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Machaeridians, Chitons, and Conchiferan Molluscs of the Mojcza Limestone

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Machaeridians, Chitons, and Conchiferan Molluscs of the Mojcza Limestone MACHAERIDIANS, CHITONS, AND CONCHIFERAN MOLLUSCS OF THE MOJCZA LIMESTONE JERZY DZIK Dzik , J. 1994 . Mac haeri dia ns, chitons , and conchiferan mo lluscs of the M6jcza Lim e­ stone. -In: J. Dzik, E. Olempska, and A. Pisera. 1994 . O rdovi cian ca rbonate plat ­ form ecosystem of the Holy Cross Mo untains. Palaeontologia Polonica 53 , 213-252. Phosph ate coated origina lly aragonitic mollu sc she lls and calcitic mach aeridi an scleri tes contribute significa ntly to foss il assemblages of the ex treme ly co ndensed M6jcza Lim estone, providin g an almos t co ntinuous record of the faunal dynami cs of these gro ups from the end of Arenig to the beginning of Ashg i11. Except for the topmost bed , which contains a relatively war m- water faun a, the M6jcza Limestone and the ove rlying Zalesie Formation co ntain mollu scs of Gondwanan affinities, pres­ umabl y co ld-water. Machaerid ians are represented by at least six lineages. Four lineages of septemchitonid polypl acoph orans, twelv e bellerophontids and gas tropods, five rostro conchs and bivalv es, and eleve n hyolith s continue across significa nt part s of the sec tion. Aulacolepos elongatum sp. n., Sarkachiton kielcensis gen. et sp. n., Solen ocarididae fam . n., Bursata santacrucensis sp. n., and Mojczatheca triangularis gen. et sp. n. are prop osed. K e y w o r d s: Machaer idia, Polypl acoph ora, gas tropods, hyolith s, Ordovician , larv al development, evo lution. Jerzy Dzik, lnstytut Paleobiologii PAN. Aleja Zwirki i Wigury 93.02--089 Warszawa. Poland. Revised version received 5th Jun e, 1993. 2 14 JERZY DZIK CONTENTS Introduction . .215 Acknowledgements . .2 16 Taxonomic descriptions .2 16 Class Machaeridia WITH ERS, 1926 .216 Order Turrilepadida PILSBRY, 1916 .2 16 Family Turrilepadidae CLARK E, 1896 .2 18 Genus Mojczalepas DZI K, 1986 .2 18 Genus De/taco/eus WITH ERS, 1926 .2 18 Family Lepidocoleidae CLARKE, 1896 .220 Genus Plicacoleus DZI K, 1986 .. .220 Genus Aulako lepos WOLBURG, 1938 .220 Phylum Mollusca CUVIER , 1797 . .22 1 Subphylum Amphineura IHERING, 1877 .. .22 1 Class Polyplacophora DE BLAI NVI LLE, 1816 .22 1 Order Septemchitonida BERGENHA YN, 1955 .221 Family Septemchitonidae BERGENH AYN, 1955 .22 1 Genus Sarkachiton gen. n. .22 1 Genus Priscochiton DALL, 1882 .222 Family Solenocarididae fam. n.. .222 Genus Bursata RHOADS, 1962 .223 Order Chelodida BERGENIl AYN, 1943 .224 Family Chelodidae BERGENIlAYN, 1943 .224 Genus Che/odes DAVIDSON et KING, 1874 .224 Subphylum Cyrtoso ma RUNNEGAR et POJETA, 1974 .224 Class Monoplacophora KNIGHT, 1952 . .226 Order Bellerophontida ULRICH et SCOFIELD, 1897 .226 Family Bucaniidae ULR ICH et SCOFIELD, 1897 .226 Genus Modestospira YOCHELSON , 1964 .226 Genus Kokenospira BASSLER, 1915 .228 Family Tropidodiscidae KNIGHT, 1956 . .228 Genus Cyr todiscus PERNER, 1903 .228 Genus Tropidodiscus MEEK et WORTHEN, 1866 .228 Order Mimospirida DZI K, 1983 . .229 Family Clisospiridae MILLER, 1889 .229 Genus Mimospira KOKEN, 1925 .229 Class Hyolitha MAREK, 1963 .23 1 Order Circothecida SYSSOI EV, 1968 . .23 1 Family Circothecidae MISSARZHEVSKY , 1968 .23 1 Genus Cir cotheca SYSSOlEV , 1958 . .231 Family ?Tetrathecidae SYSSOIEV , 1968 .232 Genus Mojczatheca gen. n. .. .. .232 Order Orthothecida MAREK , 1966 .233 Family Orthothecidae SYSSOlEV, 1958 .233 Genus Nephrotheca MAREK, 1966 .233 Genus Panitheca MAREK , 1967 .234 Genus Bactrotheca NOVA K, 1891 .235 Genus Quadrotheca SYSSOIEV, 1958 .236 Order Hyolithida MATTHEW. 