substyliform columella; remaining septa alternate in length and thickness; anastomoses common; wall septothecal. Remarks. According to Morycowa & Roniewicz (1990: 168) as well as based on the study of topotypes by the author, Cladophyllia furcifera Roemer has 3 cycles of septa in 6 systems, with the beginning of a fourth one. However, in the Jamaican specimens the number of septa can be slightly larger, corresponding to 4 complete cycles in 6 systems, but otherwise they agree well with Cladophyllia furcifera Roemer, 1888. Type locality of species. Albian of Texas. Distribution. Upper Aptian-Lower Albian of Mexico and northern Spain, Albian of Texas, Maastrichtian of Jamaica (this paper). New Material. Maastrichtian of Jamaica, NMNH, Coates, coll., sample nos.: 409; 417, (=Jerusalem Mountain Inlier); 553; J-71-34r; J-71-34s; J-71-34u; J-71-34v; J-71-34w; J-71-34x; J-71-34y; J-71-34z (=Rio Minho); J-66-43b (=Vaughnsfield).

Cladophyllia stewartae Wells, 1944 Pl. 21, figs 7a–c v*1944

Cladophyllia stewartae, new species: Wells, p. 442, pl. 74, figs 2–3.



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1982 1990 v2000 2002 v2002

Cladophyllia stewartae Wells, 1944: Kuzmicheva, p. 103, pl. 1, fig. 4. Cladophyllia stewartae Wells, 1944: Morycowa & Roniewicz, p. 168. Cladophyllia stewartae Wells, 1944: Baron-Szabo, p. 110, pl. 4, fig. 3, pl. 6, fig. 3. Cladophyllia stewartae, Wells 1944: Löser et al., p. 145 (older synonyms are cited therein). Cladophyllia stewartae Wells, 1944: Baron-Szabo, p. 181, pl. 126, fig. 1.

Dimensions. d=3–4 mm, juvenile up to 2.5 mm; s=24 (+s). Description. Phaceloid corallum; corallites diameter 3 mm on average; budding is due to ‘septal division’ (sensu Morycowa & Roniewicz, 1990) and extracalicinal; costosepta compact, straight, with lateral surfaces that are covered by sharply pointed or rounded granules, arranged in 3 complete cycles in 6 systems. Frequently, the beginning of a fourth cycle is present. Septa of the first two cycles reach the centre of the calice, where their inner ends may fuse. A trabecular, styliform, or sublamellar columella can be free or joined with oldest septa. Septa of the third cycle are distinctly thinner, reaching about three-quarters the length of the oldest ones. Youngest septa are very small and thorn-like. Anastomosis can be observed frequently. The wall is septothecal. Endotheca is made of tabulate dissepiments in the axial part of the corallite. In the peripheral region of the calice large vesicular dissepiments occur. Septal microstructure consists of simple minitrabeculae, forming dark axial lines. Remarks. In having extracalicinal gemmation and slightly exsert septa the specimens described from the Maastrichtian of the UEA/Oman border region by Baron-Szabo (2000) show characteristics of the genus Stylosmilia. However, in some corallites budding due to ‘septal division’ seems to be present, indicating that the specimens belong to the genus Cladophyllia. Type locality of species. Barremian-Aptian of Venezuela. Distribution. Barremian-Aptian of Venezuela, Aptian of Spain, Tibet, and central Uzbekistan, AlbianCenomanian of Mexico, Middle-Upper Maastrichtian of the UAE/Oman border region.

Addendum to Corals of the K/T-Boundary: Scleractinian corals of the suborders Astrocoeniina, Faviina, Rhipidogyrina, and Amphiastraeina (Baron-Szabo, 2006)

Suborder Faviina Vaughan & Wells, 1943 (=Astraeoina Alloiteau, 1952a; =Meandriina Alloiteau, 1952a)

Family Faviidae Gregory, 1900 (=Hemiporitidae Alloiteau, 1952a, p.p.) Genus Phacellocoenia Alloiteau & Tissier, 1958 Type species. Phacellocoenia bazerquei Alloiteau & Tissier, 1958, Upper Danian of France (Haute-Garonne).

Diagnosis. Colonial, phaceloid to subfasciculate. Gemmation intracalicinal and extracalicinal. Costosepta compact to subcompact, thin, finely granulated laterally. Columella trabecular, styliform-sublamellar in shape. Endothecal dissepiments thin, vesicular. Wall parathecal. Epithecal-epicostal wall present or absent.


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Phacellocoenia thomkai (Eliášová, 1991) Text-Fig. 36 v*1991

Rhabdophylliopsis thomkai sp. n.: Eliášová, p. 50, pl. 55, figs 1, 2, pl. 56, figs 1–3, pl. 58, fig. 3.

Dimensions. d (monocentric)=7–11 mm; d (max, in budding stage)=16 mm; c-c=10–20 mm; s (monocentric adult calices)=up to 48+s5. Description. Phaceloid-subfasciculate corallum; costosepta developed in 4 complete cycles with the beginning of a fifth cycle in adult corallites; endothecal dissepiments abundant throughout corallite. Remarks. In forming a phaceloid to subfasciculate colony with polyps that multiply by intra- and extracalicinal (marginal) budding, and in having mainly compact septa that are subcompact in places, a trabecular columella, and an epicostal wall, the material from the Upper Paleocene of the Czech Republic closely corresponds to the genus Phacellocoenia Alloiteau & Tissier. The genus Rhabdophylliopsis Alloiteau & Tissier, 1958, to which the Czech materal was originally assigned, shows all characteristics of Sideroseris Wells (see Remarks and illustrations of Sideroseris above). Type locality of species. Upper Paleocene of the Czech Republic. Distribution. Upper Paleocene of the Czech Republic.

Text-Fig. 36 Phacellocoenia thomkai (Eliášová, 1991), holotype SNM/Z 20655, Upper Paleocene of the Czech Republic. A, cross view, scale bar: 10 mm; B, longitudinal view, scale bar: 2 mm (photographs courtesy H. Eliášová).



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Family Mussidae Ortmann, 1890 Diagnosis. Solitary and colonial, hermatypic. Colony formation by intratentacular budding. Centres linked by lamellae or trabeculae. Wall septothecal or parathecal. Septa endocoelic, formed by several fan systems of large, simple trabeculae, each fan system producing a lobulate dentation. Endothecal dissepiments well developed. Columella trabecular.

Genus Circophyllia Milne Edwards & Haime, 1848 (=Ghirobocyathus Barta-Calmus, 1969, Type species. Ghirobocyathus lagaensis Barta-Calmus, 1969, Paleocene of Ivory Coast). Type species. Anthophyllum truncatum Goldfuss, 1826, Eocene of France.

Diagnosis. Solitary, turbinate to trochoid, or bowl-shaped to subpatellate; fixed or free; costosepta compact to fenestrate, covered with numerous rounded and pointed granules; columella spongy-papillose; endothecal dissepiments vesicular, abundant throughout the whole corallum, forming concentric rings parallel to corallite wall; wall septothecal to parathecal; epithecal wall present or absent. Remarks. Barta-Calmus (1969) stated that her newly created genus Ghirobocyathus from the Paleocene of Ivory Coast was characterized by a bowl-shaped corallum that has compact septa, the absence of pali, presence of a papillose columella, numerous endothecal (vesicular) dissepiments that occur throughout the corallum and form concentric rings parallel to the wall, and no epithecal but rather a parathecal wall. In addition, she included specimens that are also turbinate (see fig. 14 on Pl. 1 in Barta-Calmus, 1969, showing topotype). Moreover, because the holotype of Ghirobocyathus (as shown on fig. 15 on Pl. 1 in Barta-Calmus, 1969) seems to clearly show both compact and fenestrate septa, it very closely corresponds to the genus Circophyllia. Therefore, its synonymy with the latter is proposed.

Circophyllia truncata (Goldfuss, 1826) Text-Fig. 37 v*1826 1925 1969

Anthophyllum truncatum nobis: Goldfuss, p. 46, pl. 13, fig. 9. Circophyllia truncata Goldfuss sp. 1826: Felix, pars 28, p. 53 (older synonyms cited therein). Ghirobocyathus lagaensis n. g., n. sp.: Barta-Calmus, p. 827, pl. 1, figs 12–15.

Dimensions. d (max)=13.5–28 mm; d (min)=13–25 mm; d (min)/d (max)=0.85–0.96; s=96 (in corallite diameter of 13 mm and larger); h=up to 22 mm. Description. Bowl-shaped to turbinate-subpatellate corallum; costosepta arranged in 5 cycles in 6 systems in mature corallites. Remarks. The specimens assigned to Ghirobocyathus lagaensis by Barta-Calmus (1969) show 96 septa in corallites with diameters ranging between 12 and 20 mm. In the holotype of the species C. truncatum 96 septa are present when the corallum has reached a diameter of around 15 mm. Therefore, the synonymy of the two species is assumed. Type locality of species. Eocene of France. Distribution. Paleocene of Ivory Coast, Eocene of Italy, Eocene-Oligocene of France.


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Text-Fig. 37 Circophyllia truncata (Goldfuss, 1826). A, B, holotype, Goldfuss coll., no. 161, Eocene of France. A, longitudinal view of corallum; B, cross view. C, D, as figured in Barta-Calmus (1969, as holotype of Ghirobocyathus lagaensis Barta-Calmus, 1969), Paleocene of Ivory Coast. C, longitudinal of corallum; D, cross view; scale bar: 10 mm.

Family Meandrinidae Gray, 1847 Subfamily Euphyllinae Alloiteau, 1952 Genus Euphyllia Dana, 1846 (=Plocophyllia Reuss, 1868; Type species. Lobophyllia calyculata Catullo, 1852, Oligocene of Italy [Monte Grumi]). Type species. Caryophyllia glabrescens Chamisso & Eysenhardt, 1821, Recent, Pacific Ocean.

Diagnosis. Colonial, phacelo-flabellate. Gemmation intracalicinal, di- to polystomodaeal. Calicinal series laterally free. Costosepta compact, covered by spiniform and rounded granules. Columella absent but trabecular extensions of septal axial ends may form a pseudo-columella. Endothecal dissepiments vesicular, abundant. Wall septothecal and parathecal.

Euphyllia calyculata (Catullo, 1852) Pl. 21, figs 2a, b, 5a, b v*1852 v1856 v1868 v1868

Lobophyllia caliculata: Catullo, p. 28, pl. 2, fig. 7 (topotypes studied). Lobophyllia calyculata: Catullo, p. 52, pl. 4, fig. 7 (topotypes studied). Plocophyllia calyculata Catullo sp.: Reuss, p. 145, pl. 3, figs 1–5b. Plocophyllia constricta nov. sp.: Reuss, p. 146, pl. 3, figs 6a, b, pl. 4, fig. 1.



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v1868 v1872 1880 v1973 v1988 1993 v1998

Plocophyllia flabellata nov. sp.: Reuss, p. 146, pl. 4, figs 2a, b. Plocophyllia caespitosa nov. sp.: Reuss, p. 43, pl. 50, figs 2–3b, pl. 51, figs 1a–c. Plocophyllia flabellata, Reuss: Duncan, p. 39, pl. 16, figs 5, 6. Plocophyllia calyculata (Catullo) 1847: Barta-Calmus, p. 372, pl. 32, figs 1–10 (older synonmys cited therein). Plocophyllia karstica n. sp.: Drobne et al., p. 187, pl. 31, figs 1–3, pl. 32, figs 1–3. Plocophyllia calyculata (Catullo) 1847: Alvarez Perez, p. 202, pl. 16, figs 7, 8. Plocophyllia karstica n. sp.: Turnšek, in Turnšek & Drobne, p. 136, pl. 6, fig. 1.

