PALAEONTOLOGIA POLONICA — No. 62, 2004 THE EVOLUTION OF CONCH ONTOGENY IN MESOZOIC OPEN SEA GASTROPODS (EWOLUCJA ONTOGENEZY MUSZLI MEZOZOICZNYCH ŚLIMAKÓW OTWARTEGO MORZA) by ANDRZEJ KAIM (WITH 140 TEXT−FIGURES) WARSZAWA 2004 INSTYTUT PALEOBIOLOGII PAN im. ROMANA KOZŁOWSKIEGO THE EVOLUTION OF CONCH ONTOGENY IN MESOZOIC OPEN SEA GASTROPODS ANDRZEJ KAIM Kaim, A. 2004. The evolution of conch ontogeny in Mesozoic open sea gastropods. Palaeon− tologia Polonica 62, 3–183. Data on embryonic, larval, and postlarval morphology of the gastropod conchs from the Mid Jurassic to Early Cretaceous black clays from Poland are used to improve their taxonomy. The phylogenetic relationships of the fossil gastropod taxa to their Recent relatives are traced. The results show that most of the Recent groups were already present in the Mid Ju− rassic. The vetigastropods are apparently conservative and shells of some groups have not changed significantly for at least 160 Ma. The most primitive extant caenogastropods (e.g., Cerithioidea and Ampullospiridae) most probably have originated from the branches which diverged already in the Late Paleozoic. The other caenogastropods have undergone signifi− cant radiation in the Triassic. Then the Caenogastropoda had a period of relative stagnation. The significant exception is emergence and rapid radiation of Neogastropoda during the Cre− taceous. The most diversified Mesozoic family of Heterostropha, the Mathildidae, had its acme from the Late Triassic to the Early Cretaceous. Later the family gradually declines and nowadays it is a relic group. The oldest cephalaspidean opisthobranchians are known since the Early Triassic but the significant radiation has taken place in the Mid Jurassic. The siphonariids, the marine pulmonates, are recognized from the Early Cretaceous. 135 species (54 of them new) and two new genera (Dzikella and Gnaszynium) are described. Key words: Gastropoda, evolution, phylogeny, taxonomy, Jurassic, Cretaceous, Poland. Andrzej Kaim [[email protected]], Institute of Paleobiology, Polish Academy of Sciences, Twarda 51/55, PL−00−818 Warszawa, Poland. Received 7 October 2003, accepted 10 March 2004 4 ANDRZEJ KAIM CONTENTS Introduction ................................................... 6 Acknowledgements .............................................. 7 Fossil localities ................................................. 7 Methods ..................................................... 10 Taxonomy .................................................... 10 Phylum MOLLUSCA Linné, 1758....................................... 11 Class GASTROPODA Cuvier, 1797 ...................................... 11 Subclass PROSOBRANCHIA Milne Edwards, 1848 ............................. 11 Order Archaeogastropoda Thiele, 1925 .................................. 11 Suborder Vetigastropoda Salvini−Plawen et Haszprunar, 1987 ....................... 11 Superfamily Trochoidea Rafinesque, 1815 ............................... 11 Family Skeneidae Clark, 1851 .................................... 11 Genus Eudaronia Cotton, 1945 ................................... 11 Genus Adeuomphalus Seguenza, 1876 ............................... 13 Genus Aequispirella Finlay, 1924.................................. 13 Family Turbinidae Rafinesque, 1815 ................................. 15 Genus Torallochus Kiel et Bandel, 2002 .............................. 15 Family Trochidae Rafinesque, 1815 ................................. 17 Subfamily Eucyclinae Koken, 1896.................................. 17 Genus Eucycloscala Cossmann, 1895 ................................ 17 Genus Parvitomella Gründel, 2000 ................................. 21 Genus Calliotropis Seguenza, 1902 ................................. 22 Genus Turcica Adams et Adams, 1854 ............................... 26 Family Ataphridae Cossmann, 1918 ................................. 28 Genus Ataphrus Gabb, 1869 .................................... 28 Family uncertain .......................................... 29 Genus Striatoconulus Gründel, 2000 ................................ 29 Genus ?Crossostoma Morris et Lycett, 1851 ............................ 30 Order Caenogastropoda Cox, 1959 .................................... 31 Superfamily Cerithioidea Férrusac, 1822 ................................ 31 Family Cerithiidae Férrusac, 1822 .................................. 31 Genus Cryptaulax Tate, 1869.................................... 31 Genus Schroederium Kowalke, 1998 ................................ 44 Genus Rhynchocerithium Cossmann, 1906 ............................. 46 Genus Exelissa Piette, 1860 .................................... 46 Family Turritellidae Lovén, 1847 .................................. 48 Genus Turritella Lamarck, 1799 .................................. 