1899 .236 Family Hyolithidae NICHOLSON , 1872 .236 Genus .Ioachimi/ites MAR EK, 1967 . .236 MOLLUS CS OF TH E MOJ CZA LIM ESTONE 215 Genus Carinolithes SYSSOI EV, 1958 .239 Family Pauxi1litidae MAR EK , 1967 .240 Genus Recilites MAR EK , 1967 . .240 Class Gastropoda CUVI ER , 1797 . .. .240 Genus Umhonellina KOKEN, 1925 .242 Genus Gyronema ULR ICH 1897 .243 Genus Cyclonema HALL. 1859 . .244 Genus Holopea HALL, 1847 .244 Genus Straparo llina BILLI NGS, 1865 .246 Genus Pachystrophia PER NER , 1903 .246 Genus Loxonema PHILLIPS, 1841 . .247 Genus Siluriphorus COSSMANN, 1918 .247 Subphylum Diasoma RUNNEGAR et POJ ETA, 1974 .248 Class ?Rostroconchia POJETA et al., 1972 .248 Order ?Ribeiriida KOB AYASHI, 1954 .248 Incertae classis . .248 Order Jinonic ellida POK ORNY, 1978 . .248 Family Jinonicellidae POKOR NY , 1978 .248 Genus Jinonicella POKORNY, 1978 .248 Class Bivalvia LINNAEUS, 1758 . .248 Order Nuculida DALL, 1889 . .249 Family Ctenodontidae WOHRM ANN, 1893 .249 Genus Deceptrix FUCHS, 1919 . .249 Family Nuculidae GRAY, 1824 . .250 Genus Nuculoidea WILLl AM S et BREGER , 1916 .250 Order Mytilida FERRUSAC, 1822 . .250 Family Modiomorphidae MILL ER , 1877 . .250 Genus Modiolopsis HALL , 1847 .250 References . .... .... .250 INTRODUCTION The most celebrated issue in paleontology of the earliest Phanerozoic is the allegedly sudden appearance of numerous skeletal fossils at the base of the Cambrian. The reported earliest Cambrian fossil assemblages are mostly composed of secondarily phosphatized and originally phosphatic minute fossils, the so called "small shelly fossils" (see BENGTSON et al. 1990 for recent reviews). Paradoxi­ cally, the same kind of punctuated occurrence and preservation of minute skeletal fossils in the Ordovician and Silurian, known for decades, remains almost unnoticed, except for students of the Baltic Ordovician geology. The pattern of distribution of "small shelly fossils" in the whole early Paleozoic suggests that their abundance in some horizons (including the Tommotian) was controlled by preservational factors and had little to do with biological evolution. The widely accepted opinion that the suddenness of the basal Cambrian faunal change was a unique evolutionary event resulted mostly from concentration of micropaleontological research on the basal Cambrian rocks, as con­ trasted with neglection of similar faunas in the Ordovician. Occurrences of Cambrian, Ordovician, and Silurian phosphatic microfossils are generally restricted in time and space, being confined to narrow horizons in some geographic regions. The cephalopod limestone rock facie s is their most typical source. In all cases they are associated with numerous sedimentary discontinuities, at which skeletal detritus was exposed long enough to the action of sea water to undergo phosphatization (DZIK and PISERA 1994). In thi s respect the rocks of the Arenig in the Baltic region and the Tommotian of the Siberian Platform are closely similar. Phosphatic nuclei 216 JERZY DZIK of minute shells, the most co mmon way of preservation