Dimensions. d (series, min.)=3–11 mm; d (series, max.)=13–35 mm; s/mm=18–30/10. Description. Phacelo-flabellate colony with corallites arranged in short series of up to 6 calices; monocentric corallites compressed; septa straight, S1–S3 nearly equal in thickness, often regularly alternate in length; according to their length 4 septal orders can be distinguished. Remarks. In the type material of E. karstica from the Montian of Slovenia, the wall seems to be parathecal and the number of endothecal dissepiments appear to be much more numerous than in E. calyculata. However, in several places the presence of a septothecal wall can be seen in E. carstica (see Plate 21, Fig. 2b, upper margin of central corallite series). In addition, in the upper surface view of the colony of E. calyculata the presence of numerous vesicular dissepiments, which correspond to the endothecal development in E. carstica, can be seen (Plate 21, Fig. 5a, e.g., lower right corallite and corallite on the left). Therefore, their synonmy is suggested. Type locality of species. Oligocene of Italy (Monte Grumi). Distribution. Upper Danian of Pakistan, Montian of Slovenia, Middle-Upper Eocene of Spain (Catalonia), Oligocene of Italy.

Acknowledgements My thanks and gratitude go to my husband Dennis, to colleagues and friends for their encouragement and support. I am very grateful to Dragica Turšek (Ljubjana) for providing very valuable comments. For many discussions on coral taxonomy and/or stratigraphic questions of type localities of corals I wish to thank Jacob Leloux (Leiden, Netherlands), Steve Cairns (Smithsonian Institution, Washington, DC), and Wolfgang Kiessling (Natural Museum of the Humboldt University, Berlin). Type material and material from new localities were made accessible to me by Jill Darrell (The Natural History Museum, London, UK), Martin Sander (University of Bonn, Germany), Christine Perrin (Museum d'Histoire Naturelle de Paris, France), Dieter Korn and Carsten Lüter (Natural History Museum of the Humboldt University, Berlin), Franz Stojaspal, Gerhard Schubert, and Harry Lobitzer (Geologische Bundesanstalt, Wien, Austria), Heinz Kollmann (Naturhistorisches Museum, Vienna, Austria), Winfried Werner and Martin Nose (Bayerische Staatssammlung Munich, Germany), Vivianne Berg-Madsen (Museum of Evolution, Uppsala, Sweden), Christina Franzén-Bengtson (Rijksmuseum, Stockholm, Sweden), Joan Vilella i Puig (Museu Geològico del Seminari de Barcelona, Spain), Fred Collier (Museum of Comparative Zooogy, Harvard University, USA), Miguel Tellez (Universidad Autónoma de Baja California, Mexico), Jacob Leloux (Leiden, Netherlands), Christina Ifrim and Wolfgang Stinnesbeck (both Heidelberg, formerly at the Geological Institute, University Karlsruhe, Germany), Riccardo Manni (University of Rome), and Chiara Sorbini and Giovanni Bianucci (both University of Pisa, Italy). As a Research Associate of the Smithsonian Institution, Washington, DC, and Honorary Research Member of the Senckenberg Research Institute, Frankfurt/Main, Germany, the author would like to express her deep appreciation for the continuing support of both insitutions. I am very indebted to the Deutsche Forschungsgemeinschaft (DFG) for providing support by funding project Ba1830/3.


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References

Abdel-Gawad, G.L. & Gameil, M. (1995) Cretaceous and Palaeocene coral fauna in Egypt and Greece (1). Geology. Coral Research Bulletin, 4, 1–36, 21 pls. Abed, M.M. & El Asa’ad, G.M. (1981) Campanian-Maastrichtian scleractinian corals from central Saudi Arabia. Bulletin of the Faculty of Science, Mansoura University, 8, 271–295. d’Achiardi, A. (1867) Corallari fossili del terreno nummulitico dell'Alpi Venete, Catalogo delle specie e brevi note. Memorie della Societa Italiana di Scienze Naturali, 2, 1–18. d’Achiardi, A. (1875) Coralli eocenici del Friuli. Atti della Societa Toscana di Scienze Naturali resindente in Pisa, 1, 67–86, 2 pls. d’Achiardi, A. (1881) Coralli fossili di Asolo. Proc verb della Societá Toscana di Scienze Naturali. Adunanza del di 8 maggio 1881, 239–242. Adkins, W.W. (1928) Handbook of Texas Cretaceous fossils. University of Texas Bulletin, 2832, 74–77. Alloiteau, J. (1936) Polypiers fossiles de Madagascar. 1, Formes du Crétacé de la province d’Ananalava. Annales Géologiques du Service des Mines de Madagascar, 6, 41–53. Alloiteau, J. (1939a) Polypiers récoltés par M.P.Sénesse dans le Santonien de la Jouane, Commune de Sougraigne (Aude). Bulletin de la Société géologique de France (5), 9, 3–21, 1 pl. Alloiteau, J. (1939b) Deux espèces nouvelles de polypiers d'anthozoaires de l'Anatolie septentrionale. Bulletin scientifique de Bourgogne, 9, 5–11, 1 pl. Alloiteau, J. (1941) Révision de collection H. Michelin. Polypiers d’anthozoaires (1). Crétacé. Mémoires du Muséum National d’Histoire Naturelle (N.S.), 16, 1–100, pls. 1–19. Alloiteau, J. (1949) Les coraux de l'Éocène de Bojnice-les–Bains près de Prievidza dans les Karpates Slovaques. Práce atátneho heologického ústavu, 24, 1–30, 8 pls. Alloiteau, J. (1951a) Coralliaires. In: Collignon, M. (Ed.). Faune maestrichtienne de la cοte d’Ambatry (province de Betioky). Annales Géologiques du Service des Mines de Madagascar, 19, 47–49, pl. 8. Alloiteau, J. (1951b) Coralliaires. In: Collignon, M. (Ed.). Le Crétacé supérieur d’Antonibe. Annales Géologiques du Service des Mines de Madagascar, 19, 80–83 and 174, pl. 13. Alloiteau, J. (1952a) Embranchment des Coelentérés. II. Madréporaires post-paléozoiques. In: Piveteau, J. (Ed.), Traité de Paléontologie, 1, Masson, Paris, pp. 539–684. Alloiteau, J. (1952b) Sur des polypiers de Sénégal. Bulletin de la direction des mines, 14, 9–18. Alloiteau, J. (1957) Contribution à la systématique des Madréporaires fossiles. Thèse, Centre National Recherche Scientifique, Paris, 462 pp, Alloiteau, J. (1958) Monographie des Madréporaires fossiles de Madagascar. Annales Géologiques de Madagascar, 25, 1–118, 38 pls. Alloiteau, J. (1960) Nouveaux polypiers du cretacique d’Espangne. Anales de la Escuela Técnica de Peritos Agricolas y de Especialidades Agropecuarias y de los Servicios Técicos de Agricultura, 14, 80–120. Alloiteau, J. & Tissier, J. (1958) Les Madréporaires du Montien des Petites Pyrénées. Bulletin de la Societe d’Histoire Naturelle de Toulouse, 93, 243–291. Alvarez Perez, G. (1993) Cnidaria fòssils de la conca d’Igualda. Thesis. Department de Geologia Dinàmica, Geofisica i Paleontologia, Universitat de Barcelona, 261 pp, 23 pls. Angelis d’Ossat, G. de (1905) Coralli del Cretacico inferiore della Catalogna. Palaeontographica Italica, 11, 169–251. d‘Archiac, V.E.J.A.D. & Haime, J. (1853) Description des animaux fossiles du groupe nummulitique de l’Inde. Préecedée d’un réesumé géologique et’ dun monographie des nummulites. Gide & J. Baufdry, Paris, 373 pp., 36 pls. Baceta, J.I., Pujalte, V. & Bernaola, G. (2005) Paleocene coralgal reefs of the western Pyrenean basin, northern Spain, New evidence supporting an earliest Paleogene recovery of reefal ecosystems. Palaeogeography, Palaeoclimatology, Palaeoecology, 224,117–143. Baron-Szabo, R.C. (1993) Korallen der höheren Unterkreide ("Urgon") von Nordspanien (Playa de Laga, Prov. Guernica). Berliner geowissenschaftliche Abhandlungen (E), 9, 147–181. Baron-Szabo, R.C. (1997) Zur Korallenfazies der ostalpinen Kreide (Helvetikum, Allgäuer Schrattenkalk; Nördliche Kalkalpen, Brandenberger Gosau), Taxonomie, Paläökologie. Zitteliana, 21, 3–98. Baron-Szabo, R.C. (1998) A new coral fauna of the Campanian from north Spain (Torallola village, Prov. Llèida). Geologisch-Paläontologische Mitteilungen Innsbruck, 23, 127–191. Baron-Szabo, R.C. (1999) Taxonomy of Upper Cretaceous scleractinian corals of the Gosau Group (Weissenbachalm, Steiermark, Austria). In: Lobitzer, H. & Grecula, P. (Eds.), Geologie ohne Grenzen. Abhandlungen der geologischen