48 Family uncertain .......................................... 50 Genus Tomaszoviella Kaim, 2001 ................................. 50 Genus Diatrypesis Tomlin, 1929 .................................. 50 Superfamily uncertain ......................................... 52 Genus Dzikella gen. n. ....................................... 52 Suborder Ptenoglossa Gray, 1853 ..................................... 52 Superfamily Triphoroidea Gray, 1847 ................................. 53 Family Eumetulidae Golikov et Starobogatov, 1975.......................... 53 Genus Vatopsis Gründel, 1980 ................................... 53 Genus Cosmocerithium Cossmann, 1906 .............................. 56 Superfamily Janthinoidea Gray, 1853.................................. 60 Family Epitoniidae Berry, 1910 ................................... 60 Genus Claviscala de Boury, 1909 ................................. 60 Family Nystiellidae Clench et Turner, 1952 .............................. 62 Genus Opaliopsis Thiele, 1928 ................................... 62 Superfamily uncertain ......................................... 63 Family Polygyrynidae Bandel, 1993 ................................. 63 Genus Teutonica Schröder, 1992 .................................. 63 Superfamily Zygopleuroidea Wenz, 1938 ................................ 66 Family Zygopleuridae Wenz, 1938 .................................. 66 Genus Azyga Nützel, 1998 ..................................... 66 MESOZOIC GASTROPODS 5 Family Pommerozygiidae Gründel, 1999 ............................... 68 Genus Costazygia Gründel, 1999 .................................. 68 Family Pseudozygopleuridae Knight, 1930 .............................. 70 Genus Plocezyga Knight, 1930 ................................... 70 Superfamily Stromboidea Rafinesque, 1815............................... 70 Family Aporrhaidae Gray, 1850 ................................... 70 Genus Dicroloma Gabb, 1868 ................................... 71 Genus Pietteia Cossmann, 1904 .................................. 74 Genus Quadrinervus Cossmann, 1904 ............................... 77 Genus Spinigera d’Orbigny, 1850 ................................. 78 Superfamily Rissooidea Gray, 1847 ................................... 79 Family Rissoidae Gray, 1847 ..................................... 79 Genus Bralitzia Gründel, 1998 ................................... 80 Genus Buvignieria Cossmann, 1921 ................................ 85 Genus Zebina Adams et Adams, 1854 ............................... 92 Genus Pusillina Monterosato, 1884 ................................. 93 Family uncertain .......................................... 95 Genus Knightella Longstaff, 1933 ................................. 95 Superfamily Pseudomelanoidea Hoernes, 1884 ............................. 97 Family Pseudomelaniidae Hoernes, 1884 ............................... 97 Genus Pseudomelania Pictet et Campiche, 1862........................... 97 Superfamily Calyptraeoidea Lamarck, 1809............................... 100 Family Hipponicidae Troschel, 1861 ................................. 100 Genus Gnaszynium gen. n ..................................... 100 Suborder Neogastropoda Thiele, 1929................................... 101 Superfamily Muricoidea Rafinesque, 1815 ............................... 101 Superfamily uncertain ......................................... 101 Family Maturifusidae, Gründel, 2001 ................................ 102 Genus Astandes Wade, 1917 .................................... 102 Family Purpurinidae Zittel, 1895 .................................. 107 Genus Purpurina d’Orbigny, 1850 ................................. 107 Subclass HETEROBRANCHIA Gray, 1840 ................................. 110 Order Heterostropha Fischer, 1885 .................................... 110 Superfamily Pyramidelloidea Gray, 1840 ................................ 110 Family Pyramidellidae Gray, 1840 .................................. 110 Genus Chrysallida Carpenter, 1856 ................................. 110 Family Amathinidae Ponder, 1987 .................................. 111 Genus Leucotina Adams, 1860 ................................... 111 Genus Cyclothyca Stearns, 1890 .................................. 111 Family Ebalidae Warén, 1995 .................................... 114 Genus Ebala Leach in Gray, 1847 ................................. 114 Superfamily Architectonicoidea Gray, 1850............................... 115 Family Mathildidae Dall, 1865.................................... 115 Genus Mathilda Semper, 1865 ................................... 115 Genus Promathildia Andreae, 1887................................. 123 Genus Carinathilda Gründel, 1997 ................................. 127 Genus Gymnothilda Schröder, 1995................................