of micr ofossils in both region s, fre quently fill cavities in hard-grounds, and on ly locall y in areas where sedime nt was deposit ed at higher rate, can ad ult specimens of the same species be found (DZIK 1991). In so me extre me cases almost all the organic detritus seems to be enveloped in thin phosph atic lin ings, and in such stra ta any benthic macrofossils are rare or virtually lacking. Th is is the case with the Chinese Meishuc un assemblag e of the basal Camb ria n (QIAN and BENGTSON 1989), the Polish Ordovician M6jcza assemblage, but also the Silur ian Kok assemb lage of the Ca rn ic Alps. It seems crucial for better understa nding of the ea rly Phanerozoi c metazoan evolution to have more da ta on fauna l assemb lages pos t-dating the basal Cambrian "s ma ll she lly fossi ls" assemblages but correspo ndi ng to them in respect to the mode of preservation . Phosphatized and phosphate-covered m icrofossil s of molluscan and probl em atic affiniti es fro m the M6jcza Limeston e represent such a "s ma ll she lly fossils" assemblages of Ordovician age. They are the subject of the present paper. Acknowledgements. - I am ve ry grateful to Stefan BENGTSON for his thorou gh and helpful rev iew . I appreciate also the work done by an anonymous reviewer, who indi cated se veral termino­ logical mi stakes in the manuscript , even if I wa s not able to make use of his mo re gene ral comments. Mr. Wojciech SKARZYNSKI ass isted in taking SEM pictures at Nencki's Institute of Experimental Biology in Warsaw. TAXONOMIC DESCRIPTIONS Class Machaeridia WITH ERS, 1926 Order Turrilepadida PILSBRY, 1916 In the Carnbrian, along with the first true polyplacophoran mo lluscs, several other groups of foss il organisms are known which had bodies covered with an armor of sc leritized plates or scales . Th e only larger gro up of this kind in the Ordovician are the mac haeri dians, probabl e successors of the Cambrian tom mo tiids (BENGTSON 1970). Zoolog ica l affinities of the Mac hae ridia remai n unclear (possibly they are primit ive articulates, as discussed below) but an alternative molluscan co nnectio n cannot be definitely excluded (DzIK 1986). The elonga ted body of the pos t-Cambrian turrilep adi d mach aeridian s was cover ed with four rows of ca lcitic (instead of phosphat ic in the possibl y related Ca mbrian tommotiid s) sc lerites (BENGTSON 1978). At least in the most primitive family of the turrilep adids, the Plumulitidae JELL, 1979, the two first bod y segme nts had probably only the medial pair of the sclerites and the subseq uent three had their lateral sclerites morph ologically different from the follo wing thoraci c segme nts (JELL 1979). Thi s appa rent tag rnatio n may suggest affinities of the Mach aer idia to the arti cul ates rather than molluscs. In the most primitive arthro pods the anterior part of the body was tagmatized in a similar way. The two firs t segments for m the sensorial tagma in the arthro pods (oc ular and antennular segments) , while the following thr ee differ morphologicall y and functiona lly fro m the rest of the bod y rep resenting the primary oral segments (second antennae, mandibles, and first maxillae). Of special interest for und erstanding the nature of the ma ch aeridian sc ler ites is new dat a on the microm orph ology of their surfaces (PI.
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