209

Bundesanstalt Wien, Festschrift 150 Jahre Geologische Bundesanstalt, 56, 441–464. Baron-Szabo, R.C. (2000) Late Campanian-Maastrichtian corals from the United Arab Emirates-Oman border region. Bulletin of The Natural History Museum, London (Geology), 56, 91–131. Baron-Szabo, R.C. (2001) Corals of the Theresienstein reef (Upper Turonian–Coniacian, Salzburg, Austria). Bulletin of the Biological Society of Washington, 10, 257–268. Baron-Szabo, R.C. (2002) Scleractinian corals of the Cretaceous. A compilation of Cretaceous forms with descriptions, illustrations and remarks on their taxonomic position. Published by author, Knoxville, TN, 539 pp. Baron-Szabo, R.C. (2003) Taxonomie und Ontogenie von Korallen der ostalpinen Oberkreide (Hochmoos- und Grabenbachschichten Gosau Gruppe Santon). Jahrbuch der Geologischen Bundesanstalt Wien, 143(2), 107–201. Baron-Szabo, R.C. (2006) Corals of the K/T-boundary. Part 1, Scleractinian corals of the Suborders Astrocoeniina, Faviina, Rhipidogyrina, and Amphiastraeina. Journal of Systematic Palaeontology, London, 4, 1–108. Baron-Szabo, R.C., Casadio, S. & Parras, A. (2004) First shallow water scleractinian coral reef from the Danian, northern Patagonia, Argentina. Ameghiniana, 40, R79. Baron-Szabo, R.C. & González-León, C. M. (1999) Lower Cretaceous corals and stratigraphy of the Bisbee Group (Cerro de Oro and Lampazos areas), Sonora, Mexico. Cretaceous Research, 20, 465–497. Baron-Szabo, R.C. & Steuber, T. (1996) Korallen und Rudisten aus dem Apt im tertiären Flysch des Parnass-Gebirges bei Delphi-Arachowa (Mittelgriechenland). Berliner Geowissenschaftliche Abhandlungen (E), 18, 3–75. Baron-Szabo, R.C., Schafhauser, A., Götz, S. & Stinnesbeck, W. (2006) Scleractinian corals from the Maastrichtian of Mexico (State San Luis Potosí; Cardenas Formation). Journal of Paleontology, 80(6), 1033–1046. Baron-Szabo, R.C., Hamedani, A. & Senowbari-Daryan, B. (2003) Scleractinian corals from Lower Cretaceous deposits north of Esfahan (central Iran). Facies, 48, 199–216. Barrere, P. (1746) Observations sur l’origine et la formation des pierées figurées et sur celles qui, tant extéricurement, qui intéricurement, ont une figure régulière et déterminée, Paris, 67 pp. Barta-Calmus, S. (1973) Révision de collections de madréporaires provenants du Nummulitique du sudest de la France, de l’Italie et de la Yougoslavie septentrionales. Thése a l’Universite de Paris VI, C.N.R.S., A.O. 8295, 1–694, pls. 1–59. Barta-Calmus, S. (1969) Études paléontologiques et géologiques sur les Falaises de Fresco (Côte d’Ivoire). Bulletin du Muséum National d’Histoire Naturelle, 2nd series, 41(3), 817–832. Barthel, K.W. & Herrmann-Degen,W. (1981) Late Cretaceous and Early Tertiary stratigraphy in the Great Sand Sea and its SE margins (Farafra and Dakhla Oases), SW Desert, Egypt. Mitteilungen der Bayerischen Staatssammlung für Paläontologie und historische Geologie, 21, 141–182. Bataller, J. (1936) Contribución al estudio de los poliperos Cretácicos de Cataluña. Ibérica, 1103, 38–46. Bataller, J. (1937a) La fauna corallina del Cretàcic de Catalunya i regions limítrofes. Arxius de l'escola superior d'agricultura (N.S.), 3, 1–299. Bataller, J. (1937b) Primer supplement a la fauna corallina del Cretàcic de Catalunya i regions limítrofes. Arxius de l'escola superior d'agricultura (N.S.), 3, 301–310. Bataller, J. (1945) Enumeración de las especies nuevos del Cretácico de España. Memorias de la Real Academia de Ciencias y Artes de Barcelona (3), 27(11), 1–71. Bauer, J., Kuss, J. & Steuber, T. (2002) Platform Environments, Microfacies and Systems Tracts of the Upper Cenomanian - Lower Santonian of Sinai, Egypt. Facies, 47, 1–26. Beauvais, L. (1981) Sur la Taxinomie des Madréporaires Mésozoiques. Acta palaeontologica Polonica, 25, 345–360. Beauvais, L. & Beauvais, M. (1974) Studies on the world distribution of Upper Cretaceous corals. In: Cameron, A.M., Campbell, B.M., Cribb, A.B., Endean, R., Jell, J.S., Jones, O.A., Mathier, P., & Talbot, F. H. (Eds.), 2nd International Symposium on Coral Reefs, Vol. 1. Association of Australasian Palaeontologists, Brisbane, pp. 475–494. Beauvais, L. & Beauvais, M. (1975) Une nouvelle famille dans le sous-ordre des Stylinida Alloiteau, les Agatheliidae nov. fam. (Madréporaires mésozoiques). Bulletin de la Société géologique de France (7), 17, 576–581. Beauvais, M. (1960) Polypiers sénoniens des environs de Padern (Aude). Bulletin de la Société géologique de France (7), 2/6, 723–727, pl. 21. Beauvais, M. (1964) Revision Madréporaires de forme cyclolitoide des couches de Gosau de la collection F. Quenstedt. Bulletin de la Société géologique de France (7), 6, 535–544, pls. 15–16. Beauvais, M. (1982) Révision systématique des Madréporaires des couches de Gosau (Crétacé supérieur, Autriche). Travaux du Laboratoire de Paléontologie des Invertébrés, 1, 1–256; 2, 1–278; 3, 1–177; 4, atlas, 59 pls; 5, atlas, 131 figs. Bellardi, L.(1850) Monografia delle mitre fossili del Piemonte. Memorie della Reale Accademia delle scienze di Torino,


BARON-SZABO

Serie 2, 11, 34 pp., 2 pls. Bendukidze, N.S. (1956) Upper Cretaceous corals from the Godogani and Udzlouri areas. Trudy Geologicheskogo Instituta Akademiya Nauk Gruzinskoy SSR, (Seriya Geologiya), 9, 79–125. [in Russian]. Bernard, H.M. (1906) The family Poritidae (2). The genus Porites. Porites of the Atlantic and West Indies, with the European fossil forms. The genus Goniopora, a supplement to vol. IV. Catalogue of the Madreporarien in the British Museum (Natural History), 6, 173 pp, 17 pls. Bernecker, M. & Weidlich, O. (1990) The Danian (Paleocene) coral limestone of Fakse, Denmark, a model for ancient aphotic, azooxanthellate coral mounds. Facies, 22, 103–138. Bernecker, M. & Weidlich, O. (2005) Azooxanthellate corals in the Late Maastrichtian–Early Paleocene of the Danish basin, bryozoan and coral mounds in a boreal shelf setting. In: Freiwald, A. & Roberts, J.M. (Eds.), Cold-water Corals and Ecosystems. Springer-Verlag, Berlin-Heidelberg, pp. 3–25. Berryhill, H.L. Jr., Briggs, R.P. & Glover, L. (1960) Stratigraphy, sedimentation, and structure of Late Cretaceous rocks in eastern Puerto Rico: Preliminary report. AAPG Bulletin, 44, 137–155. Blainville, H. M. de (1830) Zoophytes. In: Defrance, J.L.M. (Ed.), Dictionnaire des Sciences naturelles, Vol. 60. Levrault, Paris, pp. 274–364. Blainville, H. M. de (1834) Manuel d’actinologie ou du zoophytologie. Vol. 1–2 Levrault, Paris, 694 pp, 101 pls. Bölsche, W. (1866) Die Korallen des norddeutschen Jura- und Kreide-Gebirges. Zeitschrift der Deutschen Geologischen Gesellschaft, 18, 439–486. Bölsche, W. (1867) Die Korallen des norddeutschen Jura- und Kreide-Gebirges. Thesis, published by author, Bölsche, Braunschweig, 50 pp., 3 pls. Bosellini, F.R & Russo, A. (1995) The scleractinian genus Actinacis: systematic revison and stratigraphic record of the Tertiary species with special regard to Italian occurrences. Revista Italiana di Paleontologia e Stratigrafia, 101, 215–230. Bourne, G.C. (1900) Anthozoa. In: Lankester, E.R. (Ed.), Treatise on Zoology. Vol. 2. A. & C. Black, London, pp 1–84. Bourne, G.C. (1905) Report on the solitary corals collected by Professor Herdman, at Ceylon, in 1902. Ceylon Pearl Oyster Fisheries, 4 (supplement 29), 187–242, pls. 1–4. Bowerbank, J.S. (1840) On the London Clay Formation, Charlesworth's. Magazine of Natural History, New Series, 4, 23 pp. Bronn, H.G. (1837) Lethaea geognostica (1). Schweizerbart, Stuttgart, 544 pp. and atlas. Bryan, J.R. (1991) A Paleocene coral-algal-sponge reef from southwestern Alabama and the ecology of Early Tertiary reefs. Lethaia, 24, 423–438. Bryan, J.R., Carter, B.D., Fluegeman, R.H.Jr., Krumm, D.K. & Stemann, T.A. (1997) The Salt Lake Mountain of Alabama. Tulane Studies in Geology and Paleontology, 30(1), 1–60. Budd, A., Johnson, K.G & Edwards, J.C. (1989) Miocene coral assemblages in Anguilla, B. W. I., and their implications for the interpretation of vertical succession on fossil reefs. Palaios, 4, 264–275. Budd, A., Stemann, T.A. & Stewart, R.H. (1992) Eocene Caribbean reef corals, a unique fauna from the Gatuncillo Formation of Panama. Journal of Paleontology, 66, 570–594. Budd, A., Stemann, T.A & Johnson, K.G (1994) Stratigraphic distributions of genera and species of Neogene to Recent Caribbean reef corals. Journal of Paleontology, 68, 951–977. Cairns, S.D. (1989) A revision of ahermatypic scleractinia of the Phillipine Islands and adjacent waters: Part 1, Fungiacyathidae, Micrabaciidae, Turbinoliinae, Guyniidae, and Flabellidae. Smithsonian Contributions to Zoology, 591, 1–136. Cairns, S.D. (1997) A generic revision and phylogenetic analysis of the Turbinoliidae (Cnidaria, Scleractinia). Smithsonian Contributions to Zoology, 591, 1–55. Cairns, S.D. (2001) A generic revision and phylogenetic analysis of the Dendrophylliidae (Cnidaria, Scleractinia). Smithsonian Contributions to Zoology, 615, 75 pp. Carbone, F., Matteucci, R., Pignatti, J.S. & Russo, A. (1993) Facies analysis and biostratigraphy of the Auradu Limestone Formation in the Berbera-Sheikh area, northwestern Somalia. Geologica Romana, 29, 213–235. Catullo, T.A. (1852) Cenni sopra il terreno di sedimento superiore delle Provincie Venete e descrizione di alcuni polipari fossili ch’esse rachiude. Memorie reale Ist. Veneto Scienze 4, 44 pp. [for 1847–1852]. Catullo, T.A. (1856) Dei terrini di sedimento superiore della Venzie e dei fossili Bryozoari, Antozoari e Spongiari. Angelo Sicca, 1–8, 1–88, pls. 1–19. Chamisso, L.C.A de Boncourt & Eysenhardt, C.W. (1821) De animalibus quibusdam e classe vermium Linneana. Nova Acta Physico–Medica Academiae Leopoldino–Carolinae 10 (2), 345–374, pls. 24–33.



211

Chevalier, J.-P. (1954) Contribution à la révision des polypiers du genre Heliastraea. Annales Hebert et Haug, 8, 105–190, pls. 1–8. Chevalier, J.-P. & Beauvais, L. (1987) Ordre des Scléractiniaires. In: Grasse, P.P. (Ed.), Traité d Zoologie, Cnidaires, Anthozoaires, Masson, Paris, pp. 403–764. Ciry, R. (1939) Étude géologique d’une partie des provinces de Burgos, Palenica, Léon et Santander. Societe d’historie naturelle de Toulouse, 74, 9–301. Clark, B.L. & Martin. G.C. (1901) The Eocene deposits of Maryland. Maryland Geological Survey, The Johns Hopkis Press, Baltimore, 313 pp. Collignon, M. (1931) La faune du Cénomanien à fossiles pyriteux du Nord de Madagascar. Annales de Paléontologie, 20, 41–104, 2 pls. Conrad, T.A. (1847) Observations on the Eocene formation and descriptios of one hundred and five new species of that period, from the vicinity of Vicksburg, Mississippi, with an appendix. Proceeedings of the Academy of natural Science of Philadelphia, 3, 708 pp. Coryell, N.H. & Ohlsen, V. (1929) Fossil corals of Porto Rico, with descriptions also of a few Recent species. New York Academy of Sciences, Scientific Survey or Porto Rico and the Virgin Islands, 3, 167–236. Cottreau, J. (1922). Paléontologie des Madagascar (10). Fossiles Crétacé de la Côre Oriental. Annales de Paléontologie, 11, 111–150. Dainelli, G. (1915) L’Eocene friulano. Monografia geologica e paleontologica. Memorie geografiche Firenze, 55, 1–723, pls. 1–57. Dana, J.D. (1846) Zoophytes. In: United States Exploring Expedition during the years 1838–1842 under the command of Charles Wilkes, U.S.N., Vol. 7. Lea and Blanchard, Philadelphia, 740 pp. Darga, R. (1992) Geologie, Paläontologie und Palökologie der südostbayerischen unter–priabonen (Ober-Eozän) Riffkalkvorkommen des Eisenrichtersteins bei Hallthurm (Nördliche Kalkalpen) und des Kirchbergs bei Neubeuern (Helvetikum). Münchener Geowissenschaftliche Abhandlungen A, 23, 1–66. Dietrich, W.O. (1917) Areopsammia, eine neue eupsammide Koralle aus der obersten Kreide. Sitzungsberichte der Gesellschaft der naturforschenden Freunde zu Berlin, 4, 303–307. Dietrich, W.O. (1926) Steinkorallen des Malms und der Unterkreide im südlichen Deutsch-Ostafrika. Palaeontographica, 1 (supplement 7), 43–62. Drobne, K., Ogorelec, B., Plenicar, M., Zucchi-Stolfa, M. K. & Turnšek, D. (1988) Maastrichtian, Danian and Thanetian beds in Dolenja vas (NW Dinarides, Yugoslavia). Microfacies, foraminifers, rudists and corals. Razprave IV, Razreda SAZU, 29, 147–224. Duncan, P.M. (1863) On the fossil corals of the West Indian Islands. Quarterly Journal of the Geological Society of London, 19, 406–458, pls. 13–16. Duncan, P.M. (1864) A description of, and remarks upon, some fossil corals from Sinde. .Annals and Magazine of Natural History, 13(3), 295–307. Duncan, P.M. 1868. On the fossil corals (Madreporaria) of the West-Indian Islands, Part IV. Conclusions. Quarterly Journal of the Geological Society of London, 24, 9–33. Duncan, P.M. (1869) A monograph of the British fossil corals (2). Corals from the White Chalk, the Upper Greensand, and the Red Chalk of Huntstanton. Palaeontological Society Monographs, 22, 1–26, pls. 1–9. Duncan, P.M. (1873) On the Older Tertiary formations of the West-Indian Islands. Quarterly Journal of the Geological Society of London, 29, 548–565, pls. 19–22. Duncan, P.M. (1880) A monograph of the fossil corals and Alcyonaria of Sind. Memoirs of the Geological Survey of India, Palaeontologia Indica, Series 14, 1, 1–110, pls. 1–28. Duncan, P.M. & Wall, G.P. (1865) A notice of the geology of Jamaica, especially with reference to the district of Clarendon; with descriptions of the Cretaceous, Eocene and Miocene corals of the islands. Quarterly Journal of the Geological Society of London, 21, 1–14, pls. 1–2. Durham, J.W. (1942) Eocene and Oligocene coral faunas of Washington. Journal of Paleontology, 16, 84–104. Durham, J.W. (1943) Pacific coast Cretaceous and Tertiary corals. Journal of Paleontology, 17, 196–202, pl. 32. Durham, J.W. (1949) Ontogenetical stages of some simple corals. University of California Publications, Bulletin of the Department of Geological Sciences, 28/6, 137–172, pls 4, 5, 17. Ehrenberg, C.G. (1834) Die Corallenthiere des Rothen Meeres physiologisch untersucht und systematisch verzeichnet. Köiserliche Akademie der Wissenschaften Berlin, 156 pp. El Asa’ad, G.M.A. (1990) Maastrichtian species of the coral genus Cunnolites from Saudi-Arabia. Journal of African earth sciences, 10(4), 633–642.


BARON-SZABO

Eliášová, H. (1974) Hexacorallia et Octocorallia du Paléogène des Carpates externes. Sbornik geologickych ved Paleontologie, 16, 105–156. Eliasova H. (1976) Nouvelle famille du sous-ordre Amphiastraeina Alloiteau, 1952 (Hexacorallia, Tithonien de Tchécoslovaquie). Vestnik Ústedního ústavu geologického, 51, 177–178, 2 pls. Eliášová, H. (1991) Quelques Scléractiniaires de la Slovaquie (Crétacé et Paléogène, Tchécoslovaquie). Západné Karpaty, (paleontológia), 15, 49–55. Eliášová, H. (1997) Coraux crétacés de Bohême (Cénomanien supérieur; Turonien inférieur-Coniacien inférieur), République tchèque. Vestnik Ceskeho geologickeho ústavu, 72, 245–265. Ellis, J. & Solander, D. (1786) The natural history of many curios and common zoophytes. B. White & Son, London, 208 pp, 63 pls. Etallon, A. (1859) Études paléontologiques sur le Haut-Jura. Rayonnés du Corallien. Mémoires de la Société d'émulation du département du Doubs, 3, 401–553. Faujas-Saint-Fond, B. (1799) Histoire Naturelle de la Montagne de Saint-Pierre de Maastricht. H. J. Jansen, Paris, pp. 183–214, pls. 34–42. Felix, J.P. (1884) Korallen aus ägyptischen Tertiärbildungen. Zeitschrift der Deutschen geologischen Gesellschaft, 36, 415–452, pls. 3–5. Felix, J.P. (1885) Kritische Studien über die Tertiäre Korallen-Fauna des Vicentins nebst Beschreibung einiger neuer Arten. Zeitschrift der Deutschen geologischen Gesellschaft, 37, 379–421. Felix, J.P. (1891) Versteinerungen aus der mexicanischen Jura- und Kreide-Formation. Palaeontographica, 37, 140–194. Felix, J.P. (1898) Beiträge zur Kenntnis der Astrocoeniinae. Zeitschrift der Deutschen geologischen Gesellschaft, 50, 247–256. Felix, J.P. (1899) Studien an cretaceischen Anthozoen. Zeitschrift der Deutschen geologischen Gesellschaft, 51(3), 378–387. Felix, J.P. (1900) Über zwei neue Korallengattungen aus den ostalpinen Kreideschichten. Sitzungsberichte der Naturforschenden Gesellschaft zu Leipzig, July 3, 1900, 37–40. Felix, J.P. (1903a) Studien über die korallenführenden Schichten der oberen Kreideformation in den Alpen und in den Mediterrangebieten. Palaeontographica, 49, 163–359. Felix, J.P. (1903b) Verkieselte Korallen als Geschiebe im Deluvium von Schlesien und Mähren. Centralblatt für Mineralogie, Geologie und Paläontologie (for 1903), 561–577. Felix, J.P. (1904) Polypiers du Sénonien Portugais. Communicaçoes dos Serviços Geológicos de Portugal, 5, 2, 375–388. Felix, J.P. (1906) Über eine Kreidefauna aus der Kreideformation Ost-Galiziens. Zeitschrift der Deutschen Geologischen Gesellschaft, 58, 38–52. Felix, J.P. (1914) Fossilium Catalogus. Animalia, Pars 5–7. Anthozoa palaeocretacea. Junk, Berlin, 1–273. Felix, J.P. (1925) Fossilium Catalogus. Animalia, Pars 28. Anthozoa eocaenica et oligocaenica. Junk, Berlin, 1–296. Felix, J.P. (1927). Fossilium Catalogus. Animalia, Pars 35. Anthozoa miocaenica. Junk,. Berlin. pp. 297–488. Filkorn, H.F. (1994) Fossil Scleractinian corals from James Ross Basin, Antarctica. Antarctic Research Series, 65, 1–96. Filkorn, H.F. (2003) Late Cretaceous (Maastrichtian) corals from Chiapas, Mexico. Geological Survey of America, Abstracts with Progams, 35(4), 32. Filkorn, H.F., Avendaño-Gil, J., Coutiño-José, M.A. & Vega-Vera, F.J. (2005) Corals from the Upper Cretaceous (Maastrichtian) Ocozocoautla Formation, Chiapas, Mexico. Revista Mexicana de Ciencias Geológicas, 22(1), 115–128. Floris, S. (1972) Scleractinian corals from the Upper Cretaceous and Lower Tertiary of Nûgssuaq,West Greenland. Muséum de Minéralogie et de Géologie de l’Université de Copenhague, Communications paléontologiques, 183, 1–132, 8 pls. Forbes, E. (1846) Report on the fossil Invertebrata from southern India, collected by Mr. Kaye and Mr. Cunliff. Transactions of the Royal Society of Edinburgh, 7, 97–174, pls. 7–10. Forchhammer, J.G. & Steenstrupp, J.J.S. (1850) Gaea danica. 3 pls. Fossa-Mancini, E. (1918) Catalogo dei fossili dell’Appennino centrale conservati nel Museo di Geoogia dell’Universitá di Pisa. Palaeontographica Italica, 24, 129–145. Foster, A.B. (1986) Neogene paleontology in the northern Dominican Republic. 3. The family Poritidae (Anthozoa, Scleractinia). Bulletins of American Paleontology, 90, 43–123. Fourtau, R. (1903) Contribution a l’Étude de la faune Crétacique d’Égypte. Bulletin de L’Institut Égytien, 4, 4, 349 pp. Frech, F. (1898) Charles White, On Hindeastraea, a new generic name of Cretaceous Astraeidae. Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen (1), 322. Fritsch, K. von (1878) Fossile Korallen der Nummulitenschichten von Borneo. Palaeontographica, Supplement, 3,



213

92–138, pls. 14–19. Fromentel, E. de. (1857) Description des Polypiers fossiles de létage Nèocomien. Bulletin de la Société des Sciences Historiques et Naturelles de l’Yonne. Perriquet et Rouillé, Auxerre, 78 pp. Fromentel, E. de. (1858–1861) Introduction à l´étude des Polypiers fossiles. Mémoires de la Société d’Émululation du Département du Doubs, 5, 1–357. Fromentel, E. de. (1860) Polypiers. In: Martin, J. (Ed.), Paléontologie stratigraphique de l’Infralias du départments de la Côte d’Or suive d’un aperçu paléontologique sur le même assises dans le Rhône, l’Ardèche et l’Isère. Mémoires de la Société Géologique de France, (2), 7, 1–100. Fromentel, E. de. (1862) Zoophytes, terrains crétacés (2–3). In: d’Orbigny, A. de (Ed.), Paléontologie Française. Vol. 8. Masson, Paris, pp. 49–144, pls. 1–36. Fromentel, E. de. (1863) Zoophytes, terrains crétacés (4–5). In: d’Orbigny, A. de (Ed.), Paléontologie Française. Vol. 8. Masson, Paris, pp. 145–240, pls. 37–60. Fromentel, E. de. (1864) Zoophytes, terrains crétacés (6). In: d’Orbigny, A. de (Ed.), Paléontologie Française. Vol. 8. Masson, Paris, pp. 241–288, pls. 61–70. Fromentel, E. de. (1867) Zoophytes, terrains crétacés (7). In: d’Orbigny, A. de (Ed.), Paléontologie Française. Vol. 8. Masson, Paris, pp. 289–336, pls. 73–86. Fromentel, E. de. (1870) Zoophytes, terrains crétacés (8). In: d’Orbigny, A. de (Ed.), Paléontologie Française. Vol. 8. Masson, Paris, pp. 337–384, pls. 85–96. Fromentel, E. de. (1877) Zoophytes, terrains crétacés (10). In: d’Orbigny, A. de (Ed.), Paléontologie Française. Vol. 8. Masson, Paris, pp. 433–480, pls. 109–120. Fromentel, E. de. (1884) Zoophytes, terrains crétacés (13). In: d’Orbigny, A. de (Ed.), Paléontologie Française. Vol. 8. Masson, Paris, pp. 529–560, pls. 145–156. Fromentel, E. de. (1886) Zoophytes, terrains crétacés (14–15). In: d’Orbigny, A. de (Ed.), Paléontologie Française. Vol. 8. Masson, Paris, pp. 561–608, pls. 157–180. Fromentel, E. de. (1887) Zoophytes, terrains crétacés (16). In: d’Orbigny, A. de (Ed.), Paléontologie Française. Vol. 8. Masson, Paris, pp. 609–624, pls. 181–192. Frost, S.H. & Langenheim, R.L. (1974) Cenozoic Reef Biofacies. Northern Illinois Press, Dekalb, 388 pp. Furque, G. & Camacho, H.H. (1949) El cretaceo superior de la costa Atlantica de Tierra del Fuego. Revista de la Asociacion Geologica Argentina, 4, 263–297. Gabb, W.M. (1864) Description of the Cretaceous fossils. Geological Survey of California Palaeontology, 1, 55–236. Gabb, W.M. & Horn, G.H. (1860) Description of new species of American Tertiary and Cretaceous fossils. Journal of the Academy of natural Science Philadelphia 2, 4, 375–406. Gallagher, W.B. (2002) Faunal changes across the Cretaceous-Tertiary (K-T) boundary in the Atlantic coastal plain of New Jersey, restructuring the marine community after the K-T mass-extinction event. Catastrophic Events and Mass Extinctions, Impacts and beyond. Geological Society of America Special Paper, 356, 291–301. Géczy, B. (1954) Studien über Cycloliten (Anth.). Geological Hungarica, Series Paleontologica, 24, 77–158, 15 pls. Gill, G.A. (1981) The fulturae (“compound synapticulae”), their structure and reconsideration of their systematic value. Acta palaeontologica Polonica, 25, 301–310. Götz, S. (2003) Biotic interaction and synecology in a Late Cretaceous coral-rudist biostrome of southeastern Spain. Palaeogeography Palaeoclimatology Palaeoecology, 193, 125–138. Goldfuss, A. (1826–1829) Petrefacta Germaniae. Vol. 1. Arnz, Düsseldorf, pp. 1–114. Grabau, A.W. & Shimer, H.W. (1909) North American Index Fossils, 1. New York, N.Y. 853 pp. Gray, J.E. (1842) (no title) In: Agassiz, L.J.R (Ed.), Nomenclator zoologicus, continens nomina systematica generum animalium tam viventum quam fossilium, fasc. Vol. 5. Sent et Grassmann, Soloduri, 1842–1847, p. 30. Gray, J.E. (1847) An outline of an arrangement of stony corals. Annals and Magazine of Natural History, 19, 20–128. Gregorio, A. (1890) Monographie de la faune éocénique de , et surtout de celle de Claiborne de l'étage parisien. Librarie internationale L. Pedone Lauriel de Charles Clausen, Palermo, 316 pp., 46 pls. Gregory, W. (1898) A collection of Egyptian fossil Madreporaria. Geological Magazine, (New Series 4), 5, 241–251, 2 pls. Gregory, W. (1900) On the geology and fossil corals and echinids of Somaliland. Quarterly Journal of the Geological Society of London, 56, 26–45, 2 pls. Gregory, W. (1930) The fossil fauna of the Samana Range and some neighbouring areas: Part VII. The Lower Eocene corals. Memoirs of the Geological Survey of India, Palaeontologica Indica (New Series), 15, 81–128. Guettard, M. (1770) Mémoires sur les différentes parties de la physique, de l’histoire naturelle; des sciences et des arts.


BARON-SZABO

Vol. 1. Costard, fils et compagne, Paris. Guettard, M.. (1774). Mémoires sur les différentes parties de la physique, de l’histoire naturelle; des sciences et des arts. 1–3, Atlas: 68 pls. Costard, fils et compagne, Paris. Hagenow, F. (1839) Monographie der Rügen’schen Kreide-Versteinerungen (1). Phytolithen und Poliparien. Neues Jahrbuch für Mineralogie, Geologie und Paläontologie, 251–296. Hagenow, F. (1840) Monographie der Rügen’schen Kreide-Versteinerungen (2). Radiarien und Annulaten nebst Nachträgen zur 1. Abtheilung. Neues Jahrbuch für Mineralogie, Geologie und Paläontologie, 631–672. Hansen, T.A., Farrell, B.R. & Upshaw, B. III. (1993) The first 2 million years after the Cretaceous-Tertiary boundary in east Texas: rate and paleoecology of the molluscan recovery. Paleobiology, 19/2, 251–265. Hassan, M.Y. & Salama, S.A. (1970) Contribution to the coral fauna of the Maestrichtian-Paleocene “paper shales” and “snow white chalk” of the oases of the southwestern desert of Egypt. Bulletin de l’Institut d’Egypt, 51, 73–101. Hennig, A. (1899) Faunan i Skånes yngre Krita. II. Korallerna. Kungliga Svenska Vetenskapsakademien. Bihnag till Handlingar, 24/4, 24 pp., 2 pls. Hladil, J., Otava, J. & Galle, A. (1992) Oligocene Carbonate Buildups of the Sirt Basin, Libya. In: Salem, M. (Ed.), Geology of Libya 4, pp. 1401–1420. Hoffmeister, J.E. (1929) A new fossil coral from the Cretaceous of Texas. Proceedings of the United States National Museum, 76, 3 pp, 2 pls. Hoppe, W. (1922) Jura und Kreide der Sinaihalbinsel. Zeitschrift des deutschen Palästina-Vereins, 45, 61–79, 97–219, pls. 3–6. Howe, H.J. (1960) Turbinolia Rosetta, a new coral species from the Paleocene of Alabama. Journal of Paleontology, 34, 1020–1022. Jukes-Browne, A.J. (1904) Cretaceous rocks of Britain. Memoires of the Geological Survey of the United Kingdom, 3, 520–556. Kaye, C.A. (1956) Lower Tertiary of Puerto Rico. Bulletin of the American Association of Petroleum Geologists, 40, 108–121. Kiessling, W. & Baron-Szabo R.C. (2004) Extinction and recovery patterns of scleractinian corals at the K/T-boundary. Palaeogeography, Palaeoclimatology, Palaeoecology, 214, 195–223. Klinghardt, F. (1944) Das Krönner-Riff (Gosauschichten) im Lattengebirge. Mitteilungen des Alpenländischen Geologischen Vereines, 35, 179–213. Kner, P. (1848) Die Versteinerungen des Kreidemergels von Lemberg und seiner Umgebung. Naturwissenschaftliche Abhandlungen, 3, 1–42, pls. 1–5. Knorr, G.W. & Walch, E.I. (1777) Regueil des Monumens des catastrophes que le Globe de la Terre a éssuiées, contenant des Pétrifications e d’autres Pierres curieuses. Nürnberg. 116 pp.[publisher unknown] Koby, F. (1889) Monographie des polypiers jurassiques de la Suisse (9). Mémoires de la Société paléontologique Suisse (=Abhandlungen der Schweizerischen Paläontologischen GeselIschaft), 16, 457–586, pls. 121–130. Koenen, A. von (1885) Über eine Paleocäne Fauna von Kopenhagen. Abhandlungen der königlichen Gesellschaft der Wissenschaften zu Göttingen, 32, 125 pp., 3 pls. Kossmat, F. (1907) Geologie der Insel Sokrota, Semha und Abd el Kuri. Denkschriften der Akademie der Wissenschaften Wien, 71, 1–62. Kühn, O. (1929) Beiträge zur Palaeontologie und Stratigraphie von Oman (Ost Arabien). Annalen des Naturhistorischen Museums in Wien 43, 13–33, pls. 1–3. Kühn, O. (1933) Das Becken von Isfahan-Saidabad und seine Altmiocäne Korallenfauna. Palaeontographica, Serie A, 79, 143–221. Kühn, O. & Andrusov, D. (1937) Weitere Korallen aus der Oberkreide der Westkarpathen. Vestnik Královké ceské spolecnosti nauk (Trida mathematicko–prirodovedecká), 2, 1–18. Kühn, O. & Traub, F. (1967) Die Korallen des Paleozäns von Österreich. Mitteilungen der Bayerischen Staatssammlung Paläontologue und historische Geologie, 7, 3–21, pls. 1–2. Kuzmicheva, E.I. (1974) Asexual reproduction in Scleractinians and its taxonomic significance. In: Sokolov, B.S. (Ed.), Ancient Cnidaria. Trudy (Institut geologii I geofiziki. Sibirskoe otd. AN SSSR), 201 (Nauka, Novosibirsk), pp. 228–232, 2 pls. [in Russian]. Kuzmicheva, E.I. (1975) Ranne - i srednepaleogenovye korally nekotorykh rajonov Evropejskoj chasti SSSR. In: Menner, V.V., Moskvin, M.M. & Nadyn, N. (Eds.), Razvitie I smena organicheskogo mira na rubeshemezozoya I kaynozoya. Nauka, Moskva, pp. 15–31, pls. 1–4. [in Russian]. Kuzmicheva, E.I. (1982) Corals of the Upper Aptian (Klanceja) from the central Kyzylkum. Byulleten Moskovskogo



215

obshestva ispytateley prirody, otd. geologicheskiy, 52, 98–111, 2 pls. [in Russian]. Kuzmicheva, E.I. (1987). [Upper Cretaceous and Paleogene corals of the USSR.] Verkhnenelovye paleogenovye korally SSSR. Nauka, Moskva, 187 pp. [in Russian]. Lamarck, J.B.P. de (1801) Système des animaux sans vertèbres. Deterville, Paris, 423 pp. Lamarck, J.B.P. de (1816) Histoire naturelle des animaux sans vertèbres. Verdière, Paris, 568 pp. Lang, W.D. (1909) Growth stages in the British species of the coral genus Parasmilia. Proceedings of the Zoological Society of London, 30, 285–307. Latham, M.A. (1929) Jurassic and Kainozoic corals from Somaliland. Transactions of the Royal Society of Edinburgh, 56, 273–289, pls. 1–2. Lea, I. (1833) Corals. In: Carey, E.L., Lea, I. & Blanchard, C.E. (Eds.), Contributions to Geology. Philadelphia, pp. 227 pp., 4 pls. Leloux, J. (1999) Numerical distribution of Santonian to Danian corals (Scleractinia, Octocorallia) of southern Limburg, the Netherlands. Geologie en Mijnbouw, 78, 191–195. Leloux, J. (2002) Type specimens of Maastrichtian fossils in the National Museum of Natural History, Leiden. National Natuurhistorisch Musem, Tech Bulletin, 4, 1–40. Leloux, J. (2004) Notes on taxonomy and taphonomy of two Upper Maastrichtian (Upper Cretaceous) scleractinian corals from Limburg, The Netherlands. Scripta Geologica, 127, 313–339. Lesson, R.P. (1831) Illustrations de zoologie ou recueil de figures d’animaux peintes d’après Nature. Bertrand, Paris. 60 pls. Leymerie, A. (1846) Statistique géologique et minéralogique du département de I'Aube, Atlas. Laloy, Troyes, 675 pp. Leymerie, A. (1851) Mémoire sur un nouveau type pyrénéen parallèle á craie proprement dite. Mémoires de la Société géologique de France (2), 4, 177–202. Leymerie, A. (1878) Description géologique et paléontologique des Pyrénées de la Haute-Garonne. published by author, Toulouse, pls. 1–20, A–Z5. Leymerie, A. (1881) Description géologique et paléontologique des Pyrénées de la Haute-Garonne. published by author, Toulouse. 1010 pp. Liao, Wei-hua & Xia, Jin-bao (1994) Mesozoic and Cenozoic scleractinian corals from Xizang. Palaeontologica Sinica, New Series B, 184, 1–252. [in Chinese with English summary]. Linnaeus, C. von (1758) Systema naturae per regna tria naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Vol. 1. 10th edition. Holmiae, Stockholm, 824 pp. Lonsdale, W. (1845) Account of six species of Polyparia obtained from Timber Creek, New Jersey. Quarterly Journal of the Geological Society of London, 1, 65–75. Löser, H. (2000) Catalogue of Cretaceous corals. Repertoire of Species. Vol. 1. CPress Verlag, Dresden, 135 pp. Löser, H., Barattolo, F., Badia, S.C., Chikhi-Aouimeur, F., Dhondt, A., Erlich, R.N., Fözy, I., Geister, J., Hiss, M., Kolodziej, B., Leloux, J., Lewy, Z, Minor, K.P., Mitchell, S., Moosleitner, G., Peza, L., Remane, J., Romana, R., Sikharulidze, G.Y., Sinnyovski, D., Steuber, T., Tröger, K.-A., Turnšek, D., Vecchio, E., Vilella i Puig, J. & Zítt, J. (2002) Catalogue of Cretaceous corals. List of citations. Vol. 2 (2). CPress Verlag, Dresden, 784 pp. Löser, H., & Liao, W.H. (2001) Cretaceous corals from Tibet (China): stratigraphic and palaeobiogeographic aspects. Journal of Asian Earth Sciences, 19, 661–667. Löser, H., & Raeder, M. (1995) Aptian/Albian coral assemblages of the Helicon Mountains (Boeotia, Greece), palaeontological, palaeoecological and palaeogeographical aspects. Coral Research Bulletin, 4, 37–63. Lowe, R.T. (1852) Primitiæ et novitiæ faunæ et floræ Maderæ et Portus Sancti , two memoirs on the ferns, flowering plants, and land shells of Madeira and Porto Santo , with an appendix. Transactions of the Cambridge Philosophical Society, 2 pls. Maccagno, A.M. (1942) Zoantari maestrichtiani della Tripolitania. Reale Accademia Italiana. Rendiconto della Classe di Scienze Fisiche, Matematiche e Naturali (7), 3, 786–796. Mallada, L. (1887) Sinopsis de las especies fósiles que se han encontradas en España. Sistema Cretácico inferior (1). Boletin de la Comisión del Mapa geológico de España, 14, 1–172. Mallada, L. (1892) Catálogo general de las especies fósiles encontradas en España. Boletin de la Comisión del Mapa geológico de España, 18, 1–253. Mantell, G. (1829) The fossils of the South Downs; or illustrations of the geology of Sussex. L. Relfe, London, 327 pp, 42 pls. Marenzeller, E. von (1906) Expedition S.M. Schiff "Pola" in das Rote Meer (24). Über den Septennachwuchs der Eupsammiden e und h. Denkschriften der Akademie der Wissenschaften, 80, 1–12.


BARON-SZABO

Marini, M. (1942) Revisione della fauna neocretacica della Libia, Coralli. Annali del Museo Libico di Storia Naturale, 3, 75–82, pls. 4–5. Meek, F.B. (1864) Check list of the invertebrate fossils of North America. Cretaceous and Jurassic. Smithsonian Miscellaneous Collections, 177, 40 pp. Meek, F.B. & Hayden, F.V. (1861) A report on the invertebrate Cretaceous and Tertiary fossils of the upper Missouri Country. Annual Report of the United States geological and geographical Survey of the Territories, 9, 629 pp, 45 pls. [for 1860] Merriam, J.C. (1897) The geologic relations of the Martinez group of California at the typical locality. Journal of Geology, 5(8), 770. Metwally, M.H.M. (1996) Maastrichtian scleractinian corals from the western flank of the Oman Mountains, U.A.E. and their paleoecological significance. Neues Jahrbuch für Geologie und Paläontologie, Monatshefte 1996, 375–388. Meyer, J.C. (1987) Le récif danien deVigny. Géoguide SAGA Informations, 26, 1–72. Michelin, H. (1841) Iconographie zoophytologique. Description par localités et terrains des polypiers fossiles de France. Vol 2. Bertrand, Paris, pp. 18–40. Michelin, H. (1843) Iconographie zoophytologique. Description par localités et terrains des polypiers fossiles de France. Vol 3. Bertrand, Paris, pp. 73–104. Michelin, H. (1844) Iconographie zoophytologique. Description par localités et terrains des polypiers fossiles de France. Vol. 4. Bertrand, Paris, pp. 105–144. Michelin, H. (1845) Iconographie zoophytologique. Description par localités et terrains des polypiers fossiles de France. Vol. 5. Bertrand, Paris, pp. 145–184 Michelin, H. (1846) Iconographie zoophytologique. Description par localités et terrains des polypiers fossiles de France. Vol. 6. Bertrand, Paris, pp. 185–248. Michelin, H. (1847) Iconographie zoophytologique. Description par localités et terrains des polypiers fossiles de France. Vol. 7. Bertrand, Paris, pp. 249–328. Michelotti, I. (1838) Specimen Zoophytologiae diluvianae. Heredes seb. Botta, Torino, 227 pp. Milne Edwards, H. (1857) Histoire naturelle des Coralliaires ou polypes proprement dits. Vol. 1 and 2. Librairie encyclopédique de Roret, Paris, 1–633 and atlas. Milne Edwards, H. (1860) Histoire naturelle des Coralliaires ou polypes proprement dits. Vol. 3. Librairie encyclopédique de Roret, Paris, 1–560 pp. Milne Edwards, H. & Haime, J. (1848a) Observations sur les polypiers de la famille des astréides. Comptes Rendus de l’Académie des Sciences, 27, 465–469. Milne Edwards, H. & Haime, J. (1848b) Note sur la classification de la deuxième tribu de la famille des astréides. Comptes Rendus de l’Académie des Sciences, 27, 490–497. Milne Edwards, H. & Haime, J. (1848c) Recherches sur les polypiers (2). Monographie des Turbinoliides. Annales de Sciences naturelles, 3, 211–344, pls. 7–10. Milne Edwards, H. & Haime, J. (1848d) Recherches sur les polypiers (4). Monographie des Astréides (1) Eusmiliens. Annales de Sciences naturelles, 3, 209–320, pls. 5–9.02 Milne Edwards, H. & Haime, J. (1849a) Mèmoire sur les polypiers appartenant à la famille des oculinides, au groupe intermédiaire des Pseudoastrédes et à la famille des Fongides. Comptes Rendus de l’Académie des Sciences, 29, 67–73. Milne Edwards, H. & Haime, J. (1849b) Recherches sur les polypiers (4). Monographie des Astréides (2) Astéens (4–5). Annales de Sciences naturelles, 3rd Serie, 12, 95–197. Milne Edwards, H. & Haime, J. (1850a) A monograph of the British fossil corals (1). Tertiary and Cretaceous. Monographs of the Palaeontographical Society, 3, i–lxxxv, 1–71, pls. 1–11. Milne Edwards, H. & Haime, J. (1850b) Recherches sur les polypiers (5). Monographie des Oculinides. Annales de Sciences naturelles, 3rd Serie, 13, 63–110. Milne Edwards, H. & Haime, J. (1851a) A monograph of the British fossil corals. Corals from the oolitic formations. Monographs of the Palaeontographical Society, 5, 73–146, pls. 12–30. Milne Edwards, H. & Haime, J. (1851b) Monographie des polypiers fossiles des terrains paléozoïques. Archives du muséum d’histoire naturelle, 5, 502 pp, pls. 1–20. Mitchell, S.F. (2002) Palaeoecology of corals and rudists in mixed volcaniclastic-carbonate small-scale rhythms (Upper Cretaceous, Jamaica). Palaeogeography, Palaeoclimatology, Palaeoecology, 186 (3–4), 237–259. Monsour, E. (1944) Fossil corals of the genus Turbinolia from the Gulf Coast. Journal of Paleontology, 18(2), 109–118. Moosleitner, G. (2004) Fossilien sammlen im Salzburger Land. Quelle & Meyer, Wiebelsheim, 223 pp.



217

Morren, C.F.A. (1828) Responsio ad quaestionem propositam “Quaeritur descriptio Coralliorum fossilium in Belgio repertorum”. Annales Academiae Gandavensis, 76 pp., 22 pls. Morris, J. (1843) A catalogue of British fossils; comprising the genera and species hitherto described; with references to their geological distribution and to the localities in which they have been found. 1st Edition. published by author, London, 222 pp. Morycowa, E. (1971) Hexacorallia et Octocorallia du Crétacé inférieur de Rarau (Carpathes orientales roumaines). Acta Palaeontologica Polonica, 16, 3–149. Morycowa, E. & Lefeld, J. (1966) Karolowce z wapieni urgonskich serii wierchowej Tatr Polskich. Rocznik Polskiego Towarzystwa Geologicznego, 36, 519–542. Morycowa, E. & Roniewicz, E. (1990) Revision of the genus Cladophyllia and description of Apocladophyllia gen. n. (Cladophylliidae fam. n., Scleractinia). Acta Palaeontologica Polonica, 35, 165–190. Morycowa, E. & Roniewicz, E. (1995) Microstructural disparity between Recent fungiine and Mesozoic microsolenine scleractinians. Acta Palaeontologica Polonica, 40, 361–385. Moseley, H.N. (1881) Report on certain hydroid, alcyonarian, and madreporarian corals procured during the voyage of H.M.S. Challenger, in the years 1873–1876. Challenger Reports, Zoology, 2, 1–248, 32 pls. Myers, R.L. (1968) Biostratigaphy of the Cardenas Formation (Upper Cretaceous) San Luis Potosi Mexico. Paleontologia Mexicana, 24, 1–89. Nielsen, F. (1922) Zoantharia from Senon and Paleocene in Denmark and Skaane. Biologiske skrifter, 8, 199–233, 4 pls. Nötling, F. (1897) Fauna of the Upper Cretaceous (Maëstrichtian) beds of the Mari Hills. Fauna of Baluchistan. Memoirs of the Geological Survey of India, Palaeontologia Indica (16), 1, 1–79. Nomland, J.D. (1916) Corals from the Cretaceous and Tertiary of California and Oregon. Bulletin of the Department of Geological Sciences, University of California, 9 (5), 59–76. Oppenheim, P. (1901) Die Priabonschichten und ihre Fauna, im Zusammmenhange mit gleichaltrigen und analogen Ablagerungen vergleichend betrachtet. Palaeontographica, 47, 1–344. Oppenheim, P. (1903) Ûber die Fossilien der Blättermergel von Theben. Sitzungsbereichte der Königlich-Bayerischen Akademie der Wissenschaften (Mathematisch-Physikalische Classe), 32(3), 435–456, pl. 7. Oppenheim, P. (1909) Ueber eine Eocänfaunula von Ostbosnien und einige Eocänfossilien der Herzegowina. Jahrbuch der kaiserlich-königlichen Geologischen Reichsanstalt, 58 (2), 311-344. Oppenheim, P. (1912) Neue Beiträge zur Eozänfauna Bosniens. Beiträge zur Paläontologie und Geologie ÖsterreichUngarns und des Orients, 25, 87–149. Oppenheim, P. (1930a) Die Anthozoen der Gosauschichten in den Ostalpen. published by author, Berlin-Lichterfelde, 604 pp. Oppenheim, P. (1930b) Die Korallen der obersten Kreide Palästinas. Neues Jahrbuch für Mineralogie, Geologie und Paläontologie, Beilage-Band B, 64, 307–324. d’Orbigny, A. (1849) Prodrôme de Paléontologie stratigraphique universelle. Vol. 1. Masson, Paris, 394 pp. d’Orbigny, A. (1850) Prodrôme de Paléontologie stratigraphique universelle. Vol 2. Masson, Paris, 428 pp. d’Orbigny, A. (1851) Cours élémentaire de Paléontologie (3). Polypiers ou Zoophytes. Vol. 2. Masson, Paris, pp. 151–189. Ortmann, A. (1890) Die Morphologie des Skeletts der Steinkorallen in Beziehung zur Koloniebildung. Zeitschrift der wissenschaftlichen Zoologie, 50, 278–316, pl. 11. Osasco, E. (1902) Contribuzione allo studio dei coralli cenocoici del Veneto. Palaeontographia italica, 8, 99–120. Pal, A.K., Chatterjee, A.K., Prakash, G., Thussu, J.L. & De, B. (1984) On the fossil corals (Anthozoa) from the Indus Flysch of upper Indus valley, Ladakh. Geological Survey of India (Special publication series), 15, 55–69, 3 pls. Pallas, P.S. (1766) Elenchus Zoophytorum. Hague-Comitum, xvi and 28 and 415 pp. Pfister, T. (1980) Systematische und paläoökologische Untersuchungen an oligozänen Korallen der Umgebung von San Luca (Provinz Vicena, Norditalien). Schweizerische Paläontologische Abhandlungen, 103, 121 pp. Philips, J. (1829) Illustrations of the geology or a description of the strata and organic remains of Yorkshire coast. Thomas Wilson and Sons, York, 192 pp., 14 pls. Pictet, F.-J. (1857) Traité élémentaire de paléontologie ou histoire naturelle des animaux fossile, 2nd Edition, Vol. 1–4. Genève, 110 pp. Poèta, P. (1887) Die Anthozoen der Böhmischen Kreideformation. Abhandlungen der Königlichen Böhmischen Gesellschaft der Wissenschaften, 7, 1–60. Pourtalès, L.F. de (1880) Remarks on the corals and Antipatharia. Bulletin of the Museum of Comparative Zoology, 6, 103 pp.


BARON-SZABO

Pratz, E. (1882) Über die verwandtschaftlichen Beziehungen einiger Korallengattungen mit hauptsächlicher Berücksichtigung ihrer Septalstructur. Palaeontographica, 29, 81–124. Pratz, E. (1910) A korállok léirása. In: Pethö G. (Ed.), A Péterváradi Hegység (Fruska gora) Krétaidöszaki (Hiperszenon-) faunája. Royal Hungarian Association of Natural Science, Budapest, pp. 299–315, pls. 23–24. Prever, P.L. (1921) I coralli oligocenici di Sassello nell’Appenino Ligure. Paleontographica Italica, 27, 53–100. Quenstedt, F.A. (1852) Handbuch der Petrefactenkunde, 1st Ed., Part 2. Laup & Sieber, Tübingen, pp. 529–792, pls. 43–62. Quenstedt, F.A. (1881) Petrefactenkunde Deutschlands (6). Röhren- und Sternkorallen. Vol. 3. Fues, Leipzig, pp. 913–1094. Quenstedt, F.A. (1885) Handbuch der Petrefactenkunde. 3rd Edition. Laup & Sieber, Tübingen, 1–1239 pp. Raup, D.M. & Jablonski, D. (1993) Geographic End-Cretaceous marine bivalve extinctions. Science, 260, 971–973. Reig Oriol, J.M. (1987) Revisión y validez del género Anisoria (Escleractinia Cretácica). Trabajos del Museo Geológico del Seminario C. de Barcelona, 222, 3–9, 2 pls. Reig Oriol, J.M. (1990) Madreporarios eocénicos de Castellolo y de la Sierra de Malvals. published by author, Barcelona, 71 pp. Reig Oriol, J.M. (1995) Madreporarios cretácicos. published by author, Barcelona. 62 pp. Reig Oriol, J.M. (1997) Géneros y especies nuevas de Madreporarios cretácicos. published by author, Barcelona, 45 pp. Reis, O.M. (1889) Die Korallen der Reiter Schichten. Geognostische Jahreshefte, 2, 91–162. Reuss, A.E. (1852) Ueber drei Poliparienspezies aus dem oberen Kreidemergel von Lemberg. Palaeontographica, 3(3), 117–120, 1 pl. Reuss, A.E. (1854) Beiträge zur Charakteristik der Kreideschichten in den Ostalpen, besonders im Gosauthale und am Wolfgangsee. Denkschriften der kaiserlichen Akademie derWissenschaften, Mathematisch-Naturwissenschaftliche Classe, 7, 73–133. Reuss, A.E. (1864) Die fossilen Foraminiferen, Anthozoen und Bryozoen von Oberburg in Steiermark. Denkschriften der kaiserlichen Akademie der Wissenschaften, Mathematisch-Naturwissenschaftliche Classe, 23, 1–38, 10 pls. Reuss, A.E. (1867) Palaeontologische Studien über die älteren Tertiärschichten der Alpen. Fossile Anthozoen der Schichten von Castelgomberto. Sitzungsberichte der Kaiserlichen Akademie der Wissenschaften, MathematischNaturwissenschaftliche Classe, 56, 297–303. Reuss, A.E. (1868) Palaeontologische Studien über die älteren Tertiärschichten der Alpen. 1. Theil. Die fossilen Anthozoen und Bryozoen der Schichtengruppe von Crosara. Denkschriften der kaiserlichen Akademie der Wissenschaften, Mathematisch–Naturwissenschaftliche Classe, 28(1), 129–184, pls. 1–16. Reuss, A.E. (1869) Palaeontologische Studien über die älteren Tertiärschichten der Alpen. 2. Theil. Die fossilen Anthozoen und Bryozoen der Schichtengruppe von Crosara. Denkschriften der kaiserlichen Akademie der Wissenschaften, Mathematisch-Naturwissenschaftliche Classe, 29, 215–298, pls. 17–36. Reuss, A.E. (1872) Palaeontologische Studien über die älteren Tertiärschichten der Alpen. 3. Theil. Die fossilen Anthozoen und Bryozoen der Schichtengruppe von San Giovanni, Ilarione und von Ronca. Denkschriften der kaiserlichen Akademie der Wissenschaften, Mathematisch-Naturwissenschaftliche Classe, 31, 197–270, 21 pls. Reuss, A.E. (1874) Palaeontologische Studien über die älteren Tertiärschichten der Alpen. 3. Theil. Die fossilen Anthozoen und Bryozoen der Schichtengruppe von San Giovanni, Ilarione und von Ronca. Denkschriften der kaiserlichen Akademie der Wissenschaften, Mathematisch-Naturwissenschaftliche Classe, 33, 1–60, pls. 37–56. Roemer, F.A. (1841) Die Versteinerungen des norddeutschen Kreidegebirges. Verlage der Hahn’schen Hofbuchhandlung, Hannover, 113 pp, 16 pls. Roemer, F.A. (1888) Über eine durch die Häufigkeit Hippuriten–artiger Chamiden ausgezeichnete Fauna der Oberturonen Kreide von Texas. Paleontologische Abhandlungen, 4(4), 281–296. Roniewicz, E. (1976) Les scléractiniaires du Jurassique supérieur de la Dobrogea centrale Roumanie. Palaeontologica Polonica, 34, 17–121. Roniewicz, E. (1982) Pennular and non-pennular Jurassic scleractinians: some examples. Acta Palaeontologica Polonica, 27, 157–193, pls. 52–67. Roniewicz, E. (1996) The key role of skeletal microstructure in recognizing high-rank scleractinian taxa in the stratigraphical record. Paleontological Society Papers, 1, 187–206. Rosen, B.R. (2000) Algal symbiosis, and the collapse and recovery of reef communities, Lazarus corals across the K-T boundary. In: Culver, S.J., Rawson, P.F. (Eds.), Biotic response to global change, the last 145 million years. Cambridge University Press, Cambridge, pp. 164–180. Rosen, B.R. & Turnšek, D. (1989) Extinction patterns and biogeography of scleractinian corals across the Cretaceous/



219

Tertiary boundary. Memoirs of the Association of Australasian Palaeontologists, 8, 355–370. Rossi Ronchetti, C. (1955) Revisione della fauna neocretacica della Libia coralli, Fam. Smilotrochidae, Caryophylliidae, Parasmiliidae, Eupsammidae. Revista italiana di paleontologia e stratigrafia, 63/3, 101–126. Roskowska, M. (1955) Koralowce okolic Sochaczewa z warstz z Crania tuberculata. Acta Geologica Polonica, 5, 241–272. Russo, A. (1979) Studio monografico sui Coralli dell’Eocene di Possagno (Treviso, Italia). Atti e Memorie della Accademia Nazionale di Scienze, Lettere e Arti di Modena, Ser. VI, 21, 1–87, 15 pls. Samuel, O., Borza, K. & Köhler, E. (1972) Microfauna and lithostratigraphy of the Paleogen and adjacent Cretaceous of the Middle Vah Valley (West Carpathian). Cesky Geologisky Ustav Dionyza, Bratislava, 246 pp. Sanders, D. & Baron-Szabo, R.C. (2007) Palaeoecology of solitary corals in soft-substrate habitats: the example of Cunnolites (upper Santonian, Eastern Alps). Lethaia, 10.1111/ j.1502-3931.2007.00039.x (online publication). Sars, M. (1872) On some remarkable forms of animal life from the great deeps off the Norwegian Coast. In: Sars, G.O. (Ed.), University Program from the First Half of 1869, Brogger and Christie, Chritiana, pp. 1–82. Schafhauser, A.,Götz, S., Baron-Szabo, R.C. & Stinnesbeck,W. (2003) Depositional environment of coral-rudist associations in the Upper Cretaceous Cardenas Formation (central Mexico). Geologia Croatica, 56, 187–198. Schlotheim, E.F. Baron von (1820) Die Petrefactenkunde auf ihrem jetzigen Standpunkte durch die Beschreibung seiner Sammlung versteinerter und fossiler Überreste des Thier- und Pflanzenreichs der Vorwelt erläutert. Becker’sche Buchhandlung, Gotha, 541 pp., 15 pls. Schröter, J.S. (1778) Vollständige Einleitung in die Kenntniss und Geschichte der Steine und Versteinerungen. Part III, Versteinerungen. Pentacrinites, 336 pp., pl. 4. Schuster, F. (1996) Paleoecology of Paleocene and Eocene corals from the Kharga and Farafra Oases (Western Desert, Egypt) and the depositional history of the Paleocene Abu Tartur carbonate platform, Kharga Oasis. Tübinger geowissenschaftliche Arbeiten (TGA), Reihe A, Geologie, Paläaontologie, Stratigraphie, 31, 1–96, 21 pls. Schuster, F. (2002) Scleractinian corals from the Oligocene of the Qom Formation (Esfahan-Sirjan fore-arc basin, Iran). Courier Forschungsinstitut Senckenberg, 239, 5–55. Searles Wood, W. (1844) Descriptive Catalogue of the Zoophytes from the Crag. The Annals and Magazine of Natural History, 13, 10–21. Sedgwick, A. & Murchison, R.I. (1832) A sketch of the structure of the Eastern Alps, with sections through the newer formations on the northern flanks of the chain etc. Transactions of the Geological Society, Series II, 3, 1–301. Seguenza, G. (1864) Disquisizioni paleontologiche intorno di Corallarii fossili delle rocce tertiarie del distretto di Messina. Memorie della Real Accademia di Scienze Torino (2), 21, 399–560, 15 pls. Siemiradzki, J. (1927) On the Upper Cretaceous corals of Poland. Kosmos, 51, 313–324, 1 pl. [for 1926]. Smith, A.B. (1995) Late Campanian-Maastrichtian echinoids from the United Arab Emirates-Oman border region. Bulletin of The Natural History Museum, Geology Series, 51, 121–240. Smith, A.B. Gallemi, J., Jeffery, C.H., Ernst, G. & Ward, P.D. (1999) Late Cretaceous–early Tertiary echinoids from northern Spain, implications for the Cretaceous-Tertiary boundary. Bulletin of the Natural History Museum, Geology Series, 55, 81–137. Smith, A.B. & Jeffery, C.H. (2000) Maastrichtian and Palaeocene echinoids: a key to world faunas. Special Papers in Palaeontology, 64, 1–404. Sohl, N.F. (1987) Cretaceous gastropods: contrasts between Tethys and the temperate provinces. Journal of Paleontology, 61, 1085–1111. Söhle, U. (1899) Das Ammergebirge. Geognostische Jahreshefte, 2, 1–90. Solé Sabaris, L. (1941) Fauna coralina del Eoceno Catalán. Memorias de la Real Academia de Ciencias y Artes de Barcelona (3), 26/9, 259–440, pls. 1–10. Squires, D.F. (1958) The Cretaceous and Tertiary corals from New Zealand. New Zealand Geological Survey Palaeontological Bulletin, 29, 107 pp., 16 pls. Squires, D.F. (1960) Wellsotrochus, a new name for Wellsia Squires 1958. Journal of Paleontology, 34, 1053. Squires, D.F. (1962) Additional Cretaceous and Tertiary corals from New Zealand. Transactions of the Royal Society of New Zealand (Geology), 1, 133–150, pls. 1–4. Squires, R.L. & Demetrion, R.A. (1992) Paleontology of the Eocene Bateque Formation, Baja California Sur, Mexico. Contributions in Science, 434, 1–55. Stephenson, L.W. (1916) Coelenterata. In: Clark, W.B. (Ed.), Systematic paleontology of the Upper Cretaceous deposits of Maryland. Maryland Geological Survey, Upper Cretaceous. Baltimore, Maryland, pp. 752–757, pls. 48–49. Stephenson, L.W. (1917) North American Upper Cretaceous corals of the genus Micrabacia. Professional Papers of the


BARON-SZABO

United States Geological Survey, 98, 115–131. Stephenson, L.W. (1923) The Cretaceous formations of North Carolina. 1, Invertebrate fossils of the Upper Cretaceous formations. North Carolina, Geological and Economic Survey, 5, 1, 604 pp, 102 pls. Stolarski, J. (1995) Ontogenetic development of the thecal structures in caryophylliine scleractinian corals. Acta Geologica Polonica, 40, 19–44. Stolarski, J. (1996) Paleogene corals from Seymour Islands, Antarctic Peninsula. Acta Geologica Polonica, 55, 51–63. Stolarski, J. & Roniewicz, E. (2001) Towards a new synthesis of evolutionary relationships and classification of Scleractinia. Journal of Paleontology, 75, 1090–1108. Stoliczka, F. (1873) The corals or Anthozoa from the Cretaceous rocks of South India. Memoirs of the Geological Survey of India, Palaeontologia Indica (4), 8, 130–202, pls. 1–12. Suraru, M. (1961) Contributii la Cunoasterea faunei de coralieri din cretacicul superior al bazinului Rosia–Bihor. Studia Universitates Babes–Bolyai, (Geologia–Geographia), 1, 123–135. [in Romanian]. Tchéchmédjiéva, V. (1986) Paléoécologie des Madréporaires du Crétacé supérieur dans le Srednogorié de l’Ouest (Bulgarie occidentale). Geologica Balcanica, 16, 55–81. Tchéchmédjiéva, V. (1988) Espéces nouvelles de Madréporaires du Maastrichtien en Bulgarie du Sud-Ouest. Godizhnik na Sofiyskiya Universitet «Kliment Okhridski», geologo–geografski fakultet, (1, geologie), 77 (for 1983), 236–240, 2 pls. Tchéchmédjiéva, V. (1995) Crétacé supérieur, Chaetetides (Porifera) et Anthozoaires (Coelenterate). Fossilia Bulgarica 5b, 143 pp., 26 pls. (Presses universitaires “St. Kliment Ohridski”, Sofia). Tenison-Woods, J.E. (1878) On some fossil corals from Aldinga. Transactions, Proceedings, and Reports of the Philosophica Society of Adelaide, South Australia 1877–1878, 104–119, 2 pls. Toulmin, L.D. (1977) Stratigraphic distribution of Paleocene and Eocene fossils in the eastern Gulf Coast region. Geological Survey of Alabama – Monograph, 13, 1–601. Tragelehn, H. (1996) Maastricht und Paläozän am Südrand der Nördlichen Kalkalpen (Niederösterreich, Steiermark) – Fazies, Stratigraphie und Fossilführung des ‘Kambühelkalkes’ und assoziierter Sedimente. Thesis. University of Erlangen. 216 pp. Traub, F. (1938) Geologische und paläontologische Bearbeitung der Kreide und des Tertiärs im östlichen Rupertiwinkel, nördlich von Salzburg. Palaeontographica A, 88, 1–107. Trauth, F. (1911) Die oberkretazische Korallenfauna von Klogsdorf in Mähren. Zeitschrift des Mährischen Landesmuseums, 11, 85–184. Turnšek, D. (1978) Solitary Senonian corals from Stranice and MT Medvednica (NW Yugoslavia). Razprave Slovenska Akademija Znanosti in Umetnosti (4), 21, 66–125. Turnšek, D. (1992) Tethyan Cretaceous corals in Yugoslavia. Österreichische Akademie der Wissenschaften, Erdwissenschaftliche Kommission, 9, 155–170. Turnšek, D. (1994) Upper Cretaceous reef building colonial corals of Gosau facies from Stranice near Slovenske Konjice (Slovenia). Razprave Slovenska Akademija Znanosti in Umetnosti (4), 35, 3–41. Turnšek, D. (1997) Mesozoic corals of Slovenia. Znanstvenoraziskovalni Center SAZU, Ljubljana, 512 pp. Turnšek, D. & Buser, S. (1976) Cnidarian fauna from the Senonian breccia of Banjska Planota (NW-Yugoslavia). Razprave Slovenska Akademija Znanosti in Umetnosti (4), 19, 39–88. Turnšek, D. & Drobne, K. (1998) Paleocene corals from the northern Adriatic platform. Dela-Opera SAZU, 34, 129–154. Turnšek, D. & Polšak, A. (1978) Senonian colonial corals from the biolithite complex of Orešje on Mt. Medvednica (NW Yugoslovia). Razprave Slovenska Akademija Znanosti in Umetnosti (4), 21, 129–180. Umbgrove, J.H.F. (1925) Die Anthozoa uit het Maastrichtsche tufkrit. Leidse geologische mededelingen, 1, 83–126, pls. 8–11. Vaudois-Miéja, N. (1964) Les espèces nummulitiques atttributèes au genre Trochocyathus. Annales de Paléontologie (Invertébrés), 50, 53 pp., 4 pls. Vaughan, T.W. (1899) Some Cretaceous and Eocene corals from Jamaica. Bulletin of the Museum of Comparative Zoology, 34, 227–250, pls. 36–41. Vaughan, T.W. (1900) The Eocene and Lower Oligocene coral faunas of the United States with descriptions of a few doubtfully Cretaceous species. United States Geological Survey Monograph, 39, 263 pp., 24 pls. Vaughan, T.W. (1905) A critical review of the literature of the simple genera Fungiida, with a tentative classification. Proceedings of the United States National Museum, 28, 371–424. Vaughan, T.W. (1919) Fossil corals from central America, Cuba, and Porto Rico, with an account of the American Tertiary, Pleistocene, and Recent coral reefs. Smithsonian Institution Bulletin, 103, 612 pp.



221

Vaughan, T.W. (1920) Corals from the Cannonball Marine Member of the Lance Formation. United States Geological Survey Professional Papers, 128, 61–66, pl. 10. Vaughan, T.W. (1923) Fauna of the Sooke formation, Vancouver Island. Bulletin of the Department of Geological Sciences, University of California, 14(5), 123–234. Vaughan, T.W. & Hoffmeister, J.E. (1925) New species of fossil corals from the Dominican Republic. Bulletin of the Museum of Comparative Zoology, Harvard 67(8), 315–326. Vaughan, T.W. & Popenoe, W.P. (1935) The coral fauna of the Midway Eocene of Texas. Bulletin of the University of Texas, 3301, 325–343. Vaughan, T.W. & Wells, J.W. (1943) Revision of the suborders, families and genera of the Scleractinia. Geological Society of America, Special Paper, 44, 1–363. Vecsei, A. & Moussavian, E. (1997) Paleocene Reefs on the Maiella Platform Margin, Italy, an example of the effects of the Cretaceous/Tertiary boundary events on reefs and carbonate platforms. Facies, 36, 126–140. Verrill, A.E. (1865) List of polyps and corals sent by the Museum of Comparative Zoology to other institutions in exchange, with annotations. Bulletin of the Museum of Comparative Zoology, 1, 29–60. Vetters, H. (1925) Über kretazeische Korallen und andere Fossilreste im nordalpinen Flysch. Jahrbuch der Geologischen Bundesanstalt, 75, 18 pp, 1 pl. Vidal, A. (1980) Los Scleractinia de Collades de Bastús (Con.-Sant., prepirineo de la provincia de Lérida). Publicaciones de Geología. Universidad Autónoma de Barcelona, 11, 94 pp., 12 pls. Vidal, L.M. (1874) Datos para el conocimiento del terreno garumnense de Cataluña. Boletín de la Comisión del Mapa geológico de España, 1, 209–247, pls. 1–8. Vidal, L.M. (1917) Nota paleontológica sobre el cretácico de Cataluña. Congreso de la Associación española para el progreso de las ciencia, pp. 3–19, pls. 1–4. Vidal, L.M. (1921) Contribución a la Paleontologia del Cretácico de Cataluña. Memorias de la Real academia de Ciencias y Artes de Barcelona (3), 17, 89–107, 8 pls. Wells, J.W. (1932) Corals of the Trinity Group of the Comanchean of Central Texas. Journal of Paleontology, 6, 225–256. Wells, J.W. (1933) Corals of the Cretaceous of the Atlantic and Gulf Coastal Plains and Western Interior of the United States. Bulletins of American Paleontology, 18, 1–207. Wells, J.W. (1934) Some fossil corals from the West Indies. Proceedings of the United States National Museum, Washington, 83, 71–110. Wells, J.W. (1936) The nomenclature and type species of some genera of recent and fossil corals. American Journal of Science, Series 5, 31, 97–134. Wells, J.W. (1935) Corals from the Cretaceous and Eocene of Jamaica. Annals and Magazine of Natural History, (10), 15, 183–194, pls. 10–12. Wells, J.W. (1941a) Cretaceous and Eocene corals from Peru. Bulletins of American Paleontology, 26, 304–326, pls. 44–46. Wells, J.W. (1941b) Upper Cretaceous corals from Cuba. Bulletins of American Paleontology, 26, 282–300, pls. 42–44. Wells, J.W. (1944) Cretaceous, Tertiary and Recent corals, a sponge, and an alga from Venezuela. Journal of Paleontology, 18, 429–447. Wells, J.W. (1945) West Indian Eocene and Miocene corals, Part II. Geological Society of America, Memoirs, 9, 1–25. Wells, J.W. (1956) Scleractinia. In: Moore, R.C. (Ed.), Treatise on invertebrate paleontology. Part F. Geological Society of America and University of Kansas, Press, Boulder, Colorado and Lawrence, Kansas. pp. F328–F444. Wolleben, J.A. (1977). Paleontology of the Difunta group Upper Cretaceous-Tertiary in northern Mexico. Journal of Paleontology, 51, 373–398. Wu, Wang-shi. (1975) The coral fossils from the Qomolangma Feng Region. Report of the Scientific Investigations in the Qomolangma Feng Region, Paleontology, 1, 83–113, pls. 1–10. Youssef, M.H. & Salama, S.A. (1969) Contribution to the Coral fauna of the Maastrichtian-Paleocene “Paper Shales” and “Snow White Chalk” of the Oases of the South Western Desert of Egypt. Bulletin de l’Institut Egyptien 51, 73–98.


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Appendix 1. Taxa mentioned from Maastrichtian and/or Paleocene strata without description or assigned doubtfully Actinacis ? in Barthel & Herrmann-Degen, 1981, Paleocene of Egypt, no description, no illustration. ?Aulocyathus sp. of Stolarski, 1996, Paleocene of Antarctica: incomplete preservation; the proximal part of the specimen is broken, therefore, the important generic character (reproduction by longitudinal fragmentation of parent coralla) cannot be observed. Cyathoseris ? valmondasiaca (Michelin) in Stemann, in Bryan et al., 1997, Paleocene of the USA (Alabama, Salt Mountain): no illustration, insufficient description mentioning only the cup-shaped corallum, cd: