bce eater merle: ty iy i 7 -« ale ~ os > * << -- Seren wncupere eo Iece our? eee * Se eres See = oats Sit 5 3 Beate Fivcecstones o natty ee mah i HREM iatat pT * ee Bee epee tick 2 Beier fear y ea eS pe ~ Tt aren eae Sensis EE bee Seino = = epee Dietapoe pe pre Baia ea: a5 =o ee, OU yesetzaop ods Stepeo = n Er HI uae eR ms —— - o es As . } S| > at 2 -t0.A VOLUME 52 MUSEUM NUMBER 1 27 JUNE 1996 The Bulletin of The Natural History Museum (formerly: Bulletin of the British Museum (Natural History)), instituted in 1949, is issued in four scientific series, Botany, Entomology, Geology (incorporating Mineralogy) and Zoology. The Geology Series is edited in the Museum’s Department of Palaeontology Keeper of Palaeontology: Dr L.R.M. Cocks Editor of the Bulletin: Dr M.K. Howarth Assistant Editor: Mr C. Jones Papers in the Bulletin are primarily the results of research carried out on the unique and ever- growing collections of the Museum, both by the scientific staff and by specialists from elsewhere who make use of the Museum’s resources. Many of the papers are works of reference that will remain indispensable for years to come. All papers submitted for publication are subjected to external peer review before acceptance. A volume contains about 160 pages, made up by two numbers, published in Spring and Autumn. Subscriptions may be placed for one or more of the series on an annual basis. Individual numbers and back numbers can be purchased and a Bulletin catalogue, by series, is available. Orders and enquiries should be sent to: Intercept Ltd. P.O. Box 716 Andover Hampshire SP10 1YG Telephone: (01264) 334748 Fax: (01264) 334058 Claims for non-receipt of issues of the Bulletin will be met free of charge if received by the Publisher within 6 months for the UK, and 9 months for the rest of the world. World List abbreviation: Bull. nat. Hist. Mus. Lond. (Geol.) © The Natural History Museum, 1996 Geology Series ISSN 0968-0462 Vol. 52, No. 1, pp. 1-89 The Natural History Museum Cromwell Road London SW7 5BD Issued 27 June 1996 Typeset by Ann Buchan (Typesetters), Middlesex Printed in Great Britain at The Alden Press, Oxford Bull. nat. Hist. Mus. Lond. (Geol.)52(1); 1-24 Issued 27 June 1996 Zirconolite: a review of localities worldwide, and a compilation of its chemical compositions C.T. WILLIAMS Department of Mineralogy, The Natural History Museum, Cromwell Road, London SW7 5BD, U. R. GIERE THE NATURA 27 JUN 1996 PRESENTED Mineralogisch-Petrographisches Institut der Universitat, Bernoullianum, CH-4056, Basel, Switzerland| AEONTOLOGY LIBRARY Cals Synopsis. A compilation of the chemical data and brief review of the mineral zirconolite, essentially CaZrT1,0,, is presented. A total of 321 chemical analyses, 169 previously unpublished, from 39 of the 46 known terrestrial localities, and covering 10 rock types are tabulated. A brief description of the minerals associated with zirconolite is outlined for each locality. Data from all zirconolite-bearing lunar rocks have also been compiled. The recently published nomenclature scheme for zirconolite is employed throughout. INTRODUCTION Zirconolite, although a relatively rare accessory mineral, 1s found in a wide range of rock types and _ geological environments. To date, zirconolite has been reported from 46 terrestrial localities and from 13 lunar samples: it has not been reported in meteorites. The chemical composition of natural zirconolite can vary extensively, with the main substitutions involving rare earth elements, actinide elements, niobium and iron. Synthetic zirconolite is a major component in SY NROC, a synthetic polyphase titanate ceramic designed to immobilise high-level radioactive waste. In this paper, we compile and tabulate all reported chemical data for natural zirconolites, including new, and previously unpublished analyses; we group zirconolites into specific rock types or paragenetic types, and denote those samples that are stored in the collections of the Mineralogy Department, The Natural History Museum, London. NOMENCLATURE In the literature, several minerals with stoichiometries close to CaZrTi,O,, but with different crystal structures, have been reported and this has led to confusion in the nomenclature of these minerals. The compound CaZrTi,O, can exist as three superstructures with monoclinic, orthorhombic and trigonal symmetries (Rossell, 1980), each being a polytype (White, 1984), subsequently redefined as polytypoids (Bayliss e7 al., 1989). However, the original type material, polymignite (Berzelius, 1824), zirkelite (Hussak & Prior, 1895) and zirconolite (Borodin et al., 1956) are metamict and their structures cannot be unambiguously defined. Further problems in identification and characterisation have arisen, in part because the frequent occurrence of actinide elements in the structure may render the mineral partially or totally metamict, and in part because the often small grain size does not allow for routine crystallographic techniques to be employed. These nomenclature problems have been addressed by Nickel & Mandarino (1987) and most © The Natural History Museum, 1996 recently by Bayliss et al. (1989), who summarized the crystallographic and chemical characteristics of these minerals, detailed their historical documentation, and rationalized their nomenclature. Under the Bayliss et a/. (1989) IMA-approved nomenclature scheme, zirconolite is the non-crystalline (metamict) mineral, or the mineral with undetermined polytypoid of CaZrTi,0,; zirconolite-3O is the three-layered orthorhombic polytypoid of CaZrTi,O,; zirconolite-3T is the three layered trigonal polytypoid of CaZrTi,O,; zirconolite-2M is the two-layered monoclinic polytypoid, or aristotype (White, 1984) of CaZrTi,O,; zirconolite is polymignite (metamict), and zirkelite is the cubic mineral with formula (Ti,Ca,Zr)O, ,. Smith & Lumpkin (1993) have subsequently described two additional polytypes which appear to be supercells of the zirconolite-2M and 3T structures (zirconolite-4M and -6T, respectively). CHEMICAL COMPOSITION Zirconolite has five cation-acceptor sites, these being Ca in 8-coordination, Zr in 7-coordination, and three distinct Ti sites: Ti(I) and Ti(IIl) are both 6-coordinate, and Ti(II) is 5-coordinate (Gatehouse et al., 1981; Mazzi & Munno, 1983). In natural (and synthetic) zirconolites, a wide range of cation substitutions can occur (e.g. Ringwood, 1985), ranging in ionic size from 0.051nm (Ti**) to 0.112nm (Ca?*) — ionic radii data from Shannon (1976) — and charge from 2+ (Mg) to 6+ (W). Predominant substitutions are: the rare earth elements including Y (REE) and actinide (ACT) elements for Ca; Hf for Zr; and Nb, Fe, Ta, Mg and W for Ti. In natural zirconolites the chemical variation is extensive; of the major components, CaO ranges from 1.83 to 16.54%, ZrO, from 22.82 to 44.18%, and TiO, from 13.56 to 44.91% (Table 1). Up to 79% of the Ca site can be replaced by other cations (e.g. analyses A4, L11, Table 3), and up to 65% of the Ti site (analysis C69, Table 3). TIT er | HISTORY MUSEUM | M417 ’ DETAILS ee) ee from terrestrial and lunar a hd 3 in) of their host rock (or parce Des analysis numbers, together with references relating both to the first report for that occurrence, and to the sources of the analytical data. Full chemical analyses, where available, are given in Table 3. Kimberlite. Raber & Haggerty (1979) report zirconolite from three localities in South Africa. All their microprobe analyses are presented in Table 3 (K1 to K3); however, two analyses (K1 and K2) have significantly higher ZrO, values than all other zirconolites reported. The number of Zr cations for these analyses, recalculated on the basis of 7 oxygens, exceed the theoretical value by >50% and >100%. It seems probable therefore, that these analytical data are in error, or that the minerals analysed are not zirconolite. Calzirtite has been suggested as a possible alternative mineral for one of these questionable phases (Kogarko et a/., 1991). Analyses K1 and K2 are thus omitted from comparative data of zirconolites (such as in Table 1). Zirconolite described by Raber & Haggerty (1979) is very fine-grained, associated with baddeleyite, + zircon, ilmenite, armalcolite and calcite, and is considered to have been formed as a secondary mineral due to infiltration of, and reaction with, a carbonatitic fluid. Ultrabasic Rocks. Zirconolite has been described from two ultrabasic cumulate complexes — Laouni, Algeria (Lorand & Cottin, 1987) and Rhum, Scotland (Williams, 1978). At Laouni zirconolite occurs as compositionally homogeneous, discrete grains up to 200um in diameter, in plagioclase-rich adcumulates. Although baddeleyite also occurs in this intrusion, it is located in cumulates richer in trapped intercumulus liquid. Analyses Ul—U2 (Table 3) are from Lorand & Cottin (1987). In the layered, ultrabasic complex of Rhum zirconolite occurs as a rare, late-stage accessory mineral, associated with apatite, baddeleyite and zircon, predominantly in olivine-rich mesocumulates — i.e. those cumulates with a relatively high proportion of trapped (and fractionated) magma. The microprobe analysis (U3, Table 3) is from Fowler & Williams (1986). Gabbro Pegmatite. Harding et al. (1982, 1984) describe accessory zirconolite (reported as ‘zirkelite’), with acicular habit, from a gabbro pegmatite of Tertiary age at St. Kilda, Scotland. The pegmatite consists essentially of ferroaugite, ferroedenite, chlorite, magnetite, Mn-rich ilmenite, quartz and alkali feldspar. Associated with zirconolite are the accessory minerals biotite, epidote, allanite, titanite, apatite and zircon. The pegmatite is considered to have formed from the last residues of basaltic (tholeiitic) liquid from which the major Mg-minerals and feldspars of the Glen Bay Gabbro had previously precipitated. The zirconolite analysis tabulated here (G1, Table 3) is from Fowler & Williams (1986). Harding et al.’s (1982) orisinal analysis totals only 91%, as it excludes many of the heavy REE, Hf and Ta. Since the EREE** exceeds 50% of the Ca site cations, it is sensu strictu a rare earth mineral, and following the Bayliss & Levinson (1988) nomenclature guidelines, this mineral can be classified as zirconolite-(Y) [subject to approval from the CNMMN]. C.T. WILLIAMS AND R GIERE Syenite. Zirconolite has been described from four syenite localities. At Glen Dessarry, Scotland (Fowler & Williams, 1986), zirconolite occurs as an accessory mineral in a rock consisting of aegirine augite, edenitic amphibole, hypersolvus alkali-feldspar, orthoclase, albite and biotite. Zirconolite, typically <10pm in diameter, is enclosed by alkali feldspar phenocrysts and associated with Fe-Ti oxides, titanite, allanite, apatite and zircon. The microprobe analysis (S1, Table 3) is from Fowler & Williams (1986). Zirconolite is reported in the alkaline intrusions of the Arbarastakh massif, Aldan, Russia, where segregations of zirconolite occur in fa] ‘...micatized large-grained pyroxenite .. .’ with accessory apatite and ilmenite (Borodin er al., 1960). In Table 3, analyses $2 (Borodin et al., 1960) and S3 (Gaidukova ef al., 1962) are wet chemical determinations on mineral separates; analysis S4 (Wark et a/., 1973) is a microprobe analysis of a separate zirconolite grain. Zirconolite (described as ‘polymignite’) was reported from the syenite pegmatites of Fredicksvarn, S. Norway by Berzelius (1824), and was a mineral separately analysed by Brogger (1890) using ‘classical’ wet chemical techniques (S5, Table 3). It also occurs at Langesundfjord, near Larvik in coarse-grained syenite pegmatites (S6 and S7, Table 3 — previously unpublished microprobe data). Zirconolite (reported as ‘polymignite’) was described from a ‘sanidinite’ at Campi Flegrei, Italy (Mazzi & Munno, 1983). No analysis is included in Table 3, because the total of the reported analysis is low. Material is no longer available for analysis (Munno pers. comm., 1992). Nepheline Syenite. At Pine Canyon, Utah, USA, zirconolite was found as an accessory mineral associated with hibonite, perovskite and pseudobrookite (Agrell et al., 1986). The rock-forming minerals include corundum, nepheline and Mg-hercynite. Analyses (NSI-NS6, Table 3) are previously unpublished wavelength-dispersive microprobe data (CTW). At the Elk Massif, Poland, zirconolite and Nb-zirconolite are reported in an association with apatite, fluorite and pyrochlore in agpaitic nepheline syenite pegmatites (Dziedzic, 1984). The major minerals of the pegmatites are microcline, nepheline, aegirine, arfvedsonite, and more rarely eudialyte. No analytical data are given for the zirconolite. At Chikala, Chilwa Alkaline Province, Malawi, zirconolite occurs also as an accessory mineral, up to 0.3mm in size, in a nepheline syenite (sample number BM1980,P23(1), Platt ef a/., 1987 — microprobe analyses NS7-NS1I1, Table 3). The rock-forming minerals are alkali feldspar, nepheline, biotite, apatite and an opaque oxide phase. At Tchivira, Angola, niobian zirconolite is reported as intimately crystallized with wohlerite from nepheline syenite rocks of the alkaline complex (Mariano & Roeder, 1989). Analyses NS11 to NS16 are previously unpublished microprobe data of zirconolite-(Y) [see above] from Tchivira. At Pilanesberg, Transvaal, South Africa (Lurie, 1986), zirconolite occurs as an accessory mineral (A.N. Mariano, personal communication, 1993). No analytical data is reported. At Tre Croci, near Vetralla in the Vico Volcanic complex of the Roman Comagmatic Province, Latium, Italy, crystalline epitaxial zirconolite occurs as an accessory mineral associated with baddeleyite, zircon and rare thorian hellandite in a sanidinitic eyjectum with-nepheline and sodalite (G.C. Parodi, personal communication, 1993). Analyses NS1I7 to NS21 are previously unpublished microprobe data from this locality. ZIRCONOLITE Carbonatite. Carbonatites, with sixteen reported occurrences of zirconolite, seem to be the most common host rock type for this mineral. In the Kola Peninsula, Russia, zirconolite occurs in four separate carbonatite complexes where detailed descriptions, including studies on crystal morphology and crystal chemistry, is given in Bulakh & Ivanikov (1984). In the Afrikanda complex, Kola, zirconolite is described from the amphibolitized and fenitized pyroxenites (Borodin er al., 1956) In Table 3 analyses C1—C3 are wet chemical analyses; Cl-C2 from Borodin et a/. (1956) and C3 from Bulakh et al. (1960). C4 is a microprobe analysis (Wark et al., 1973). At Vuoriyarvi, Kola, zirconolite was first described as zirkelite (Bulakh et a/., 1960), then *... tentatively described as a niobium variety, niobozirconolite’ (Borodin et a/., 1960). It is associated with apatite-magnetite rocks (accompanying carbonatites, Zhuravleva ef al., 1976), accumulating predominantly in apatite, and was also observed replacing hatchettolite (Kapustin, 1980). Analyses C5—C7 (Table 3) are unpublished wavelength-dispersive microprobe data (CTW) on separate grains (BM1970,39); C8—-Cll are wet chemical analyses: C8—C9 from Borodin et al. (1960), C8 and C11 quoted in Kapustin (1980); C10 is from Bulakh et al. (1960). Bulakh et al. (1960) refer to zirconolite from the Sayan Province, Russia. Analysis C12 (Table 3) is a wet chemical analysis from Bulakh et al. (1960). At Seblyavr, Kola, niobozirconolite was initially identified as zirkelite (Bulakh et a/., 1960) and is described as. . . the typical mineral of the process of amphibolization-dolomitization confined to carbonatite ...’ (Kapustin, 1980). The mineral is partly metamict, it has good symmetry habit and displays complicated twinning (Bulakh et a/., 1960). Associated minerals are apatite, clinohumite, tetraferriphlogopite, pyrrhotite and richterite. Analysis C13 (Table 3) is a wet chemical analysis from Bulakh et al. (1960). At Kovdor, Kola, zirconolite is associated with zones of carbonatization, Kapustin (1980). Cl14-C16 (Table 3) are wet chemical analyses: Cl14-C15 from Kapustin (1980), and C16 from Kukharenko eft al. (1965). Analyses C17—-C34 are unpublished wavelength-dispersive micropobe data (CTW), on chemically-zoned zirconolite grains in a thin section of carbonatite. Associated minerals are baddeleyite and U-Ta-rich pyrochlore (Williams, in press). At Schryburt Lake, Ontario, Canada, zirconolite occurs intergrown with calzirtite, baddeleyite, and U-rich pyrochlore (Williams & Platt, in preparation). Microprobe analyses (C35—CS58, Table 3), show significant variations in PREE,O, and Nb,O;. Some of the grains have SREE** >50% of the Ca site, and, with Nd as the most abundant REE, this mineral can be classified as zirconolite-(Nd), following Bayliss & Levinson (1988), and subject to approval from the CNMMN. At Santiago Island, Cape Verde Republic, non-metamict Zirconolite-2M occasionally up to 2mm in diameter, occurs as an accessory mineral, often associated with pyrochlore, in apatite-rich s6vite, beforsite and glimmerite rocks of the Canafistula carbonatitic plug (Silva, 1979; Silva & Figueiredo, 1980). Analysis C59 (Table 3) is the wavelength-dispersive microprobe analysis from Silva & Figueiredo (1980). At Phalaborwa, South Africa, zirconolite was first described by Verwoerd (1986) from the carbonatite. Analyses C60—C64 (Table 3) are unpublished wavelength-dispersive microprobe analyses by CTW on sample BM1988,260, kindly provided by Prof. G. Bayer (ETH, Ziirich). In this rock, zirconolite is associated with baddeleyite and zircon, the latter mineral probably having crystallized at a later stage. Analyses C65—C66 3 are unpublished microprobe data (from Bochon University) from Prof. G. Bayer. At Sokh, Finland, zirconolite (reported as zirkelite) was originally described by Vartiainen (1980) from hydrothermal phoscorites. The crystals have apparently formed ‘... at the expense of pyrochlore and occur around and as inclusions in pyrochlore . . .’, and are also found as separate prisms. Analyses C67—-C70 (Table 3) are unpublished wavelength-dispersive microprobe data from Dr. I. Hornig-Kjarsgaard (pers. comm., 1992), analysed at the University of Mainz. At Kaiserstuhl, Germany, zirconolite occurs with calzirtite, baddeleyite, Nb-perovskite and pyrochlore (Keller, 1984). Analyses C71 and C72 are wavelength-dispersive microprobe analyses (Keller, 1984; Sinclair & Eggleton, 1982). In the Hegau volcanic province, Germany, zirconolite (described as “Nb-zirconolite’) is a typical accessory mineral in the carbonatitic tuffs (Keller et a/., 1990). Analyses C73—C88 (Table 3), are unpublished wavelength-dispersive microprobe analyses (CTW) of eight grains from a heavy mineral separate provided by Prof. J. Keller (Freiburg). At Prairie Lake, Ontario, Canada, niobian zirconolite is reported in association with wohlerite, pyrochlore, betafite and niobian perovskite (Mariano & Roeder, 1989). A microprobe analysis provided by Dr A.N. Mariano of six major elements is shown in Table 3 (C89). At the Cummins Range Carbonatite, Kimberley Area, Western Australia, accessory zirconolite occurs in an apatite-amphibolite rock. Qualitative analysis of the zirconolite showed the presence of Ca, Zr, Ti and minor Fe, but with Nb absent (Dr. A.N. Mariano, personal communication, 1993). At Howard Creek, British Columbia, Canada (Woolley, 1987; p.16), zirconolite is associated with zircon, magnetite and diopside in an apatite calcite carbonatite (Dr. A.N. Mariano, personal communication, 1993). There is no analytical data. At Catalao, Goias, Brazil (Woolley, 1987; p.179), zirconolite is associated with apatite and phlogopite in a calcite carbonatite (Dr. A.N. Mariano, personal communication, 1993). There is no analytical data. At Araxa, Minas Gerais, Brazil (Woolley, 1987; p.66), zirconolite occurs as prismatic crystals with anastase in a glimmerite, and also with pyrochlore, baddeleyite and apatite in a calcite carbonatite (Dr. A.N. Mariano, personal communication, 1993). There is no analytical data. Metasomatic Rocks. Zirconolite has been reported from metasomatic rocks at nine localities, although analyses from only seven of these have been published. In the Mt. Melbourne Volcanic Field, Victoria Land, Antarctica, zirconolite occurs as isolated grains with a maximum grain diameter of 0.08mm within ultra-potassic veins in a mantle xenolith from a basanite host (Hornig & Worner, 1991). The major vein-forming minerals are leucite, plagioclase, nepheline, Mg-ilmenite, apatite and titaniferous mica. Analyses MI1-M7 (Table 3) are selected from Hornig & Worner (1991) as being the least ‘contaminated’ by adjacent silicate minerals. At the contact between granodiorite with gneisses and marbles in the Bergell aureole, Switzerland/Italy, chemically discontinuously-zoned zirconolite is observed, typically 30—40um in diameter, associated with allanite and titanite, in a skarn (Gieré, 1986; Williams & Giere, 1988). The major minerals of the skarn are calcite, spinel, phlogopite and anorthite. The microprobe analyses M8—M22 (Table 3) are unpublished data (CTW) of eleven grains from three discrete zones, the averages of which are published in Williams & Gieré (1988). In the O6etztal-Stubai complex, Austria, within polymetamorphic metacarbonates, zirconolite and baddeleyite occur in several mineral assemblages consisting of chlorite, ilmenite, apatite, spinel, phlogopite, titanian clinohumite, olivine, calcite, dolomite and diopside (Purtscheller & Tessadri, 1985). Analyses M23—M27 (Table 3) are the wavelength-dispersive microprobe analyses given by Purtscheller & Tessadri (1985). In the Adamello contact aureole, Italy, compositionally-zoned and corroded zirconolite occurs in two zones within a Ti-rich vein in dolomite marbles at the contact with a tonalite intrusion (Gieré, 1990a). In the phlogopite zone, zirconolite is always found associated with phlogopite and calcite (tdolomite), and occasionally with geikelite, rutile and fluorapatite. In the titanian clinohumite zone, zirconolite occurs with titanian clinohumite, spinel, calcite, dolomite, pyrrhotite, geikelite, fluorapatite and minor secondary chlorite. A detailed mineralogical and chemical description is given in Gieré & Williams (1992), and analyses M28-M66 (Table 3) are wavelength-dispersive microprobe analyses from Gieré (1990b). At Koberg Mine, Bergslagen, Sweden, yttrian zirconolite occurs as anhedral grains, predominantly 20-30pm in diameter, in an altered phlogopite-rich sample associated with a marble skarn (Zakrzewski et al., 1992). The low analytical totals (analyses M67-M94, Table 3) suggest the zirconolite is hydrated (Zakrzewski et al., 1992). At the agpaitic alkaline syenite complex of Lovozero, Kola, zirconolite (reported as ‘zirkelite’) has been described in a mineral assemblage including rosenbuschite, from the contact metasomatic rocks of the massif (Semenov ef al., 1963). Analyses M95-M96 (Table 3) are the wet chemical data of Semenov et al. (1963). In a dolomitic marble from the Neichi mine, Iwate Prefecture, Japan zirconolite is associated with geikelite and baddeleyite, forsterite and spinel (Kato & Matsubara, 1991). The composition of zirconolite is close to the theoretical composition (analyses M97-M98, Table 3, from Kato & Matsubara (1991). At Ser Rondane, Antarctica, Grew et al. (1989) report zirconolite (qualitative analysis only, cf. p. 119) from a marble affected by metasomatic processes which had introduced rare metals. Associated minerals include dissakisite-(Ce) (Grew et al., 1991), calcite, dolomite, phlogopite, chlorite, ilmenite-geikelite and spinel. Rekharskiy & Rekharskaya (1969) discovered zirconolite (reported as zirkelite) intergrown with jordisite and abundant metasomatic pyrite, as veins in zones of altered trachytic to rhyolitic volcanic rocks. Locality details are not reported. Kinny & Dawson (1992) report the occurrence of zirconolite, as a rare accessory phase associated with zircon and baddeleyite, in a veined and metasomatised harzburgite xenolith from Kimberley, southern Africa. The metasomatism is MARID-related (Kinny & Dawson, 1992) and considered to be associated with kimberlite magma. Analysis M99, Table 3, is the unpublished mean of 6 microprobe analyses (Prof. J.B. Dawson, pers. comm., 1993). Rubin et al. (1993) report zirconolite occurring as inclusions in phlogopite in one sample of a complex skarn from the Ertsberg District of Irian Jaya, Indonesia. No analytical details are given. Placer Deposit. Zirconolite is reported as millimetre-sized crystals from two placer deposits. At Jacupiranga, Sao Paulo, Brazil, zirconolite (named as the C.T. WILLIAMS AND R GIERE new mineral ‘zirkelite’), is found with baddeleyite and perovskite in the heavy mineral fraction from pyroxene sands of “. . . the decomposed magnetite-pyroxenite of Jacupiranga . . .’ (Hussak, 1895; Hussak & Prior, 1895; see also Pudovkina et al., 1974). Analyses Pl, P2 are unpublished wavelength-dispersive microprobe analyses (CTW) of separate grains (80142). The wet chemical analysis given in Hussak & Prior (1895) is not included here, as the chemical separation techniques employed in the analysis could only provide qualitative data for ZrO, and TiO,. In Sri Lanka, zirconolite (reported as ‘zirkelite’) was observed from two ‘gem gravel’ localities in the Sabaragamuwa Province: at Walaweduwa in the Bambarabotuwa district, and in southern Sabaragamuwa (Blake & Smith, 1913). Analyses P3, P4 (Table 3) are from Bambarabotuwa, and P5-P7 from Sabaragamuwa; analyses P8, P9 are microprobe data from Lumpkin et al. (1986); analysis P10 is an unpublished (CTW) wavelength-dispersive microprobe analysis of several grains (BM1905,361). ‘Other’ Rock Types. Sapphirine Granulite: Zirconolite occurs as acicular grains in a mineral assemblage including sapphirine, spinel, enstatite and minor phlogopite from a sapphirine granulite nodule sampled from a xenolith-rich norite wall zone in the Archaean Vestfold Hills, east Antarctica (Harley, 1994). The zirconolite is considered to have been a relatively early crystallising phase during the melt crystallisation history of the entrapped granulite xenolith. Analyses SG1-SG3 (Table 3) are from Harley (1994). Alnoite: Thin (1 um) rims of zirconolite (no compositional details given) are reported as overgrowing a baddeleyite crystal from the ile Bizard alnodite, Québec, Canada (Heaman & Le Cheminant, 1993). Although the baddeleyite crystals are considered to be mantle-derived xenocrysts, the associated overgrowths, which include perovskite and melilite as well as zirconolite, are considered to have formed after exposure to the alnoite magma. Lunar. Zirconolite has been observed in several lunar samples, (cf. review by Frondel, 1975). Data from the literature are presented as analyses LI-L13 (Table 3). It occurs in coarse-grained basalts (Apollo 11, 15 and 17), in a feldspathic Table 1 Range of chemical variation in natural zirconolite Terrestrial* Lunar Maximum Minimum Maximum Minimum _ Theoretical composition CaO 16.54 01.83 10.70 02.63 16.5 REE,O, 23.66 0.00 31.98 04.74 PbO 00.80 00.00 02.19 00.00 TOs 2228 00.00 02.34 00.00 UO, 23.98 00.00 01.16 00.00 ZrO, 44.18 DR? 45/40 29.80 36.3 HO, O13 00.00 01.34 00.00 TiO, 44.91 13.56 34.60 25.48 47.2 MgO _— 03.04 00.00 01.15 00.00 ALO, 03.47 00.00 01.60 00.35 FeO 10.20 00.00 11.40 04.23 Fe,0, 09.58 01.08 x és Nb,O;, 27.00 00.19 04.34 00.00 Ta,O, 05.83 00.00 00.40 00.00 wo, 01.44 00.00 z = *Excluding kimberlite analyses K1 and K2 (see text) ZIRCONOLITE Table 2 Details of zirconolites from terrestrial and lunar occurrences Nn Analysis Reference Reference Rock type Sample locality Country number (Occurrence) (Analytical data) Kimberlite Monastery South Africa K] Raber and Haggerty (1979) Raber and Haggerty (1979) Kimberlite Mothae South Africa K2 Raber and Haggerty (1979) Raber and Haggerty (1979) Kimberlite Kimberley South Africa K3 Raber and Haggerty (1979) Raber and Haggerty (1979) Ultrabasic Laouni Algeria U1-U2 Lorand and Cottin (1987) Lorand and Cottin (1987) Ultrabasic Rhum (+) Scotland U3 Williams (1978) Fowler and Williams (1986) Gabbro Pegmatite St. Kilda Scotland Gl Harding er al. (1982) Fowler and Williams (1986) Syenite Glen Dessarry Scotland Sl Fowler and Williams (1986) Fowler and Williams (1986) Syenite Arbarastakh, Aldan Russia S2-S4 Borodin et al. (1960) Borodin er al. (1960, S2); Gaidukova et al. (1962), $3); Wark e7 al. (1973, S4) Syenite Fredericksvark (*) Norway S5 Berzelius (1824) Brogger (1890) Syenite Langesundfjord (+) Norway S6-S7 Brogger (1890) CTW (unpubl. data) Syenite Campi Flegrei Italy - Mazzi and Munno (1983) = Nepheline Syenite Pine Canyon, Utah U.S.A. NSI-NS6_ Agrell e¢ al. (1986) CTW (unpubl. data) Nepheline Syenite — Chilwa Island (+) Malawi NS7-NSI11_ Platt et a/. (1987) Platt et al. (1987) Nepheline Syenite Elk Massif Poland ~ Dziedzic (1984) No analytical data Nepheline Syenite Tchivira Angola NS12—-NS17 Mariano and Roeder (1989) CTW (unpubl. data) Nepheline Syenite _ Pilanesberg, Transvaal South Africa = Mariano (pers. comm., 1993) No analytical data Nepheline Syenite Tre Croci, Latium Italy NS18—NS22 Parodi (pers. comm., 1993) CTW (unpubl. data) Carbonatite Afrikanda, Kola (**) Russia Cl-C4 Borodin et al. (1956) Borodin e7 al. (1956, C1, C2); Bulakh et al. (1960, C3); Wark er al. (1973, C4) Carbonatite Vuoriyarvi, Kola(+) Russia C5-Cl1 Borodin er al. (1960) CTW (unpubl. data, CS—C7); Borodin etal. (1960, C8, C9); Bulakh er al. (1960, C10); Kapustin (1964, C11) Carbonatite Sayan Province Russia eZ Gaidukova et al. (1962) Gaidukova et al. (1962) Carbonatite Seblyavr, Kola Russia (GjI8) Bulakh et al. (1960) Bulakh et al. (1960) Carbonatite Kodvor, Kola (+) Russia C14-C34 Kukharenko et al. (1965) Kapustin (1980, C14, C15); Kukharenko etal. (1965, C16); CTW (unpubl. data, C17-C34) Carbonatite Schryburt Lake (+) Canada C35-C58 Williams and Platt (in prep.) CTW (unpubl. data) Carbonatite Santiago Island Cape Verde Republic C59 Silva (1979) Silva and Figueiredo (1980) Carbonatite Phalaborwa(+) South Africa C60-C66 ~—- Verwoerd (1986) CTW (unpubl. data); G. Bayer (unpubl. data) Carbonatite Sokli Finland C67-C70 _—~ Vartianen (1980) Hornig-Kjarsgaard (unpubl. data) Carbonatite Kaiserstuhl Germany C71-C72 Keller (1984) Keller (1984, C71); Sinclair & Eggleton (1982, C72) Carbonatite Hegau Germany C73-C88 Keller et a/. (1990) CTW (unpubl. data) Carbonatite Prairie Lake, Ontario Canada C89 Mariano and Roeder (1989) Mariano (unpubl. data) Carbonatite Howard Creek, B.C. Canada C90-C135 Mariano (pers. comm., 1993) CTW (unpubl. data) Carbonatite Cummins Range, Kimberly __ W. Australia - Mariano (pers. comm., 1993) No analytical data Carbonatite Catalao, Goias Brazil ~ Mariano (pers. comm., 1993) No analytical data Carbonatite Araxa, Minas Gerais Brazil C136—-C161 Mariano (pers. comm., 1993) CTW (unpubl. data) Metasomatic Mt. Melbourne Antartica MI1—-M7 Hornig and Worner (1991) Hornig and Worner (1991) Metasomatic Bergell (+) Switzerland/Italy M8-—M22 ~~ Gieré (1986) Williams and Gieré (1988) Metasomatic Oetzal-Stubai Austria M23—-M27 Purtscheller and Tessadri (1985) Purtscheller and Tessadri (1985) Metasomatic Adamello (+) Italy M28—-M66_ Gieré (1990a) Gieré (1990b) Metasomatic Koberg, Bergslagen (+) Sweden M67—-M94_ Zakrzewski et al. (1992) Zakrzewski et al. (1992) Metasomatic Lovozero, Kola Russia M95-M96_ Semenov e¢ al. (1963) Semenov et al. (1963) Metasomatic Neichi, Iwate Prefecture Japan M97-M98___ Kato and Matsubara (1991) Kato and Matsubara (1991) Metasomatic Sor Rondane Antarctica ~ Grew et al. (1989) No analytical data Metasomatic ? Former USSR - Rekharskiy and Rekharskaya (1969) No analytical data Metasomatic Kimberley South Africa M99 Kinny and Dawson (1992) Dawson (unpubl. data) Metasomatic Irian Jaya Indonesia — Rubin e7 al. (1993) No analytical data Metamorphic Vestfold hills East Antarctica SGI-SG3 Harley (1994) Harley (1994) Alnoite ile Bizard, Quebec Canada - Heaman and LeCheminant (1993) | No analytical data Placer (1) Jacupiranga (***) (+) Brazil P1—P2 Hussak (1895); CTW (unpubl. data) (3) Hussak and Prior (1895) Placer (2) Sabaragamuwa Province(+) Sri Lanka P3-P10 Blake and Smith (1913) Blake and Smith (1913, P3—P7); Lumpkin e7 al. (1986, P8—P9): CTW (unpubl. data, P10) Lunar Apollo 11 Landing Site Moon LI-L4 Lovering and Wark (1971) Wark et al. (1973) Lunar Apollo 12 Landing Site Moon LS Busche et al. (1972) Busche et al. (1972) Lunar Apollo 14 Landing Site Moon L6-L7 Busche er a/. (1972) Busche et al. (1972); Wark et al. (1973) Lunar Luna 20 Landing Site Moon L8 Roedder and Weiblen (1973) Roedder & Weiblen (1973) Lunar Apollo 15 Landing Site Moon L9-L11 Brown et al. (1972) Brown et al. (1972); Wark et al. (1973) Lunar Apollo 16 Landing Site Moon = Lovering and Wark (1974) No analytical data Lunar Apollo 17 Landing Site Moon L12-L13. Meyer and Boctor (1974) Meyer and Boctor (1974) De (*) type POLY MIGNITE ) (**) type ZIRCONOLITE j (***) type ZIRKELITE (+) In BMNH collection now all renamed as zirconolite (see Bayliss ef a/., 1989) (1) Heavy mineral fraction from pyroxene sand; (2) Heavy mineral fraction from alluvial deposits: two separate localities; (3) Analysis from Hussak & Prior (1895) not included as method used could not adequately distinguish Ti from Zr. a ezle peo vy SPO ry Ly0'v 960 7 Lvov 1SO'v LcO'¥ SPO 096° 6207 000 sop £80 p cv0 v 0SO0'y bZO'v 600 7 00 yO6€ S80’ WLOL 4) SG) SELZ 7002 £46 1 8002 £L07~ 086 t 086 1 790°2 cl? 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LoL 92°0 ceo bE L SOL 3 co'0 66€ Sty ee 9F OL Le os Le oo'cL os Zl 1466 oe OL © elo elly 09°92 02°82 vS'OL Sez SLé 4€0 620 v0 L4€0 WOILSYOSHL eh (4a 648°0 Z€z0 6L LOL eL6e 988'L 000 €200 8z00 vv 0 600°0 8ze lL Soc L 9200 6e7 1 e280 so0'0 s00'0 g00°0 o4s0 ez 0 suabAxo J 0} suoneod 2066 61 WLol +¢!1 WNS +9M +Ge1 +SON +019 +EIV +€94 +294 +cUlN +Z6W +49!S +Hll +¥JZ WNS +93H +pJZ +289 WNS +9 +p +24d +€S55N+A +289 WLOL »SJBYIO,, O2H OzO1'eN) zon zouL Odd fOM sozel ZcOJH £0cn) €07gA cocwL pieyZE| £€0Z0H e0zd eozql €0ZPD e€07ng COZWS €OZPN £0dd £0299 €0ze1 SOZQN fOUZ £OZA £0294 0283 OUW LeyAye) zOllL Ok8D zo!ls £Oclv 2D. peridotite (Apollo 12), in lithic fragments (Apollo 14, LUNA 20), and in a metamorphosed breccia (Apollo 16). Zirconolite is often associated with baddeleyite as small discrete crystals, no larger than 50um in diameter, and is considered to have crystallised at a late stage from interstitial liquids in the lunar basalts (e.g. Busche ef al., 1972). Lunar zirconolites are generally rich in Y and heavy-REE when compared with terrestrial zirconolites (Lovering & Wark, 1974; Kochemasov, 1980; Fowler & Williams, 1986). The majority of the lunar zirconolites have EREE** > 50% of the Ca site, and with Y being the dominant REE, these may be considered as zirconolite-(Y). DISCUSSION AND CONCLUSION Zirconolite occurs as an accessory mineral only, generally less than 0.1mm in diameter, but from a wide variety of rock types. Its small size and low modal abundance means that it can be easily overlooked using traditional optical microscopy. However, with the increasing accessibility of analytical scanning electron microscopes (usually with a backscatter electron detector attached), zirconolite, even if present at a very low modal abundance, will be readily observed, because its backscatter component is considerably higher than the majority of the rock-forming minerals. It is probable therefore, that the number of zirconolite occurrences will increase significantly in the near future. It is also evident that zirconolite is often zoned, and/or finely intergrown with other minerals, and early bulk chemical analyses were unable to characterise fully the chemical variability of this mineral. Microprobe analyses, together with a detailed SEM investigation, is therefore essential in any study. It is generally recommended that microprobe analysis is performed using wavelength-dispersive means, because zirconolite can accommodate more than 30 elements at the 0.1 to 1.0 wt.% concentration level (which in energy-dispersive electron microprobe analysis is close to, or below, the detection limit), However, quantitative analysis of sub-micron zones has been successfully undertaken using an energy-dispersive analytical transmission electron microscope (Lumpkin et al., 1994). As can be seen from the data for natural zirconolite, the range of elements substituting, and the degree of substitution are extensive. The most commonly occurring elements, and therefore the minimum that should be reported in any microprobe analysis of zirconolite are: Mg, Al, Si, Ca, Ti, Mn, Fe, Y, Zr, Nb, Hf, Ta, W, Pb, Th, U and of the REE, La, Ce, Pr, Nd, Sm, Gd. It should also be noted that Cr and Zn are present in some zirconolites: Cr predominantly from lunar samples, and Zn occasionally from metasomatic samples (e.g. Zakrzewski et al., 1992). H,O has been reported in wet chemical analyses of separated grains (e.g. Borodin et a/., 1960; Bulakh et al., 1960), and has also been inferred from low analytical totals of microprobe data (e.g. Platt et al., 1987; Zakrzewski et al., 1992). Na and K, although also quoted in some wet chemical analyses of separated grains, have not been observed in microprobe analyses. It is probable therefore, that Na and K are not present to any significant extent in zirconolite. It is of note also, that Sr and Ba generally do not occur in natural zirconolite, and Pb only rarely does so. These elements might have been expected to substitute more readily for Ca, but it appears that the Ca C.T. WILLIAMS AND R GIERE structural site does not easily accommodate 2+ cations larger than Ca. The valency state of Fe in zirconolite is unclear: where measured directly on mineral separates, both FeO and Fe,O, are present. It is evident that zirconolite, although invariably occurring only as an accessory or ‘trace’ mineral in a range of rock types, is able to accommodate many incompatible elements, such as © REE, ACT, Nb, Zr, Hf, Ta to concentration levels whereby it can become a major repository for these elements. As such, it has the potential for playing a _ significant role in the petrological/geochemical evolution of those rock-types in which it crystallizes. Several studies have provided evidence that | zirconolite can reflect changes in the composition of the fluid during its evolutionary history, both in metasomatic systems - (Williams & Gieré, 1988; Gieré & Williams, 1992), and in magmatic fractionation processes (Platt et a/., 1987). It is hoped that this review and compilation will prove useful | as a comparative database for geologists who discover zirconolite in their samples, and also to material scientists working on various SYNROC projects, in order that they can compare laboratory-based experiments on synthetic zirconolite with studies of the natural forms of zirconolite. This database is available in a computerised format from | CTW. We would be grateful also to receive any additional | analytical data and/or material from new occurrences of zirconolite, in order to periodically update this database. ACKNOWLEDGEMENTS. We are very grateful to Professor G. Bayer (ETH, | Zirich) for providing us with zirconolite samples from Phalaborwa, and — also for some unpublished data, to Alf Olav Larsen (Porgrunn, Norway) | for samples from Langesundfjord, to Dr. S.L. Harvey Edinburgh, Scotland for providing information on zirconolite from East Antarctica, to Professor J. Keller (Freiburg, Germany) for permission to analyse zirconolite from Hegau, to Dr I. Hornig-Kjarsgaard (University of Mainz, Germany) for allowing us to include her unpublished data from Sokli, to Professor J.B. Dawson (Edinburgh, Scotland) for unpublished | data, to Dr E.S. Grew (University of Maine, USA) for providing material from Ser Rondane, Antartica, to Dr. A.N. Mariano (Carlisle, Massachusetts, USA) for providing samples and information on several zirconolite localities, and for some unpublished data, to Dr G.C. Parodi (University of Rome, Italy) for samples from Latium, Italy, and to the | Kovdor Mining Museum, Kola Peninsula for material from Kovdor. We l further wish to thank Drs M. Welch, A.R. Woolley and R.F. Symes and | other colleagues at The Natural History Museum, London, also to | Professor Andrei Bulakh (University of St. Petersburg) for comments | and suggestions which have improved the manuscript, and to Greg | Lumpkin (ANSTO, Australia) for discussions regarding synthetic | zirconolite. This study forms part of a British/Swiss Joint Research | Programme, and we gratefully acknowledge funding provided by the | Schweizerischer Nationalfonds and the British Council (Grant No. 83BC-033381). REFERENCES Agrell, S.O., Charnley, N.R. & Rowley, P.D. 1986. The occurrence of hibonite, perovskite, zirconolite, pseudobrookites and other minerals in a metamorphosed hydrothermal system at Pine Canyon, Piute County, Utah, USA. Abstract. Mineralogical Society Bulletin, 72: 4. Bayliss, P. & Levinson, A.A. 1988. A system of nomenclature for rare-earth mineral species: revision and extension. 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British Museum (Natural History), 216pp. ) Zakrzewski, M.A., Lustenhouwer, W.J., Nugteren, H.J. & Williams, C.T. 1992. — Rare-earth minerals yttrian zirconolite and allanite-(Ce) and associated minerals from Koberg mine, Bergslagen, Sweden. Mineralogical Magazine, 56: 27-35. Zhuravleva, L.N., Berezina, L.A. & Gulin, Ye. N. 1976. Geochemistry of rare and radioactive elements in apatite-magnetite ores in alkali-ultrabasic complexes. Geochemistry International, 13: 147-166. Bull. nat. Hist. Mus. Lond. (Geol.)52(1): 25-49 Issued 27 June 1996 A review of the stratigraphy of Eastern Paratethys (Oligocene—Holocene) R.W. JONES BP Exploration, Uxbridge One, 1 Harefield Rd., Uxbridge, Middlesex, UBS 1 PD, U.K. M.D. SIMMONS Department of Geology and Petroleum Geology, University of Aberdeen, Meston Building, King’s College, Aberdeen, AB9 2UE, U.K. CONTENTS Absolute @hronostrati rap liv? wsreet ean meet se tee cc eee eee caster cere sucsunceetht testa cn cs eeeae re ate one EEE TST Ter ago te nsviste eee 2 Micropalaeontological Biostratigraphy and Palaeoenvironmental Interpretation BIOS ELAbT SAP y: Wein e.ces secte cee erase crac coset nae ea cee eaee deat sath ow ctlcg decison atbeacavosteaeteta ses oaeenteste ts bene cousenceeneis ras eduevunerstennsensi vas Ralacoenvironmental interpretations meta -tcccceccets cet ececsatceteetae oetacce sae t ert Tee ee Reece ee oT 2 INOnAVIGT INE ENV IFONIMCNiSimettetece ete eee eee ee Quasi- Marine and Marine Environments Glimatosinaticnaph yg cc. ceeeeetence eee eee a= IMMA PNELOStratlOrap ly: cc eeccesseacsescexeces << cee sues eve duev sess ussacace'es ities a Geuavesaeesosee ine ia sutdea sack fivede dane sultcusasavesoece thant saventbecuensabeans 29 Oxo PenilSotoperS trratsl oirraapo ba yseeceere aes seer ee wearers cesar ve suse hsonie hanes TREE Saba vas ate eg aseda sede denies gedlee dua cnua teesbieele taeieees 29 Sequences tratipraphyjandibalacOPeo pirap liye este eee e sence cere ane ee eee ele ree enn ne ne o e 29 (GLY eee ene reece eee eae ra eee aac dos sae svn Sawen ust acabsctonadavsdlte Mek anu sen cet tu sc tees vith sev svstaseved aides vissacdeiteaes SO Gea reA SAINT EAM teres tence Se Se MRRP ccc Saas css oeuid wove sda ice Stale dle tawaucd ames datu'an sce seessaneenaceass = ee aes ceessnves Sdewteiedens« duseeesuceut oD Konkian SO GRYSTAETSONT _ aasaacsshobepaséeseonasce Josaacanecgecooase add ac SUSDEeSEDE ECE ana caESe SE UeEiceo aoc ccEsnERE cada CoSoReica gee aToC oan on ao nes ac Cone teceuccooo eee ESCORTS INTC O Gleam mmeri ee sere tere eee em ne ese aes Says sctectsenes i anesuite ce ta Suvae ete astsetae caseeead ee uraenrenetacteten coh wor aes Secon angie O TROSEQTENT. . . cocoscse nto ERea ese antcncontoe cases EAS a ie en ene EEE EERE DOERR E CE cape cone CaciicacracceccE Cha RE Ge sccERSECE Recerca eek EERE EEEer ES ts LO TTOTITETHEND), «carob oeetaeoecatds tsocbbeGanen Eb Pad onos coe e OOSERES EEE SeecCECEA Spacer Coa SCo eS ERS CR EE creator cece RRC cae RT Goce eREcenn cori eee ee rE Ramee A Akchagylian to Khvalynian (Caspian Sea) JANI) FASS /IVEANT “Sepeeanoncnasaceceticeacedcdodect Ceonceo abe a oLca ce cea EEEROTiEnY er DESEO RODRE DO Ce Be AGO OREEE coe EEC ECS a GARACER es ReescROEEESET noes FOE ESE ED OSEE PANTS ROLOMMAM wae eecoachee redone te naa states tect cares rou bat dares Sia erate Sapa Sa eae eae ete ae eavawsaze i seteeuas Vakay oA aa TR EMIRATE 42 [BATT gadsaganteptese ar acaronespcee casa eS CGARDOnES HO UEEEERBE EBS EEE EEE cERCe SSE cOcRE Sao Beer cotocc ao neo coca RE Sct Reco een eee eee ceeCooccs 42 Kehazanians Girkaniandikehwvaliynianteencececscsse-cotcccescecaceuesenesaenerc snes see cre steretent atrl-cea os cress sccesseenc-raerynsmekanentee seaecees ss 42 KenvalniktanktopNeoeuxiniany (BlacksSea) esses. seesecccesecexscaet ces eesteenc snare nectar screenees etretcny cevsccstetssessunaneenneenestes 42 GUY ANT AGL eAN UN Sess osene Se ae race Sic ner Secs eee ys Se ntaes Peron seat aah a aD ntsc tos ating cevacsieldntoaanes: dactszanttoien sacheedtatss 42 Gurianepee ers sc ex2ce Chaudian Wannlaniana karan gatiantand’Neoeuxiiiamees eesti cer tes strc caseestenntetntes eoeietes ae neetaeosenncnannar tar cirneatenteehamednadirseseat cet 43 ING RIOR TOTES: - cocececcoseccsoedbenda99006839600035gS CCOREE Sac ORGEORECICCCOBEEBOE-BORCCEGEEADUS Seb anc BEELO5050oq06 60nd donobononone oaaSaccenoCKESasoosImoueNG 43 IRGICISINSES) ccesasqosoonenecooscbocosoeee ee aeebeD PN scat eae MERA SSSGR Fe ooo CAA Minn iss sn don tbatie re Mee d tnaee nee v nee Ca eee GEERT eee eee se toes 43 Synopsis. All available data pertaining to the regional stratigraphy of Eastern (Ponto-Caspian) Paratethys, much of it in sources not freely available in the west, is reviewed. Particular emphasis is placed on the South Caspian. Where possible, regional datums are calibrated against global standards. An attempt is made to place the regional stratigraphy in a (global) sequence stratigraphic framework for the first time. Palaeogeographic reconstructions are given for selected time-slices. © The Natural History Museum, 1996 26 INTRODUCTION The Tethyan Ocean began to close in the Eocene as a result of plate collisions along the southern margin of the Eurasian Supercontinent that ultimately gave rise to the formation of the mountain chain extending from the Alps in the west to the Himalayas in the east. The initial response to these plate collisions was the formation of a suite of east-west trending sedimentary basins extending from Austro-Hungary in the west to Central Asia in the east, collectively constituting the intracontinental Paratethyan Sea (Fig. 1). Subsequent tectonic uplift (enhanced by eustatic shallowing) through the Mio-Pliocene led to widespread marginal- to non- marine sedimentation. Ultimate severance of connections to the world’s oceans led to the evolution of largely endemic faunas and floras (in particular in Eastern Paratethys, which was more isolated than Central Paratethys). This renders stratigraphic correlation between established Mediterranean and Paratethyan stages extremely difficult. The problem is locally compounded by confusion between chronostratigraphic and lithostratigraphic nomenclature. In this paper we review all available data, much of it in sources not freely available in the west, on the regional stratigraphy of Eastern Paratethys (Fig. 1). We place particular emphasis on the South Caspian, an area in which the western oil industry is showing a growing interest, and one with which we, through our industrial work and our academic contacts in the Former Soviet Union, are particularly familiar. We give an indication of the palaeontology of each regional stage. For the sake of brevity and because of their stratigraphic utility, we concentrate on various groups of microfossils, though we acknowledge that macrofossils, especially molluscs, also have stratigraphic value (see, for instance, Ali-Zade (1954), Azizbekov (1972), Ali-Zade et al. (1986), (Azerbaijan); Lupov et al. (1972) (Turkmenia); Andreescu (1981) (Dacic Basin); and Ozsayar (1985) and Taner (1985) (Turkey)). A forthcoming paper (Simmons ef a/. in press) will document in detail the micropalaeontological (including nannopalaeontological and palynological) and macropalaeontological zonation of the Paratethyan in Bas Black Sea Fig. 1 R.W. JONES AND M.D. SIMMONS Neogene to Pleistogene sediments of Azerbayan. We attempt to place the regional stratigraphy in a global framework by calibrating biostratigraphic and magnetostratigraphic datums against Central Paratethyan and Mediterranean standards, and by _ suggesting possible calibrations between regional sequence boundaries and flooding surfaces, and the global sequence stratigraphic framework and eustatic sea-level curve of Haq et al. (1988). ABSOLUTE CHRONOSTRATIGRAPHY Notwithstanding the efforts of such authors as Steininger & Papp (1979), Chumakov er al. (1984, 1988, 1992a—b) and Vass (1985), there is no established comprehensive absolute chronostratigraphic time-scale for Eastern Paratethys. Thus, in Eastern Paratethys, absolute chronostratigraphic dating is often only possible by calibration of regional stratigraphic datums against global standards. We have attempted to calibrate regional datums against the Haq et al. (1988) timescale, which is the most up-to-date timescale that conveniently integrates bio-, magneto- and sequence- stratigraphic data. The confidence with which this sort of calibration can be made varies considerably with stratigraphic interval (see below). MICROPALAEONTOLOGICAL BIOSTRATIGRAPHY AND PALAEOENVIRONMENTAL INTERPRETATION Biostratigraphy Those groups of planktonic organisms traditionally used in the biostratigraphic zonation of the Cenozoic (planktonic foraminifera and calcareous nannoplankton) are restricted in their development in Paratethys because of the isolated & O Aral Sea Possible Connection to Amu Darya and Tarim Basins Geological sketch map of the Paratethyan Basin in the Oligocene. Modified after Steininger & Papp (1979). C = Central Paratethys; E = Eastern Paratethys. Eastern Paratethys can be considered as including the Pontian (Black Sea) Basin and the Caspian Basin. Throughout most of the Miocene, the eastern limit of the Paratethyan Basin was probably the Caspian Basin, but in the Late Pliocene at least it was probably further east once more. REVIEW OF STRATIGRAPHY OF EASTERN PARATETHYS geological evolution of the region. Only locally or periodically (as in the Maykopian, Maeotian and Kuyalnikian/Akchagylian (see section below on sequence stratigraphy)) are they sufficiently well developed to enable ties to global biostratigraphic zonation schemes (Blow (1969) for planktonic foraminifera; Martini (1971) for calcareous nannoplankton). Biostratigraphic zonation in Eastern Paratethys relies largely on facies-dependent benthonic foraminifera and, especially in the marginal- to non- marine environments of the Mio-Pliocene, benthonic ostracods and terrestrially-derived pollen and spores (see section below on sequence stratigraphy). Important ostracod references include those of Livental (1929), Sveier (1949), Agalarova (1956, 1967), Suzin (1956), Agalarova et al. (1961), Mandelstam ef al. (1962), Faridi (1964), Imnadze (1964, 27 1974), Sheydayeva-Kuliyeva (1966), Rozyeva (1971), Gramann (1971), Karmishina (1975), Vekua (1975), Krstic (1976), Olteanu (1978), Imnadze & Karmishina (1980), de Deckker (1981), Jiricek (1984), Mamedova (1984, 1985, 1988), Dzhanelidze et al. (1985), Aliyulla et al. (1985) and Yassini (1986). Important pollen and spore and associated palynomorph references include those of Ramishvili (1969), Dzhabarova (1973, 1978, 1980), Grichuk (1973, 1984), Wall & Dale (1973), Ananova (1974), Shikmus-et al. (1977), Koreneva & Kartashova (1978), Traverse (1978), Shchekina (1979), Abramova (1982, 1985), Yakhimovich et al. (1983), Grichuk et al. (1984), Khotinskiy (1984), Mamedov & Rabotina (1984a—b), Shatilova (1984), Ananova et al. (1985), Ivanova (1985), Sirenko & Turlo (1986), Bludorova et al. (1987), Naidina (1988, 1990a—b, Fig.2 Vegetation belts in the former Soviet Union. After Knystautas (1987). A.D. = Amu Darya; S.D. = Syr Darya. 28 1991la—c) and Malaeva & Kulikov (1991). Other locally stratigraphically useful fossil groups include otoliths (Brzobohaty, 1983), Problematica (Bolboforma (Szezechura, 1985; Spiegler & Rogl, 1992)), siliceous microfossils (diatoms (Shishova, 1955; Ushakova & Ushko, 1971; Gasanova, 1965; Rasulov, 1986), radiolarians (Slama, 1983), silicoflagellates (Dumitrica, 1985) and sponge spicules (Riha, 1983)), and, in non-marine environments, vertebrate remains (Camelopardis, Felis, Gazella, Hipparion, Hyaena, Mastodon, Mesopithecus, Rhinoceros, etc.) (Kretzoi, 1985; Steininger, Rabeder & R6gl, 1985; Bernor et al., 1987, 1993; Lindsay et al., 1989; Régl et al., 1993) and charophytes (R6gl et al., 1993). Ali-Zade et al. (1994a, 1994b, 1995, in press), Reynolds et al. (in press) and Simmons et a/. (in press) give further details of the Neogene biostratigraphy of Eastern Azerbaijan. Palaeoenvironmental Interpretation Non-Marine Environments. Non-marine environments are characterised by fresh-water ostracods such as Aglaiocypris, Candona, Candonella, Cyclocypris, Cypria, Eucypris, Ilyocypris, Pseudostenocypria and Zonocypris, and terrestrially-derived pollen and spores. Pennate diatoms, fresh-water gastropods and terrestrial vertebrate remains may also be found. Palaeoclimate can be inferred from the distribution of vegetation types as inferred from pollen and spores. At the present-day, the distribution of vegetation types is determined chiefly by climatic factors (temperature (latitude, altitude), aridity). Thus, for instance, birches characterise the cold ‘forest-tundra’ of the extreme north, diverse coniferous and deciduous types the ‘taiga’ of the central area, and grasses and shrubs the arid treeless ‘steppe’ and semi-desert to the extreme south (Figs 2-3). Quasi- Marine and Marine Environments. Deposition in oligo- to meso- haline (hereafter ‘quasi-marine’ (brackish, reduced salinity)) environments prevailed in the Paratethyan Basin (especially in the Caspian) throughout much of its geological evolution because of its restricted connection to the open ocean (see above). However, water depths and sedimentary regimes may have been similar to those of the normal marine realm, and, moreover, deposition under normal or near-normal marine conditions did take place at times (e.g., Maykopian and Akchagylian). Palaeosalinity can be inferred from diatoms (e.g., Ushakova & Ushko, 1971; Schrader, 1979) or from foraminifera and ostracods ranging through to the Recent (see below). Palaeosalinity and/or palaeotemperature curves are presented by Semenenko (1979), Chepalyga (1985) and Demarcq (1985). Quasi-marine environments in the Black Sea and Caspian Sea are characterised by the benthonic foraminiferal genus Florilus, and some species of the genera Ammobaculites, Ammoscalaria, Ammonia and Elphidium (salinity tolerance range 1—5 parts per thousand (ppt)), and Miliammina, Haynesina and Rosalina and some species of Nonion s.l. and Quinqueloculina (1—26ppt) (Macarovici & Cehan-Ionesi, 1962; Tufescu, 1968, 1973; Murray, 1973, 1991; Gheorghian, 1974; Yassini & Ghahreman, 1977; Yanko, 1990b), the ostracod genera Cyprideis (2—-14ppt), Maetocythere (4-14ppt), Loxoconcha (5—14ppt), Bakunella, Caspiolla and Cytherissa (11-13/l4ppt) and Graviacypris (12-13ppt) (Gofman, 1966; Yassini, 1986; Boomer, 1993a), and the calcareous nannofossil genus Emiliania (1|ppt) (Bukry, 1974). Normal or environments are near-normal marine R.W. JONES AND M.D. SIMMONS characterised by the benthonic foraminiferal genera Discorbis, Textularia, Bolivina, Bulimina, Brizalina, Cibicides, Gavelinopsis and Trifarina and some species of the genera Ammonia, Nonion s.. and Quinqueloculina (salinity tolerance range 11—26ppt) (Macarovici & Cehan-Ionesi, 1962; Tufescu, 1968, 1973; Murray, 1973, 1991; Yassini & Ghahreman, 1977; Yanko, 1990b). CLIMATOSTRATIGRAPHY Zubakov & Borzenkova (1990) defined a _ series of climatostratigraphic units called ‘climathems’ (some conceptual, some stratotypified (and with representative pollen spectra documented)) which they used in the regional correlation of Eastern Paratethys (see also Zubakov, 1993). Of these, ‘superclimathems’ (SCTs), with an average duration of 200,000 years, are the most useful. SCTs are correlated with half the 370,000-425,000-year cycle of orbital eccentricity, and reflect changes in climate (alternating between ‘cryo-’ and ‘thermo-’ meric (cool and warm respectively)). Zubakov & Borzenkova interpreted pollen spectra dominated by steppe and semi-desert vegetation as being of ‘warm’ aspect (whereas, in fact, they are more characteristic of aridity than high temperature) and those dominated by forest vegetation as being of ‘cool’ aspect (whereas they are in fact more characteristic of humidity than of low temperature). In the Caspian, they found the former to characterise regressions and the latter to characterise transgressions, and therefore correlated regressions with ‘warm’ phases (interglacials) and transgressions with ‘cool’ phases (glacials) (Fig. 4A). Regression during interglacials is possible if sediment supply and subsidence are in equilibrium but evaporation exceeds precipitation and run-off. Transgression during glacials is possible if sediment supply (reduced by rivers freezing) fails to fill the accommodation space created by subsidence. Note in this context that the level of the Caspian has fallen by some 5m since records were first taken in the 1760’s. However, it also appears to have been rising over the last fifty years (currently at a rate of 20cms/year in the South Caspian). Historical records are probably unreliable in a geological context because of the influence of man on the environment. This is particularly true in the case of the Aral Sea, whose level has fallen and whose salinity has risen drastically since the 1960’s owing to abstraction of the headwaters of the feeder rivers (the Amu Darya and Syr Darya) to irrigate the cotton fields of Uzbekistan (see, for instance, Boomer, 1993a—b). Incidentally, as a result of this catastrophic ecological change, eleven species of fresh-water ostracod known to have been living in the Aral Sea thirty years ago no longer live there. Only the quasi-marine species Cyprideis torosa lives there today. The evidence for regressions during interglacials and transgressions during glacials appears somewhat equivocal. An equally strong case, and one more in keeping with a priori expectation from experience in other parts of the world, can be made for correlating regressions with glacials and transgressions with interglacials (Fig. 4B). One key observation in support of this case is the apparent correlation of the major transgressions not only with warm phases (see, for instance, Skalbdyna, 1985), but also with global transgressions (see, for instance, Haq et al., 1988). Pollen spectra of ‘arid’ aspect in glacial sediments and of ‘humid’ aspect in interglacial sediments are explicable in terms of, respectively, contractions and expansions of the forest belt | REVIEW OF STRATIGRAPHY OF EASTERN PARATETHYS Vegetation Type Vegetation Belt (N-S) Ferns, Mosses Tundra Taiga (Coniferous) Taiga (Deciduous) cS Ss) = jaa) D £& o = O x< So 2 Cc & oO = o) ne) © ® = Broad-Leaved Grasses Shrubs & Semi Shrubs Fig.3 Distribution of principal vegetation types in relation to vegetation belts. Compiled from various sources. Bar width provides a measure of abundance. (the former in response to permafrost development). This is also indicated by palaeoclimatic reconstructions for the Late Valdai Glacial and Mikulino Interglacial (Grichuk, 1984; Savina & Khotinskiy, 1984; Velichko, 1984b). An alternative explanation envisages a time lag between the onset of glaciation and the response of vegetation (belts being displaced by up to 2000km.). Similar disequilibrium phenomena have been described from interstadial complexes in the United Kingdom. MAGNETOSTRATIGRAPHY Much magnetostratigraphic data is available from Eastern Paratethys (e.g., Zubakov & Kochegura (1971), Trubikhin (1977), Semenenko (1979), Semenenko & Pevzner (1979), Grishanoy et al. (1983), Rogl & Steininger (1984), Steininger & Rogl (1984), Chepalyga (1985), Chepalyga et al. (1985), lossofova (1985), Pevzner & Vangengeim (1985a, 1993), Senes (1985), Skalbdyna (1985), Vass (1985), Zubakov & Borzenkova (1990) and Trubikhin et a/. (1991a—b)). Theoretically, this sort of data ought to enable a correlation between Eastern Paratethys and the rest of the world (which, as noted above, is difficult to do using the available biostratigraphic data). However, in practice the process is complicated by apparently inconsistent definition and usage of magnetostratigraphic units (polarity epochs). It is beyond the scope of this paper to address this problem in any more detail (instead, we simply quote the published magnetostratigraphic (polarity epoch) ranges for the various regional stages). It is nonetheless evident that the potential exists for a refined magnetostratigraphic subdivision of critical intervals using short-lived polarity reversal ‘episodes’ within the longer-term epochs. OXYGEN ISOTOPE STRATIGRAPHY Theoretically, the ages of the Plio-Pleistogene sediments of Eastern Paratethys are resolvable using oxygen isotope stratigraphic techniques. However, 1n practice, what data there is exists in widely disseminated form and is not particularly useful. SEQUENCE STRATIGRAPHY AND PALAEOGEOGRAPHY Introduction No published sequence stratigraphic schemes exist for the Oligocene-Holocene of Paratethys, although relative sea-level changes and associated aspects of sequence stratigraphy are discussed by Rog] & Steininger (1983), Chepalyga (1985, 1991), Demarcq (1985), Krhovsky (1985), Nevesskaya et al. (1985), Pogacsas (1985), Pogacsas & Revesz (1985), Skalbdyna (1985), Andalibi (1991), Zubakov & Borzenkova (1990) and Klopovotskaya (1991). This section attempts to place the regional stratigraphy of Eastern Paratethys in a (global) sequence stratigraphic framework. It is written in the form of a geological history. Stratigraphic (bio-, climato-, magneto- and sequence- stratigraphic) data are summarised on Figs 5-6. Palaeogeographic reconstructions for selected time-slices are given on Figs 7-12. These are based in part on previously published maps (Podobina et al, 1956; Muratov, 1960; Sheydayeva-Kuliyeva, 1966; Ushakova & Ushko, 1971; Senes, 1973; Azizbekova, 1974; Senes & Marinescu, 1974; Luttig & 30 R.W. JONES AND M.D. SIMMONS A GLACIAL - TRANSGRESSION INTERGLACIAL - REGRESSION e Rivers frozen. Extensive Permafrost. e River Systems Reactivated. ¢ Sediment Supply < Subsidence. ¢ Evaporation > Precipitation / Runoff. e Expansion of 'Cool' Zone. e Expansion of 'Warm' Zone. MONTANE EMI VEGETATION SERT B GLACIAL - REGRESSION INTERGLACIAL - TRANSGRESSION e Water locked up in Ice Sheet. e Water released from melting Ice Sheet ¢ Pollen Spectra of Arid Aspect. e Expansion of Forest Zone * Contraction of Forest Zone Fig.4 Response to glaciation — alternative models for the Caspian. A: glacial transgression/interglacial regression. B: glacial regression/interglacial transgression. B better follows the palaeoclimatic reconstruction data given by Grichuk (1984) (see text). REVIEW OF STRATIGRAPHY OF EASTERN PARATETHYS 3] REGIONAL STRATIGRAPHY REGIONAL SEQUENCE GLOBAL SEQUENCE STRAT. CENTRAL = RES.} cyci¢ | | COASTAL ONLAP parnrenns|aacxseal casran [9020 To] § [POT curve CHRONOSTRAT. BIOSTRAT. CONTROL SEE FIGURE 6 FOR DETAIL = = L_NANTS J /20 Punta] km | s coon 7 17 | 6 | PONTIAN | __NOVORUSSIAN__| o PANNONIAN MAEOTIAN | omacovian | CHERSONIAN Fis] SARMATIAN BESSARABIAN VOLKHYNIAN KONKIAN mv

|[h2||o= OY — [rm] o i BELOGLINIAN KUMIAN Fig.5 Stratigraphic summary (Eocene-Holocene). Chronostratigraphy, magnetostratigraphy, global sequence stratigraphy and calibration after Haq et al. (1988) (see also chronostratigraphic schemes of Chumakov et a/., 1984, 1988, 1992a—b and others). Sequence boundary ages in Ma. Biostratigraphy after Blow (1969) (planktonic foraminifera) (prefixed P and N) and Martini (1971) (calcareous nanoplankton) (prefixed NP and NN). Regional stratigraphy from this paper. Regional sequence stratigraphy modified after Chepalyga (1985). Cycles are regressive (mega) sequences. II-III are equivalent to the ‘Eoparatethyan’, IV to the ‘Mesoparatethyan’ and V—VI to the ‘Neoparatethyan’ of Nevesskaya er al. (1985). Curve shows extent of open marine connection (function of sea-level) as inferred from salinity data from palaeontological analyses (S%vo = salinity parts per thousand). Res. pot. = Resource potential (S = source, R = reservoir, C = caprock). The correlation of Eastern Paratethyan sequence stratigraphy with the global coastal onlap curve of Haq et al. (1988) is tentative. Where biostratigraphic control constrains the correlation this is indicated. 32 Steffens, 1976; Steininger, Rogl & Martini, 1976; Hsu, in Ross et al., 1978; Régl et al., 1978; Steininger & Papp, 1979; Hsu, 1983; Baldi, 1984; Rog] & Steininger, 1983, 1984; Steininger & Rogl, 1984; Voronina & Popov, 1984; Iossofova, 1985; Popescu, 1985; Rusu, 1985; Steininger, Rabeder & Rogl, 1985; Steininger, Rogl & Nevesskaya, in Steininger, Senes, Kleeman & Régl, 1985; Voicu, 1985; Bernor et al., 1987; Nevesskaya et al., 1987; Panakhi & Buare Mamadu Lamin, 1987; Veto, 1987; Mamedov, 1989: Olteanu, 1989; Adamia et al., 1990; Dercourt et al., 1990; Tchoumatchenko et a/., 1990; Kerimov et al., 1991; Spiegler & Régl, 1992; Pevzner & Vangengeim, 1993), and in part on previously unpublished maps. It should be noted that the calibration against the global sequence stratigraphic framework and eustatic sea-level curve of Haq et al. (1988) is tentative. Fig. 5 demonstrates where biostratigraphic control exists in order to constrain the calibration. In the absence of such constraint, calibration is made by matching patterns of transgression and regression within a looser stratigraphic framework. The correlation between Eastern Paratethyan and global sequence stratigraphy and eustatic sea-level appears good, with all of the global eustatic sea-level trends finding their expression in Eastern Paratethys. It could be argued that this apparent correlation is entirely fortuitous. However, the stratigraphic signature of the mid-late Cenozoic appears remarkably consistent throughout the world, presumably because at this time it was an “ice-house’ world characterised by over-riding glacio-eustacy (Vail et al., 1991). Indeed, it may be that not only third-order but also higher frequency sea-level oscillations are recognisable in areas characterised by a high sedimentation rate such as Eastern Paratethys (and the Gulf of Mexico (see, for instance, Beard er al., 1982; Lamb et al., 1987; Pacht et a/., 1990; Wornardt & Vail, 1990; see also Fig. 6)). General features of Eastern Paratethyan stratigraphy have been discussed by, among others, Bogdanowicz (1947), Muratov (1960), Subbotina et al. (1960), Dzhanelidze (1970), Mamedova (1971, 1987), Stocklin & Setudehnia (1971, 1972), Azizbekov (1972) (and authors cited therein), Lupov et al. (1972), Cicha et al. (1975), Jiricek (1975), Ross et al. (1978), Nikiforova & Dodonov (1980), Verisharin et al. (1982) (and authors cited therein), Alekseyev & Nikiforova (1984), Iossofova (1985), Popov & Voronina (1985), Semenenko & Lulieva (1985), Skalbdyna (1985), Volkova et al. (1985), Yakhemovich ef al. (1985), Muratov & Nevesskaya (1986), Kereudren & Thibault (1987), Nigarov & Fedorov (1987), Benyamovskoy et al. (1988), Steininger et al. (1989), Yanko (1990a—b, 1991), Zubakov & Borzenkova (1990), Ghanbari (1991), Ali-Zade et al. (1994a, 1994b, 1995, in press), Jones (1996), Reynolds et a/. (in press) and Simmons ef al. (in press). Correlations within Eastern Paratethys and between Eastern and Central Paratethys have been discussed by Papp (1969), Chelidze (1973), Rogl, Steininger & Muller (1978), Paramonova et al. (1979), Semenenko (1979, 1984), Semenenko & Pevzner (1979), Steininger & Papp (1979), Steininger & Rogl (1979, 1984), Baldi (1980), Semenenko & Lulieva (1982), Rogl & Steininger (1983, 1984), Nevesskaya et al. (1984, 1985, 1987), Velichko (1984a), Yakhimovich, Bludorova, Zhidovinoy et al. (1984), Chepalyga et al. (1985), Nevesskaya & Nosovsky (1985), Nosovsky (1985), Pevzner & Vangengeim (1985), Rogl (1985a), Senes (1985a—b), Senes & Steininger, in Steininger et al. (1985), Yakhimovich, Bludorova, Chiguryaeva et al. (1985), Zosimovich et al. (1985), Yassini (1986), Mekhtiev & Pashaly (1987), Muzylev & Golovina (1987), Steininger et al., in Royden & Horvath (1987), Olteanu (1989), Rog] et al. (1991), Fedorov (1994), Markova & Mikhailesku (1994) and Jones R.W. JONES AND M.D. SIMMONS (1996). A comprehensive bibliography of general stratigraphic references (to 1984) is given by R6gl (1985b). Petroleum geological aspects have been discussed by, among others, Khain et al. (1937), Ismailov & Idrisov (1963), Ali-Zade et al. (1966), Shilinski (1967), Ismailov et al (1972), Buryakovsky (1974, 1993), Alikhanov (1977), Nikishin (1981), Ulmishek & Harrison (1981), Babayan (1984), Panakhi & Buare Mamadu Lamin (1987), Bagir-Zade et al (1988), Akramkhodzhaev et al. (1989), Kerimov et al. (1991), Kleschev et al. (1992), Narimanov (1993) and Reynolds et ai. (in press). Additional comments on petroleum geology are inserted as appropriate in the succeeding sections. Maykopian (Figs 7-8) The Maykopian takes its name from a town in the Caucasus (Likharev, 1958). The term Maykopian refers to essentially argillaceous rocks of Oligocene to Early Miocene age. The Zeivar Formation of Northern Iran and Lower Red Formation (predominantly clastics) of Central Iran (the latter locally contains age-diagnostic lepidocyclinid and nummulitid larger benthonic foraminifera) appear correlative, as does the Qom [Qum] Formation (predominantly carbonates) of Central Iran (which contains numerous age-diagnostic species of alveolinid, lepidocyclinid and miogypsinid larger benthonic foraminifera (Rahaghi, 1973)) (St6cklin & Setudehnia, 1971, 1972). The Maykopian (in particular the Khadumian) is an important regional source rock (e.g., Veto, 1987). It also constitutes a minor reservoir in the Kobustan-Kura region of the South Caspian (Ali-Zade et al., 1966). Details of Maykopian stratigraphy have been discussed by Muratov (1960), Ali-Zade (1966), Ali-Zade & Mamedov (1970), Mamedova & Mamedova (1970), Azizbekov (1972), Lupov et al. (1972), Khalilov & Mamedova (1973), Bolli & Krasheninnikov (1977), Ali-Zade & Atayeva (1982), Krasheninnikov & Muzylev (1975), Krasheninnikov, Muzylev & Ptukhian (1985), Nevesskaya & Nosovsky (1985), Bugrova (1986), Koshkarly (1986, 1993), Krasheninnikov (1986), Krasheninnikov & Ptukhian (1986), Koshkarly & Baldi-Beke (1987), Gasanov & Kyazamov (1988), Nagymarosy (1992) and Koshkarly & Alekperov (1993). Microbiostratigraphic study of the Maykopian is hindered by massive reworking, reflecting deposition in a foreland basin in front of the emerging Caucasus. Maykopian samples can contain >90% reworked (especially Eocene) microfossils. The Maykopian has been divided into five sub-stages by various authors (see Fig. 5). In ascending stratigraphic order, these are the Khadumian, Roshenian, Caucasian, Sakaraulian and Kozakhurian. The Khadumian is dated as Early Oligocene on the evidence of planktonic foraminifera and calcareous nannofossils and can therefore be calibrated against global standard biostratigraphic zonation schemes and absolute chronostratigraphic time-scales (see below). In contrast, the Roshenian is dated as Late Oligocene and the Caucasian to Kozakhurian as Early Miocene essentially only on the evidence of benthonic foraminifera (see, for instance, Nevesskaya & Nosovsky, 1985). The youngest sub-stages of the Maykopian appear to be absent in the Dacian Basin in the Western Black Sea region (Steininger et a/., in Royden & Horvath, 1987). Micropalaeontology and Nannopalaeontology. The Khadumian | has been dated as Early Oligocene on both planktonic foraminiferal and calcareous nannoplankton evidence. Ali-Zade | REVIEW OF STRATIGRAPHY OF EASTERN PARATETHYS CHRONOSTRAT. YAR BIO- STRAT. BLACK SEA GURIAN KUYALNIKIAN 5 KIMMERIAN CASPIAN KHVALYNIAN GIRKAN KHAZARIAN BAKUNIAN = = PLEISTOCENE i = H z BD bai, UNDIFF. PLIOCENE LATE EARLY LATE pm fo Surakhany Sabunchi Balakhany Pereriva N18 N19/20 HA i z| 2 eS PONTIAN KIMMERIAN AKCHAGYLIAN REGIONAL STRATIGRAPHY APSHERONIAN Akchagyl Suite THA Productive Series z z S = Pre-Pereriva 5 ] A 55 = z | BOSPORIAN Babajan | ® Gia] 3.3 i < fea} 30 3) r|s 5 § = ey) = 2 Shemakha | 5 @ NOVORUSSIAN ae, 34 35 33 REGIONAL SEQ. STRAT REGIONAL TRANSGRESSION CLIMATO- STRAT SCT GLOBAL SEQUENCE STRAT. CYCLE COASTAL ONLAP CURVE a? w (>) ioe) N ié>) cop) TB3 ie) to o : to) a ie) Ls) Le} ae) PS ine) a G |Go] ed ee) DP ie) = rs a re an ea ee Fig.6 Stratigraphic summary (Miocene-Holocene). Chronostratigraphy after Beard e¢ al. (1982) and Lamb et al. (1987) (conceptual Gulf of Mexico deep-water stage nomenclature) (Wis. = Wisconsinian; San. = Sangamonian; Ill. = Illinoisian; Yar. = Yarmouthian; Kan. = Kansan; Aft. = Aftonian; Neb. = Nebraskan). Biostratigraphy after Blow (1969) (planktonic foraminifera) (prefixed N) and Martini (1971) (calcareous nanno- plankton) (prefixed NN). Magnetostratigraphic polarity epochs and absolute age values are after Haq er al. (1988). Regional stratigraphy from this paper (upper case = chronostratigraphic, lower case = lithostratigraphic units). Regional sequence stratigraphy (transgressions) after Skalbdyna | (1966) | Krasheninnikov (1986) (1985). Global sequence stratigraphy after Haq er al. (1988) (third-order cycles TB3.2-TB3.8), and Beard er al. (1982) and Lamb et al. (1987) (fourth-order cycles Q2-Q8). Sequence boundary and maximum flooding surface ages in Ma. Climatostratigraphy (superclimathems or SCTs) after Zubakov & Borzenkova (1990). The correlation of Eastern Paratethyan sequence stratigraphy with the global coastal onlap curve of Haq et al. (1988) is tentative. The extent of biostratigraphic control constraining the correlation is indicated on Fig. 5. recorded the planktonic foraminifer Globigerina officinalis (Middle Eocene to Oligocene, Zones P14—P22 of Blow, 1969) from the basal ‘Planorbella Horizow in Azerbaijan. recorded planktonic foraminifera indicative of Early Oligocene (P18) from a stratigraphically similar position (immediately below the ‘Ostracod-Horizon’) in the Kuban-Kuma Interfluve (North Caucasus). Krasheninnikov et al. (1985) recorded Globigerina tapuriensis (Oligocene, P18—P20) associated with the larger benthonic foraminifer Nummulites intermedius (Early Oligocene) from the Khadumian of Armenia. Later, Krasheninnikoy & Ptukhian (1986) recorded Globigerina sellii (Oligocene, P19/20 to ‘early’ P22) associated with Nummulites intermedius (also Oligocene) and the calcareous nannofossil Helicosphaera reticulata (Eocene to Early Oligocene, Zones NP17—NP22 of Martini, 1971) from the Khadumian of Armenia. Koshkarly (1986) recorded the calcareous nannofossils Reticulofenestra umbilica (Eocene to Early Oligocene, NP16—-NP22), Chiasmolithus oamaruensis (Eocene to Early Oligocene, NP18—NP22), Isthmolithus recurvus (Eocene to Early Oligocene, NP19-NP22), Sphenolithus pseudoradians (Eocene to Early Oligocene, NP20—NP23) and Ericsonia subdisticha (Eocene to Early Oligocene, NP20—NP21) from the Khadumian of Azerbaijan. Later, Koshkarly & Baldi—Beke (1987) recorded R. umbilica, C. oamaruensis, I. recurvus and S. pseudoradians, and Koshkarly & Alekperov (1993) H. reticulata and E. subdisticha from the Early Maykopian of Azerbaijan. Palynology. Although only non-age-diagnostic palynomorphs were recorded from the Maykopian of the Middle Kura Depression by Dzhabarova (1973), recent observations suggest that some age-diagnostic dinocysts, and pollen and spores do 34 exist (in the Early, and Middle to Late Maykopian respectively). These will be reported in detail in a future publication. Palynological evidence indicates that the Maykopian is a regressive unit characterised by upwardly-increasing terrestrial input (upwardly-increasing pollen and spore content). Micropalaeontological and sedimentological evidence also indicates shallowing upward. Tarkhanian to Konkian (Figs 8—10) These stages have collectively been correlated with the Badenian of Central Paratethys (see, for instance, Steininger et a/., in Royden & Horvath, 1987). The Badenian is Middle Miocene (planktonic foraminiferal zones N8—?N12 (see, for instance, Papp et al., 1968, Rogl et al., 1978 and Papp & Schmid, 1985); calcareous nannoplankton zones NN5—NN7 (see, for instance, Rog] et al., 1978, Papp & Schmid, 1985 and Meszaros, 1992)). Palynologically, it is locally characterised by mangrove elements (Nagy & Kokay, 1991). The base of the Badenian (the Moravian sub-stage of Papp et al. (1978) (Lagenid Zone)) is defined at the first appearance of the planktonic foraminifer Praeorbulina (Zone N8) (Papp et al., 1968). The first appearance of the ancestral form Globigerinoides bisphericus, which defines the base of Zone N8 (Blow, 1969), falls within the underlying Karpatian (Cicha et al, 1967). This biostratigraphic control indicates that the base of the Badenian can be correlated with the 16.5Ma (glacio-eustatic) sea-level low-stand of Haq et al. (1988) (Fig. 5). The middle part of the Badenian (the Wielician sub-stage of Papp et al. (1978) (Sandschaler Zone)) is characterised by marginal marine sediments (including evaporites). This R.W. JONES AND M.D. SIMMONS regressive sub-stage can be tentatively calibrated against planktonic foraminiferal Zones NIO-N12 or calcareous nannoplankton zones NNS—NN6 (see, for instance, R6gl er al. (1978) and Papp & Schmid (1985)). The onset of regressive conditions can be tentatively correlated with the 15.5Ma (glacio-eustatic) sea-level low-stand of Haq et al. (1988) (Fig. 5). The regressive coarse clastics of the Chokrakian and Karaganian in Eastern Paratethys also appear to be associated with this event (though they could be associated with a separate tectonic event). These clastics constitute important reservoirs in the Indol Kuban and Terek Caspian Foredeeps (Ulmishek & Harrison, 1981) and in eastern Azerbaijan (Ali-Zade et al., 1986). Chepalyga (1985) calibrates the Chokrakian and | Karaganian against magnetostratigraphic polarity epochs 15-12, while Zubakov & Borzenkova (1990) calibrate them against polarity epochs 16-14. The top of the Badenian (the Kosovian sub-stage of Papp er al. (1978) (Buliminid-Bolivinid Zone)) is defined below the last appearances of Globorotalia mayerilsiakensis (N14) and Globigerina druryi (N15) (Papp et al., 1978; Papp & Schmid, 1985). Details of Tarkhanian to Konkian stratigraphy have been | discussed by Andrusov (1884), Bogdanowicz (1950a—b, 1965), Shishova (1955), Gasanova (1965), Mamedova (1971), Azizbekov (1972), Lupov et al. (1972), Dzhabarova (1973), Cicha et al. (1983) and Ali-Zade et al. (1986). A monograph of polymorphinid foraminifera of this age from Georgia was published by Dzharelidze (1977). In eastern Azerbaijan the Karaganian and Konkian, together with the overlying Sarmatian and Maeotian, are collectively referred to as the Diatom Suite (because of the presence of we Regional (Khadumian) Source Early Oligocene Platform Carbonates in Armenia Fig.7 Palaeogeographic reconstruction, Early Maykopian (Early Oligocene). Solid line indicates relatively well constrained, dashed line poorly constrained shoreline. Ticks on landward side. The location of the Maykopian stratotype is indicated. REVIEW OF STRATIGRAPHY OF EASTERN PARATETHYS diatomaceous limestones). The individual Eastern Paratethyan stages can be recognised within the Diatom Suite on the basis on fish otoliths and benthonic microfossils (particularly molluscs and foraminifera (e.g., Azizbekov, 1972; Ali-Zade et al., 1986) and diatoms (E.Z. Ateava, pers. comm., 1994)). The work of Dzhabarova (1973) suggests that palynology may also be used to recognise the various stages (see notes below). In eastern Azerbaijan parts of the Diatom Suite are considered to have hydrocarbon source potential. Tarkhanian (Fig. 8) The Tarkhanian takes its name from a promontory in the Crimea (Likharev, 1958). The stratotype section yields Middle Miocene (NNS5) calcareous nannoplankton (F. Régl, pers. comm., 1994). Micropalaeontology. Only non-age-diagnostic quasi-marine, smaller benthonic and rare planktonic foraminifera were recorded by Bogdanowicz (1950a) from the Tarkhanian of Kuban and later by Mamedova (1971) and Azizbekov (1972) from the Tarkhanian of Azerbaijan. These include Rotalia [Ammonia] ex gr. beccarii (smaller benthonic), which has a cosmopolitan distribution and probably ranges no older than Middle Miocene (RWJ’s unpublished observations), and Nonion [Florilus) boueanum (smaller benthonic) and Globigerina — tarchanensis (planktonic), both of which have also been recorded in the Badenian of Central Paratethys (Papp ef al., 1978; Papp & Schmid, 1985). Chokrakian (Fig. 9) The Chokrakian takes its name from a lake in the Crimea (Likharev, 1958). It is of Middle Miocene age on regional evidence (see above). Direct biostratigraphic evidence is lacking. The “Vindobonian Marls’ of Norfhern Iran appear correlative (Stocklin & Setudehnia, 1971, 1972). Micropalaeontology. Only non-age-diagnostic, quasi-marine, smaller benthonic foraminifera were recorded by Bogdanowicz (1950b) from the Chokrakian of the western Precaucasus and later by Mamedova (1971) and Azizbekov (1972) from the Chokrakian of Azerbaijan and Popkhadze (1983) for the Chokrakian of western Georgia. These include Rotalia [Ammonia] ex gr. beccarii (smaller benthonic), which has a cosmopolitan distribution and probably ranges no older than Middle Miocene (RWJ’s unpublished observations), Nonion [Florilus] boueanum and Miliolina [Quinqueloculina] akneriana sspp., both of which have also been recorded in the Badenian of Central Paratethys (Papp & Schmid, 1985), and Miliolina caucasica, Sigmoilina tschokrakensis and Tschokrakella longiuscula, all of which are endemic to Eastern Paratethys. The ostracod, Leptocythere bardrakensis was recorded by Popkhadze (1984) from the Chokrakian of western Georgia. Palynology. Only non-age-diagnostic palynomorphs were recorded by Dzhabarova (1973) from the Chokrakian of the Middle Kura Depression. Pollen spectra are characterised by relatively high incidences of herb and shrub taxa including presence of Chenopodiaceae and Ephedra. The | Fig.8 Palaeogeographic reconstruction, Late Maykopian to Tarkhanian (Late Oligocene to early Middle Miocene). Key as for Fig. 7. The locations of the Maykopian and Tarkhanian stratotypes are indicated. 36 Fig.9 Palaeogeographic reconstruction, Chokrakian to Karaganian (Middle Miocene). Key as for Fig. 7. The location of the Karaganian strato- type is indicated. Chenopodiaceae probably indicates the local development of salt-marshes. Karaganian (Fig. 9) The Karaganian takes its name from a locality on the Mangyshlak Peninsula in Kazakhstan (Likharev, 1958). It is Middle Miocene on regional evidence (see above). Direct biostratigraphic evidence is lacking. The Spaniodontella Beds of Northern Iran appear correlative (St6cklin & Setudehnia, 1971, 1972). Micropalaeontology. Only non-age-diagnostic, quasi-marine, smaller benthonic foraminifera were recorded by Mamedova (1971) and Azizbekov (1972) from the Karaganian of Azerbaijan. These include Nonion bogdanowiczi. The fish otoliths Rhombus corius and R. corius binagadinica are regarded as index-species for the Karaganian in Azerbaijan (E.Z. Ateava, pers. comm., 1994). Palynology. Only non-age-diagnostic palynomorphs were recorded by Dzhabarova (1973) from the Karaganian of the Middle Kura Depression. Pollen spectra are characterised by relatively high incidences of tree taxa, which indicates a forested hinterland. The predominance of Betula (birch) indicates a climatic regime similar to that of the present-day taiga or forest-tundra. Konkian (Fig. 10) The Konkian takes its name from a river in the Ukraine (a tributary of the Dniepr) (Likharev, 1958). It is of Middle R.W. JONES AND M.D. SIMMONS Chokrakian to Karaganian Reservoirs in Indol Kuban and Terek Caspian Foredeeps and Eastern Azerbaidzhan Miocene age on regional evidence (see above). Direct — biostratigraphic evidence is lacking. The Pholas Beds of — Northern Iran appear correlative (St6cklin & Setudehnia, 1971, 1972). Micropalaeontology. Only non-age-diagnostic, quasi-marine, — smaller benthonic foraminifera were recorded by Bogdanowicz — (1965) from the Konkian of the western Precaucasus and by — Mamedova (1971) and Azizbekov (1972) from the Konkian of — Azerbaijan. These include Rotalia [Ammonia] ex gr. beccarii — (smaller benthonic), which has a cosmopolitan distribution and _ probably ranges no older than Middle Miocene (RWJ’s unpublished observations), Articulina gibbosa and Miliolina [Quinqueloculina] haidingerii, both of which have also been recorded in the Badenian of Central Paratethys (Papp & © Schmid, 1985), and Articulina elongata konkensis, Bulimina : konkensis and Elphidium nachischevanicus, all of which are endemic to Eastern Paratethys. Bulimina konkensis and Elphidium kudakoense, together with the fish otolith Trigla | konkensis, are regarded as index-species for the Konkian in Azerbaijan (Podobina et al., 1956; Mamedova, 1971). Shishova (1955) and Gasanova (1965) recorded the following E diatoms from the Konkian of Eastern Azerbaijan: Actinocyclus ehrenbergi, A. rafsii, Asterolampra marylandica, Cocconeis — placentula lineta, C. scutelum, Coscinodiscus radiatus, C. oculus — and Melosira sulcata. Coscinodiscus radiatus was considered particularly typical. — Palynology. Only non-age-diagnostic palynomorphs were recorded by Dzhabarova (1973) from the Konkian of the Middle Kura Depression. Pollen spectra are characterised by relatively | high incidences of tree taxa, which indicates a forested j REVIEW OF STRATIGRAPHY OF EASTERN PARATETHYS 37 Sarmatian to Maeotian Reservoirs in Indol Kuban Foredeep Sarmatian Evaporites in Nakchichevan Depression - Fig.10 Palaeogeographic reconstruction, Konkian to Maeotian (late Middle to early Late Miocene). Key as for Fig. 7. The location of the Konkian stratotype is indicated. hinterland. The predominance of Taxodiaceae (cypresses and swamp-cypresses) indicates a warm-temperate (possibly even subtropical) climatic regime. Sarmatian (Fig. 10) The Sarmatian of Eastern Paratethys is probably equivalent to the stratotypical Sarmatian of Central Paratethys (late Middle to early Late Miocene (calcareous nannoplankton zones NN7-NN9) (Meszaros, 1992), planktonic foraminiferal zones N13?-N15), but may also be equivalent to the lower part of the Pannonian (Slavonian) of that area (Late Miocene) (see, for instance, Papp et al., 1974, 1985). Chepalyga (1985) calibrates the Sarmatian of Eastern Paratethys against magnetostratigraphic polarity epochs 10-7, while Zubakov & Borzenkova (1990) calibrate it against epochs 14-9, and Pevzner & Vangengeim (1993) calibrate it against polarity epochs 10-7. The regressive events which characterise the Sarmatian suggest that (within the limits of biostratigraphic control) its base can be correlated with the 10.5Ma (glacio-eustatic) sea-level low-stand of Haq etal. (1988). _ The Sarmatian of Eastern Paratethys is characterised by areally restricted regressive marginal marine sediments including coarse clastics, and, in the Nakchichevan Depression, evaporites (though transgressive black shales (with source potential) also occur locally). The ‘Sarmatian’ of Northern Iran and part of the Upper Red Formation of Central Iran, also characterised by clastics and evaporites, appear correlative (Stocklin & Setudehnia, 1971, 1972). The Sarmatian of Eastern Paratethys has been divided into three sub-stages, which are, from oldest to youngest, Volkhynian, Bessarabian and Chersonian. The ‘mid’ Sarmatian (Bessarabian) represents the culmination of a major regressive phase that began in late Konkian or ‘early’ Sarmatian (Volkhynian) times (see, for instance, Chepalyga, 1985), and resulted in the first isolation of the South Caspian Basin. Details of the Sarmatian stratigraphy of Eastern Paratethys have been discussed by Gasanova (1965), Maisuradze (1971), Mamedova (1971, 1987), Azizbekov (1972), Lupov et al. (1972), Dzhabarova (1973), Ali-Zade & Aleskerov (1974), Azizbekova (1974), Paramonova ef al. (1979) and Pevzner & Vangengeim (1993). Micropalaeontology. Only non-age-diagnostic (and largely endemic), quasi-marine, smaller benthonic foraminifera and ostracods were recorded from the Sarmatian by Podobina et al. (1956), Mamedova (1971, 1987), Voroshilova (1971) and Azizbekoy (1972) from Azerbaijan, by Azizbekova (1974) from the Nakchichevan Depression, and by Paramonova et al. (1979) from various sites in the Ponto-Caspian region. These include rare cosmopolitan species such as Streblus [Ammonia] beccarii and Elphidium macellum (foraminifera), which probably range no older than Middle Miocene (RWJ’s_ unpublished observations), Elphidium reginum (foraminifer) (lower part only), which is also found in the Sarmatian of Central Paratethys (Steininger et al, 1976) and (?reworked) in the Pliocene of the Black Sea (Gheorghian, in Ross et al., 1978), Nonion div. spp., Porosononion div. spp. and Quinqueloculina consobrina (foraminifera) and Cyprideis littoralis, Cythere multistriata, Leptocythere stabilis, Loxoconcha eichwaldi and Xestoleberis lutrae (ostracods). Podobina et al. (1956), Mamedova (1971) and Voroshilova (1971) have indicated that the three subdivisions of 38 the Sarmatian in Azerbaijan can be recognised on the basis of foraminifera and ostracods. The alga Ovulites sarmaticus and the fish otoliths Gadidarum minusculus and Gobius sarmaticus are regarded as index-species for the Sarmatian in Azerbaiyan (Ateava, personal communication, 1994). The Sarmatian part of the “Diatom Suite’ in Azerbaijan also contains 23 species of diatoms, chiefly Coscinodiscus, Licmophora and Navicula (Gasanova, 1965). Palynology. Only non-age-diagnostic palynomorphs were recorded from the Sarmatian of the Middle Kura Depression (Azizbekov, 1972; Dzhabarova, 1973). Pollen spectra from the middle (Bessarabian) sub-stage are characterised locally by (as in the “Cryptomactra-Horizon) relatively high incidences of Pinaceae (pine) pollen and locally (higher in the section) by relatively high incidences of broad-leaved tree pollen (Alnus (alder), Betula (birch), Carya (hickory), Carpinus (hornbeam), Corylus (hazel), Juglans (walnut)) and angiosperm (flowering plant) pollen (Azizbekov, op. cit.). Pollen spectra from the upper (Chersonian) sub-stage are also characterised locally by relatively high incidences of broad-leaved tree pollen (Fagus (beech), Quercus (oak), Taxodium (swamp-cypress), Tilia (lime) and Ulmus (elm)), though locally Polypodiaceae predominate (Azizbekoy, op. cit.). Maeotian (Fig. 10) The Maeotian of Eastern Paratethys is equivalent to the Pannonian (Late Miocene) of Central Paratethys (though possibly only the upper part thereof (Serbian) (see, for instance, Papp et al. (1974, 1985) and Pevzner & Vangengeim (1985b)). Rogl (1985a) calibrates it against calcareous nannoplankton Zone NNI1O. Chepalyga (1985) calibrates it against magnetostratigraphic polarity epochs 6-5, while Zubakov & Borzenkova (1990) calibrate it against polarity epochs 10-7. Krakhmalnaya et al. (1993) have recently described a succession at Novaya Emetovka on the Black Sea coast with a good Maeotian mammal fauna which they calibrate against polarity epochs 6 and 5. The Maeotian is represented by a transgressive-regressive cycle. It is locally characterised by black shales (with hydrocarbon source potential). The Maeotian sea was probably characterised by reduced salinity. Similar environmental conditions evidently obtained in the Pannonian of Central Paratethys, where benthonic foraminiferal assemblages are of quasi-marine (brackish water) aspect (Ammobaculites, Ammomarginulina, Miliammina, Trochammina) (Papp et al., 1985). Details of Maeotian stratigraphy have been discussed by Shishova (1955), Gasanova (1965), Bogdanowicz (1967, 1969, 1974), Mamedova (1971), Voroshilova (1971), Azizbekov (1972), Lupov et al. (1972), Popkhadze (1977), Paramonova et al. (1979), Ananova et al. (1985), Ali-Zade et al. (1986), Rasulov (1986), Maisuradze (1988), Naidina (1988) and Ateava (personal communication, 1994). Micropalaeontology and Nannopalaeontology. Only non-age-diagnostic, | quasi-marine, smaller _ benthonic foraminifera were recorded by Bogdanowicz (1967, 1969) from the Maeotian of the Kuban and Western Precaucasus and by Paramonova et al. (1979) from the Maeotian of various sites in the Ponto-Caspian. These include Elphidium macellum, which has a cosmopolitan distribution and probably ranges no older than Middle Miocene (RWJ’s unpublished observations) and Nonion diy. spp. Some _ stratigraphically and/or palaeoenvironmentally R.W. JONES AND M.D. SIMMONS significant ostracods were recorded by Azizbekov (1972) from Azerbaijan and by Popkhadze ef al. (1980) from Abkhazia. These include Leptocythere biplicata, L. meotica, Loxoconcha meotica, L. tamarindus and L. viridis (quasi-marine). Leptocythere meotica and Loxoconcha meotica, together with the foraminifera Quinqueloculina sulacensis (lower part) and Q. ludwigi (upper part) and the fish otoliths Clupea gidjakensis and Percidarum sigmalinoides are regarded as index-species for the Maeotian in Azerbaijan (Podobina et al., 1956; Mamedova, 1971). Rich diatom floras including Coscinodiscus gigas, C. oclisirides, Grammatophora azens, Cocconeis heteroidea, Melosira archotecturallis and M. sulcata were recorded by Shishova (1955) from the Maeotian part of the “‘Diatom Suite’ of — the Apsheron Peninsula in Azerbaijan. Actinocyclus ehrenbergii, Cymatosira sovtchenkoi and Rhaphoneis maeotica were recorded by Gasanova (1965). The stenohaline (normal marine) forms Asterolampha marylanica, Coscinodiscus asteromphalus and C. lewisanus sensu lato, the euryhaline (quasi-marine) forms Actinocyclus ehrenbergii and Rhapolodia musculus, and the freshwater forms Amphora ovalis and Diatoma vulgare were recorded by Rasulov (1986). Coscinodiscus lewisianus sensu stricto is a Middle Miocene species. A range of diatoms, calcareous nannofossils and ostracods were recorded by Ananova et al. (1985) from the Maeotian of | the Black Sea. These include Actinocyclus ehrenbergi, Rhaponeis maeotica and Thalassiosira maeotica (diatoms), | Braarudosphaera spp. (calcareous nannofossil) and Cyprideis | torosa and Leptocythere spp. (ostracods). Palynology. Only non-age-diagnostic palynomorphs have been | recorded from the Maeotian, and only the acritarch | Micrhystridium sp. was recorded by Ananova et al. (1985) from | the Maeotian of the Black Sea. Pollen spectra from the Late | Maeotian of the Task-Sunzhenskii Region are characterised by | relatively high incidences of Asteraceae and Polygonaceae | (herbs), Ephedra (shrubs) and Gramineae (grasses) (Naidina, | 1988). This indicates an open, sparsely forested hinterland and an arid climatic regime similar to that of the present-day steppe or semi-desert. The locally relatively high incidences of Gleicheniaceae (ferns) and Lycopodiaceae (club-mosses) indicate local development of conditions similar to those of the present-day tundra, forest-tundra or mountain belt. The | presence of Chenopodiaceae probably indicates local development of salt-marshes. Pontian The Pontian takes its name from the ancient name for the Black Sea. It is essentially regressive (though it also includes an overstepping transgressive unit at the base), and is characterised regionally by marginal marine coarse clastics and shallow marine carbonates and locally (Babajan Formation, Bosporian Sub-Stage) by evaporites. Chepalyga (1985) calibrates the Pontian against magnetostratigraphic polarity epoch 4 (Gilbert), while Zubakov & Borzenkova (1990) calibrate it against polarity epochs 6-5. We correlate the Pontian evaporites against those of the stratotypical Messinian, which can be calibrated against magnetostratigraphic polarity epoch 5 (Zubakov & Borzenkova, 1990). Details of the Pontian stratigraphy of Eastern Paratethys have been discussed by, among others, Sveier (1949), Mandelstam et } al. (1962), Sheydayeva-Kuliyeva (1966), Agalarova (1967), | Vekilov et al. (1969), Ramishvili (1969), Rozyeva (1971), REVIEW OF STRATIGRAPHY OF EASTERN PARATETHYS Azizbekov (1972), Lupov et al. (1972), Chelidze (1973), Karmishina (1975), Vekua (1975), Krstic (1976), Shchekina (1979), Imnadze & Karmishina (1980), Ali-Zade et al. (1986), Sirenko & Turlo (1986) and Yagmurlu & Helvaci (1994). Micropalaeontology. Only the non-age-diagnostic, quasi-marine, smaller benthonic foraminifer Elphidium stellatum was recorded by Imnadze & Karmishina (1980) from the Late Pontian (Bosporian sub-stage) of the Black Sea. Stratigraphically and/or palaeoenvironmentally significant ostracods recorded by Sheydayeva-Kuliyeva (1966) and Azizbekov (1972) from the Pontian of Azerbaijan (Western Caspian), by Karmishina (1975) from the Northern Precaspian and South-Eastern Kalmyk (Northern Caspian) and Prechernomore (Northern Black Sea), and by Vekua (1975) from Abkhazia (north-eastern Black Sea) include Bakunella dorsoacuata, Caspiolla acronasuta, Pontoniella acuminata and P. loczyi. Bakunella dorsoacuata and Caspiolla acronasuta are quasi-marine species (Yassini, 1986). Species of Leptocythere, Loxoconcha and Xestolebris also occur. It is possible to recognise three ostracod zones in the Pontian of Eastern Azerbaijan (Sheydayeva-Kuliyeva, 1966). It is also possible to recognise three corresponding mollusc zones. The lower part of the Continental (Cheleken) Series of Northern Iran and the Maragheh Bone Beds of Central Iran can be correlated with the Pontian on vertebrate palaeontological, limited malacological and ostracod evidence (Faridi, 1964; Stocklin & Setudehnia, 1971, 1972). | Palynology. Only non-age-diagnostic palynomorphs have been recorded from the Pontian. Pollen spectra from the Pontian of Georgia are characterised by relatively high incidences of tropical elements such as Nypa (palm) (Ramishvili, 1969). Those SS Palaeo- Danube Fig. 11 39 from the Late Pontian (Bosporian sub-stage) of the Ukraine are characterised initially by thermophilic (warm-temperate) and hydrophilic (moisture-loving) elements such as Taxodiaceae (cypresses and swamp-cypresses) and later by arid steppe and semi-desert elements (Shchekina, 1979; Sirenko & Turlo, 1986). Kimmerian (Fig. 11) The Kimmerian takes its name from an ancient tribe who lived on the shores of the Black Sea (Likharev, 1958). The Dacian (a stage name sometimes used in the Black Sea region) and the ‘Eoakchagylian’ (a stage name used in the Caspian Sea region by Zubakov & Borzenkova (1990)) appear synonymous. Semenenko (1979), Pevzner & Vangengeim (1985), Skalbdyna (1985) and Zubakov & Borzenkova (1990) calibrate the Kimmerian against magnetostratigraphic polarity epochs 54, while Chepalyga (1985) and Chepalyga et al. (1985) calibrate it against polarity epoch 4 (Gilbert). Essentially on the basis of magnetostratigraphic evidence (including the calibration of the underlying Pontian against polarity epoch 5 (see above)), we have tentatively calibrated the major unconformity at the base of the Kimmerian against the 5.5Ma glacio-eustatic sea-level low-stand of Haq er al. (1988) and apparently coincident uplift around and subsidence within the Caspian (leading to a massive sea-level fall (of the order of 1000m) within the Caspian and the severance of the connection between the Caspian and the Black Sea). Additional unconformities in the Caspian succession cannot be confidently calibrated against any of the third-order glacio-eustatic sea-level low-stands on the Haq et al. chart, and may be associated with higher frequency glacio-eustatic low-stands or local tectonic events. Due partly to tectono-eustatic effects (see above), and partly to climatic effects Palaeo-Ural Palaeo-Volga fae [ ee Palaeo-Don — Palaeo-Kura Kimmerian Reservoirs in South Caspian Palaeogeographic reconstruction, Dacian/Kimmerian (latest Miocene to Pliocene). Key as for Fig. 7. 40 (an excess of evaporation over precipitation) and a change in the drainage system (with rivers formerly flowing into Paratethys captured by rejuvenated rivers flowing into the Mediterranean, where base-level had fallen considerably during the Messinian salinity crisis (see, for instance, Hsu (1983)), the level and the areal extent of the Caspian ‘athalassic lake’ were drastically reduced during the Kimmerian. Palaeontological evidence points to a stratigraphically upward reduction in salinity. The basal Kimmerian contains a quasi-marine ostracod fauna (characterised by Cyprideis littoralis/torosa) that appears correlative with that of the Messinian “Lago Mare’ facies in the Mediterranean. In Azerbaijan, the Kimmerian is characterised by several thousand metres (thicknesses of around 4km are typical around the Apsheron Peninsula) of principally Palaeo-Volga-derived, regressive marginal and non-marine coarse clastics comprising the main (‘Productivnaya Tolsha’ = ‘Productive Series’) reservoirs in several billion-barrel oil-fields (Khain et al., 1937; Ismailov and Idrisovy, 1963; Ismailov et al., 1972; Nikishin, 1981; Babayan, 1984; Kerimov ef al., 1991; Narimanov, 1993). The sedimentology of this succession is discussed in detail by Reynolds et al. (in press). Constituent lithostratigraphic units include, in ascending stratigraphic order, the Kalin or Kala, Podkirmakina (Pre-Kirmaky Sand (PK )), Kirmakina (Kirmaky Sand (KS)), Nadkirmakina (Post-Kirmaky Sand and Post-Kirmaky Clay (NKP and NKG)), Pereriva, Balakhany, Sabunchi and Surakhany Suites. Details of Kimmerian stratigraphy have been discussed by, among others, Ali-Zade (1946), Khalilov (1946), Sveier (1949), Agalarova (1956), Mandelstam er al. (1962), Rozyeva (1971), Azizbekov (1972), Lupov et al. (1972), Ali-Zade (1974), Karmishina (1975), Vekua (1975), Ananova et al. (1985), Sirenko & Turlo (1986) and Yassini (1986). Micropalaeontology Only non-age-diagnostic, quasi-marine, smaller benthonic foraminifera were recorded by Karmishina (1975) from the Kimmerian of Prechernomore (Northern Black Sea). These include Ammonia beccarii and Elphidium incertum. Siratigraphically and/or palaeoenvironmentally significant ostracods recorded by Karmishina (1975) and Vekua (1975) from the Kimmerian of Prechernomore (Northern Black Sea) and Abkhazia (North-eastern Black Sea) respectively include Bakunella_ dorsoacuata, Caspiocypris labiata, — Caspiolla acronasuta, Cyprideis littoralis/torosa, Leptocythere bosqueti, Mediocythereis apatoica and Pontoniella acuminata. Bakunella dorsoacuata, Caspiolla acronasuta and Cyprideis torosa are quasi-marine species (Yassini, 1986). Caspiocypris labiata is fresh-water. Ostracods recorded by Karmishina (1975) from the Kimmerian of the Northern Precaspian and South-Eastern Kalmyk (Northern Caspian) include Candona angulata, C. neglecta, C. rostrata, Cyclocypris laevis, Cypria arma, Eucypris naidinae, Ilyocypris bradyi, I. gibba, I. serpulosa and Zonocypris membranae. These are all fresh-water forms. Microbiostratigraphic study of the Kimmerian ‘Productive Series’ in Azerbaijan is hindered by massive reworking (Khalilov, 1946; Agalarova, 1956). However, the general pattern of reworking indicates unroofing, which in itself is of some stratigraphic value (with pulses of reworking related to sea-level low-stands). Moreover, the potential exists for biostratigraphic subdivision on the basis of ostracods (see, for instance, Khalilov, 1946; Agalarova, 1956; Azizbekov, 1972). The Kala Suite contains Cythere [Leptocythere] camellii and Paracypris liventalina (quasi-marine), the Kirmakina Suite Cythere [Leptocythere] cellula, C. [Cyprideis] littoralis/torosa, C. olivina, R.W. JONES AND M.D. SIMMONS C. [Leptocythere] praebacuana, Hemicythere pontica and Loxoconcha djabaroffi (quasi-marine), the Pereriva Suite Cyprideis littoralis/torosa (quasi-marine), the Balakhany Suite Cythere [Leptocythere| praebacuana, Loxoconcha alata, L. eichwaldi and Xestoleberis lutrae (quasi-marine), the Sabunchi Suite no in situ ostracods, and the Surakhany Suite Eucypris (fresh-water) and Leptocythere and Loxoconcha (quasi-marine). The Krasnotsvetnaya (“Red-Coloured’) Series of Turkmenia can be correlated with the Kimmerian on ostracod evidence (Agalarova, 1956; Lupov et al., 1972). The lowermost part contains Bakunella, Caspiolla, Pontoniella and Xestoleberis and | is of Pontian aspect, while the uppermost part contains Leptocythere and Limnocythere and is of Akchagylian aspect. The upper part of the Continental (Cheleken) Series of Northern Iran can be correlated with the Kimmerian on vertebrate palaeontological, limited malacological and ostracod evidence (Faridi, 1964; Stocklin & Setudehnia, 1971, 1972). The Lacustrine Fish Beds of Central Iran, locally characterised by volcanic ash bands, can also be tentatively correlated with the Kimmerian (St6cklin & Setudehnia, op. cit.). Nannopalaeontology. Stratigraphically significant calcareous nannofossils recorded by Zubakov & Borzenkova (1990) from i! the Kimmerian include Discoaster quinqueramus (NN11) (?reworked), Ceratolithus acutus (‘late’ NN12) and C. rugosus | (NN13-2?NN19). Palynology. Only non-age-diagnostic palynomorphs have been recorded from the Kimmerian. However, at least theoretically, climatostratigraphy ought to have some utility in the stratigraphic subdivision of the “Productive Series’ (with a number of superclimathems discernable on the basis of different | pollen spectra). Indeed, the results of preliminary analyses have | shown pollen spectra from ‘Thermo’-SCT27 in the Ukraine to be characterised by broad-leaved and subtropical forest elements (Castanea (chestnut), Rhus (rue) and Taxodiaceae (cypress and swamp-cypress) (Sirenko & Turlo, 1986), and those from “Cryo’-SCT26 and ‘Cryo’-SCT24 to be characterised by steppe elements (Ananova et al., 1985; Zubakov & Borzenkova, 1990). ‘Thermo’-SCT23 has been shown to be characterised by the quasi-marine acritarch Sigmailina (and the quasi-marine diatom Actinocyclus ehrenbergi) (Ananova et al, Zubakov & Borzenkova, opp. cit. ). Akchagylian to Khvalynian (Caspian Sea) (Fig. 12) The Akchagylian to Khvalynian of the Caspian Sea appear | equivalent to the Kuyalnikian to Neoeuxinian of the Black Sea | (Late Pliocene to Holocene). Details of Akchagylian to | Khvaklynian stratigraphy have been discussed by, among others, | Livental (1929), Sveier (1949), Mandelstam et al (1962), | Yakhimovich et al. (1965, 1984), Dzhabarova (1966), Kuliyeva (1968), Dmitriyev er al. (1969), Rayevskiy (1969), Sultanov & Sheydayeva-Kuliveya (1969), Aliyev & Aliyeva (1970), Kopp (1970), Rozyeva (1971), Ushakova & Ushko (1971), Zubakov & + Kochegura (1971), Azizbekov (1972), Lupov et al. (1972), Ananova (1974), Karmishina (1975), Kaplin (1977), Trubikhin (1977), Fedkovich (1978), Veliyev (1980), Abramova (1982, | 1985), Semenenko & Lulieva (1982), Mamedov & Rabotina (1984a), Aliyulla et al (1985), Ivanova (1985), Mamedov & Aleskerov (1985, 1986), Sultanov (1985), Yassini (1986), | Aleskerov et al. (1987), Bludorova et al. (1987), Zhidovinov et al. (1987), Naidina (1988), Ali-Zade & Aliyeva (1989), Yanko | (1990a—b, 1991), Trubikhin et a/. (1991) and Mamedova (1993). REVIEW OF STRATIGRAPHY OF EASTERN PARATETHYS Akchagylian (Fig. 12) The Akchagylian takes its name from a locality near Krasnovodsk on the southern shores of the Gulf of Karabogaz in Western Turkmenia (Likharey, 1958). It appears equivalent to the Kuyalnikian of the Black Sea. Zubakoy & Borzenkova (1990) calibrate it against magnetostratigraphic polarity epochs 4 (Gilbert) or 3 (Gauss) to 2 (Matuyama). The Akchagylian is transgressive and is represented by marine shales (constituting an important regional seal in the South Caspian). The transgressions within the Akchagylian temporarily re-established marine connections to the world’s oceans. -Calcareous nannoplankton data (see below) indicates that they may correspond to the 3.4Ma, 2.7Ma and 2.0Ma maximum flooding surfaces of Haq et al. (1988). Details of Akchagylian stratigraphy have been discussed by, among others, Ali-Zade (1936), Yakhimovich et al. (1965, 1983), Dzhabarova (1966), Ali-Zade (1969), Dmitriyev et al. (1969), Sultanov & Sheydayeva-Kuliveya (1969), Aliyev & Aliyeva (1970), Ushakova & Ushko (1971), Zubakov & Kochegura (1971), Azizbekov (1972), Lupov et al. (1972), Ananova (1974), Karmishina (1975), Trubikhin (1977), Fedkovich (1978), Sultanov (1979), Semenenko & Lulieva (1982), Mamedov & Rabotina (1984a), Bludorova et al. (1987), Zhidovinov et al. (1987), Naidina (1988) and Benyamovskoy & Naidina (1990). Micropalaeontology. Only non-age-diagnostic —_ smaller benthonic foraminifera were recorded by Yakhimovich ef al. (1965), Sultanov & Sheydayeva-Kuliyeva (1969) and Karmishina (1975) from the Akchagylian of the Volga-Urals Region, Azerbaijan, and the Northern Precaspian and South-Eastern Kalmyk (Northern Caspian) respectively. These floridana, 4] include Ammonia beccarii, Bolivina ex gr. advena, B. aksaica, B. Buccella sp., Cassidulina crassa, C. oblonga, Cassidulinita prima, Cibicides lobatulus, Discorbis arculus, D. multicameratus, D. orbicularis, D. pliocenicus, Elphidium incertum, E. kudakoense, E. macellum, E. subarcticum, Nonion aktschagylicus and Quinqueloculina spp. Bolivina, Cibicides and Discorbis are near-normal marine genera, while the remainder are quasi-marine. Stratigraphically and/or palaeoenvironmentally significant ostracods recorded by Sultanov & Sheydayeva-Kuliyeva (1969) and Karmishina (1975) from the Akchagylian of Azerbaijan, and the Northern Precaspian and South-Eastern Kalmyk (Northern Caspian) respectively include Aglaiocypris chuzchievae, Candona_ convexa,_ Ilyocypris gibba and Leptocythere verrucosa. Aglaiocypris, Candona and Ilyocypris are fresh-water forms, while Leptocythere is quasi-marine. In Azerbaijan, the Akchagylian (Akchagyl Suite) contains a diverse marine ostracod and molluscan fauna (Livental, 1929: Ali-Zade, 1954; Sultanov & Sheydayeva-Kuliyeva, 1969). On the Apsheron Peninsula, it can be divided into three units on the basis of ostracods. Freshwater forms (Candona, Eucypris, Ilyocypris, Limnocythere) characterise the lower unit, marine and quasi-marine forms the middle and upper units. Stratigraphically and/or palaeoenvironmentally significant diatoms recorded by Ushakova & Ushko (1971) from the Akchagylian to Apsheronian of the Krasnovodsk Peninsula in Western Turkmenia include Actinocyclus ehrenbergi ssp. ralfsii, Campylodiscus noricus, C. noricus ssp. hibernica, Cocconeis scutellum, Coscinodiscus argus, Cymbatopleura elliptica, Diploneis bombus, D. domblittensis, D. fusca ssp. pervasta, D. rombica, Epithemia turgida, Grammatophora oceanica, Melosira | Top-Seal in South Caspian } Fig. 12 Palaeogeographic reconstruction, Akchagylian/Kuyalnikian (Late Pliocene). Key as for Fig. 7. The location of the Akchagylian stratotype is indicated. 42 arenaria, M. scabrosa, M. sulcata, Navicula lyra ssp. elliptica, Nitzchia cocconeiformis, Rhopalodia parallella, — Surirella striatula, Terpsinoe musica and Triceratium sp. Campylodiscus noricus, C. noricus ssp. hibernica, Diploneis domblittensis, D. rombica, Melosira arenaria, M. scabrosa, M. sulcata and Rhopalodia parallella are exclusively fresh-water forms, while the remainder are quasi-marine. The Akchagyl Beds of Northern Iran can be correlated with the stratotypical Akchagylian on ostracod evidence (Faridi, 1964; St6cklin & Setudehnia, 1971, 1972). Various volcanics in Central Iran can also be tentatively correlated with the Akchagylian (St6cklin & Setudehnia, op. cit.). Nannopalaeontology. Stratigraphically significant calcareous nannofossils recorded by Semenenko & Lulieva (1982) from the Akchagylian include Discoaster brouweri (NN8?-NN18) (no younger than 1.89Ma), D. pentaradiatus (NN9?-NN17) (no younger than 2.33-2.43Ma) and Reticulofenestra pseudoumbilica (NN7?-NN15). These are similar to nannoflora found in the Kuyalnikian of the Black Sea region by Semenko & Pevzner (1979), and suggest a link between the two basins at this time. Palynology. Only non-age-diagnostic palynomorphs have been recorded from the Akchagylian. Pollen spectra from ‘Cryo’-SCT14 (and ‘Thermo’-SCT15) are characterised by high incidences of Abies (fir), indicating a forested hinterland similar to that of the present-day taiga, and a cool, wet climate (Bludorova et al., 1987). Those from ‘Thermo’-SCT13 are characterised by Abies (fir), Acer (maple), Carpinus (hornbeam) and Ulmus (elm), indicating a warmer climate (Bludorova et al., op. cit.). Pollen spectra from “Cryo’-SCT14 in the Volga-Urals Region are characterised by relatively high incidences of Lycopodiaceae (club-mosses), again indicating a cool, wet climate (Yakhimovich et al., 1965, 1983, 1984). Pollen spectra from ‘Cryo’-SCTs 12 and 10 are characterised by alternations of Alnus (alder), Betula (birch) and Pinus (pine), indicating a cool, dry climate, and Tsuga (hemlock), indicating a warmer, wetter climate (Ananova, 1974; Bludorova et al, 1987). Those from ‘Thermo-SCT11 in the Volga Basin are characterised by relatively high incidences of Tsuga (hemlock) (Zhidovinoy et al., 1987). Pollen spectra from the undifferentiated Akchagylian of the Tersko-Sunzhenskii Region are characterised by high incidences of broad-leaved tree taxa such as Fagus (beech) and Quercus (oak), indicating a forested hinterland similar to that of the present-day taiga (Broad-Leaved Forest Zone), and a warm-temperate climatic regime (Naidina, 1988). Apsheronian The Apsheronian takes its name from the Apsheron Peninsula in Eastern Azerbayan (Likharev, 1958). It appears equivalent to the Gurian of the Black Sea (Pleistocene). Zubakov & Borzenkova (1990) calibrate it against magnetostratigraphic polarity epoch 2 (Matuyama). The Apsheronian is essentially regressive in character. Marine connections between the Caspian and the Black Sea were probably only intermittently developed. Details of Apsheronian stratigraphy have been discussed by Livental (1929), Sultanov (1964), Kuliyeva (1968), Ushakova & Ushko (1971), Azizbekov (1972), Lupov et al. (1972), Ali-Zade (1973), Ivanova (1985), Zhidovinovy et al. (1987) and Mamedova (1988). Micropalaeontology. Stratigraphically and/or palaeoenviron- R.W. JONES AND M.D. SIMMONS mentally significant ostracods recorded by Kuliyeva (1968) and Mamedova (1984, 1988) from the Apsheronian of the Baku Archipelago in Azerbaijan include Bakunella dorsoacuata, Candona_ albicans, Caspiocypris lyrata, C. rotulata, C. sinistrolyrata, Caspiolla acronasuta, C. gracilis, Eucypris membranae, Leptocythere andrusovi, L. caspia, L. fragilis, L. multituberculata, L. propinqua, L. quinquetuberculata, L. turquianica, L. verrucosa, Loxoconcha eichwaldi, L. gibboida, L. impressa and Trachyleberis azerbaidzhanica. Bakunella, Caspiolla, Leptocythere, Loxoconcha and Trachyleberis are quasi-marine forms, while Candona, Caspiocypris and Eucypris are fresh-water (see, for instance, Yassini, 1986). At least six assemblage zones can be recognised on the basis of ostracods (D.N. Mamedova, pers. comm., 1994). These will be discussed in a forthcoming paper (Mamedova, in press) (see also Mamedova, 1988). Stratigraphically and/or palaeoenvironmentally significant diatoms recorded by Ushakova & Ushko (1971) from the Akchagylian to Apsheronian of the Krasnovodsk Peninsula in Western Turkmenia are listed under ‘Akchagylian’ above. They include exclusively fresh-water, fresh-water to brackish, and brackish to near-normal marine forms. The Apsheron Beds of Northern Iran can be correlated with the stratotypical Apsheronian on ostracod evidence (Faridi, 1964; Stocklin & Setudehnia, 1971, 1972). Palynology. Only non-age-diagnostic palynomorphs have been recorded from the Apsheronian. Pollen spectra described by Ivanova (1985) from the Apsheronian of Western Turkmenia and by Zhidovinov et al. (1987) from the Apsheronian of the Volga Basin are characterised by relatively high incidences of non-tree taxa. This indicates an open, sparsely forested hinterland similar to the present-day steppe or forest-steppe, and a cool-temperate climatic regime. Bakunian The Bakunian takes its name from the city of Baku in Eastern Azerbaijan (Likharev, 1958). It appears equivalent to the Chaudian of the Black Sea (Pleistocene). Zubakovy & Borzenkova (1990) calibrate it against magnetostratigraphic polarity epochs 2 (Matuyama) to | (Brunhes). Details of the Bakunian stratotype have recently been discussed by Mamedova (1984, 1985, 1993, and in press) and Aliyulla et al. (1985). | Stratigraphically and/or palaeoenvironmentally significant ostracods recorded by Aliyulla et al (op. cit.) include Bakunella dorsoacuata, Caspiocypris filona, Graviacypris elongata, Leptocythere delicata, L. lunata, L. propinqua, Loxoconcha endocarpa, Pseudostenocypria asiatica and Xestoleberis ementis. Bakunella, Graviacypris, Leptocythere, Loxoconcha and Xestoleberis are quasi-marine forms, while Caspiocypris and Pseudostenocypria, are fresh-water (see, for instance, Yassini, 1986). Four assemblage zones can be recognised on the basis of ostracods (Mamedova, in press). The Baku Formation of Northern Iran can be correlated with the stratotypical Bakunian on ostracod evidence (Faridi, 1964; Stocklin & Setudehnia, 1971, 1972). Khazarian, Girkan and Khvalynian The Khazarian takes its name from an ancient tribe who lived in the area between the Don and Volga Rivers, and the Girkan(ian) and Khvalynian take theirs from ancient names for the Caspian REVIEW OF STRATIGRAPHY OF EASTERN PARATETHYS Sea (Likharey, 1958). The Khazarian, Girkan and Khvalynian appear equivalent to the Uzunlarian, Karangatian and Neoeuxinian respectively of the Black Sea (Pleistocene-Holocene). Zubakov & Borzenkova (1990) calibrate the Khazarian, Girkan and Khvalynian against magnetostratigraphic polarity epoch | (Brunhes). Kuyalnikian to Neoeuxinian (Black Sea) (Fig. 12) The Kuyalnikian to Neoeuxinian of the Black Sea appear equivalent to the Akchagylian to Khvalynian of the Caspian (Late Pliocene to Holocene). Details of Kuyalnikian to Neoeuxinian stratigraphy have been discussed by Wall & Dale (1973), Karmishina (1975), Vekua (1975), Kaplin (1977), Shikmus et al. (1977), Schrader (1979), Semenenko & Pevzner (1979), Shatilova (1984), Sirenko & Turlo (1986), Yanko (1990a—b, 1991), Balabanov et al. (1991) and Trubikhin et al. (1991a—b). Kuyalnikian (Fig. 12) The Kuyalnikian takes its name from a locality on the Danube Delta (Likharev, 1958). The Romanian appears synonymous. It appears equivalent to the Akchagylian of the Caspian. Zubakov & Borzenkova (1990) calibrate it against magnetostratigraphic polarity epochs 3 (Gauss) to 2 (Matuyama). The Kuyalnikian/Akchagylian represents a major transgressive episode when Eastern Paratethys was reconnected with the world oceans (Fig. 12). During this time, marine plankton were introduced into Eastern Paratethys enabling calibration to global datums. Details of Kuyalnikian stratigraphy have been discussed by Karmishina (1975), Vekua (1975), Semenenko & Pevzner (1979), Shatilova (1984) and Sirenko & Turlo (1986). Micropalaeontology. Only the non-age-diagnostic, quasi-marine, smaller benthonic foraminifera Ammonia beccarii and Elphidium incertum were recorded by Karmishina (1975) from the Kuyalnikian of Prechernomore (Northern Black Sea). Stratigraphically and/or palaeoenvironmentally significant ostracods recorded by Karmishina (1975) and Vekua (1975) from the Kuyalnikian of Prechernomore (Northern Black Sea) and Abkhazia (North-eastern Black Sea) respectively include Bakunella dorsoacuata, Caspiocypris labiata, _ Caspiolla acronasuta, Cryptocyprideis bogatschovi, Cypria arma, Leptocythere andrusovi, L. circumsulcata, Loxoconcha eichwaldi, L. petasa, Mediocythereis apatoica and Pontoniella acuminata. Bakunella dorsoacuata, Caspiolla acronasuta and Loxoconcha petasa are quasi-marine species (Yassini, 1986). Caspiocypris acronasuta and Cypria arma are fresh-water. Nannopalaeontology. Stratigraphically significant calcareous nannofossils recorded by Semenenko & Pevzner (1979) from the Kuyalnikian include Discoaster pentaradiatus (NN9?-NN17) and Reticulofenestra pseudoumbilica (NN7?-NN15). Palynology. Only non-age-diagnostic palynomorphs have been recorded from the Kuyalnikian. Pollen spectra from ‘Thermo’-SCT15 are characterised by polydominance (Shatilova, 1984: Sirenko & Turlo, 1986). Gurian The Gurian takes its name from an ancient province of what is now Western Georgia (Likharev, 1958). It appears equivalent to 43 the Apsheronian of the Caspian (Pleistocene). Zubakov & Borzenkova (1990) calibrate it against magnetostratigraphic polarity epoch 2 (Matuyama). Chaudian The Chaudian takes its name from a promontory on the Kerch Peninsula (Crimea) (Likharev, 1958). It appears equivalent to the Bakunian of the Caspian (Pleistocene). Zubakov & Borzenkova (1990) calibrate it against magnetostratigraphic polarity epochs 2 (Matuyama) to 1 (Brunhes). Uzunlarian, Karangatian and Neoeuxinian The Uzunlarian takes its name from a lake, the Karangatian takes its name from a promontory on the Kerch Peninsula (Crimea), and the Neoeuxinian its name from an ancient name for the Black Sea (Likharev, 1958). The Uzunlarian, Karangatian and Neoeuxinian appear equivalent to the Khazarian, Girkan and Khvalynian respectively of the Caspian (Pleistocene-Holocene). Zubakov & Borzenkova (1990) calibrate the Uzunlarian, Karangatian and Neoeuxinian against magnetostratigraphic polarity epoch | (Brunhes). Details of Uzunlarian, Karangatian and Neoeuxinian stratigraphy have been discussed by Schrader (1979) and Markova & Mikhaleska (1994). Schrader (1979) recorded the palaeoenvironmentally significant diatoms Actinocyclus divisus, A. ochotensis, Cymatopleura solea and Stephanodiscus astraea (which indicate a palaeosalinity range of 0.5-3.0ppt). Markova & Mikhaleska (1994) recorded the ostracods Amnicythere cymbula, Callistocythere alfera, C. lopalica, C. mediterrarea, C. quinquetuberculata and Loxoconcha immodalata in the stratotypical Uzunlarian. ACKNOWLEDGEMENTS. We wish to acknowledge our colleagues at the Azerbaijan Academy of Sciences, Baku, in particular Drs. Elmira Ateava, Dilara Mamedova and Reyhan Koshkarly, for their assistance in procuring publications, for facilitating fieldwork, and for sharing their extensive knowledge with us. We also wish to acknowledge Dr. Fred Rog! of the Naturhistorisches Museum, Vienna, for his constructive critical review of the manuscript. BP Exploration provided drafting facilities and is thanked for permission to publish. REFERENCES Abramoya, T.O. 1982. Cyclicity of spore-pollen spectra in Quaternary deposits of the west coast of the Caspian Sea. /n, Kovalev, S.A. et al. (editors), Sea-coasts: 32-39. Moscow (in Russian). — 1985. 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(editor): The Problems of Correlation of the Latest Sediments of the Eurasian North. Leningrad (Geogr. Soc. USSR). Bull. nat. Hist. Mus. Lond. (Geol.)52(1): 51-59 Issued 27 June 1996 A new protorichthofenioid brachiopod (Productida) from the Upper Carboniferous of the Urals, Russia C.H.C. BRUNTON Department of Palaeontology, The Natural History Museum, Cromwell Road, London SW7 5BD Synopsis. A new genus from the mid Upper Carboniferous of the southern Urals, Russia, is described and interpreted as a surprising, aspinose early form of the Richthofenioidea. An undescribed Proteguliferina? from the Upper Carboniferous of northern Spain and two previously described Permian Russian species are possibly congeneric, but thereafter the stock probably died out. The new genus Za/vera contains the new species Z. sibaica, specimens from Spain and the two described Permian species. INTRODUCTION In 1982 Dr S. S. Lazarev, of the Palaeontological Insitute, Moscow, led an expedition to the southern Urals where brachiopods were collected and passed to me for comment and description. The brachiopods include seven taxa which can be assigned to productid families with little difficulty. However, most specimens belong to a distinctive species which initially _ was difficult to identify, even at phylum level. It now seems | clear that these subconical, thin-shelled specimens also belong within the Productida, although they display bizarre _ characteristics. | | | Material | The material was all collected from a dense, finely-grained | buff-coloured limestone within the Kordailovskaya Formation _ (probably equivalent to the Verejan Horizon in the Moscow | area) dated as late Bashkirian to Lower Moscovian (mid Upper | Carboniferous) and occurring at the Sibay stream, on the left bank of the Ural River, 6 km up-stream from Pokrovka, in the southern Urals, Russia. The fragments containing brachiopods amount to about 40 in total, several being multi-parts and counterparts of larger pieces that have been broken down to between 20 and 50mm in greatest dimension. Some pieces include more than one brachiopod specimen. From the total of 40 fragments, about half contain the unusual subconical species. Of the rest: 1. Four contain a species reaching about 15mm in length, with fine spines, densely covering the ventral valve only, but with dorsal valve dimples, and having a low lateral profile. Superficially the species resembles Stipulina (Fig. 1). | 2. Eleven contain a Thomasella-like species, but differ in the | absence of rugae over the spinose body region and in having a | __ relatively more strongly ribbed and flanged trail (Figs 2 and 3): | 3. Two contain small (ca. 7mm wide) specimens more like | Thomasella than the above in that they are distinctly rugose up to the start of the flange (Fig. 4a, b). | ©The Natural History Museum, 1996 4. One part and counterpart contains a strongly rugose species (only ca. 9mm wide) resembling a small plicatiferid, but with a widely flanged and ribbed trail (Fig. 5). 5. There are two incomplete specimens of a rugose and less clearly ribbed (reticulate) species with spines near the umbo and on the ventral exterior which resembles Pectenoproductus proprius Likharev, a species reported from the Lower Permian of the Caucasus. Muir-Wood & Cooper (1960) were unsure as to whether this genus was a pectinid mollusc, but the presence of spines on these two specimens confirms their affinity with the Productidina (Fig. 7). 6. There is one incomplete ventral exterior, seen also in section, of a deep bodied, small, rounded species apparently lacking rugae or ribs, but with relatively stout spines, which somewhat resembles a Lower Carboniferous leiproductid, such as genus Magnumbonella Carter (Fig. 7). 7. Two specimens like the conical shells described below have variably developed radial ribs, starting within 5 mm of the apex and with a low profile. The cone apex seems to have some attachment spines and others are arranged widely on the cone. The species resembles Planispina armata (Girty) from the mid Carboniferous of the USA (Fig. 8). 8. There are, in addition, two pieces of a strongly ribbed thynchonellid and a _ section through a _ probable reticulariacean. These identifiable productids may prove to be new taxa. However, this paper deals only with the more common (ie. in the collection at hand) subconical species. Of these 25 fragments, 21 are of the subconical valve while four include separate parts of what is clearly a different valve and is interpreted as the second valve of the same species. 52 C.H.C. BRUNTON METHODS The shell material on most specimens is thin, somewhat laminar in appearance and takes the buff colour of the matrix. In order to establish the nature of this shell, slivers from two specimens were studied by qualitative energy dispersive x-ray microanalysis using scanning electron microscopy in the Department of Mineralogy, The Natural History Museum, London. The results show a strong dominance of calcium, with no magnesium or phosphorus present (Fig. 9). The mineralogy indicates dominance of calcium carbonate (lacking magnesium) and the lack of phosphate precludes the presence of apatite. The shell is not calcophosphatic so articulate brachiopods could have been the animals which secreted this shell material. It seemed unlikely that more material would become available, so it was important to preserve what we had. For this reason only two specimens were sectioned in an attempt to determine what internal structures were present. However, some internal details could also be seen on broken surfaces which cut across specimens while the rock was being broken, as well as on naturally weathered surfaces cutting across the partial interiors of two specimens. Careful examination of all specimens under a binocular microscope, commonly making drawings using a Wild drawing arm, gradually allowed the recognition of some consistent features on several specimens, which provided a form of orientation. The recognition of the same structures in different views and sections has allowed a general picture of the morphology of these specimens to be built up. Portions of shell from near the apex and the distal regions of the cone, and from the supposed dorsal valve have been studied uncoated in the environmental chamber of an ISI ABTS55 scanning electron microscope and coated using a $2500 Hitachi machine, both in The Natural History Museum. Fig. 1 Stipulina-like species, ventral valve exterior, BD9673, <4. Figs 2,3 Two views of anon-rugose Thomasella-like species. 2, exfoli- ated dorsal valve interior and the flanged and geniculated trail, BD9677. 3, oblique view of a complete specimen viewed anterolater- oil ally showing the strongly ribbed trail, BD9676b, x3. Fig.4 Two views of cf. Thomasella showing the posterior rugae, flanged trail and ventral spines, BD9673(0), x5. | Fig.5 A Plicatiferinid showing the exfoliated dorsal valve interior, BD9674, x4. i Fig.6 Pectenoproductus ventral valve exterior, BD9675, x2. | Fig.7 The incomplete leioproductine ventral valve, BD9688(1), <3. Fig.8 Cf. Planispina sp. viewed almost apically, BD9688(0), <1.5. Fig.9 Plot from qualitative energy dispersive X-Ray microanalysis of non-coated fragments using scanning electron microscopy. The plot displays a major calcium peak (plus a secondary peak to its right), but no phosphorus and only minor amounts of elements with lower atomic weights, to the left. PROTORICHTHOFENIOID BRACHIOPOD FROM URALS SYSTEMATIC DESCRIPTIONS The material described here is housed in the BMNH collections of The Natural History Museum, London. Specimens are uniquely recognised by BD registration numbers. Order PRODUCTIDA Waagen, 1883 Suborder STROPHALOSIIDINA Schuchert, 1913 DIAGNOSIS. Productida retaining ventral commonly, toothed articulation. interareas and, Superfamily RICHTHOFENIOIDEA Waagen, 1885 DIAGNOsIs. Ventral valve conical or sphenoidal, dorsal valve recessed below the ventral margins. Normally attached by cicatrix and/or spines. Family ZALVERIDAE nov. DIAGNOsIs. Richthofenioids lacking external body spines, with shallow body cavities and short subparallel ventral ridges associated with a near apical chamber involved in the articulation of the valves; marginal ventral valve protective structures absent. COMMENTS. Currently only the one genus is known. The family differs from the Richthofeniidae most clearly in its lack of external spines and attachment to the substrate. Although Collumatus Cooper & Grant, 1969, lacks spines this Permian genus is attached to the substrate by concentric sheets of shell | surrounding the base of the ventral valve. Genus ZALVERA nov. /DIAGNOsIs. Zalveridae retaining small juvenile ventral valve at japex and with strong brachial impressions. \EtyMo.ocy. Anagram of the letters of the name Lazarev. as Teguliferina(?) uralica and by Likharev, (1932) as Keyserlingina caucasica. Unfortunately, little is known of the uncertain. Likharev (1931, 1932) specifically noted the absence of internal dorsal valves, wondering if they ever existed. xternally both these species seem to fit better here than in rorichthofenia, which is a true teguliferinine with external pines. In addition, two specimens referred to as Proteguliferina? y Winkler Prins in Sanchez de Posada et al. (1993) belong in alvera. See further discussion under the species description. alvera sibaica sp. nov. Figs 10-24 YPE SPECIMEN. Holotype, BD9653, from the Kordailovskaya ormation of late Bashkirian to early Moscovian, mid Upper arboniferous age, 6km up-stream from Pokrovka, Ural River nthe southern Urals, Russia (Figs 20a—c). 53 Figs 10-13 Zalvera sibaica gen. et sp. nov. from late Bashkirian to early Moscovian rocks in the southern Urals, Russia. 10, apical view of a slightly crushed specimen including the small regular looking initial growth stages of the ventral valve (arrow). The umbo is to the left. BD9671, <3. 11, side view of a specimen showing the inferred subparallel ridges as white lines on the shell (arrow) and rounded apex. BD9669, <1.5. 12, part external mould of a specimen viewed towards the apex, in which parts of the subparallel ridges are visible. The rock surface to the left cuts a nodose outgrowth (arrow), BD966la, 2. 13, apical view of a specimen associated with the exter- nal mould of figure 12 showing partially exposed subparallel ridges, BD9661b, x2. DIAGNOSIS. Za/vera with irregular rounded dorsal outline and apex profile, rugae relatively prominent. ETYMOLOGY. Species named from the Sibay stream, where the specimens were found. MATERIAL. Allspecimens of Z. sibaica are from the one locality in the southern Urals (BMNH BD9653 (the holotype), BD9654-9672). A few other fragments occur with specimens of other species. DESCRIPTION. In one specimen (BD9671) the earliest growth stage of the subconical valve resembles a tiny (ca. Smm diameter) productid valve (Fig. 10). This consists of a subtriangular, gently convex valve exterior, which appears to be covered by closely spaced, but fine spine bases. There is, therefore, a question as to the reliability of this small valve being part of the species. The specimen was cut medianly and appears to show continuous shell between the small spinose valve, its thick-shelled gutter-like surround and the remaining 54 Figs 14,15 Zalvera sibaica gen. et sp. nov. from late Bashkirian to early Moscovian rocks in the southern Urals, Russia. 14, oblique apical view of a partially exfoliated specimen showing the fine internal tuberculation on the internal mould (arrow) and positions of the subparallel ridges (arrows), BD9664, <7. 15, apical view of a specimen with a xenomorphic apex associated with a bryozoan and showing the subparallel ridge positions, BD9663, x5. thin-shelled cone. A small apical spinose valve was also noted by Tschernyschev (1902) in his original description of Tegulifera(?) uralica. The material has been checked recently by Dr Lazarev (personal communication) who confirmed its presence, so for now it 1s accepted as part of Z. sibaica. Most of the material consists of broken parts of a subconical to subcylindrical structure with a flattened, weakly rounded, ‘base’ (Fig. 11), extending directly from the initial tiny spinose valve. The cone expanded rapidly to about 15mm in diameter and then expanded gradually to as much as 25mm in diameter. The preserved length is variable, but no specimen exceeds 25mm; some are much more squat in shape. The exteriors are smooth apart from irregularly developed rugae (Fig. 11) and very rare non-hollow, nodose outgrowths on the subcylindrical area (Fig. 12). Growth lines are commonly visible. The shell material is thin except for restricted areas in which there are internal ridges or blunt, rounded endospines. A few specimens display two layers of thin shell in the rugose cone region, each separated by a narrow (less than Imm) layer of sediment. The place from which the inner, younger, shell layer originated can be found rarely, but indicates that these lamellose layers were growth structures resulting from a mobile mantle epithelium. On the weakly rounded ‘bases’, beyond the small initial spinose valve, the shell material appears to be finely pseudopunctate, with small tubercles on internal surfaces (Fig. C.H.C. BRUNTON 14). Breakage and shell exfoliation is common in this area, with the result that complete undamaged exteriors are rare. However, this does allow recognition of some structures from the outside. Most obvious isa pair of plates or ridges (Figs 11, 13), about 2 to 3mm apart distally, diverging slightly from within about Imm to between 5 and 10mm from the apex, and extending to or beyond the growth stage at which the cone expansion slowed to give a more subcylindrical profile. The presence of drusy crystals between these plates indicates an original cavity in that region. Similar crystals are commonly present apically and externally to} these plates in specimens from which the base of the subconical valve appears to have been broken. More complete specimens show signs of shell growth distortion over this basal area; in one example a fenestellid bryozoan is adpressed to the surface (Fig. 15). | The cutting of two relatively complete subconical valves failed} to reveal a second valve within the cone, although one specimen has a platform-like 2.8mm extension into the ‘internal’ spaced originating about 5mm from the apex. This specimen was cut between the pair of subparallel plates, allowing reference to position in the shell, and these plates are probably connected.) thus forming a chamber. More information about interiors was obtained from broken sections through three specimens, one of which is additionally weathered and displays parts of a flat, thi structure lying near the base of the cone, which can be PROTORICHTHOFENIOID BRACHIOPOD FROM URALS Fig. 16 Drawing of a natural section through a specimen with its apex uppermost and in which part of a dorsal valve is preserved (B). A = ventral valve; C = crystaline cavity infilling; D = part of the ventral valve internal articulatory structure; E = external mould of rugae near the apex. From specimen BD9663(2). interpreted as a second valve (Fig. 16). The available specimens all display a consistent relationship between the conical valve and internal plate-like structures, indicating that the latter are remnants of a second valve in their positions of articulation, or the support structures for a second valve. Four different-looking specimens in the collection are taken as being conspecific. These are fragments of nearly complete, oval-shaped valves, 12 to 15mm wide, and are interpreted as interior surfaces, one of which has a counterpart. Their size and general outline would allow them to have fitted into the more apical region of the conical valves. One edge, seemingly parallel to the maximum width, shows a slight median flattening and is taken as being posterior. The most obvious feature is a pair of weak, crescent-shaped ridges (Fig. 17a, b), which bound shallow depressions, follow close to the edges of the valve and originate from posteromedian positions on each side of a low broad median ridge. Surfaces on either side of the crescent-shaped ridges are finely endospinose (Fig. 18). Posteriorly, close to the straighter sector of the margin, is a pair of shallow pits, between which is the posterior end of the wide median ridge. Posteromedianly, a transverse ridge is situated just anterior to the posterior flattened sector. This expands widely anterior to the median ridge, leaving a T-shaped pair of lateral protuberances, 2mm wide (Fig. 19). The lateral and anterior margins of these valves are of thin shell substance, which display growth lines, and seem to be reflexed away from the internal surface, resembling a valve trail. SHELL STRUCTURE. Scanning electron microscopy of shell sections (Figs 22, 23) and fractured fragments (Fig. 24) show the shell to be composed of somewhat recrystallised laminae, probably originally made up of thin laths. Macroscopic examination revealed rather fine pseudopunctation, especially in the apical regions of the ventral and dorsal valves (Fig. 18), and thus confined mostly to the body region. The conical trail, beyond the body cavity, has fewer internal tubercles (Fig. 22) indicating a reduction in the pseudopunctation. INTERPRETATION. The scant and poorly preserved characters available indicate a subconical valve, which is interpreted as 55 ventral, within which lies a dorsal valve close to the apex of the shell, resulting in a very shallow body cavity. Distal to this was a relatively long subcylindrical skirt of thin shell. The posterior transvere ridge in the dorsal valve articulated with a pair of grooves associated with the subparallel plates in the ventral valve (Fig. 21). Additionally, the dorsal valve bent dorsally around its lateral and anterior margins forming a short trail against the longer internal surface of the ventral valve (Figs 19, 21). There is no direct evidence for the dorsal trail being as long as that of the ventral valve, but it is possible that there was an extremely thin layer of dorsal shell supporting the mantle in this trail region. On the other hand, a long dorsal trail might have hindered the opening of the shell, performed by contraction of diductor muscles attached to the poorly differentiated cardinal process situated between the ventral articulatory ridges. A short dorsal trail would have left the internal mantle epithelium of the ventral valve open to the sea and vulnerable. Cowan (1970) suggested similar exposed epithelia in lyttonioids, and similarly exposed epithelia occurs in other Productida, including teguliferinids. However, the evidence for this in the present material is insufficient to allow further discussion. The internal morphology of the few dorsal valves available indicates a standard anatomy in which the lophophore was probably a schizolophe, strung from brachial ridges close to the edges of the body cavity (Fig. 17). Dorsal adductor muscles were attached posteromedianly, anterior to the articulation ridge and between the median ridge and posterior ends of the brachial ridges. Diductor muscles were probably attached at a small posteromedian boss, which hardly deserves the term cardinal process. Externally, the ventral apical region is somewhat distorted on some specimens, which might indicate that initial growth closely followed the substrate. However, the apparent lack of external body spines, which might have been used to fix specimens, and the variable nature of this ‘basal’ deformation seems to indicate that specimens were not cemented or closely adpressed to a hard substrate, but occurred on, or partially buried in, a relatively soft substrate. Unfortunately, field observations are lacking on both the orientation of specimens in the rock and on the nature of adjacent lithologies. Drusy fillings in body cavity regions could have developed whichever way up the specimens were entombed in the sediment, provided both valves were preserved and the shell was closed. However, the rare specimens showing these features invariably have drusy crystals within the body cavity, not dorsal to the dorsal valve, indicative of an apex down position in life. The rock appears not to include potential hard substrates, other than the fossils themselves, although algae could have caused surfaces to become firm. Although probably not fixed to a hard substrate, these specimens appear to have resembled richthofenioids lacking adult spines, and lived in areas of soft, fine sediment. If they sat, rather cup-like, with their bases buried in sediment, the ventral trails would have raised the inhalent areas well above the sediment surface. To have achieved this position with only a thin-shelled ventral valve and no anchoring structures indicates that the environment was probably one of quiet water and fine-grained sedimentation. DISCUSSION. Specimens somewhat resembling this material were described by Tschernyschev (1902) from Lower Permian, Asselian, rocks on the river Yuresan, also in the Urals, Russia, as Tegulifera (?) uralica. One specimen figured by Tschernyschev (1902, fig. 85, pl. 60, fig. 14) has a small triangular, regular-looking productid valve at its apex like that on Z. sibaica C.H.C. BRUNTON 56 PROTORICHTHOFENIOID BRACHIOPOD FROM URALS 2a 2b 5mm Fig. 21 Reconstruction of the apical area of Za/vera in median longi- tudinal section (1a), between the subparallel ridges, and an apical view of the ventral valve at the arrow position (1b). In 2a the section cuts one of the subparallel ridges in which the articulatory groove is situated (2b). Ventral valve black; dorsal valve stippled. (BD9671) (Fig. 10). Also, the species seem to be of comparable size and both have similar concentric ornamentation. Dr Lazarev has inspected the Tschernyschev specimen and confirms the similarity. Another similar-looking species was first described by Likharev in 1931 as Teguliferina? (Chaoella) caucasica and again in 1932 as Keserlingina caucasica. It came from Permian rocks in the Caucasus, originally described as P,,, but now thought to be of Kazanian, early Upper Permian age. Likharev also recorded related specimens from the Sim river in the southern Urals. Likharev (1932) gives a measure of 9mm for the maximum width, so his specimens are about half the size of the new material. Otherwise the illustrations closely resemble the Carboniferous specimens and one figure (1932, pl. 2, fig. 9b) seems to show similar subparallel marks near the apex. In describing caucasica Likharev briefly referred to uralica, but other than discussing the apparent small ventral valve at the apex he wrote little to differentiate between the two species. Thus, somewhat similar specimens to Z. sibaica occurred in the Urals at Bashkirian to Moscovian boundary times and again in the Permian. Neither Tschernyschev’s JT. uralica nor Likharev’s K. caucasica belong in Teguliferina or Keyserlingina, the latter belonging with the Lyttonioidea on account of its lobate interiors. Teguliferina and Proteguliferina belong in the Richthofenioidea and are thus more closely related to the new material than is Keyserlingina. For instance, Proteguliferina displays a weakly concave dorsal valve within a gently convex ventral valve which does not reach the internal margins of the spinose ventral valve. Its ventral marginal epithelium may, therefore, have been exposed. Girty (1908) described two species, Tegulifera armata and T. kansasensis, from rocks of late Missourian (mid Kasimovian) 57 age, from localities in Illinois and Kansas respectively. These species differ in important characters from the new Urals specimens described here. The American specimens are less deeply conical, less rugose, are radially ornamented, and are attached by cementation and external spines, as well as having internal ventral spines. These species were placed by Muir-Wood & Cooper (1960) into Planispina Stehli, a genus assigned to the Teguliferininae. The radial ribbing and possible spinose exteriors of two specimens in the Urals collection (Fig. 8) resemble P. armata (Girty, 1908, pl. 20, fig. 10). Sutherland (1989) reported another species, well preserved as silicified specimens, from early Upper Carboniferous (Morrowan [= Bashkirian]) rocks in Oklahoma, USA. This material as yet remains un-named, but is probably closely related to Teguliferina, with somewhat similar dorsal valve interiors and similar external rhizoid fixing spines. These Morrowan specimens, therefore, appear to be the earliest known true teguliferinids, and are the earliest known Richthofenioidea. The material from the Urals is of similar age to that described by Sutherland (1989) but is very different in character, resembling more closely the specimens described by Tschernyschev (1902) and Likharev (1931). The brachiopod relationship of the new Urals material is no longer in doubt and the general form of the shell is highly indicative of a richthofenioid relationship. However, other than in the Permian genus Collumatus Cooper & Grant, 1969, from Texas, the richthofenioids have external spines to aid the fixing of specimens to hard substrates, commonly within reef environments, while Z. sibaica has no such spines. Apart from this difference, the general architecture of the shell fits with that of richthofenioids; a conical ventral valve with a dorsal valve recessed below its margins, the two valves articulating, not by true ventral teeth and dorsal sockets, but by dorsal protuberances fitting into ventral cavities. In detail the Urals specimens differ from the general richthofenioid pattern: they lack external spines, other than perhaps at the earliest stages in ontogeny when ventral valves show signs of fine spines for about 5mm of growth; the body cavity is shallow, with the dorsal valve deeply recessed below the ventral margin; the dorsal valve has internal structures producing relatively thick shelly ridges, the brachial impressions and wide median ridge; the dorsal valve has short, geniculated, thin-shelled margins extending a short distance up the ventral valve interior; the ventral valve interior has a simple posteromedian structure of subparallel plates acting as supports for the dorsal valve; the ventral valve cone interior has sparcely distributed blunt, well rounded, endospines protruding into the space above the dorsal valve exterior; there is no indication of any complete protection for the opening to the subconical valve margins, as in most true richthofenioids. In a biostratigraphical description of regions in Cantabria, northern Spain, Sanchez de Posada et al. (1993) listed ‘Proteguliferina? n. sp. from “Kasimovian’ rocks. Dr C. F. Winkler Prins has kindly lent me the two specimens from which this reference was made, which he now dates as Moscovian, possibly Podolsky age. They do appear to be congeneric with the new Urals specimens. Specimens (also seen) determined by Figs 17-20 Zalvera sibaica gen. et sp. nov. from late Bashkirian to early Moscovian rocks in the southern Urals, Russia. 17, part and counterpart of an incomplete dorsal valve showing probable brachial impressions and the median ridge. The valve margins curve away into the rock in 17a, BD9659, x4. 18, scanning electron micrograph showing the finely endospinose surface within the dorsal valve brachial impression, BD9659a, x60. 19, a rather elongate, deformed and partly exfoliated dorsal valve interior; posterior is to the top, showing the transverse ridge (exfoliated and arrowed), interpreted as the dorsal articulation structure, and one brachial ridge, on the right, BD9670, x10. 20, holotype viewed apically and ‘anteriorly’ (1.5) and obliquely apically (x5) showing the positions of the subparallel ridges ‘posteriorly’. BD9653. C.H.C. BRUNTON Fig. 25 Zalvera sp. from early Upper Moscovian rocks north of Bran- cosera, Palencia, Spain. 25a,b, apical and iateral views of the speci- men (<1) showing an ‘umbonal’ region (arrowed). 25e, an enlargement (<3) of 25b showing pseudopunctate shell (arrowed) and the layered nature of the valve, each with thin sediment between. National Museum of Natural History, Leiden, WAG 77. Winkler Prins (personal communication, September 1993) as Proteguliferina sp. from Austria are not Za/vera, but are more like an early form of the Permian teguliferinid Acritosia. The two Moscovian specimens from northern Spain resemble Z. sibaica closely, but appear to differ in having well flattened apices, suggesting closer attachment to the substrate than seen in the specimens from the Urals. Both specimens show signs of the subparallel internal ridges, the apical internal microtuberculation and both have comparable external irregular rugae. These similarities between the Spanish and Russian specimens reinforce the concept of widespread marine faunas in Eurasia during Upper Carboniferous times. Present evidence, therefore, indicates two almost contemporaneous stocks in the early to mid Upper Carboniferous, one in North America which evolved into the Richthofeniidae, the other in Russia which, although considered _ as belonging in the Richthofenioidea, was an _ early ‘experimental’ stock which lost its spines early in ontogeny and lived on relatively soft substrates, unlike the American forms : Figs 22-24 Zalvera sibaica gen. et sp. nov. from late Bashkirian to early Moscovian rocks in the southern Urals, Russia. 22, a cut and lightly etched section through a ventral valve just distal to the inter- nal thickening against which the dorsal valve rested. Apex to the top right, exterior to the left. Irregularities in the laminae are oblique sec- tions through pseudopunctae, apparently with taleolae (arrows), BD9669a, 160. 23, same section, closer to the apex, x1200. 24, a fracture flake of shell from a thickened area of a dorsal valve. Signs of original laths can be seen centrally, BD9670, x5000. PROTORICHTHOFENIOID BRACHIOPOD FROM URALS which were attached and retained spines. While some stratigraphically younger examples of this aspinose stock, in addition to those from northern Spain, may yet be discovered, it seems it did not persist after the Kazanian, when the Caucasian species of Likharev died out. The Permian genus Col//umatus would appear not to have been derived from this stock, but to be a true richthofenioid which had also lost its spines. A feature of taxonomical importance in the specimens described by Sutherland (1989) from Oklahoma, but not seen in this new material, is the juvenile ventral interarea. A small ventral interarea is also reported in the Teguliferininae (Muir-Wood & Cooper, 1960), which belongs in the Richthofenioidea. It is the presence of interareas in these Carboniferous species that places the Richthofenoidea in the Strophalosiidina, the structure being lost in the more common Permian richthofenioids. In Zalvera it seems the earliest growth of the ventral valve was normal for productids, but whether it involved the growth of a juvenile interarea is unknown. If present it might have become incorporated into the later subconical valve growth. In any event, once growth changed, after the initial 4-S5mm, from being a small finely spinose subtriangular valve to a rapidly expanding rounded cone with a spineless exterior, shell growth was holoperipheral. In the North American species there was a short period when the ventral interarea grew before the posterolateral mantle margins grew posteromedianly to continue the style of shell secretion seen for the rest of the valve. This left the juvenile interarea preserved, but with a short suture line posteromedianly where the two mantle margins had grown together and fused, allowing the typical teguliferinid cone to grow. In the Urals material this posterior fusion of the mantles occurred earlier in growth so that no interarea has been preserved. This raises the question as to whether Zalvera should be assigned to the Strophalosiidina. The alternative is to consider Za/vera within the Productidina as a unique aberrant offshoot showing tendencies towards the morphology of the Richthofenioidea. This seems less likely than placing Zalvera in the Richthofenioidea and accepting that the interarea never developed in these unusually-shaped brachiopods. It is hoped that more of this material will become available so as to allow more complete preparation and a better insight into the way in which this strange brachiopod grew. There is no clear evidence for the origin of the Zalveridae, but it is possible to suggest that the ancestral stock was within the Strophalostidina. The cone development seems to have been from exaggerated, and posteriorly fused, growth of the ventral 59 trail. Within strophalosiidines the Aulostegoidea includes many groups with elaborate trails, and as they lack a toothed articulation and have variably developed interareas, they provide possible ancestors for Zalvera. ACKNOWLEDGEMENTS. I thank Dr S. Lazarev, Palaeontological Institute, Moscow, for information about this material and other specimens in Russian collections; Dr Cor Winkler Prins, National Museum of Natural History, Leiden, for the loan of comparable material from Spain and Austria; Dr Patrick Sutherland, University of Oklahoma, for information about his specimens, and Dr R. Cocks for reading and commenting constructively on a draft version of this paper. REFERENCES Cooper, G. A. & Grant, R. E. 1969. New Permian brachiopods from Texas. Smithsonian Contributions to Paleobiology, 1: 1-20 Cowen, R. 1970. Analogies between the Recent bivalve Tridacna and the fossil brachiopods Lyttoniacea and Richthofeniacea. Palaeogeography, Palaeoclimatology and Palaeoecology, 8: 329-344. Girty, G. H. 1908. On some new and old species of Carboniferous fossils. Proceedings of the United States National Museum, Washington, 34: 281-303, pls 14-21. Likharey, B. K. 1931. Uber eine problematische Brachiopode aus den unterpermischen Ablagerungen des Nordlichen Kaukasus. Annuaire des Société Paleontologique de Russie, Leningrad, 9: 157-161, figs 1—S. 1932. Fauna of the Permian deposits of northern Caucasus, 2 Brachiopoda, Lyttonidae. Trudy Vsesoyuznogo geologo-razvedchnogo ob'edineniya NKTP [Transactions of the Geological Prospecting Service of USSR], Leningrad & Moscow, 215: 55—84 (in Russian), 85—111 (in English), Spls. Muir-Wood, H. M. & Cooper, G. A. 1960. Morphology, classification and life habits of the Productoidea (Brachiopoda). Memoirs of the Geological Society of America, New York, 81: 447pp, 135 pls. Sanchez de Posada, L. C., Martinez Chacon, M. L., Mendez, C. A., Menendez Alvarez, J. R., Truyols, J. & Villa, E. 1993. El Carbonifero de las regiones de Picos de Europa y Mantuo del Ponga (zona Cantabrica, N de Espana): Fauna y biostratigrafia. Revista Espanola de Paleontologia, No. Extraordinario: 87-108. Schuchert, C. 1913. Class 2. Brachiopoda, in Zittel, K. A., Text book of Palaeontology, 1: 355-420. London. Sutherland, P. K. 1989. Possible ancestor to the late Carboniferous/early Permian Teguliferiniid brachiopods in the Middle Carboniferous of Oklahoma, USA. XTe Congres International de stratigraphie et de géologie du Carbonifeére, Beijing 1987, Compte Rendu, 2: 355-360, pl.1. Tschernyschev, T. 1902. Die obercarbonischen Brachiopoden des Ural und des Timan. Memoires du Comite géologique, St Petersburg, 16 (2): 749pp, 63 pls (in Russian & German). Waagen, W. H. 1882-1885. Salt Range Fossils, Pt. 4 (2), Brachiopoda. Palaeontologia Indica, Memoirs of the Geological Survey of India. Calcutta. Pe. Se ry a { te bb ~ — a ee al = iad Wee vans iia emma sg " se 2: a y Bull. nat. Hist. Mus. Lond. (Geol.)52(1): 61-89 Issued 27 June 1996 The Upper Cretaceous ammonite Vascoceras Choffat, 1898 in north-eastern Nigeria P.M.P. ZABORSKI Department of Geology and Mining, University of Jos, P M.B. 2084, Jos, Nigeria CONTENTS Introd UC On mmew ss Sete. cece hie: Mon bee ante Rages os) idea Sd OL OD See et tee SV Stehlatic descnptlonspeeis-e-sees ee eee eee Family Acanthoceratidae Grossouvre ............. Subfamily Acanthoceratinae Grossouvre .... Genus Paravascoceras Furon. ............20000020+ Paravascoceras cauvini(Chudeau)_ ........... (CEIUIS JANA DNERATRAIES IMSINON'E cosncocceseeqacncocncconasee see sc0ce caccceecevoconcceakeouseecnoosconeeboreoceoonen: Pseudovascoceras nigeriense (Woods) RamilvavascoccratidaciD ouvillevenss..tusteee set. ets, .aciesiobc donates Saas SOE SP en peo: RE ot ole oe Subfamily Vascoceratinae Douvillé Gonusmyascocerasl @lio tit tae memento nce ccc cre, Rea eee eae oe ER RO ic Ee Vas GOGEN ASV OOASIS DMIOW Maree cere tree eeen cee ore scan 2 soe Reyer te ere oe RN 72 Vascoceras bullatum Schneegans Vascoceras globosum (Reyment) Vascoceras globosum costatum (Reyment) Vascoceras globosum globosum (Reyment) Vascoceras globosum proprium (Reyment) Vascoceras obscurum Barber VOScOcerasiiaricii (Gay Att ieee trace anette ne Mecca ae occa econ wks cose Ge nade ore en nen Stratigraphical and phylogenetic discussion Acknowledgements INGTSRSNEES socrsscosaeeoonsc Appendix Synopsis. Large collections of ammonites that have been referred at one time or another to Vascoceras Choffat, can be made under tight stratigraphical control in north-eastern Nigeria. The following forms are present, in order of stratigraphical appearance: Paravascoceras cauvini (Chudeau); Vascoceras woodsi sp. nov.; V. bullatum Schneegans, V. globosum costatum (Reyment), V. globosum globosum (Reyment) and Pseudovascoceras nigeriense (Woods); V. globosum proprium (Reyment); V. obscurum Barber; and V. harttii (Hyatt). Only the last three occur in the Lower Turonian; the remainder are restricted to the Upper Cenomanian, the earliest appearing above the level of the European Metoicoceras geslinianum Zone. Paravascoceras Furon (type species Vascoceras cauvini Chudeau) is retained as a separate genus for forms derived from Nigericeras Schneegans. Pseudovascoceras gen. nov. (type species Vascoceras nigeriense Woods) is proposed for ribbed and multituberculated forms thought to have arisen from Cunningtoniceras Collignon. Paravascoceras and Pseudovascoceras are most properly referred to the subfamily Acanthoceratinae since they have an origin separate from that of Vascoceras. Several of the taxa present show a high degree of individual variation. Palaeoecological factors played an important role in their geographical distribution and probably also in their potential for polymorphism. Separate lineages converged on a ‘Vascoceras morphology’ in north-eastern Nigeria as a response to the particular environmental conditions prevailing there during Late Cenomanian and Early Turonian times. Its ammonites have been described by Woods (1911), Reyment INTRODUCTION In Tethyan regions ammonites referred to the genus Vascoceras Choffat, 1898, are often present in large numbers in the Upper \Cenomanian and Lower Turonian. The upper Benue Trough a in north-eastern Nigeria is a classic region for such faunas. © The Natural History Museum, 1996 (19546), Barber (1957, 1960), Meister (1989), Zaborski (1990a, 1993, 1995) and Courville (1992). Species of Vascoceras have been widely employed in biostratigraphical analysis in Nigeria but the taxonomic treatment applied to them has varied widely from author to author. The north-eastern Nigerian faunas are of particular interest since large collections can be made under 62 P.M.P. ZABORSKI WESTERN INTERIOR ZONES 20m PINDIGA ASHAKA Pseudaspidoceras flexuosum TURONIAN 15 CENOMANIAN Nigericeras scotti | 10 he ere aa Neocardioceras juddii to Burroceras clydense ——— 5 ——— a —— Sciponoceras gracile BIMA SANDSTONE o-: $e a ee YOLDE Fm. Fig. 1 Stratigraphical sections through the limestone-bearing parts of the Pindiga Formation at Ashaka and Pindiga with letter codes identifying limestone units mentioned in the text. Approximate biostratigraphical correlations with the ammonite biozones of the western interior of the United States (after Cobban et a/. 1989, Hancock 1991) are also indicated. tight stratigraphical control, especially at Ashaka quarry. Furthermore, dissection of adults is frequently successful in recovering well-preserved inner whorls which in some cases are invaluable for identification purposes, as well as for analysing ontogenetic development. These attributes have allowed a revised taxonomy to be presented here for the Nigerian faunas. The family Vascoceratidae as a whole has been discussed by Spath (1925), Furon (1935), Schneegans (1943), Reyment (19546, 1955, 1956), Barber (1957), Wiedmann (1960), Cooper (1978) and Wright & Kennedy (1981). From the time of its proposal it has been recognized that the genus Nigericeras Schneegans, 1943 is morphologically intermediate between the subfamilies Acanthoceratinae and Vascoceratinae. More recently, a number of additional genera of intermediate character have been decribed, a notable feature being the combination of a vascoceratine-type suture pattern with an acanthoceratine-type ornament. Such intermediates | include Microdiphasoceras Cobban, Hook & Kennedy (1989: 53), Rubroceras Cobban, Hook & Kennedy (1989: 54), Fikaites | Zaborski (1993: 362) and Pseudovascoceras, described herein. It j is believed here that the family Vascoceratidae is polyphyletic, | including homeomorphic derivatives of various acanthoceratine } genera. In Nigeria at least the same is true of forms previously referred to the genus Vascoceras. | UPPER CRETACEOUS AMMONITE _ SYSTEMATIC DESCRIPTIONS Repositories. Unless otherwise stated all the specimens referred to below are housed in the Department of Palaeontology, The Natural History Museum, London, their register numbers being prefixed with the letter C. In addition to those specifically listed, large numbers of Paravascoceras cauvini, Vascoceras woodsi, V. bullatum, V. globosum costatum, V. globosum globosum and Pseudovascoceras nigeriense have also been studied. Provenance of material. The ammonite-bearing horizons at the / two main localities in north-eastern Nigeria, Ashaka and Pindiga, are shown in Fig. 1. A fuller description of these sections and lists of their ammonite faunas were given by Wozny & Kogbe (1983), Popoff et a/. (1986), Meister (1989) and ' Courville (1992). The Pindiga section has been described by _ Barber (1957), Carter et al. (1963), Wozny & Kogbe (1983) and Popoff et al. (1986). The whereabouts of other ammonite localities mentioned in the text were shown by Zaborski (1990a: fig. 1). Dimensions (in mm). D, diameter; Wb, whorl breadth; Wh, whorl height; U, umbilical diameter. Figures in parentheses are dimensions as a percentage of the total diameter. Family ACANTHOCERATIDAE Grossouvre, 1894 Subfamily ACANTHOCERATINAE Grossouvre, 1894 | Genus PARAVASCOCERAS Furon, 1935 Me Pardcanthoceras Furon, 1935; Pachyvascoceras Furon, 1935; | Broggiiceras Benavides-Caceres, 1956) | i | | Superfamily ACANTHOCERATACEAE Grossouvre, 1894 TYPE SPECIES. Vascoceras cauvini Chudeau, 1909; by the subsequent designation of Reyment, 1955. REMARKS. Furon (1935: 60) proposed Paravascoceras as a subgenus of Vascoceras and it has subsequently been treated as such by, for example, Schneegans (1943), Cooper (1978), Howarth (1985) and Meister er al. (1992). Others, for example Reyment (1955), Barber (1957), Wright (1957), Freund & Raab | (1969), Schobel (1975) and Meister (1989), have regarded it as a |distinct genus while recently it has been widely listed as a _}synonym of Vascoceras (see, for example, Berthou er al. 1985, | Kennedy et al. 1987, Luger & Gréschke 1989, Cobban et al. 1989). | Furon’s original diagnosis of Paravascoceras specified non-globular forms characterized by a simple suture pattern which was said to distinguish it from Paracanthoceras Furon (1935: 59) (type species, by monotypy, Vascoceras (Paracanthoceras) chevalieri Furon, 1935). Both these forms _ show strong ventral ribbing in their later growth stages. Furon included V. (P.) cauvini, V. (P.) cauvini var. semiglabra Furon (1935) and V. (P.) chudeaui Furon (1935) in Paravascoceras. The last two are here regarded as synonyms of P. cauvini. Schneegans (1943: 127-128) showed that sutural differences between Paravascoceras and Paracanthoceras were insignificant and demonstrated the latter to be a synonym of the former. Indeed, V. (Paracanthoceras) chevalieri itself is a synonym of Paravascoceras cauvini. Schneegans gave a revised diagnosis of Paravascoceras stressing its vascoceratid suture pattern, ovoid to globular whorl section, lack of tubercles and possession of simple ventral ribs or folds in the adult stages. The absence of 63 umbilical tubercles has since been cited as a chief distinguishing feature of Paravascoceras (see, for example, Freund & Raab 1969, Sch6bel 1975, Meister 1989, Meister et al, 1992). Berthou et al. (1985), however, regarded the presence or absence of umbilical tubercles in Vascoceras as an inadequate basis for generic and subgeneric diagnosis, a conclusion accepted by Kennedy ef al. (1987) and Cobban et al. (1989). This view is supported here. There is great inconsistency in this feature even within individual species of Vascoceras. Meister et al. (1992: 70: see also below) further showed that P cauvini may itself show umbilical tubercles at certain growth stages. As pointed out by Schneegans (1943: 127), the juvenile stages are often of greater value in taxonomic subdivision of Vascoceras than the often highly variable middle and adult whorls. Morphological and stratigraphical evidence from north-eastern Nigeria indicates that Paravascoceras was derived from Nigericeras (type species Nigericeras gignouxi Schneegans, 1943: 119, pl. 5, figs 10-15 = N gadeni (Chudeau); by the subsequent designation of Reyment 1955: 62), an origin separate from that of Vascoceras (see below). In recognition of this probability Paravascoceras is here treated as a distinct genus. In view of its ornament and suture pattern Nigericeras should be included in the subfamily Acanthoceratinae (see also Kennedy et al. 1989, Cobban et al. 1989, Kennedy & Wright in press). Paravascoceras, therefore, cannot be maintained within the Vascoceratidae but should be transferred to the Acanthoceratinae also. There remain problems in providing a reliable and unambiguous morphological diagnosis of Paravascoceras. Its members are generally compressed, moderately involute, without umbilical tubercles and with strong regular ribbing on the outer flanks and venter in the later growth stages. The last two features are not, however, consistent while certain rather depressed forms may belong in the genus. Vascoceras (Pachyvascoceras ) Furon (1935: 58) (type species Vascoceras ( Pachyvascoceras ) crassus Furon 1935: 58, pl. 3, figs 2a, b; by the subsequent designation of Reyment 1954b: 257) was proposed on the basis of its globular shape, deep narrow umbilicus and lack of adult ornament. None of these morphological features is sufficient to distinguish Pachyvascoceras from Vascoceras. Whorl breadth is often particularly variable within individual species of that genus. The phylogenetic affinities of V. (P.) crassum, however, may lie with Paravascoceras rather than Vascoceras. Meister et al. (1992) described topotype material which they regarded as variants of Paravascoceras cauvini with which they are transitional (see also Schneegans 1943). Pachyvascoceras is accordingly treated here as a probable synonym of Paravascoceras (see also below under Vascoceras bullatum and V. globosum). The genus Broggiiceras Benavides-Caceres (1956: 469-470) was proposed for the Peruvian forms B. olssoni Benavides-Caceres (1956: 471, pl. 55, figs 1-4), the type species, and B. humboldti Benavides-Caceres (1956: 471, pl. 56, figs 3-6). These forms have smooth inner whorls and an adult ornament of strong ventral ribs matching that in P. cauvini. B. olssoni has whorls a little broader than high. While the opposite condition may prevail on the body-chamber of B. humboldti, the two are probably synonyms. In the absence of any significant recorded differences from Paravascoceras, Broggiiceras is best regarded as a synonym. Schébel (1975) and Meister et al. (1992), indeed, considered both B. olssoni and B. humboldti as synonyms of P. cauvini. P.M.P. ZABORSKI 64 ‘ “1x “J Wun ‘uonewio0y esipulg ‘ ‘Tx “P9T16'D ‘dAyereg “esIpulg ‘W Mun ‘uonewsoj esiputd ‘q ‘e] | “S14 7x 0966€6'D ‘adAiesed ‘q ‘eQ] “S14 “7x ‘e96SE6'D ‘adAjeIed ‘q ‘v6 “31 ‘aqeyY egeq ‘uONRULIO4 BSIPUl_ ‘01 “6 S814 ‘AOU ‘ds Ispoom spsaz0ISN4 | |-6 SBA OPSE6'D ‘esIpuld *H HUN ‘voNeUIO, BSIpUld “q “PY “BIL 1x “E1EEG'O “GP L'Blq ‘1x ‘8ISE6'D ‘q ‘RO “BLY exLYSY ‘O HUN ‘uoNeUIO; edIpuld “L “9 sdly [x “egcce6'D ‘BXeYSW q ‘eg “Bld 1x “8EEE6 O'9'Pp Bld 1x “LEEES'O E “BIA “1x “9ECEGD “47 “BLA “BAPYSY “Y WUN ‘UOMO eIpUld ‘p-Z S314 (NeapNyYD) MANDO spiad09svADIDg g-T SBLY UPPER CRETACEOUS AMMONITE Paravascoceras cauvini (Chudeau, 1909) 1909 1921 1933 1935 1935 1935 1935 1943 1943 1943 1943 1957 1957 11957 71965 1969 1969 1975 1981 1992 1992 Figs 2-8 Vascoceras cauvini Chudeau: 68, pls 1, 2; pl. 3, figs 1, 2. Thomasites cauvini (Chudeau) Chudeau: 463, fig. 1. Vascoceras cauvini Chudeau; Furon: 268, pl. 9, fig. 9. Vascoceras (Paracanthoceras) Chevalieri Furon: 59, pl. 4, figs la, b. Vascoceras ( Paravascoceras ) Cauvini Chudeau Furon: 60, pl. 5, figs la, b. Vascoceras ( Paravascoceras) Chudeaui Furon: 61, pl. 4, fig. 2. Vascoceras (Paravascoceras) Cauvini Chudeau nov. var. semiglabra Furon: 61, pl. 4, fig. 3. Paravascoceras cauvini (Chudeau); Schneegans: 128, pl. 4, fig. 2. Paravascoceras cauvini var. evoluta Schneegans: 130, pl. 8, fig. 2. Paravascoceras cauvini vat. inflata Schneegans: 131. Paravascoceras chevalieri Furon Schneegans: 132, pl. 4, fig. 7. Vascoceras bulbosum (Reyment) Barber: 19, pl. 6, figs 6, 8; pl. 27, figs 1-6. Vascoceras depressum Barber: 19, pl. 6, fig. 5; pl. 27, figs 7-9. Paravascoceras aff. cauvini (Chudeau); Barber: 37, pl. 14, figs 2, 3; pl. 32, figs 8, 9. Paravascoceras aff. cauvini (Chudeau); Collignon: 183. Paravascoceras cauvini (Chudeau); Freund & Raab: 20, pl. 3, figs 1-3; text-figs 5a, b. Paravascoceras tavense (Faraud) Freund & Raab: 23, pl. 2, fig. 9, text-figs 5e—g. Paravascoceras cauvini (Chudeau); Schobel: 119, pl. 4, fig. 3; pl. 5, figs 1-4. Paravascoceras cauvini (Chudeau); Collignon & Roman (in Amard, Collignon & Roman): 51, pl. 3, fig. o) Paravascoceras_ chevalieri (Furon); Collignon & Roman (in Amard, Collignon & Roman): 52, pl. 6, figs il, 2 Nigericeras barcoicense (Choffat) Collignon & Roman (in Amard, Collignon & Roman): 54, pl. 4, figs 16a, b. Vascoceras cauvini Chudeau; Luger & Gréschke: 374, pl. 40, figs 3, 6, 8, 9; pl. 41, figs 1-4; pl. 42, fig. 1; text-figs 6G, H, 8C. Nigericeras gadeni (Chudeau) /amberti Schneegans; Meister: 10, pl. 3, figs 13; text-fig. 6. Nigericeras jacqueti Schneegans; Meister: 11, pl. 2, figs 3, 4; pl. 4, fig. 1; text-fig. 7. Paravascoceras aff. nigeriense? (Woods) Meister: 16, pl. 5, fig. 3. Vascoceras cauvini Chudeau; Zaborski: figs 8, 12—15. Vascoceras bulbosum (Reyment); Zaborski: fig. 11. Vascoceras (Paravascoceras) cauvini (Chudeau); Meister, Alzouma, Lang & Mathey: 71, pl. 4, fig. 6; pl. 5, fig. 1, pl. 6, fig. 2. Vascoceras (Paravascoceras) cauvini forme lisse Meister, Alzouma, Lang & Mathey: 72, pl. 5, fig. 2; pl. 6, figs 1, 3. Vascoceras ( Paravascoceras) cauvini forme comprimée Meister, Alzouma, Lang & Mathey: 72, pl. 5, fig. 3; pl. 6, fig. 4. Vascoceras gr. cauvini Chudeau; Courville: pl. 4, figs 1-3. 65 MATERIAL AND OCCURRENCE. Thirty-six specimens, C.91304, Pindiga Formation, unit E, Ashaka; C.93556a, b, C.93557-9, C.93932, Pindiga Formation, unit F, Ashaka; C.93336-8, Pindiga Formation, unit K, Ashaka; C.91271-4, C.93304, C.93313, C.93517-8, Pindiga Formation, unit O, Ashaka; C.91278-84, C.93540-2, C.93933, Pindiga Formation, unit H, Pindiga; C.91285-9, C.93539, Pindiga Formation, unit J, Pindiga; C.91312, Pindiga Formation, unit N, Pindiga. The species has a known stratigraphical range from unit E (upper half) to unit O at Ashaka and from unit G to unit N at Pindiga. DIMENSIONS. See Fig. 12. REMARKS. In north-eastern Nigeria Paravascoceras cauvini includes forms showing whorls slightly to distinctly higher than broad, with rounded to slightly flattened venters and an umbilicus representing 16-29% of the total diameter. The species has a relatively long stratigraphical range here but successive assemblages show some variation. Material from unit F at Ashaka reaches a maximum diameter of some 100 mm. The adult whorls are smooth or with weak, irregular, crease-like ventral ribbing. The inner whorls, however, may show alternating long and short ribs (Fig. 5). The long ribs arise at umbilical tubercles. All ribs bear vague ventrolateral swellings but there are no siphonal tubercles. Umbilical tubercles or bulges may persist into the middle growth stages. This umbilical ornament is especially pronounced in certain specimens collected from the equivalent horizon (unit H) at Pindiga. Material from unit K at Ashaka is the oldest found to show the strong ventral adult ribbing which characterizes the species (Fig. 3; Meister 1989: pl. 3, fig. 1). Material from unit O can be regarded as fully typical of P cauvini. The inner whorls (Fig. 6) are completely smooth. This is usually the case with the middle growth stages also but rare individuals show bullate to clavate umbilical tubercles. Even less frequently there are broad, low, ventrolateral swellings but such features disappear by a diameter of 45 mm. Ventral ribbing is commonly displayed in the later growth stages (see Zaborski 1990a: fig. 14) but this ornament appears at a diameter varying from 60 mm to over 100 mm and is sometimes lacking altogether. Adults reach a maximum diameter of over 160 mm. In general whorl proportions P. cauvini is a very close match for Nigericeras gadeni (Chudeau). The middle and adult whorls of the two may be difficult to distinguish unless sutures are visible; N. gadeni has square saddles, a narrow L and a distinctly bifid E/L, P cauvini has more rounded and evenly frilled saddles. The material from Ashaka referred to Nigericeras by Meister (1989) in fact belongs in P. cauvini. The early whorls of N. gadeni are distinct, showing a typically acanthoceratine ornament with long and short ribs and seven rows of tubercles (see Schneegans 1943, Zaborski 1990a, Meister et al. 1992). Vestiges of a similar ornament, but without siphonal tubercles however, occur in early P. cauvini from unit F at Ashaka. The middle whorls of specimens from unit H at Pindiga sometimes show the strong bulge-like umbilical tubercles that are common at the same growth stage in N. gadeni. Meister et al. (1992: 70) and Courville (1992: 415) have also drawn attention to similarities between the juvenile ornament and in some cases suture pattern of P. cauvini and Nigericeras. Nigericeras gadeni characterizes the basal ammonite-bearing beds in north-eastern Nigeria, occurring in unit D at Ashaka and unit A at Pindiga. It therefore predates P cauvini. It is probable that P cauviniis derived from N. gadeni by a progressive 66 0.5 Fig.12 Shell proportions in Paravascoceras cauvini (Chudeau), Pseudovascoceras nigeriense (Woods), Vascoceras bullatum Schneegans, V. globosum costatum (Reyment) and V. globosum globosum (Reyment) from unit O at Ashaka. These individuals are the product of collecting carried out over | the course of two hours specifically for the purpose of comparing dimensions. This figure also gives a good indication of the relative abundances of the taxa concerned. loss of juvenile ornamentation (peramorphosis), simplification of suture pattern and development of adult ribbing. Forms from unit F at Ashaka and unit H at Pindiga are transitional in nature. Schneegans (1943: 119-125) proposed three additional species of Nigericeras, N. gignouxi, N.lamberti and N. jacqueti which show an increasingly weaker ornament but which all display the typically acanthoceratine suture pattern of the genus. It may be noted that in north-eastern Nigeria there is a variation in the strength of ornament of Nigericeras collected from place to place. Individuals from Teli and from north-east of the Biu Plateau (see Zaborski 1990a: figs 4, 5) have a weak juvenile ornament while those from the Hinna region and from between Kanawa and Wayari (see Zaborski 1990a: figs 6, 7) have stronger ornament. There is, however, no available evidence of this variation having a stratigraphical significance. In Niger, forms of N. jacqueti type occur alongside typical N. gadeni (Meister et al. 1992). In north-eastern Nigeria N. gadeni occurs in the equivalent of the Geslinianum Zone in north-west Europe and the Gracile Zone in the western interior of the United States (Zaborski 1990a). P. cauvini ranges through the probable equivalents of the Juddii Zone in north-west Europe and the Clydense to Scotti zones in the western interior (see below). Lewy et al. (1984) described P cauvini in association with Metoicoceras geslinianum (d’Orbigny) in Israel. Their material, however, Wb/Wh P.M.P. ZABORSKI ® P cauvini P nigeriense 9 V. bullatum © V.globosum costatum o 4 V.globosum globosum shows broad flank ribbing and umbilical bulges (Lewy et al. } 1984: fig. 41) in its middle whorls, features more typical of } Nigericeras from which it appears to be transitional. Its suture is } unknown. | Cooper (1979) and Kennedy & Wright (in press) proposed an origin for Nigericeras within Pseudocalycoceras Thomel, 1969. } 10-16; text-figs 83B, C, 84D-H), a lowest Upper Cenomanian } Guerangeri Zone species. Nigericeras may be a peramorphic derivative. The small Nigerian specimens from low in the Pindiga section!} referred to Vascoceras bulbosum (Reyment) and V. depressum sp. nov. by Barber (1957) are here regarded as P. cauvini as is the} material from Ashaka included in Nigericeras by Meister (1989)./} The P. cauvini from Nigeria are compressed forms which fall! into a distinct morphological group within the ammonite fauna|} from unit O at Ashaka (Fig. 12). Individuals from Niger.) however, develop a broader whorl section (see Meister et al.|} 1992: pl. 6, fig. 2). These specimens seem to be of the same age as! those from unit O at Ashaka. The latter are associated with very, large numbers of more inflated ammonites referrable to! Vascoceras globosum ___costatum, V. bullatum and Pseudovascoceras nigeriense. These three forms are of markedly\} UPPER CRETACEOUS AMMONITE less importance in Niger, if they are present at all. Their possible absence may have allowed populations of P. cauvini in Niger to develop a greater range of morphotypes due to lack of competition. Meister et al. (1992: 72-76) described a number of additional forms from Niger as Vascoceras (Paravascoceras) cauvini forme crassum (Furon) and V. (P.) cauvini forme de transition entre forme crassum et V. (P.) proprium (Reyment) . They are further discussed below under Vascoceras globosum. Also of interest in regard to their general whorl proportions are Paravascoceras rumeaui Collignon (1957: 122, pl. 16, fig. 2; _ Freund & Raab 1969: 21, pl. 3, figs 4, 5; text-figs 6c, d; Luger & | Groschke 1989: 380, pl. 41, figs 5, 6; pl. 42, figs 3, 4; text-fig. 8D) from Algeria, Egypt and Israel and Vascoceras costellatum Collignon & Roman (in Amard et al. 1981: 51, pl. 2, figs 6a, b) from Algeria. These species have adult ventral ribbing like that in P. cauvini but are more inflated. They may, like the Niger forms, be regional variants of P cauvini. Luger & Groéschke (1989: 375-376) discussed the question of whorl breadth in P cauvini but, unlike Schébel (1975), regarded P. rumeaui as distinct from P. cauyini. They further separated individuals with depressed whorls but which were otherwise similar to P cauvini as Vascoceras cf. cauvini (Luger & Gréschke 1989: 376, pl. 42, fig. 2: pl. 43, fig. 3; text-figs 6F, 8B). The P. cauvini of Collignon & Roman (in Amard et al. 1981: 51, pl. 3, fig. 9) have whorls only a little broader than high and probably belong here. Their Paravascoceras chevalieri (Furon) (Collignon & Roman in Amard et al. 1981: 52, pl. 6, figs 1, 2) and Nigericeras barcoicense (Choffat) (Collignon & Roman in Amard etal. 1981: 54, pl. 4, figs 16a, b) are similar to and may be conspecific with P. cauvini. The Paravascoceras aff. chevalieri of Reyment (1955: 63, pl. 14, figs 1a, b), however, shows three rows of tubercles upon the ventral ribs and more closely resembles early Thomasites gongilensis from unit O at Ashaka (see Figs 41-44). The Vascoceras (Paravascoceras) cf. cauvini from Angola described by Cooper (1978: 130, figs 6C-H, 35-37) is a Nigericeras. Berthou et al. (1985: 72) speculated that Vascoceras barcoicense Choffat (1898: 67, pl. 17, fig. 1; pl. 22, fig. 5; Berthou et al. 1985: 70, pl. 4, figs 1-3) might turn out to be a senior synonym of P. cauvini. The strong adult ribbing of the latter species is, however, unknown in V. barcoicense. Whorl proportions in the two are similar but nothing is known of the early growth stages in V. barcoicense. The species may belong in Paravascoceras or alternatively it may be an involute, weakly ornamented variant of Vascoceras gamai Choffat, according to Berthou et al. (1985: 71). V. barcoicense exile Cobban, Hook & Kennedy (1989: 47, figs 47, 87Q-S, 89M-GG) from New Mexico resembles P. cauvini in whorl proportions but is more involute. Specimens from low in the Pindiga section may be similar in this respect (Fig. 8) but V barcoicense exile has a different juvenile ornament of rather strong ventral ribs. The V. (V.) cauvini of Kennedy et al. (1989: 82, figs 9G, 20C-G) from Texas are similar to and probably conspecific with V. barcoicense exile. | Further involute compressed forms are the Nigericeras ‘|Jacqueti involutum Meister, Alzouma, Lang & Mathey (1992: 68, pl. 4, figs 3-5; text-fig. 14) from Niger. Again, these show similarities with P cauvini from unit H at Pindiga but are consistently more involute. Their suture pattern (Meister et al. 1992: fig. 14) is incompletely known but seems to be intermediate _|between that of Nigericeras and Paravascoceras. Meister et al. '|(1992) regarded N. jacqueti involutum as an offshoot of N. gadeni derived through N. jacqueti jacqueti. It may be the product of a 67 local lineage independent of that giving rise to P. cauvini. Genus PSEUDOVASCOCERAS gen. nov. TYPE SPECIES. Vascoceras nigeriense Woods, 1911. DIAGNOsIS. Moderately evolute to moderately involute, moderately compressed to moderately depressed ammonites. Whorls rounded to subpentagonal. Ornament of umbilical, inner and outer ventrolateral and siphonal tubercles which may be borne upon transverse to concave ribs of varying strength. Additional ventral ribs frequently present. Ornamental elements of highly variable persistence during ontogeny, sometimes extending onto the body-chamber, in other cases confined to the earliest growth stages. Suture line simple with evenly frilled elements; saddles often elongate and rectangular in outline, lateral lobe fairly broad. REMARKS. Of all the ammonites from north-eastern Nigeria showing ‘vascoceratid’ suture patterns it is the multituberculated forms which have proved most problemmatical and which have received the most varied taxonomic treatment. This is not surprising given the huge range of morphotypes that are represented within assemblages from the same stratigraphical horizon, at Ashaka unit O. In fact three multituberculated genera are present therein, end members of which are not always easy to differentiate. Forms attributable to Rubroceras Cobban, Hook & Kennedy occur as rarities (Zaborski 1993); a larger number of individuals belong in Fikaites Zaborski (1993); but the greatest number are here referred to Pseudovascoceras nigeriense (Woods). In his treatment of this last group Barber (1957) assigned them to three genera, Vascoceras, Nigericeras and Paramammites Furon, and no less than seven species. The last two generic determinations can easily be disposed of. The type species of Paramammites (by the subsequent designation of Reyment 1954b: 225), Vascoceras polymorphum Pervinquiére (1907: 336, pl. 21, figs 2, 6; text-fig. 126) (see also Renz 1982: 84-85; Chancellor e¢ a/, in press) has a juvenile ornament of varying strength, often with large spinose tubercles, but always lacks siphonal tubercles. The present material has nothing to do with Paramammites. Forms with strong adult costae, interrupted ventrally, have often been referred to this genus without knowledge of their ontogenetic development (see also Cobban et al. 1989: 51; Zaborski 1990b: 574-575) thus creating a rather confused situation. Nigericeras resembles the present material in only one real respect, the presence of seven rows of tubercles. In detail its ornament is more regular and in all genuine members of the genus it is confined to the early whorls (see Schneegans 1943). The suture in Nigericeras, although simple, is of a distinctly acanthoceratine pattern, unlike that in the present material and other forms mentioned below also previously referred to Nigericeras. Nor can the present material be referred to Vascoceras. Its ornament is unlike than in any known species of the genus and quite distinct from that in the type species V. gama. Cobban er al. (1989: 51) pointed out that Barber’s (1957) Paramammites needed a new generic name. They suggested that these forms were in part ribbed and tuberculated derivatives of Vascoceras. The genus Pseudovascoceras is here proposed to include this material, the name alluding to the homeomorphy between smooth members of the type species and true Vascoceras. The origin of the genus, however, is thought to lie in 68 an earlier acanthoceratine genus, probably Cunningtoniceras Collignon, 1937 as detailed below. Nigericeras scotti Cobban (1971: 18, pl. 9, figs 1-4; pl. 18, figs 1-9; text-figs 15-19) from the terminal Cenomanian of the United States western interior may be a Pseudovascoceras. It lacks the suture pattern typical of Nigericeras but resembles the more strongly ornamented examples of P. nigeriense. The unnamed specimen from Turkestan figured by Kler (1909: pl. 8, figs 3a, b; text-fig. 6) may also be a Pseudovascoceras. The English specimen (C.82287) from the high Cenomanian referred to Nigericeras cf. gignouxi Schneegans by Wright & Kennedy (1981: 85, pl. 15, figs 6a, b) is a fragment, the ornament and suture pattern of which cannot be made out clearly. It might be best referred to Pseudovascoceras. It occurs alongside Thomasites gongilensis. In Nigeria Thomasites occurs well above the stratigraphical level of Nigericeras, but its earliest members are coeval with P. nigeriense. Pseudovascoceras nigeriense (Woods, 1911) Figs 14-24, 36, 37 21909 ~~ Vascoceras cauvini Chudeau: pl. 3, figs 4a, b (only). 1911 Vascoceras nigeriense Woods: 281, pl. 21, fig. 6; pl. 22, figs 2, 3. 21943 + Vascoceras nigeriense Woods; Schneegans: 133, pl. 4, fig. 1. 21943 Paravascoceras cf. barcoicense (Choffat) Schneegans: 134, pl. 8, fig. 1. 1954b Vascoceras nigeriense Woods; Reyment: 256. 21955 Nigericeras ogojaense Reyment: 62, pl. 13, fig. 6; pl. 14, fig. 3; text-fig. 28. 1957 —_ Vascoceras nigeriense Woods; Barber: 15, pl. 4, fig. 2; pl. 26, figs 1, 2. 1957 = Nigericeras costatum Barber: 29, pl. 10, figs 3, 4; pl. 11, fig. 3; pl. 30, figs 1-7. 1957 = Nigericeras glabrum Barber: 29, pl. 10, figs 1, 2; pl. 30, fig. 8. 1957 Nigericeras? intermedium Barber: 31, pl. 11, figs 1, 2; pl. 30, figs 9, 10. 1957 =Paramammites tuberculatus Barber: 31, pl. 12, fig. 1; pl. 13, fig. 2; pl. 31, figs 1-3, 9. 1957 Paramammites raricostatus Barber: 33, pl. 12, fig. 3; pl. 31, figs 4, 6, 7. 1957 Paramammites inflatus Barber: 33, pl. 12, fig. 2; pl. 13, fig. 1; pl. 31, figs 5, 8. Paramammites laffitei Collignon: 186, pl. A, fig. 2. Paramammites subtuberculatus Collignon: 187, pl. A, fig. 3. 1965 + Vascoceras nigeriense Woods; Reyment: pl. 2, fig. 2. 1965 Nigericeras costatum Barber; Reyment: pl. 3, fig. 13. 21965 21965 21965 — Nigericeras ogojaense Reyment; Reyment: pl. 3, fig. 14. 1965 Paramammites tuberculatus Barber; Reyment: pl. 3, figs 15a, b. 1980 Nigericeras costatum Barber; Wright & Kennedy: figs 10a, b. 1989 Paravascoceras nigeriense? (Woods); Meister: 14, pl. 5, fig. 1; pl. 6, fig. 1; text-fig. 11. 1989 - Vascoceras costatum (Barber) Meister: 23, pl. 10, figs 3, 5; pl. 11, figs 1, 2, 5; text-figs 16a—d. 1989 Vascoceras costatum glabrum (Barber) Meister: 23, pl. 9, figs 2, 4; pl. 10, fig. 4; text-figs 16e—g. 1989 _Vascoceras ellipticum Barber; Meister: 28, pl. 12, figs 1, 3; text-fig. 18. 1989 Paramammites subconciliatus (Choffat) Meister: 30, pl. P.M.P. ZABORSKI 12, figs 4, 5; pl. 13, figs 1-4; pl. 14, figs 1, 2; pl. 15, figs 1, 4; text-fig. 21. 1989 Paramammites polymorphus (Pervinquiére); Meister: 36, pl. 14, figs 3, 4; text-fig. 24. 1990a_ Vascoceras nigeriense Woods; Zaborski: fig. 25. 1992 Vascoceras sp. gr. costatum (Barber) sensu Meister, | 1989; Courville: pl. 5, fig. 3; pl. 6, figs 2, 3. LECTOTYPE. Specimen B3237, Sedgwick Museum, Cambridge | (see Woods 1911: pl. 22, figs 2, 3); from Kunini, north-eastern Nigeria (selected by Berthou, Chancellor & Lauverjat 1985: 69). | PRESENT MATERIAL AND OCCURRENCE. Sixty-one specimens, | C.93305-8, C.93311, C€.93315-21, C.93370-93, C.93494a-d, C.93495a-f, C.93496a—d, C.93497-507, Pindiga Formation, unit O, Ashaka. | DIMENSIONS. See Fig. 12. REMARKS. P. nigeriense is generally a moderately evolute | species having rather compressed to moderately depressed whorls with a rounded to subpentagonal outline. In overall shell proportions it overlaps with both Vascoceras bullatum and V. | globosum costatum; smooth individuals are often especially | difficult to distinguish from the last form. The adult diameter varies from about 85 to 120 mm when the body-chamber makes up two-thirds of the final whorl. | It is in its ornamentation that P nigeriense shows its greatest variation, from almost entirely smooth to highly decorated end members. A variation series is shown in Figs 14—24, and there is | also abundant figured material in the previous literature (see synonymy list). Dissection of numerous individuals, including those with smooth outer whorls, shows that siphonal tubercles are consistently developed but they may have already disappeared by a diameter of 10 mm. Outer ventrolateral | tubercles are also commonly developed while inner ventrolateral _ and umbilical tubercles may or may not be present. One | combination or another of tubercle rows may persist throughout the length of the septate whorls or disappear at any stage in | ontogeny. Umbilical tubercles, when present, are the most’ persistent ornamental features and siphonal tubercles are the | least, with the result that numerous individuals show six rows of | tubercles in their middle growth stages. Strongly tuberculated | forms may in addition display rectiradiate to concave ribs | connecting the tubercles. The ribs may branch across the venter| while additional ribs with inner and/or outer ventrolateral and! siphonal tubercles may be intercalated. Ornamental strength is! initiated very early in ontogeny. The ornament of the’ phragmocone may persist onto the adult body-chamber or this) part of the shell may be smooth. Most frequently, however, there are irregularly developed ribs upon the flanks and the venter! which vary from strong, broad fold-like structures to fine, dense crease-like features recalling those in adult Vascoceras woodsi and V. bullatum. Suture patterns are of a simplified type but the saddles tend to) be elongated, especially in strongly ornamented forms. The lateral lobe is fairly wide and often subdivided by a distinct median element. Meister (1989) separated members of P nigeriense from Ashaka into six taxa, Paravascoceras nigeriense (Woods). Vascoceras. ellipticum Barber, Paramammites aff. gr polymorphus (Pervinquiére), P subconciliatus (Choffat). Vascoceras costatum (Barber) and V. costatum glabrum (Barber) Nevertheless, he showed how the ornamental variation betweer! the last three could easily be interpreted in terms ol| || Fig.13 Vascoceras bullatum Schneegans. Pindiga Formation, unit O, Ashaka. C.93513, x1. _\ Figs 14-19 Pseudovascuceras nigeriense (Woods). Pindiga Formation, unit O, Ashaka. Fig. 14a, b, C.93382, x1. Fig. 15a, b, C.93383, x1. Fig. 16a, I b, C.93379, x1. Fig. 17a, b, C.93380, x1. Fig. 18, C.93321, x1. Fig. 19a, b, C.93307, x0.75. Figs 20-24 Pseudovascoceras nigeriense (Woods). Pindiga Formation, unit O, Ashaka. Fig. 20a, b, C.93375, x0.75. Fig. 21, b, C.93374, x1. Fig. 22, C.93305, x0.75. Fig. 23a, b, C.93371, x0.75. Fig. 24a, b, C.93306, x0.75. UPPER CRETACEOUS AMMONITE heterochronic ontogenies (Meister 1989: 34, 36, text-fig. 23). Concerned, however, that the faunas from unit O at Ashaka were condensed and might contain chronologically successive taxa, he refrained from placing them in synonymy. Unit O at Ashaka is the product of a ‘slow rate of sediment accumulation associated with a marine flooding phase. It contains the most diverse marine fauna found at Ashaka including numerous bivalves, gastropods and echinoids as well as a large number of ammonite species (see also Courville 1992: fig. 2). Encrustations of Plicatula occur throughout while many of the ammonites show oyster and serpulid overgrowths. There is, however, no significant phosphatization or reworking, and glauconite is rare or absent. There is no reason to believe that this unit represents any greater condensation than several other limestones at Ashaka and elsewhere in north-eastern Nigeria. Phosphatic matter, glauconite and reworked ammonites are common components in the limestone beds of the region, particularly in the upper parts of those interbedded with shales. It is the upper surface of unit O at Ashaka that marks the most significant break in sedimentation, but P nigeriense is found throughout the unit below this level. Both Meister (1989) and Courville (1992) believed they could differentiate between faunas from different levels in unit O (their Niveau 21 and Niveau 22 ), though their reported successions differ. No significant stratigraphical variation in the nature of the ammonite faunas from this unit | has been detected in the present work, apart from the restriction of strongly ornamented Thomasites to its upper surface. In the cases of the other ammonites present intraspecific variation 1s by far the most important factor. No morphometric or ornamental evidence has been obtained which allows objective taxonomic subdivision of P. nigeriense. There is a complete intergradation from smooth to strongly ornamented individuals while the latter vary considerably among themselves. In view of these factors all _ these morphotypes are regarded as conspecific, despite the great differences between end members. Courville (1992: 419-420) came to a similar conclusion and favoured the name Vascoceras costatum (Barber) which was used by Meister (1989) for individuals of intermediate ornamental strength. Priority, however, belongs to Vascoceras nigeriense Woods, 1911. The | lectotype is a smooth end member of the species, as are the individuals referred by Meister (1989) to Paravascoceras | nigeriense? (Woods) and Vascoceras ellipticum Barber. Smooth examples of Pseudovascoceras nigeriense have in the past been compared with Vascoceras gamai (Barber 1957: 15; Hancock & Kennedy 1981: 357). Their similarity concerns only the outer whorls, however, and is homeomorphic in nature. Ornamented examples of P nigeriense share similarities with | various genera. Meister (1989) referred forms in which the siphonal tubercles disappear early in ontogeny to Paramammites, which he regarded as a senior synonym of Spathites (Jeanrogericeras) Wiedmann, 1960 (type species Ammonites reveliereanus Courtiller, 1860). As mentioned above, |\this material cannot be referred to Paramammites or Jeanrogericeras as neither shows siphonal tubercles at any growth stage (see Choffat 1898: 64; Pervinquiére 1907: 336; Wiedmann 1960: 741; Renz 1982: 84; Berthou et al. 1985: 62) and resemblances are superficial only. Some of the present specimens have the appearance of giant Protacanthoceras proteus (compare Fig. 23 and Wright & Kennedy 1980: fig. 5). Others with dense, multiple ventral ribbing resemble Kamerunoceras Reyment, 1954b (type species Acanthoceras eschii Solger, 1904) or Euomphaloceras Spath, 1923 (type species | Ammonites euomphalus Sharpe, 1855), though they lack the typically euomphaloceratine constrictions upon their early 71 whorls. In its style of ribbing and the generally coarse nature of the tuberculation, the present material most closely resembles Cunningtoniceras Collignon, 1937 (type species Ammonites cunningtoni Sharpe, 1855). This is mainly a Middle Cenomanian genus (see, for example, Kennedy 1971, Zaborski 1985, Kennedy & Cobban 1990a) but it ranges into the Upper Cenomanian (Wright & Kennedy 1987, Cobban et al. 1989, Kennedy & Cobban 1990h). Pseudovascoceras may be a descendant of Cunningtoniceras. Yntroduction into north-eastern Nigeria produced peramorphic individuals losing their ornament early in ontogeny and coming to resemble Vascoceras. Interestingly there is a morphological overlap between P nigeriense and Nigericeras ogojaense Reyment (1955: 62, pl. 13, fig. 6; pl. 14, fig. 3; text-fig. 28); the two are probably conspecific. The latter comes from the southern, oceanward, end of the Benue Trough where smooth individuals are unknown; the holotype (C.47401) and newly collected material (C.93578-61) all show prominent ornament. Reyment (1979, 1988) has remarked upon the extraordinary polymorphism that may be displayed by vascoceratid species. He believed that in the changeable environment of the Cenomanian-Turonian intracontinental sea in west and Saharan Africa selection would have favoured forms with genetic or phenotypic flexibility. Such taxa would have been capable of responding to environmental fluctuations, each morphotype being best suited to a particular kind of environment. Meister et a/. (1992) took up this issue in respect of the Niger ammonites. They noted that particular stratigraphical horizons there commonly yield monospecific faunas or assemblages dominated by one species. They speculated that taxa able to occupy niches in the exacting environments prevailing during the Late Cenomanian and Early Turonian faced virtually no competition, the result being a high degree of polymorphism. In unit O at Ashaka a number of ammonite taxa co-exist, largely as a result of introduction of species during a marine flooding episode. While there is some overlap, however, each of the four main taxa described here, Paravascoceras cauvini, Vascoceras bullatum, V. globosum costatum and Pseudovascoceras nigeriense, occupies a particular part of the morphological spectrum (Fig. 12). Variation in gross shell proportions to the extent of that suggested by Meister er al. (1992) for P. cauvini in Niger is not seen in these taxa. On the other hand, within P nigeriense there seems to have been virtually no selection pressure favouring any particular strength of ornamentation. In this respect the polymorphic potential of the species was capable of wide expression. Much the same can be said of Fikaites varicostatus Zaborski, the other strongly ornamented form found in some numbers in unit O at Ashaka. This species shows a significant variation in the strength of its ribbing and tuberculation (see Zaborski 1993). Family VASCOCERATIDAE Douville, 1912 Subfamily VASCOCERATINAE Douvillé, 1912 Genus VASCOCERAS Choffat, 1898 (= Discevascoceras Collignon, 1957; Greenhornoceras Cobban & Scott, 1972; Provascoceras Cooper, 1979) TYPE SPECIES. Vascoceras gamai Choffat, subsequent designation of Roman, 1938. 1898: by the REMARKS. Choffat (1898: 51-53) had a broad concept of Vascoceras as encompassing forms basically united by the 72 possession of a simple suture pattern. Some of these original members are now referred to Spathites Kummel & Decker, 1954. The type species, V. gamai, was regarded as part of a group characterized by a wide umbilicus and the possession of a single (umbilical) row of tubercles. Recent discussions of Vascoceras have been given by Wright & Kennedy (1981) and Berthou et al. (1985). It is commonly suggested that Paravascoceras, Pachyvascoceras, Paracanthoceras, Broggiiceras, Discovascoceras, Provascoceras and, sometimes, Greenhornoceras should be regarded as strict synonyms of Vascoceras without even subgeneric distinction (see Berthou et al. 1985, Kennedy, Wright & Hancock 1987, Luger & Gréschke 1989, Kennedy, Cobban, Hancock & Hook 1989, Cobban et al. 1989). The oldest known species included in Vascoceras is the micromorph Ammonites diartianus d’Orbigny, the type material of which was redescribed by Kennedy & Juignet (1977). Further examples were subsequently described by Wright & Kennedy (1981: 86, pl. 17, fig. 1; text-figs 29A—F), Forster et al. (1983: 133, pl. 3, figs 1-5) and Cobban et al. (1989: 47, figs 48, 88TT-AA). V. diartianum occurs most frequently in the Geslinianum Zone or equivalents but Kennedy et al. (1989: 80) reported examples in New Mexico from equivalents of the underlying Guerangeri Zone. Kennedy & Wright (1985) and Wright & Kennedy (1987) drew attention to the morphological similarity between V. diartianum and Protacanthoceras of the P. proteus Wright & Kennedy group (see Wright & Kennedy 1980: 95, figs 49-S1, 57-58; 1987: 216, pl. 55, figs 4, 9, 17, 18, 21-23; text-figs 82B, 83G, M, 84P). They believed the former to have been derived from the latter. Accordingly, V. diartianum would constitute the root stock of Vascoceras and the above-mentioned suggested synonyms. Cooper (1979) was reluctant to admit the Late Cenomanian age of Vascoceras and pointed to a number of differences between Ammonites diartianus and V. gamai, the most significant of which regarded their ornamentation. He proposed the genus Provascoceras for A. diartianus but regarded the species as ancestral to both Vascoceras and Paravascoceras. A. diartianus 1s clearly transitional between Acanthoceratinae and Vascoceratidae, retaining the bifid first lateral saddle of the former. Its ornament consists of rounded to twisted to distinctly bullate umbilical tubercles which may envelop practically the whole of the flanks and fine bundled ribbing extending across the venter. Relatively little is known of the inner whorls of topotype material of V. gamai. Choffat (1898: pl. 7, figs 3, 4; pl. 8, fig. 4; pl. 10, fig. 2) figured a number of juveniles which show approximately 8 umbilical bullae and 20 coarse, regularly developed major and minor ribs which cross the venter (see also Berthou et al. 1985: 67). The umbilical tubercles may be persistent but the ornament generally disappears on the later whorls. A similar juvenile ornament has been described in material referred to V. gamai from Egypt (Luger & Gréschke 1989: 378, pl. 40, figs 5, 7) and V. cf. gamai from New Mexico (Cobban er al. 1989: 45, figs 87W-AA, EE-RR). There are, however, other juveniles from Portugal with rather different ornamentation. V. silvanense Choffat (1898: 57, pl. 8, fig. 5; pl. 21, fig. 9) shows massive umbilical bullae but no definite ribbing. Berthou et al. (1985: 68) regarded V. silvanense as a nomen dubium and almost certainly the inner whorls of one or another P.M.P. ZABORSKI of the Portugese species of Vascoceras. Another individual (Berthou ef al. 1985: 68, pl. 3, figs 4, 8, 9) displays about 10 | umbilical bullae intermediate in strength between those found in V. gamai and V. silvanense and about twice as many low ventral ribs mostly arising in pairs from these bullae. Berthou et al. (1985: 68) compared this specimen with V. adonense Choffat which they regarded as a synonym of V. gamai. Its ornament is reminiscent of that in Ammonites diartianus. Numerous juvenile whorls of V. woodsi are available from north-eastern Nigeria (see below). They show a considerable variation in ornament. | Although no comparable variation series is available for the Portugese Vascoceras, it is possible that certain V. gamai could © show an early ornament approaching that in Ammonites diartianus. Despite its transitional nature, A. diartianus is here regarded as belonging in Vascoceras and Provascoceras is therefore a synonym. Discovascoceras Collignon (type species Vascoceras ( Discovascoceras ) tesselitense Collignon 1957: 125, pl. 1, figs la, b; by original designation) was originally proposed by Collignon (1957: 123) as a subgenus of Vascoceras but was later raised to the status of a separate genus following an amended diagnosis © (Collignon 1965: 179). Its essential characters were given as its triangular whorls, presence of three carinae on the middle whorls, variation in spacing and degree of indentation of the sutures, depth of the umbilicus, egression of the adult whorl and tendency for apertural constriction. Berthou er al. (1985: 75) regarded the holotype of D. tesselitense as an indeterminate Vascoceras, the species as invalid and Discovascoceras Collignon, 1957 as a synonym of Vascoceras. In view of its ventral carinae they compared the material later described as D. tesselitense by Collignon (1965: 181, pl. G, figs la, b) with Pseudotissotia Peron (see also Hirano 1983: 69-70). They proposed that Collignon’s second account be taken as the first valid one of Pseudotissotia? tesselitense. Collignon’s (1965) material, however, shows similarities with Nigerian forms intermediate between Vascoceras globosum costatum and Thomasites which are described below. Here both the Collignon 1957 and 1965 descriptions are regarded as dealing with Vascoceras but the 1965 material could alternatively be assigned to Thomasites. Greenhornoceras Cobban & Scott (type species Vascoceras ( Greenhornoceras ) birchbyi Cobban & Scott 1972: 85, pl. 22; pl. 23, figs 1-13; pl. 24, figs 1-12; pl. 25; pl. 26, figs 5—8, 11, 12; pl. 27, figs 1-6; text-figs 43-47; by original designation) is amongst the stratigraphically youngest examples of Vascoceras. Cobban & Scott (1972: 84-85) distinguished the subgenus Greenhornoceras only on the basis of being more involute than V. ( Vascoceras) and in maintaining a square to rectangular whorl section. Its juvenile ornament of strong, regularly developed long and short ribs gives way to smooth later whorls. There is no compelling reason to regard Greenhornoceras as anything other than a strict synonym of Vascoceras. As mentioned above, Paravascoceras (= Paracanthoceras, Pachyvascoceras, Broggiiceras) is here regarded as a distinct genus with an origin separate from that of Vascoceras and is most properly included in the Acanthoceratinae. Figs 25-30 Vascoceras woodsi sp. nov. Figs 25-27, Pindiga Formation, unit M, Ashaka. Fig. 25a, b, paratype, C.93341, x1. Fig. 26a, b, paratype, C.93339, x1. Fig. 27a, b, holotype, C.93342, x1. Fig. 28a, b, Pindiga Formation, unit M, Pindiga. Paratype, C.91263, x1. Fig. 29a, b, Pindiga For- mation, unit N, Pindiga. Paratype, C.93351, x1. Fig. 30, Pindiga Formation, Deba Habe. Paratype, C.91257, x1. UPPER CRETACEOUS AMMONITE 73 74 Vascoceras woodsi sp. nov. Bigs 911, 25—32 1957. Vascoceras sp. juv. Barber: 27, pl. 6, figs 2, 4, 7; pl. 27, figs 10-15. 21965 Vascoceras gamai Choffat; Collignon: 185, figs 5—7. 1989 Plesiovascoceras aff. gr. thomi (Reeside) ou sp. nov. Meister: 11, pl. 4, figs 2, 3, 5; text-fig. 8. 1989 Paravascoceras gr. evolutum Schneegans; Meister: 14 (pars), pl. 5, fig. 4 (only); text-fig. 10. 1990a_ Vascoceras gr. evolutum (Schneegans); Zaborski: 7. 1990a_ Vascoceras sp. Zaborski: figs 9, 10. 1990a_ Vascoceras sp. juv. Zaborski: figs 16-18, 20, 21. 1992 Vascoceras gr. thomi (Reeside) ou evolutum (Schneegans); Courville: pl. 5, fig. 1. 1993 Vascoceras sp. nov. aff. gamai Choffat; Zaborski: 365. 1995 Vascoceras sp. nov. aff. gamai Choffat; Zaborski: 54, 5): HOLOTYPE. C.93342 (Fig. 27), Pindiga Formation, unit M, Ashaka. PARATYPES. Thirty-four specimens, C.93339, ©C.93341, C.93343-4, C.93543, Pindiga Formation, unit M, Ashaka; C.91262-70, Pindiga Formation, unit M, Pindiga; C.91224-5, C.91311, C€.91313-4, C.93351, Pindiga Formation, unit N, Pindiga; C.91256-61, C.93355, C.93596a-f, C.93597, Pindiga Formation, Deba Habe. DIMENSIONS. D Wb Wh U €.93597 94 - 31 (33) 39 (41.5) C.93351 60 34 (57) 22 (37) DKS) C.91264 53 32 (60) 20 (38) 18 (34) €.93355 Sy) 31 (60) 18 (35) 20 (38.5) C.91256 50 29 (58) 20 (40) 17 (34) C.91263 40 25 (62) 15 (37.5) NSIS 2E5)) C.91257 39 24 (61.5) 16 (41) 11 (28) C.91262 34 20 (59) 13 (38) 10 (29) DERIVATION OF NAME. After the late H. Woods who first described ammonites from north-eastern Nigeria. DIAGNOSIS. Evolute Vascoceras with whorls broader than high. Middle whorls with rounded or more normally highly bullate umbilical tubercles fusing with inner ventro-lateral bullae to cover the flanks. Adult body-chamber smooth or with umbilical tubercles and/or relatively weak ventral ribbing. DESCRIPTION. The shell is evolute, the umbilicus widening during growth from about one-third to 40% or more of the overall diameter. The maximum diameter attained is about 120 mm, when the body-chamber makes up two-thirds of the final whorl. In all but the very earliest growth stages the whorls are distinctly broader than high. Two nuclei are available. In C.93596f the whorls are initially smooth and tubular with a broadly rounded venter. At a diameter of 3 mm broad bullate swellings enveloping the inner half of the flanks appear and give rise to low ribs which cross the Fig. 32 €.93514, x1. Figs 36-37 Pseudovascoceras nigeriense (Woods). Pindiga Formation, unit O, Ashaka. Fig. 36a, b, C.93499, x1. Fig. 37a, b, C.93494d, x1. Vascoceras woodsi sp. nov. Pindiga Formation, Deba Habe. Paratype, C.93597, x1. Figs 33-35 Vascoceras bullatum Schneegans. Pindiga Formation, unit O, Ashaka. Fig. 33a, b, C.93512, x1. Fig. 34a, b, C.93516a, x1. Fig. 35a, b, P.M.P. ZABORSKI l 1 cm j Fig. 31 Suture in Vascoceras woodsi sp. nov. Holotype, C.93342. Pindiga Formation, unit M, Ashaka. venter. The suture shows a direct transition from an entire to an ~ evenly frilled E/L; this saddle is never bifid. In C.93596e about 10 umbilical bullae have developed by a diameter of 6 mm but the venter lacks ribbing. In the succeeding growth stages (Figs 9, 10) the characteristic ornament consists of 8-10 lateral bullae in each whorl, upon which discrete umbilical and inner ventrolateral swellings can sometimes be made out. Most of the bullae give rise to a narrow rounded rib which crosses the venter and bears outer ventrolateral tubercles. No definite siphonal tubercles can be | made out. There may be a single ventral rib bearing only outer ventrolateral tubercles alternating with each major rib. In other | specimens no well-developed ribbing exists. In all cases sharply defined ribbing disappears at diameters of 10-15 mm though the | lateral bullae persist. At these diameters the venter becomes | flattened and the whorls increasingly depressed. At diameters of 20-60 mm the main ornament consists of 6-8 | umbilical tubercles in each whorl which are of variable shape | and strength. They are commonly highly bullate but may be | rounded, clavate or paired in nature. At first the more bullate | types may partially fuse with bullate inner ventrolateral swellings | but the latter features quickly fade during growth. Broad, vague fold-like ribs which cross the venter may issue from the umbilical tubercles or the venter may be smooth. Such ribs, however, rarely persist beyond diameters of 40 mm. Umbilical tubercles may persist onto the adult body-chamber which is frequently compressed. Ventral ribbing may develop | here taking the form of irregular closely-spaced plicae at one | extreme and moderately strong fairly evenly-spaced ribs at the » other. The suture (Fig. 31) is of the typically simple type found in Vascoceras. REMARKS. Ina previous account (Zaborski 1990a: 5) doubt was | expressed about whether a number of juvenile Vascoceras | collected from Deba Habe and Pindiga were conspecific. V. woodsi has been found at these two localities and at Ashaka. At | Pindiga adult specimens are not found, only the middle septate | whorls being found in units L, M and N. At Ashaka the species is abundant in units K and, especially, M but the material consists almost entirely of poorly preserved adult body-chambers. At Deba Habe, however, specimens representing all growth stages | occur ina 10 cm limestone less than 1 m below the level at which } Vascoceras globosum costatum and Pseudovascoceras nigeriense appear. Occasional juveniles from Ashaka fit comfortably within the morphological range exhibited by the Pindiga and Deba Habe material. There is little doubt that all these specimens are conspecific. | 75 UPPER CRETACEOUS AMMONITE 76 Adult V. woodsi are closely similar to V. gamai Choffat (1898: 54, pl. 7, figs 1-4; pl. 8, fig. 1; pl. 10, fig. 2; pl. 21, figs 1-4; see also Berthou et al. 1985 for a revision of the species). The strong regular ribbing of the early whorls in V. gamai figured by Choffat (1898: pl. 7, figs 3, 4; pl. 8, fig. 4; pl. 10, fig. 2), however, differs from the ornament at the same stages in V. woodsi. It should be noted that certain juvenile specimens from Portugal show similarities with some Nigerian individuals: compare C.91264 (Fig. 11) with V. silvanense Choffat (1898: pl. 8, fig. 5), and C.93351 (Fig. 29) with Berthou et a/. (1985: pl. 3, figs 4, 8, 9). A more complete knowledge of the early whorls in V. gamai is necessary for full comparison with V. woodsi. Closer to the Nigerian juveniles is V. diartianum, particularly material from Germany which reaches diameters of over 30 mm (see Forster et al. 1983). This collection includes individuals with rather rounded umbilical tubercles (Forster et al. 1983: pl. 3, fig. 1; compare with Zaborski 1990a: fig. 20) and others with highly bullate umbilical tubercles similar to those common in V. woodsi (compare Forster et al. 1983: pl. 3, figs 2-5 and C.91263, C.91257, Figs 28, 30 herein). V. woodsi is a little younger than V. diartianum and derivation from the latter can easily be imagined by peramorphosis and further simplification of suture pattern. The material from Ashaka described by Meister (1989: 11, pl. 4, figs 2, 3, 5; text-fig. 8) as Plesiovascoceras aff. gr. thomi (Reeside) belong in V. woodsi; Vascoceras thomi Reeside (1923) is a synonym of Fagesia catinus (Mantell) (see also Wright & Kennedy 1981: 88, 97). Those he referred to Paravascoceras gr. evolutum Scheegans are partly V. woodsi (Meister 1989: pl. 5, fig. 4) and probably partly Pseudaspidoceras pseudonodosoides (Choffat) (Meister 1989: pl. 5, fig. 2). Paravascoceras cauvini vat. evoluta Schneegans (1943: 130, pl. 8, fig. 2) is here considered to be a strict synonym of Paravascoceras cauvini. Specimens of Vascoceras described by Zaborski (1990a: 5, figs 9, 10) are further examples of V. woodsi. They were incorrectly reported as having come from the Gadeni Zone at Pindiga but are from large exotic blocks derived from unit N upstream and not from the immediately adjacent unit A. Specimens from the Algerian Sahara referred to V. gamai by Collignon (1965: 185, figs 5—7) are a very close match for adult V. woodsi and may be conspecific. Unfortunately their inner whorls are completely unknown. Vascoceras bullatum Schneegans, 1943 Figs 13, 33-35 1943 Parayvascoceras crassus (Furon) var. bullata Schneegans: 131, pl. 8, figs 3, 4. 1989 Paravascoceras crassum (Furon); Meister: 18, pl. 6, figs 2, 3; text-fig. 12. 1989 Paravascoceras carteri (Barber); Meister: 21 (pars), pl. 9, fig. 1 (only). 1992 Vascoceras gr. crassum (Furon) ou costellatum Collignon; Courville: pl. 5, fig. 2; pl. 6, fig. 1. MATERIAL AND OCCURRENCE. Eleven specimens, C.93508-15, C.93516a-c, Pindiga Formation, unit O, Ashaka. DIMENSIONS. See Fig. 12. P.M.P. ZABORSKI REMARKS. Members of this species are relatively evolute and generally show markedly depressed whorls with a rounded to subtriangular outline. Umbilical bullae may or may not be present. The evenly frilled sutures are characterized by a broad low E/L and a narrow L. Most individuals are readily recognizable due to the development of regular ribbing on the flanks and venter during | their middle ontogenetic stages. The ribs may be coarse and — rounded but vary to finer, denser structures in other individuals. Although this ribbing may be developed at diameters of less — than 20 mm, some specimens remain smooth throughout | ontogeny (see Fig. 13; Meister 1989: pl. 9, fig. 1). Regularly | developed ribbing may be a transient feature which disappears | or weakens greatly on the later septate whorls. The more involute — members of the species overlap in shell proportions with certain — Vascoceras globosum costatum and Pseudovascoceras nigeriense (see Fig. 12) and such individuals may be difficult to differentiate if they lack ribbing. The juvenile whorls are not distinctive. They are often only moderately depressed and are smooth or ornamented with umbilical bullae alone (Fig. 34). The adult body-chamber makes up between two-thirds and three-quarters of the final whorl. It may be smooth but generally © shows an irregularly developed ornament of coarse, rounded fold-like to denser, sharper, narrow crease-like ribbing on the venter and outer flanks. Meister (1989: 18) pointed out that the body-chamber becomes constricted but the adult aperture itself is flared (Fig. 35). This modification occurs at diameters between 73 mm and 110 mm. There is no clear evidence of size dimorphism, however; other individuals showing a flared aperture do so at diameters of 80, 82, 82, 85, 85, 85, 86, 88, 95, 97, 98, 100 and 104 mm. Cobban & Hook (1983) described a large population sample of Neoptychites cephalotus (Courtiller) from New Mexico which shows similar adult body-chamber modifications. They too found that adult sizes were highly variable with no discernible bimodal pattern. Paravascoceras crassus var. bullata Schneegans (1943: 131, pl. 8, figs 3, 4) has depressed whorls and ribbing of a closely similar style to that in the present material. Its umbilicus, representing about 25% of the overall diameter, is narrower than the average in the Nigerian forms described here but in this respect there is an overlap with the more involute individuals (see Fig. 12). Schneegans (1943) regarded his material as a variety of Vascoceras ( Pachyvascoceras ) crassus Furon (1935: 58, pl. 3, figs 2a, b; text-fig. 17), a slightly less depressed and less evolute form. V. (P.) crassum shows only a very weak ornament of fine riblets though its suture pattern is close to that in the present material. As mentioned above, V. (P.) crassum may be most properly referred to Paravascoceras. In view of the clear similarities between the present material and Pachyvascoceras crassum vat. bullatum, however, the name Vascoceras bullatum Schneegans is | applied to it. The Nigerian material is distinct from coeval Paravascoceras cauvini and is therefore referred to Vascoceras rather than Paravascoceras, although with some uncertainty. Paravascoceras costatum multicostatum Barber (1960: 60, pl. 13, fig. 3; pl. 14, figs 1, 2) combines the relatively open umbilicus Figs 38-40 Vascoceras globosum costatum (Reyment). Pindiga Formation, unit O, Ashaka. Fig. 38a, b, C.93527, x1. Fig. 39a, b, C.93536b, x1. Fig. 40a, b, C.93536c, x1. Figs 41-44 Thomasites gongilensis (Woods). Pindiga Formation, unit O, Ashaka. Fig. 41, C.93582, x1. Fig. 42, C.93579, x1. Fig. 43, C.93580, x1. Fig. 44a, b, C.93583, x1. Figs 45-47 Vascoceras globosum proprium (Reyment). Pindiga Formation, unit T2, Ashaka. Fig. 45a, b, C.93548, x1. Fig. 46a, b, C.93551, x1. Fig. 47a, b, C.93547, x1. Til UPPER CRETACEOUS AMMONITE 78 and dense ribbing style of V. bullatum with the whorl proportions and suture pattern of V. globosum costatum. Material of this kind has not been found in the present study and its precise affinities are uncertain. Vascoceras rumeaui Collignon (1957) has an ornament of strong ribs similar to that in V. bullatum but is less depressed. As discussed above, it may be more closely related to Paravascoceras cauvini. V. durandi Choffat and the doubtfully distinct V kossmati Choffat (see Berthou et al. 1985 for a review) are both depressed, relatively evolute species but lack the marked ribbing and adult apertural modifications seen in V. bullatum. Vascoceras globosum (Reyment, 19545) REMARKS. This species was discussed by Kennedy et al. (1987: 46). They brought into synonymy a large amount of material described from Nigeria and elsewhere including Pachyvascoceras — globosum Reyment (19545) and Pachyvascoceras proprium Reyment (19545) under the name Vascoceras proprium (Reyment, 19546). Barber (1957: 21-27), however, had already treated the above two forms as conspecific and selected Vascoceras globosum (Reyment, 19545) as the species name. V. globosum accordingly has priority over V. proprium and should replace it. Nigerian specimens assigned to the species are here referred to three subspecies as detailed below. The V. globosum group shows a wide morphological variation and complex phylogenetic relationships. Early members of the group seem to include the ancestors of Thomasites gongilensis (Woods). Later members are characterized by rather complex sutures; V. obscurum and probably also Neoptychites Kossmat were derivatives. The precise origin of V. globosum is obscure. There are no obvious ancestors within the north-eastern Nigerian sections. V. woodsi is a possibility but is more evolute and has a stronger, more persistent juvenile ornamentation. Paravascoceras cauvini is markedly more compressed while the inner whorls of its later members are entirely smooth, unlike those in V. globosum costatum. Berthou et al. (1985: 72) suggested that Vascoceras (Pachyvascoceras ) crassus Furon, 1935 was a senior synonym of Pachyvascoceras costatum Reyment, 1954b (= V. globosum costatum). Meister et al. (1992: 72, pl. 7, figs 1, 2, 4, 5) described formes from sections at Tanout Aviation and Birgimari in Niger as Paravascoceras cauvini of V. crassum type and others transitional from V. crassum to V. proprium type. The latter group could probably include their V. gr. ellipticum Barber (Meister et al. 1992: 76, pl. 7, fig. 3; pl. 8, figs 1, 2) from the same sections. These formes are associated with Paravascoceras cauvini of typical aspect. Schneegans (1943) also reported passage forms from P. cauvini to V. crassum in Niger. Meister et al. (1992: 74) discussed the possible relationships within the Niger faunas. They remarked that undoubted V. proprium (= V. globosum) had not been found in Niger, and speculated that the horizons with P cauvini contained essentially monospecific ammonite faunas; this species produced a wide range of morphotypes with variable degrees of whorl compression and ornamental strength. According to this interpretation V crassum is a homeomorph of V. globosum costatum, the first a member of the P cauvini group, the latter having a separate origin. In support it may be noted that unit O at Ashaka, in which V. globosum appears, also contains the abrupt first occurrences of V. bullatum and Pseudovascoceras nigeriense. The base of unit O is a marine flooding surface suggesting that its fauna is largely an introduced one, the ancestors of which lie P.M.P. ZABORSKI outside north-eastern Nigeria. Vascoceras (Pachyvascoceras) crassum and V. globosum are here regarded as distinct from one another; the former, as stated above, is probably best referred to Paravascoceras. Vascoceras globosum costatum (Reyment, 1954b) Figs 38-40, 50 Pachyvascoceras costatum Reyment: 257, pl. 3, fig. 6; pl. 4, fig. 3; pl. 5, fig. 2; text-figs 3a, b, 5. 1955 Pachyvascoceras costatum Reyment; Reyment: 65, pl. 19546 14, figs 2, 4. 21957 Vascoceras robustum Barber: 15, pl. 5, fig. 1; pl. 26, figs 5, 6: 1957 Vascoceras polygonum Barber: 17, pl. 5, fig. 2; pl. 29, figs 1-3. 1957 Paravascoceras costatum costatum (Reyment) Barber: 35, pl. 14, fig. 1; pl. 32, figs 1-3. 1957 Paravascoceras costatum quadratum Barber: 35, pl. 16, fig. 2; pl. 32, figs 10, 11. 1957 Paravascoceras costatum tectiforme Barber: 37, pl. 14, fig. 4; pl. 15, figs 1, 3; pl. 16, fig. 2; pl. 32, figs 4-7. / 1965 Pachyvascoceras costatum Reyment; Reyment: pl. 3, fig. 17. 1976 Vascoceras robustum Barber; Offodile & Reyment: 54, figs 23a, b. 1976 —-Vascoceras ellipticum Barber; Offodile & Reyment: 55, figs 25a, b. 1976 Paravascoceras costatum (Reyment); Offodile & Reyment: 55, figs 26a, b. 1976 Paravascoceras tectiforme Reyment: 55, figs 29a, b. 1989 Paravascoceras tectiforme Barber; Meister: 21, pl. 7, figs 1, 2; pl. 8, figs 1—S; text-fig. 13. 1992 Vascoceras tectiforme (Barber) Courville: pl. 7, figs 1, 2. Barber; Offodile & sensu. Meister; MATERIAL AND OCCURRENCE. Twenty-six specimens, C.93310, | C.93519-34, C.93535a—d, C.93536a-e, Pindiga Formation, unit | O, Ashaka. | DIMENSIONS. See Figs 12, 48. REMARKS. The phragmocone in V. globosum costatum reaches a diameter in excess of 130 mm, making it the largest member of | the genus known in north-eastern Nigeria. Whorl breadth is | slightly to distinctly greater than whorl height while the | umbilicus represents 15-28% of the total diameter. In overall shell proportions V. globosum costatum overlaps with V. bullatum and Pseudovascoceras nigeriense and smoother individuals of | these two species may be difficult to distinguish from it, especially in their middle growth stages. At diameters of less than 30 mm (Figs 39, 40) the whorls in V. globosum costatum are weakly ornamented. Some forms are | virtually smooth, others display umbilical tubercles but most commonly there are weak, broadly rounded ribs, most | pronounced ventrally and sometimes with traces of bullate | ventrolateral tubercles. The whorls tend to be more compressed | in the early than in the later growth stages. In the middle whorls ornament may be lacking or there may be | broad, low ventral ribs. Umbilical tubercles persist in some | individuals. In the adult stages irregular fold-like ribs may appear, especially upon the venter. The range of shell shapes and | ornamentation is well displayed by the abundant previously described material (see synonymy list). UPPER CRETACEOUS AMMONITE Suture patterns are of the typically simple type characteristic of Vascoceras. Barber (1957), Meister (1989) and Courville (1992) separated forms of V. globosum costatum with subtriangular to triangular whorl sections as Paravascoceras costatum tectiforme Barber, P. tectiforme Barber and Vascoceras tectiforme (Barber). In unit O t Ashaka there is a complete intergradation of forms with ounded and triangular whorl sections. The latter themselves how rounded venters in their early growth stages. This shape ariation is attributed no taxonomic significance here. Of greater interest is the fact that V. globosum costatum shows gradation into Thomasites at the same stratigraphical level at shaka. A variation series is shown in Figs 41-44, 50. This sncompasses more or less smooth forms with ovoid to subtriangular whorls (Figs 41, 50) through forms with weak but efinite ventral tubercles and incipient carinae (Figs 42, 43), into strongly ornamented individuals (Fig. 44) occurring in the upper part of the unit and well within the morphological range of Thomasites gongilensis (Woods) (see faunas described by Barber 1957, Meister 1989). The Neoptychites cephalotus (Courtiller) of Meister (1989: 12, pl. 4, fig. 4) and Thomasites sp. nov.? of ourville (1992: 420, pl. 10, fig. 4; pl. 12, fig. 1) are further 2xamples of early Thomasites. V. globosum costatum seems to 2ontain the ancestors of 7: gongilensis, a species which reaches ts acme in unit R at Ashaka where it forms the bulk of the mmonite fauna. The Paravascoceras aff. chevalieri (Furon) of Reyment (1955: 53, pl. 14, figs 1a, b) from southern Nigeria also shows three rows of ventral tubercles and resembles the early Thomasites from Ashaka. The same can be said of the material of Discovascoceras tesselitense described by Collignon (1965: 181, pl. G, figs la, b) which shows three ventral carinae, subtriangular whorls and a eep, fairly narrow umbilicus. Hirano (1983) and Berthou et al. (1985) compared this material with Pseudotissotia, but it ppears closer to the Nigerian forms transitional from Pascoceras to Thomasites described here. Vascoceras globosum globosum (Reyment, 1954b) igs ol yo2 Pachyvascoceras globosum Reyment: 259, pl. 3, fig. 3; pl. 4, fig. 4; text-figs 3e, 7. | 1957 Vascoceras globosum globosum (Reyment) Barber: 21 (pars). 1957 —- Vascoceras globosum carteri Barber: 25, pl. 8, fig. 2; pl. 28, figs 8, 9. | 1965 Vascoceras carteri Barber; Reyment: pl. 3, fig. 12. 30 number of individuals 20.6 23.4 U/D(%) globosum globosum (Reyment) (see also Fig. 12). ee 79 1976 Paravascoceras carteri (Barber) Offodile & Reyment: 55, figs 27, 28. 1989 Paravascoceras carteri (Barber); Meister: 21 (pars), pl. 9, fig. 3 (only); pl. 10, figs 1, 2; text-fig. 14. 1992 Vascoceras tectiforme (Barber) sensu Courville: pl. 7, fig. 3 (only). 1992 Vascoceras gr. globosum (Reyment) ou Fagesia sp. Courville: pl. 8, figs 1, 2. Vascoceras sp. aff. obscurum Barber; Courville: pl. 10, fig. 3 (only). Meister; 71992 MATERIAL AND OCCURRENCE. Three specimens, C.93544-6, Pindiga Formation, unit R, Ashaka. The form also occurs in unit O at Ashaka and unit O at Pindiga. DIMENSIONS. See Figs 12, 48. REMARKS. V. globosum globosum is characterized by its highly depressed whorls which are at least twice as broad as high. Although such forms occur throughout the range of V. globosum in north-eastern Nigeria there are morphological and stratigraphical reasons for considering the earlier individuals as a separate subspecies. In unit O at Ashaka highly depressed examples of V. globosum are rare but those that occur fall well outside the morphological range of V. globosum costatum (see Figs 12, 48). In both unit R at Ashaka and unit O at Pindiga V. globosum globosum is the only member of the species that has been found. It is fairly frequent at these levels where the more compressed part of the morphological spectrum is occupied by large numbers of Thomasites gongilensis. The juvenile whorls in V. globosum globosum show sharper ribbing than in V. globosum costatum. The ribs cross the flank, unlike in V. globosum proprium in which they are largely confined to the venter and outer flanks. Specimen C.93544 (Fig. 51) is a very close match for the holotype of Pachyvascoceras globosum (C.47408, see Reyment 19545: pl. 3, fig. 3; pl. 5, fig. 4) which is also a juvenile. Since details of the inner whorls are useful in identification it is difficult to determine which of the specimens described by Barber (1957: 21) as V. globosum globosum should be referred to that subspecies and which are depressed examples of V. globosum proprium (see below). His material appears to include both. The sutures in V. globosum globosum are deeply incised; this appears to be a general feature of highly depressed Vascoceras which is shared with V. harttii (see below). Courville (1992: 421) drew attention to the suture pattern in V. globosum globosum and suggested that it may be better referred to Fagesia Pervinquiete. 30 1.31 1.89 2.47 Wb/Wh ig. 48 Shell proportions in Vascoceras globosum (Reyment) occurring in unit O at Ashaka. Forms with a high Wb/Wh ratio are here assigned to V. 80 This genus, however, as well as being generally more evolute, shows a different ontogeny (see Zaborski 1987: 43 for a discussion). Its juvenile whorls characteristically show strong ribs arising in twos or threes from umbilical tubercles, elements of this ornament persisting to varying stages in later ontogeny. The present material shows an ontogenetic development typical of Vascoceras rather than Fagesia. Barber (1957) proposed two species of Fagesia from north-eastern Nigeria but pointed out that they were both intermediate with Vascoceras. These forms are problematical, being based on a very few specimens of unknown stratigraphical provenance. F. simplex Barber (1957: 27, pl. 8, fig. 1; pl. 29, figs 4, 5) has a simple suture and is best regarded as an indeterminate Vascoceras. F. involuta Barber (1957: 27, pl. 9, fig. 3; pl. 29, figs 6, 7) has a complex suture, narrow umbilicus and highly depressed whorls; the early growth stages are unknown. It may be most closely related to V. globosum globosum. Vascoceras globosum proprium (Reyment, 19545) Figs 45-47, 49, 55—57, 63, 64 1920 Vascoceras angermanni Bose: 217, pl. 16, figs 1, 3 (only); pl. 17, fig. 1. 1920 Neoptychites aff. cephalotus (Courtiller); Bose: 221, pl. 18, figs 3, 10, 13. 1931 Thomasites sp. Adkins: 56, pl. 2, figs 16, 17. 1954b Pachyvascoceras proprium Reyment: 258, pl. 5, figs la, b; text-fig. 3d. 1954b Pachyvascoceras proprium plenum Reyment: 258, pl. 5, fig. 5; text-figs 3c, 6. 1954b Gombeoceras? bulbosum Reyment: 263, pl. 4, figs 2a, b; Fig. 49 Sutures in Vascoceras globosum proprium (Reyment). A, C.93550 at a diameter of 40 mm. B, C.93906 at a diameter of 54 mm. C, C.93905 at a diameter of 18 mm. D, C.93904 at a diameter of 30 mm. E, C.93549 at a diameter of 38 mm. All specimens from the Pindiga Formation, unit T2, Ashaka. 21957 1957 1957 1957 1957 1963 1963 1978 1982 1982 1987 1989 1989 1992 MATERIAL AND OCCURRENCE. Eleven specimens, C.93365, P.M.P. ZABORSK] text-figs 3g, 9. Vascoceras ellipticum Barber: 17 (pars), pl. 6, figs la, b: pl. 26, fig. 11 (only). Vascoceras globosum globosum (Reyment) Barber: 21 (pars), pl. 7, fig. 1. Vascoceras globosum plenum (Reyment) Barber: 23, pl. 7, fig. 2; pl. 9, fig. 2; pl. 28, figs 3—S. Vascoceras globosum proprium (Reyment) Barber: 25, pl. 7, fig. 3; pl. 28, figs 6, 7. Vascoceras globosum compressum Barber: 25, pl. 7, fig. 4; pl. 9, fig. 1; pl. 28, figs 10, 11. Pachyvascoceras compressum (Barber) Powell: 321, pl. 32, figs 2-4, 7; pl. 34, figs 8, 10; text-figs 3b—d, f. Pachyvascoceras globosum Reyment; Powell: 321, pl. 34, figs 7, 11; text-fig. 3s. Paravascoceras carteri (Barber); Chancellor, Reyment & Tait: 92, figs 15-17. Paravascoceras carteri (Barber); Chancellor: 102, figs 35-37. Paravascoceras compressum (Powell Chancellor: 106, figs 49, 50. Vascoceras proprium (Reyment); Kennedy, Wright & Hancock: 46, pl. 4, figs 1-15, 18, 19; pls 5, 6; text-figs 8A-C, 9. | Vascoceras (Vascoceras) proprium (Reyment);): Kennedy, Cobban, Hancock & Hook: 80, figs 20A, B. Vascoceras sp. juv. indet. Meister: pl. 11, fig. 3. Vascoceras sp. aff. obscurum Barber; Courville: pl. 10, fig. 2 (only). not Barber) L_5 mm_4 UPPER CRETACEOUS AMMONITE C.93547-51, C.93904-8, Pindiga Formation, unit T2, Ashaka. The subspecies also occurs in unit T1 at Ashaka. DIMENSIONS D Wb Wh U C.93365 64 48 (75) 32 (50) 13 (20) 93550 49 45 (92) 21 (43) = C.93549 46 30 (65) 21 (46) = C.93551 4) 34 (83) AL (SIV) 10.5 (26) 93548 37 23 (62) 17 (46) 6 (16) REMARKS. V. globosum proprium is generally a distinctive form on account of its narrow umbilicus, representing 15-28% of the total diameter, its deeply incised sutures often showing elongate saddles (Fig. 49), and its juvenile ornament of sharp ribbing, almost always confined to the outer flanks and venter. Associated with the juvenile ribs there may be pronounced onstrictions which persist until diameters of about 25 mm (Figs 53, 64). Umbilical bullae may or may not be present in the early 2rowth stages. The material from unit T2 at Ashaka varies from oderately compressed (Fig. 45) to highly depressed forms (Fig. 7). The latter resemble the stratigraphically younger V. zlobosum globosum. Their juvenile ribbing style is closer to that n Ke globosum proprium, however, and they are here treated as *nd members of that subspecies; similar variants occur in an issemblage of V. globosum proprium from Texas (Kennedy et al. 987). Some individuals have subdued ornamentation. The holotype f Gombeoceras? bulbosum Reyment (C.47295, Reyment 1954b: 1. 4, figs 2a, b) isa smooth V. globosum proprium. The holotype of Vascoceras ellipticum Barber (C.47679, 3arber 1957: pl. 6, figs la, b; pl. 26, fig. 11) is probably a further xample of V. globosum proprium. Other individuals referred to ‘ ellipticum by Barber (C.47680-4) are of uncertain affinities. whe example from Dukul (C.47633, Barber 1957: pl. 14, figs la, ); pl. 26, figs 3, 4) is an involute abraded specimen difficult to Jentify with certainty. After dissection many such forms found dose at Dukul prove to be Thomasites gongilensis. Courville (1992: 424, pl. 10, fig. 2) reported a specimen of V. Vobosum proprium (his V. gr. obscurum Barber) from unit U (his iveau 32) at Ashaka. The associated fauna described therein, owever, is that typical of unit T2 (= upper part of his Niveau 30 at Ashaka. In the present work V. globosum proprium has been und only in unit T. In its complex sutures and constricted inner whorls V. lobosum proprium resembles V. venezolanum Renz (1982: 80, pl. 3, figs S—11; pl. 24, figs 1-10; pl. 25, figs 1-8; text-fig. 61). The itter, highly variable species, however, generally shows denser, 1ore persistent ribs which cross the flanks, although certain idividuals may have subdued ribbing. V. venezolanum is known om southern Nigeria (Zaborski 1990b) but from beds Intaining Mammites nodosoides (Schliiter) which are younger 1an unit T at Ashaka. Jascoceras obscurum Barber, 1957 Figs 53, 54, 59, 61, 62 57 Vascoceras obscurum Barber: 19, pl. 6, figs 3, 9; pl. 27, figs 16-18. 289 Vascoceras obscurum Barber; Meister: 28, pl. 12, fig. 2; text-fig. 20. 81 MATERIAL AND OCCURRENCE. Five specimens, C.93552-3, C.93909-10, Pindiga Formation, unit T2, Ashaka; C.93326, Pindiga Formation, unit X, Ashaka. DIMENSIONS. D Wb Wh U C.93909 64 29 (45) 31 (48) 6(9) C.93553 48 24 (50) 26 (54) 4 (8) C.93326 44 21 (48) 24 (54.5) 4 (9) C.93552 38 19 (50) 20 (53) 4 (10.5) REMARKS. V. obscurum is a highly involute compressed species with a flattened to tabulate venter in its early stages which becomes rather more rounded during growth. The juvenile whorls bear strong regular ribs, some of which reach the umbilicus but which are mainly confined to the ventral region. The sutures are complex for the genus with fairly elongated highly frilled saddles (Fig. 59). V. obscurum appears in unit T2 at Ashaka, occurring there alongside V. globosum proprium. A similar style of ribbing and complex suture pattern is present in these two forms. V. obscurum could be considered as an end member of V. globosum proprium. The consistently high degree of involution, the compressed whorls and the flattened venter, however, set V. obscurum apart while it has a different stratigraphical range than the latter form, being found in unit X at Ashaka. For these reasons V. obscurumis here treated as a discrete species, though it is clearly very closely related to V. globosum proprium from which it is probably derived. The early whorls in V. obscurum resemble those in V. pioti (Peron & Fourtau) (see Freund & Raab 1969: 28, pl. 4, figs 1-9; text-figs 6d—g). The later growth stages, however, are unknown in the former, precluding comparison with the Neoptychites-like body-chamber in V. pioti. In what is known of its morphology V. obscurum shows a close similarity to Neoptychites. Pronounced constrictions have not been seen in the available material but are found in the related V. globosum proprium. The V. globosum proprium-V. obscurum lineage may contain the root stock of Neoptychites. Very similar to V. obscurum are the Neoptychites sp. juv. of Freund & Raab (1969: 47, pl. 8, figs 3-6) from Israel which occur well below the main stratigraphical range of the genus there. Material from unit O at Ashaka referred to Neoptychites by Meister (1989: 12, pl. 14, text-fig. 4) is, as mentioned above, an early Thomasites. That described by Courville (1992: 421, pl. 12, fig. 2) from unit R as Neoptychites aff. cephalotus (Couttiller) is probably a member of the V. globosum group. Vascoceras harttii (Hyatt, 1870) Figs 58, 60 1870 Ceratites harttii Hyatt; 386. 1875 Buchiceras hartti (Hyatt) Hyatt: 370. 1887 Ammonites (Buchiceras) harttii (Hyatt); White: 226, pl. 19, figs 1, 2; pl. 20, fig. 3. 1903 Vascoceras hartti (Hyatt) Hyatt: 103, pl. 14, fig. 16. 1936 Vascoceras hartti (Hyatt); Maury: 247, pl. 22, figs 1, 2. 1978 Paravascoceras hartti (Hyatt) Chancellor, Reyment & Tait: 96, fig. 20. 1982 Paravascoceras hartti (Hyatt); Chancellor: 98, figs 28C, 29-33. 1985 Vascoceras ( Paravascoceras ) harttii (Hyatt); Howarth: 82 P.M.P. ZABORSK UPPER CRETACEOUS AMMONITE 83 LL 5 mm_y Fig.59 Sutures in Vascoceras obscurum Barber. A, C.93910 at a diameter of 32 mm. B, C.93909 at a diameter of 45 mm. C, C.93552 at a diameter of 30 mm. D, C.93326 at a diameter of 34 mm. All specimens from the Pindiga Formation, unit T2, Ashaka except C.93326 which is from unit X at the same locality. 100, fig. 25 (with synonymy). Vascoceras hartti (Hyatt); Cobban, Hook & Kennedy: 49, figs 49, 91A-D, G-K. 1989 Fagesia superstes var. levis Renz; Meister: 37, pl. 16, fig. 2; text-fig. 26. 1992. Vascoceras gr. globosum (Reyment) ou Fagesia sp.; Courville: pl. 9, fig. 1. 21989 MATERIAL AND OCCURRENCE. Two specimens, C.93554-5, Pindiga Formation, unit X, Ashaka. DIMENSIONS. | D Wb Wh U | 93555 79 83 (105) 33 (42) 21 (26.5) C.93554 49 50 (102) 21 (43) 12 (24.5) | |REMARKS. These moderately evolute cadicones show sharp umbilical shoulders, which are undulating in C.93555, and steeply sloping umbilical walls. There are sharply rounded ventral ribs and occasional constrictions in C.93554 (Fig. 58) but they fade by a diameter of 40 mm leaving the internal mould | smooth but for transverse growth striae. Neither specimen shows | umbilical tubercles. Kennedy et al. (1987: 51) pointed out the difficulty in distinguishing V. hartti from globose V. globosum. They regarded a more evolute coiling and a steeply sloping umbilical wall as most useful in identifying the former. In these respects the similar V. globosum globosum. In the present work V. harttii has been found only in unit X at Ashaka. Meister (1989: 37-38) and Courville (1992: 424) also reported examples from unit U there which they referred to or compared with Fagesia superstes (Kossmat). As with V. globosum globosum (see above) these forms have an ontogenetic development characteristic of Vascoceras not Fagesia. The globose shape and complex suture pattern are not in themselves diagnostic of Fagesia. STRATIGRAPHICAL AND PHYLOGENETIC DISCUSSION The oldest ammonite-bearing beds in north-eastern Nigeria yield no vascoceratid taxa. They are characterized by Nigericeras gadeni, Metengonoceras dumbli (Cragin), Placenticeras (Karamaites) cumminsi Cragin and Metoicoceras geslinianum (d’Orbigny), the last species allowing correlation with the Geslinianum Zone in north-western Europe and the Gracile Zone of the western interior of the United States (see Kennedy 1984, Cobban 1984, Cobban ef al. 1989). This ‘Nigericeras fauna’ is widely recognizable in West and Saharan Africa (see Lefranc 1978, Meister et al. 1992). Paravascoceras cauvini appears in unit E at Ashaka and becomes common in unit F there and in unit H at Pindiga. In the last two horizons it is associated with Burroceras? sp. (Zaborski 1995). Although not identifiable to species level this material may indicate correlation with the Burroceras clydense Zone of New Mexico. P. cauvini ranges through units K and M at Ashaka wherein Vascoceras woodsi occurs. These units also contain Pseudaspidoceras pseudonodosoides, on which basis they can be correlated with the Juddii Zone in the western interior. In south-western New Mexico a gap exists between the Juddii Zone and the basal Turonian Flexuosum Zone (Cobban et al. 1989). Pseudaspidoceras flexuosum Powell occurs in unit T2 at Ashaka. Units N to T1 at Ashaka belong to the Upper Cenomanian but Fig.50 Vascoceras globosum costatum (Reyment). Pindiga Formation, unit O, Ashaka. C.93523, x1. Figs 51,52 Vascoceras globosum globosum (Reyment). Pindiga Formation, unit R, Ashaka. Fig. 51, C.93544, x1. Fig. 52a, b, C.93545, x1. Figs 53,54 Vascoceras obscurum Barber. Pindiga Formation, unit T2, Ashaka. Fig. 53a, b, C.93552, x1. Fig. 54, C.93553, x1. Figs 55-57 Vascoceras globosum proprium (Reyment). Pindiga Formation, unit T2, Ashaka. Fig. 55a, b, C.93365, x1. Fig. 56a, b, C.93549, x1. Fig. : material is more properly referred to V. harttii than the ‘] 57a, b, C.93550, x1. | Fig.58 Vascoceras harttii (Hyatt). Pindiga Formation, unit X, Ashaka. C.93554, x1. 84 Fig. 60 Vascoceras harttii (Hyatt). Pindiga Formation, unit X, Ashaka. C.93555, x1. Figs 61,62 Vascoceras obscurum Barber. Fig. 61a, b, Pindiga Formation, unit X, Ashaka. C.93326, x1. Fig. 62a, b, Pindiga Formation, unit T2, Ashaka. C.93909, x1. Figs 63,64 Vascoceras globosum proprium (Reyment). Pindiga Formation, unit T2, Ashaka. Fig. 63a, b, C.93904, x1. Fig. 64a—-c, C.93905, x1. lack equivalents in south-western New Mexico. Unit O at Ashaka, which contains the youngest Paravascoceras cauvini, Pseudovascoceras nigeriense, Vascoceras bullatum, V. globosum costatum and V. globosum globosum is probably at least partially equivalent to beds with Nigericeras scotti in south-eastern Colorado. Unit T2 at Ashaka contains an ammonite assemblage including Pseudaspidoceras flexuosum and Vascoceras globosum proprium. These forms occur together in the basal Turonian Flexuosum Zone in west Texas (Kennedy et al. 1987), Thomasites and Wrightoceras munieri (Pervinquiére) also being associated in both places. V. globosum proprium is further recorded from New Mexico and Hancock (1991) suggested that it may serve as a better marker for the base of the Turonian than Pseudaspidoceras flexuosum. In the Ashaka section, however, it actually appears in unit T1 just below the first occurrence of P flexuosum. A minor discontinuity representing a marine P.M.P. ZABORSKI flooding surface separates units T1 and T2. The fauna of unit T2— seems to have in large part been introduced during this flooding event which may complicate the order of occurrence of these taxa at Ashaka. Vascoceras obscurum ranges from the basal Turonian unit T2 into unit X at Ashaka. Units U to X at Ashaka, which also represent the known range of V. harttii, are of Early Turonian age. They cannot, however, be dated more precisely on the basis” of their ammonite faunas which are almost entirely composed of Pseudotissotia nigeriensis (Woods) and Eotissotia simplex Barber. V. harttii has been assigned to the Lower Turonian in Angola (Howarth 1985), Brazil (Bengtson 1983) and Mexico (Chancellor 1982) but material from the Upper Cenomanian of New Mexico has also been referred to the species (Cobban et al. 1989: 49, figs 49, 91A—D, G—K). With the exception of a few taxa ‘vascoceratid’ ammonites | have not proved to be of great value in detailed correlation, UPPER CRETACEOUS AMMONITE levels of immigrant faunas especially inter-regionally. A number of problems complicate ‘their use including: the variable taxonomic treatment authors have employed; the difficulty of identifying poorly preserved ‘material, especially if the inner whorls are not available; the lack of an exact stratigraphical provenance for many species; and the problem of polymorphism. In regard to the last of these factors Meister (1989) and Meister et al. (1992) have made the important point that in certain regions only a portion of the potential morphological range of a species may be expressed on account of palaeoecological factors. It has long been appreciated that ‘vascoceratid’ ammonites are predominantly a Tethyan group. Less attention has been paid to the potential palaeoenvironmental influences on their regional distribution. In this regard it is of interest to compare the faunas lof north-eastern Nigeria and Niger, the stratigraphical distributions of which are now well understood. Despite their geographical proximity correlation between these two areas is not as straightforward as might be expected. Ther is little problem with the horizons of Geslinianum Zone 85 — = = oe & = £ o Qa = S alee ” a 5 ” 5 a a A € fe) 3) fe) = 5 ra 8 = a) (2) oO 3 = Sg ® 7) fe} = oO x2) Lu ° FSH agaee Oo cod S = = 1 3 g = Oo. -10 ew ie aah | : () ae ) ee | 5 BH loacore DB re) ® o no 6 2 @ 0S P= ) E = 9 = Ss oO a oO = S Q @ B ~ (e} pes iD § o oD “3 8: = © 88 o = oE Bie D , @ = 122 | ee eee (oe le Onna Me of Lasts ° to) > COML aa. pises . ae BS -OoD ® : SO Gide) S70\Te oD : . > 0S) On i x =o ome) : : 2 ic (oe) : cit = nO : : z os : : a > © ; 2) i © ° _ % : oO 2 : oO = 8 . D : ® . g . a e < : _ o fal 2) o = oo , : =O . : '3s Y v 5 2 Vascoceras Cunningtoniceras? diartianum Fig.65 Stratigraphical ranges in the Ashaka section of taxa mentioned in the text and their suggested phylogenetic relationships. age; in both places assemblages of Nigericeras, Metoicoceras and Metengonoceras are found. Similarly, in the upper part of the Turonian Coilopoceras is present in both areas. There are, however, faunal differences in the intervening sequences. Vascoceras woodsi is unknown in Niger. In Nigeria it occurs alongside Pseudaspidoceras pseudonodosoides which was compared with P tassaraense Meister, Alzouma, Lang & Mathey (1992) from the Monts Iguelela area of Niger by Zaborski (1995). The two may be of comparable age. P tassaraense occurs alongside Nigericeras jacqueti involutum, a form unknown in Nigeria. Slightly lower in the same section there occur large numbers of Cibolaites(?) africaensis Meister, Alzouma, Lang & Mathey (1992) which has not been found further to the south. At Tanout Aviation and Birgimari there are horizons dominated by compressed individuals of Paravascoceras cauvini. In their ornament or lack of it they match the variation shown by the species in unit O at Ashaka. The Niger faunas, however, include more inflated individuals of Vascoceras crassum type. None of the species associated with P 86 cauvini in unit O at Ashaka is found in the described Niger horizons. Higher in the sequence Thomasites gongilensis, so abundant in unit R at Ashaka and unit O at Pindiga, and with an overall range from unit O to unit T2 at the former locality, is not found in any of the Niger sections. Pseudotissotia nigeriensis, on the other hand, occurs in large numbers at Tanout Aviation but none of the associated taxa in Nigeria accompany it there. Biostratigraphical comparison between Niger and Nigeria is complicated by the fact that ammonites are restricted to limestone horizons which are, in the main, thin units within dominantly argillaceous sequences. The presence or absence of particular faunas, therefore, may in some cases be related to the occurrence of calcareous beds (see also Meister et al. 1992: 91). The possibility of control over ammonite distributions by transgressive pulses of the trans-Saharan sea during the Late Cenomanian and Early Turonian has long been discussed, most recently by Courville et a/. (1991). Meister et al. (1992: 94-95), however, have speculated that local palaeoenvironments were a strong influence on ‘vascoceratid’ distributions, their morphological polymorphism and _ their consequent evolutionary potential. In support of the latter hypothesis it may be noted that members of evolutionary lineages ‘indigenous’ to north-eastern Nigeria (Nigericeras, Paravascoceras cauvini and Pseudotissotia nigeriensis) extend into Niger. Introduced taxa do not. Among these may be mentioned Burroceras? from unit F at Ashaka; Vascoceras woodsi from units K and M; the greater part of the fauna from unit O including Pseudaspidoceras footeanum (Stoliczka), Fikaites, Rubroceras, Pseudovascoceras nigeriense and, probably, Vascoceras globosum costatum; Pseudaspidoceras paganum from unit R; and Pseudaspidoceras flexuosum, Watinoceras, Choffaticeras and Wrightoceras munieri from unit T2. The appearances of these taxa are probably related to transgressive pulses, the influences of which did not fully extend into Niger. As suggested by Meister et al. (1992) the absence of these forms and consequent lack of competition may have permitted local intraspecific variants and evolutionary lineages to develop in Niger. Examples would be the inflated variants of Paravascoceras cauvini and the lineage leading to Nigericeras jacqueti involutum. Rather than the overall extent of the trans-Saharan sea as such, more localized influences such as water depth and temperature may have controlled the distribution of taxa. If these factors did apply they would place important constraints on the use of ‘vascoceratid’ species in long distance correlation. Associated with the above matter is the probability that a number of acanthoceratine taxa independently gave rise to vascoceratid-like forms during Late Cenomanian times. Reyment (1979: 111) suggested that the family Vascoceratidae was polyphyletic, the morphological similarities between its members being due to adaptation to the same kind of epicontinental palaeoenvironments rather than to close phylogenetic affinities. The Late Cenomanian transgression brought several forms into north-eastern Nigeria which show elements of the ‘vascoceratid’ morphology, notably simplified sutures. Rubroceras and Fikaites, the latter probably derived from Eucalycoceras Spath, are examples, as is Pseudovascoceras which, as mentioned above, may be a descendent of Cunningtoniceras. Reyment (1955: 62, text-fig. 27) regarded Nigericeras as the root stock of the entire family Vascoceratidae while Cooper (1979) suggested that Vascoceras diartianum gave rise to both Paravascoceras and the younger Vascoceras. It is suggested here that Paravascoceras is derived from Nigericeras and belongs to a lineage separate to that leading to Vascoceras. The earliest Vascoceras in north-eastern Nigeria, V. woodsi, P.M.P. ZABORSKI appears to be a peramorphic derivative of V. diartianum. The immediate origins of V. bullatum, V. globosum and V. harttii are obscure. It may be mentioned, however, that V. globosum costatum probably contains the progenitors of Thomasites gongilensis. This species in turn gave rise to Pseudotissotia nigeriensis in terminal Cenomanian times (see also Barber 1957, Cooper 1979, Meister 1989) from which Eotissotia simplex originated as a paedomorph during the Early Turonian (Zaborski 1993). The youngest member of the V. globosum group, IY, globosum proprium, straddles the Cenomanian-Turonian boundary. It gave rise to V. obscurum — and probably also Neoptychites. Kennedy & Wright (1979: 681) believed the latter genus to have been derived from Paravascoceras but used this name in the sense of Vascoceras without umbilical tubercles. The suggested phylogenetic relationships of forms from north-eastern Nigeria are shown in Fig. 65. Several converging lineages are believed to exist, their frequently homeomorphic similarities being due to colonization of the same palaeoenvironment. ACKNOWLEDGEMENTS. Thanks are due to Drs M. K. Howarth and H. G. Owen, and Mr. S. Baker for help in many ways. Dr. N. Morris kindly allowed access to a collection of ammonites from Niger. Mr. M. Baku, Quarry Manager, Ashaka Cement Co. kindly allowed easy access to the Ashaka quarry. Photographs were provided by the Natural History Museum (London) Photographic Unit. Dr. W. J. Kennedy kindly made useful suggestions concerning the manuscript. REFERENCES Adkins, W. S. 1931. Some Upper Cretaceous ammonites in western Texas. Bulletin of the University of Texas Bureau of Economic Geology and Technology, Austin, | 3101: 35-72, pls 2-5. Amard, B., Collignon, M. & Roman, J. 1981. Etude stratigraphique et | paléontologique du Crétacé supérieure et Paléocene du Tinrhert-W et | Tademait-E (Sahara Algérien). Documents du Laboratoire de Géologie de la Faculté des Sciences de Lyon, (H.S.) 6: 15-173, pls 1-17. Barber, W. 1957. Lower Turonian ammonites from north-eastern Nigeria. Bulletin of the Geological Survey of Nigeria, Kaduna, 26: 1-86, pls 1-35. — 1960. 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Monographs of the United States Geological Survey, Washington, 44: 351 pp., 47 pls. Kennedy, W. J. 1971. Cenomanian ammonites from southern England. Special sahariennes. Annales de 87 Papers in Palaeontology, London, 8: 1-133, pls 1-64. 1984. Ammonite faunas and the ‘standard zones’ of the Cenomanian to Maastrichtian stages in their type areas, with some proposals for the definition of the stage boundaries by ammonites. Bulletin of the Geological Society of Denmark, Copenhagen, 33: 147-161. — & Cobban, W. A. 1990a. Cenomanian ammonite faunas from the Woodbine Formation and the lower part of the Eagle Ford Group, Texas. Palaeontology, London, 33: 75-154, pls 1-17. —— & —— 1|990b. Cenomanian micromorphic ammonites from the western interior of the USA. Palaeontology, London, 33: 379-422, pls 1-7. % , Hancock, J. M. & Hook, S. C. 1989. Biostratigraphy of the Chispa Summit Formation at its type locality: a Cenomanian through Turonian reference section for Trans-Pecos Texas. Bulletin of the Geological Institution of the University of Uppsala, (NS) 15: 39-119. —— & Juignet, P. 1977. Ammonites diartianus d’Orbigny, 1850, Vascoceratidae du Cénomanien supérieur de Saint-Calais (Sarthe). Geobios, Lyon, 10: 583-595, pls 1, 2. —— & Wright, C. W. 1979. Vascoceratid ammonites from the type Turonian. Palaeontology, London, 22: 665-683, pls 82-86. —— & —— 1985. Evolutionary patterns in Late Cretaceous ammonites. Special Papers in Palaeontology, London, 33: 131-143. & In press. The affinities of Nigericeras Schneegans, 1943 (Cretaceous, Ammonoidea). Geobios, Lyon. & Hancock, J. M. 1987. Basal Turonian ammonites from west Texas. Palaeontology, London, 30: 27-74, pls 1-10. Kler, M. O. 1909. [Neoceratites of eastern Bukhara.]. Trudy Geologicheskago i Mineralogicheskago Muzeya, St. Petersburg, 2 (for 1908): 157-174, pls 6-8. [In Russian]. Kummel, B. & Decker, J. M. 1954. Lower Turonian ammonites from Texas and Mexico. Journal of Paleontology, Tulsa, 28: 310-319, pls 30-33. Lefranc, J. P. 1978. Etat des connaissances actuelles sur les zonations biostratigraphiques du milieu du Crétacé (Albien 4 Turonien) au Sahara. Annales du Muséum d'Histoire Naturelle de Nice, 4 (X1X): 1-19. Lewy, Z., Kennedy, W. J. & Chancellor, G. R. 1984. Co-occurrence of Metoicoceras geslinianum (d’Orbigny) and Vascoceras cauvini Chudeau (Cretaceous Ammonoidea) in the southern Negev and its stratigraphic implications. Newsletters on Stratigraphy, Leiden, 13: 67-76. Luger, P. & Gréschke, M. 1989. Late Cretaceous ammonites from the Wadi Qena area in the Egyptian eastern desert. Palaeontology, London, 32: 355-407, pls 38-49. Maury, C. J. 1936. O Cretaceo de Sergipe. Monografias Servico Geologico e Mineralogico do Brasil, Rio de Janeiro, 11: 283 pp., 28 pls. Meister, C. 1989. Les ammonites du Crétacé supérieure d’Ashaka (Nigéria). Bulletin des Centres Recherches Exploration-Production Elf Aquitaine, Boussens, 13 (supplement): 1-84, pls 1-28. ——, Alzouma, K., Lang, L. & Mathey, B. 1992. Les ammonites du Niger (Afrique Occidentale) et la transgression transsaharienne au cours du Cénomanien-Turonien. Geobios, Lyon, 25: 55-100, pls 1-11. Offodile, M. E. & Reyment, R. A. 1976. Stratigraphy of the Keana-Awe area of the middle Benue region of Nigeria. Bulletin of the Geological Institution of the University of Uppsala, (NS) 7: 37-66. Pervinquiére, L. 1907. Etudes de paléontologie tunisienne. 1, Céphalopodes des terrains secondaires. 438 pp., 27 pls. Paris, Carte géol. Tunis. Popoff, M., Wiedmann, J. & de Klasz, I. 1986. The Upper Cretaceous Gongila and Pindiga Formations, northern Nigeria: subdivisions, age, stratigraphic correlations and paleogeographic implications. Eclogae geologicae Helvetiae, Basel, 79: 343-363. Powell, J. D. 1963. Cenomanian-Turonian (Cretaceous) ammonites from Trans-Pecos, Texas and north-eastern Chihuahua, Mexico. Journal of Paleontology, Tulsa, 37: 309-332, pls 31-34. Reeside, J. R. 1923. A new fauna from the Colorado Group of southern Montana. Professional Papers of the United States Geological Survey, Washington, 132-B: 25-33, pls 11-21. Renz, O. 1982. The Cretaceous ammonites of Venezuela. 132 pp., 40 pls. Basel. Reyment, R. A. 1954a. New Turonian (Cretaceous) ammonite genera from Nigeria. Colonial Geology and Mineral Resources, London, 4: 149-164, pls 1-4. 1954b. Some new Upper Cretaceous ammonites from Nigeria. Colonial Geology and Mineral Resources, London, 4: 248-270, pls 1—S. — 1955. The Cretaceous Ammonoidea of southern Nigeria and the southern Cameroons. Bulletin of the Geological Survey of Nigeria, Kaduna, 25: |—112, pls 1-25. — 1956. On the stratigraphy and palaeontology of the Cretaceous of Nigeria and the Cameroons, British West Africa. Geologiska Foreningens i Stockholm Forhandlungar, 78: 17-96. 1965. Aspects of the geology of Nigeria. 145 pp., 18 pls. Ibadan. 1979. Variation and ontogeny in Bauchioceras and Gombeoceras. Bulletin of the Geological Institution of the University of Uppsala, (NS) 8: 89-111. 88 1988. Does sexual dimorphism occur in Upper Cretaceous ammonites? Senck. lethaea, Frankfurt, 69: 109-119. Roman, F. 1938. Les ammonites jurassiques et crétacées. Essai de genera. 554 pp., 53 pls. Paris. Schneegans, D. 1943. Invértébres du Crétacé supérieure du Damergou (Territoire du Niger). Bulletin de la Direction Fédérale des Mines et de la Géologie, Afrique Occidentale Frangaise, Dakar, 7: 87-150, pls 1-8. Schobel, J. 1975. Ammoniten der Familie Vascoceratidae aus dem Unterturon des Damergou-Gebietes, République du Niger. Special Publications of the Palaeontological Institution of the University of Uppsala, 3: \-136, pls 1-6. Sharpe, D. 1855. Description of the fossil remains of Mollusca found in the Chalk of England. 1, Cephalopoda: 27-36, pls 11-16. Monographs of the Palaeontographical Society, London. Solger, F. 1904. Die Fossilien der Mungokreide in Kamerun und ihre geologische Bedeutung, mit besonderer Beriicksichtigung der Ammonitiden. Jn, Esch, E., Solger, F, Oppenheim, M. & Jakel, O., Beitraége zur Geologie von Kamerun: 83-242, pls 3-5. Stuttgart. Spath, L. F. 1923. Appendix II. On the ammonite horizons of the Gault and contiguous deposits. Summary of Progress of the Geological Survey of Great Britain, 1922: 139-149. 1925. On Upper Albian Ammonoidea from Portugese East Africa. With an appendix on Upper Cretaceous ammonites from Maputoland. Annals of the Transvaal Museum, Pretoria, 11: 179-200, pls 28-37. White, C. A. 1887. Contribuicoes 4 paleontologia do Brazil. Archivos do Museu Nacional Rio de Janeiro, 7: 1-273, 28 pls. Wiedmann, J. 1960. Le Crétacé supérieure de |’Espagne et du Portugal et ses céphalopodes. Jn, Colloque sur le Crétacé supérieure Frangais (Dijon, 1959). Compte rendu Congres des Sociétés Savantes de Paris et des Départments, Section des Sciences, Paris, 1959: 709-764, 8 pls. Woods, H. 1911. The palaeontology of the Upper Cretaceous deposits of Northern Nigeria. Jn, Falconer, J. D., The geology and geography of Northern Nigeria: 273-286, pls 19-24. London. APPENDIX P.M.P. ZABORSKI Wozny, E. & Kogbe, C. A. 1983. Further evidence of marine Cenomanian, Turonian and Maastrichtian in the Upper Benue Basin of Nigeria (west Africa). Cretaceous Research, London, 4: 95-99. Wright, C. W. 1957. Mollusca 4; Cephalopoda, Ammonoidea. Jn, Moore, R. C. (ed.), Treatise on Invertebrate Paleontology, L: L80-L490. Lawrence, Kansas. — & Kennedy, W. J. 1980. Origin, evolution and systematics of the dwarf acanthoceratid Protacanthoceras Spath, 1923 (Cretaceous Ammonoidea). Bulletin of the British Museum (Natural History), London, (Geology), 34 (2): 65-107. —— & —— 1981. The Ammonoidea of the Plenus Marls and the Middle Chalk. 148 pp., 32 pls. Monographs of the Palaeontographical Society, London. —— & —— 1987. The Ammonoidea of the Lower Chalk. 2: 127-218, pls 41-55. Monographs of the Palaeontographical Society, London. Zaborski, P. M. P. 1985. Upper Cretaceous ammonites from the Calabar region, south-east Nigeria. Bulletin of the British Museum (Natural History), London, (Geology), 39 (1): 1-72. — 1987. Lower Turonian (Cretaceous) ammonites from south-east Nigeria. Bulletin of the British Museum (Natural History), London, (Geology), 41 (2): 31-66. — 1990a. The Cenomanian and Turonian (mid-Cretaceous) ammonite biostratigraphy of north-eastern Nigeria. Bulletin of the British Museum (Natural History), London, (Geology), 46 (1): 1-18. — 1990b. Some Upper Cretaceous ammonites from southern Nigeria. Journal of African Earth Science (and the Middle East), Oxford, 10: 565-581. — 1993. Some new and rare Upper Cretaceous ammonites from north-eastern Nigeria. Journal of African Earth Science (and the Middle East), Oxford, 17: 359-371. 1995. The Upper Cretaceous ammonite Pseudaspidoceras Hyatt, 1903 in north-eastern Nigeria. Bulletin of the British Museum (Natural History), London, (Geology), 51: 53-72, 24 figs. A list of previously described material from Nigeria representing taxa discussed herein is given below with revised taxonomic determinations. The page and, where necessary, plate and figure numbers quoted are those in the original publications. Woods (1911) 281 Vascoceras nigeriense sp. nov. Reyment (1954) 256 Vascoceras nigeriense Woods 257 Pachyvascoceras costatum sp. nov. 258 Pachyvascoceras proprium sp. Nov. 258 Pachyvascoceras proprium plenum subsp. nov. 259 Pachyvascoceras globosum sp. nov. 263 Gombeoceras? bulbosum sp. nov. Reyment (1955) 63 Paravascoceras aff. chevalieri (Furon) 65 Pachyvascoceras costatum Reyment Barber (1957) 15 Vascoceras nigeriense Woods 15 Vascoceras robustum sp. nov. 17 Vascoceras polygonum sp. nov. 17, pl. 4, fig. 1 Vascoceras ellipticum sp. nov. 17, pl. 6, fig. 4 Vascoceras ellipticum sp. nov. 19 Vascoceras bulbosum (Reyment) 19 Vascoceras depressum sp. nov. 19 Vascoceras obscurum sp. nov. 21 Vascoceras globosum globosum (Reyment) 23 Vascoceras globosum plenum (Reyment) 25 Vascoceras globosum proprium (Reyment) 25 Vascoceras globosum compressum subsp. nov. 25 Vascoceras globosum carteri subsp. nov. 27 Vascoceras sp. Juv. 27 Fagesia simplex sp. nov. Revised determination Pseudovascoceras nigeriense (Woods) Pseudovascoceras nigeriense (Woods) Vascoceras globosum costatum (Reyment) Vascoceras globosum proprium (Reyment) Vascoceras globosum proprium (Reyment) Vascoceras globosum globosum (Reyment) Vascoceras globosum proprium (Reyment) Thomasites Vascoceras globosum costatum (Reyment) Pseudovascoceras nigeriense (Woods) 2 Vascoceras globosum costatum (Reyment) ? Vascoceras globosum costatum (Reyment) ?Thomasites gongilensis (Woods) ? Vascoceras globosum proprium (Reyment) Paravascoceras cauvini (Chudeau) Paravascoceras cauvini (Chudeau) Vascoceras obscurum (Barber) Vascoceras globosum globosum (Reyment) (part) and V. globosum proprium (Reyment) (part) Vascoceras globosum proprium (Reyment) Vascoceras globosum proprium (Reyment) Vascoceras globosum proprium (Reyment) Vascoceras globosum globosum (Reyment) Vascoceras woodsi sp. nov. indeterminate Vascoceras UPPER CRETACEOUS AMMONITE 27 Fagesia involuta sp. nov. 29 Nigericeras costatum sp. nov. 29 Nigericeras glabrum sp. nov. 31 Nigericeras(?) intermedium sp. nov. 31 Paramammites tuberculatus sp. nov. 33 Paramammites raricostatus sp. Nov. 33 Paramammites inflatus sp. nov. 35 Paravascoceras costatum costatum (Reyment) 35 Paravascoceras costatum quadratum subsp. nov. 37 Paravascoceras costatum tectiforme subsp. nov. 37 Paravascoceras aff. cauvini (Chudeau) Meister (1989) 10 Nigericeras gadeni (Chudeau) — /amberti Schneegans 11 Nigericeras jacqueti Schneegans 11 Plesiovascoceras aff. gr. thomi (Reeside) ou sp. nov. 12 Neoptychites cephalotus (Courtiller) 14, pl. 5, fig. 2 Paravascoceras gr. evolutum Schneegans 14, pl. 5, fig. 4 Paravascoceras gr. evolutum Schneegans 14 Paravascoceras nigeriensis(?) (Woods) 14 Paravascoceras aff. nigeriensis (?) (Woods) 18 Paravascoceras crassum Furon 21 Paravascoceras tectiforme (Barber) 21, pl. 9, fig.1 Paravascoceras carteri Barber 21, pl. 10, figs 1, 2 Paravascoceras carteri Barber 23 Vascoceras costatum (Barber) 23 Vascoceras costatum (Barber) glabrum (Barber) 28 Vascoceras ellipticum Barber 28 Vascoceras silvanense Choffat 28 Vascoceras obscurum Barber 30 Paramammites subconciliatus (Choffat) 36, pl. 14, figs 3, 4 Paramammites polymorphus (Pervinquiére) 36, pl. 15, figs 2, 3 Paramammites aff. gr. polymorphus (Pervinquiére) 37 Fagesia superstes var. levis Renz Pl. 16, fig. 1 Thomasites? Zaborski (1990a) Figs 8, 12-15 Vascoceras cauvini Chudeau Figs 9, 10 Vascoceras sp. Fig. 11 Vascoceras bulbosum (Reyment) Figs 16-18, 20, 21 Vascoceras sp. juv. Fig. 25 Vascoceras nigeriense Woods Courville (1992) Pl. 4, figs 1-3 Vascoceras gr. cauvini Chudeau Pl. 5, fig. 1 Vascoceras gr. thomi (Reeside) ou evolutum (Schneegans) Pl. 5, fig. 2 Vascoceras gr. crassum (Furon) ou costellatum Collignon Pl. 5, fig. 3; pl. 6, figs 2, 3 Vascoceras sp. gr. costatum (Barber) Pl. 7, figs 1, 2 Vascoceras tectiforme (Barber) Pl. 7, fig. 3 Vascoceras tectiforme (Barber) PI. 8, figs 1, 2 Vascoceras gr. globosum (Reyment) ou Fagesia sp. PI. 9, fig. 1 Vascoceras gr. globosum (Reyment) ou Fagesia sp. Pl. 10, fig. 1 Vascoceras sp? Pl. 10, figs 2, 3 Vascoceras sp. aff. obscurum Barber 89 ? Vascoceras globosum globosum (Reyment) Pseudovascoceras nigeriense (Woods) Pseudovascoceras nigeriense (Woods) Pseudovascoceras nigeriense (Woods) Pseudovascoceras nigeriense (Woods) Pseudovascoceras nigeriense (Woods) Pseudovascoceras nigeriense (Woods) Vascoceras globosum costatum (Reyment) Vascoceras globosum costatum (Reyment) Vascoceras globosum costatum (Reyment) ? Paravascoceras cauvini (Chudeau) Paravascoceras cauvini (Chudeau) Paravascoceras cauvini (Chudeau) Vascoceras woodsi sp. nov. Thomasites ? Pseudaspidoceras pseudonodosoides (Choffat) Vascoceras woodsi sp. nov. Pseudovascoceras nigeriense (Woods) Paravascoceras cauvini (Chudeau) Vascoceras bullatum Schneegans Vascoceras globosum costatum (Reyment) Vascoceras bullatum Schneegans Vascoceras globosum globosum (Reyment) Pseudovascoceras nigeriense (Woods) Pseudovascoceras nigeriense (Woods) Pseudovascoceras nigeriense (Woods) indeterminate Vascoceras Vascoceras obscurum Barber Pseudovascoceras nigeriense (Woods) Pseudovascoceras nigeriense (Woods) Fikaites varicostatus Zaborski Vascoceras harttii (Hyatt) Fikaites varicostatus Zaborski Paravascoceras cauvini (Chudeau) Vascoceras woodsi sp. nov. Paravascoceras cauvini (Chudeau) Vascoceras woodsi sp. nov. Pseudovascoceras nigeriense (Woods) Paravascoceras cauvini (Chudeau) Vascoceras woodsi sp. nov. Vascoceras bullatum Schneegans Pseudovascoceras nigeriense (Woods) Vascoceras globosum costatum (Reyment) Vascoceras globosum globosum (Reyment) Vascoceras globosum globosum (Reyment) Vascoceras harttii (Hyatt) ?Fikaites varicostatus Zaborski Vascoceras globosum proprium (Reyment) Bulletin of The Natural History Museum Geology Series Earlier Geology Bulletins are still in print. The following can be ordered from Intercept (address on inside front cover). Where the complete backilist is not shown, this may also be obtained from the same address. Volume 33 No. | An account of the Ordovician rocks of the Shelve Inlier in west Salop and part of north Powys. W.F. Whittard, E.R.s. (Compiled by W.T. Dean). 1979. Pp. 1-69, 38 figs, frontispiece, coloured map, folded, in pocket. Map available separately No. 2 Miscellanea A new, possibly algal, microproblematicum from the Lower Carboniferous of England. G.F. Elliott, 8 Figs. Acanthopleurella Groom 1902: origin and life-habits of a miniature trilobite. R.A. Fortey & A.W.A. Rushton. 21 figs. Pleistocene bird remains from Tornewton Cave and the Brixham Windmill Cave in south Devon. C.J.O. Harrison. | fig. The succession of Hyracotherium (Perissodactyla, Mammalia) in the English early Eocene. J.J. Hooker, 6 figs. Salenia trisuranalis sp. nov. (Echinoidea) from the Eocene (London Clay) of Essex, and notes on its phylogeny. D.N. Lewis & R.P:S. Jefferies. 5 figs. Tertiary and Cretaceous brachiopods from Seymour, Cockburn and James Ross Islands, Antarctica. E.F. Owen. 33 figs. Revision of the rugose coral Diphyllum concinnum Lonsdale, 1845, and historical remarks on Murchison’s Russian coral collection. B.R. Rosen & R.F. Wise. 3 figs. Neuroptera (Insecta) in amber from the Lower Cretaceous of Lebanon. P-E.S. Whalley. 12 figs. 1980. Pp. 71-164. £12.00 No. 3 The Caradoc faunal associations of the area between Bala and Dinas Mawddwy, north Wales. M.G. Lockley. 1980. 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Revision of the microproblematicum Prethocoprolithus Elliott, 1962. G.F. Elliott. 4 figs. Basilicus tyrannus (Murchison) and the glabellar structure of asaphid trilobites. R.A. Fortey. 12 figs. A new Lower Ordovician bivalve family, the Thoraliidae (? Nuculoidea), interpreted as actinodont deposit feeders. N.J. Morris. 7 figs. Cretaceous brachiopods from northern Zululand. E.F. Owen. 13 figs. Tupus diluculum sp. nov. (Protodonata), a giant dragonfly from the Upper Carboniferous of Britain. P.E.S. Whalley. | fig. Revision of Plummerita Brénniman (Foraminiferida) and a new Maastrichtian species from Ecuador. J.E. Whittaker. 34 figs. 1980. Pp. 217-297. £11.00 Volume 35 No. 1 Lower Ordovician Brachiopoda from mid and south-west Wales. M.G. Lockley & A. Williams. 1981. Pp. 1-78, 263 figs, 3 tables. £10.80 No. 2 The fossil alga Girvanella Nicholson & Etheridge. H.M.C. Danielli. 1981. 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Hippoporidra edax (Busk 1859) and a revision of some fossil and living Hippoporidra (Bryozoa). P.D. Taylor & P.L. Cook. 6 figs. 1981. Pp. 109-252. £20.00 No. 4 The English Upper Jurassic Plesiosauroidea (reptilia) and a review of the phylogeny and classification of the Plesiosauria. D.S. Brown. 1981. Pp. 253-347, 44 figs. £13.00 Volume 36 No. 1 Middle Cambrian trilobites from the Sosink Formation, Derik-Mardin district, south-eastern Turkey. W.T. Dean. 1982. Pp. 1-41, 68 figs. £5.80 No. 2 Miscellanea British Dinantian (Lower Carboniferous) terebratulid brachiopods. C.H.C. Brunton. 20 figs. New microfossil records in time and space. G.F. Elliott. 6 figs. The Ordovician trilobite Neseuretus from Saudi Arabia, and the palaeogeography of the Neseuretus fauna related to Gondwanaland in the earlier Ordovician. R.A. Fortey & S.F. Morris. 10 figs. Archaeocidaris whatleyensis sp. nov. (Echinoidea) from the Carboniferous Limestone of Somerset and notes on echinoid phylogeny. D.N. Lewis & P.C. 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Pp. 303-332, 36 figs. £4.00 Volume 37 No. 1 Taxonomy of the arthrodire Phlyctaenius from the Lower or Middle Devonian of Campbellton, New Brunswick, Canada. V.T. Young. 1983. Pp. 1-35, 18 figs. £5.00 No. 2 Ailsacrinus gen. nov., an aberrant millericrinid from the Middle Jurassic of Britain. P.D. Taylor. 1983. Pp. 37-77, 48 figs, 1 table. £5.90 No. 3 Miscellanea Glossopteris anatolica Sp. nov. from uppermost Permian strata in south-east Turkey. S. Archangelsky & R.H. Wagner. 14 figs. The crocodilian Theriosuchus Owen, 1879 in the Wealden of England. E. Buffetaut. | fig. A new conifer species from the Wealden beds of Féron-Glageon, France. H.L. Fisher & J. Watson. 10 figs. Late Permian plants including Charophytes from the Khuff formation of Saudi Arabia. C.R. Hill & A.A. El-Khayal. 18 figs. British Carboniferous Edrioasteroidea (Echinodermata). A.B. Smith. 52 figs. A survey of recent and fossil Cicadas (Insecta, Hemiptera-Homoptera) in Britain. P.E.S. Whalley. 11 figs. The Cephalaspids from the Dittonian section at Cwm Mill, near Abergavenny, Gwent. E.I. White & H.A. Toombs. 20 figs. 1983. Pp. 79-171. £13.50 No. 4 The relationships of the palaeoniscid fishes, a review based on new specimens of Mimia and Moythomasia from the Upper Devonian of Western Australia. B.G. Gardiner. 1984. Pp. 173-428. 145 figs. 4 plates. 0 565 00967 2. £39.00 Volume 38 No. | New Tertiary pycnodonts from the Tilemsi valley, Republic of Mali. A.E. Longbottom. 1984. Pp. 1—26. 29 figs. 3 tables. 0 565 07000 2. £3.90 No. 2 Silicified brachiopods from the Viséan of County Fermanagh, Ireland. (111) Rhynchonellids. Spiriferids and Terebratulids. C.H.C. Brunton. 1984. Pp. 27-130. 213 figs. 0 565 07001 0. £16.20 No. 3 The Llandovery Series of the Type Area. L.R.M. Cocks. N.H. Woodcock, R.B. Rickards, J.T. Temple & P.D. Lane. 1984. Pp. 131-182. 70 figs. 0 565 07004 5S. £7.80 No. 4 Lower Ordovician Brachiopoda from the Tourmakeady Limestone, Co. Mayo, Ireland. A. Williams & G.B. Curry. 1985. Pp. 183-269. 214 figs. 0 565 07003 7. £14.50 No. 5 Miscellanea Growth and shell shape in Productacean Brachiopods. C.H.C. Brunton. Palaeosiphonium a problematic Jurassic alga. G.F. Elliott. Upper Ordovician brachiopods and trilobites from the Clashford House Formation, near Herbertstown, Co. Meath, Ireland. D.A.T. Harper, W.I. Mitchell, A.W. Owen & M. Romano. Preliminary description of Lower Devonian Osteostraci from Podolia (Ukrainian S.S.R.). P. Janvier. Hipparion sp. (Equidae, Perissodactyla) from Diavata (Thessaloniki, northern Greece). G.D. Koufos. } Preparation and further study of the Singa skull from | Sudan. C.B. Stringer, L. Cornish & P. Stuart-Macadam. Carboniferous and Permian species of the cyclostome bryozoan Corynotrypa Bassler, 1911. P.D. Taylor. Redescription of Eurycephalochelys, a trionychid turtle from — the Lower Eocene of England. C.A. Walker & R.T.J. Moody. Fossil insects from the Lithographic Limestone of Montsech (late Jurassic-early Cretaceous), Lérida Province, Spain. P.E.S. Whalley & E.A. Jarzembowski. 1985. Pp. 271-412, 162 figs. 0 565 07004 5. £24.00 | \ Volume 39 No. | Upper Cretaceous ammonites from the Calabar region, south-east Nigeria. P.M.P. Zaborski. 1985. Pp. 1-72. 66 figs. 0 565 07006 1. £11.00 No. 2 Cenomanian and Turonian ammonites from the Novo Redondo area, Angola. M.K. Howarth. 1985. Pp. 73-105. 33 figs. 0 565 07006 1. £5.60 No. 3 The systematics and palaeogeography of the Lower Jurassic insects of Dorset, England. P.E.S. Whalley. 1985. Pp. 107-189. 87 figs. 2 tables. 0 565 07008 8. £14.00 No.4 Mammals from the Bartonian (middle/late Eocene) of the Hampshire Basin, southern England. J.J. Hooker. 1986. Pp. 191-478. 71 figs. 39 tables. 0 565 07009 6. £49.50 Volume 40 No. | The Ordovician graptolites of the Shelve District, Shropshire. I. Strachan. 1986. Pp. 1-58. 38 figs. 0 565 07010 Xe Lo} 00 No. 2 The Cretaceous echinoid Boletechinus, with notes on the phylogeny of the Glyphocyphidae and Temnopleuridae. i D.N. Lewis. 1986. Pp. 59-90. 11 figs. 7 tables.056507011 8. £5. 60) No. 3 The trilobite fauna of the Raheen Formation (upper Caradoc), Co. Waterford, Ireland. A.W. Owen, R.P. Tripp & S.F. Morris. 1986. Pp. 91-122. 88 figs. 0 565 07012 6. £5.60, No. 4 Miscellanea I: Lower Turonian cirripede—Indian coleoid Naefia—Cretaceous—Recent Craniidae—Lectotypes of Girvan trilobites—Brachiopods from Provence—Lower Cretaceous cheilostomes. 1986. Pp. 125-222. 0 565 07013 4. £19. No. 5 Miscellanea II: New material of Kimmerosaurus—Edgehills Sandstone plants—Lithogeochemistry of Mendip rocks— ~ Specimens previously recorded as teuthids—Carboniferous ° lycopsid Anabathra—Meyenodendron, new Alaskian lepidodendrid. 1986. Pp. 225-297. 0 565 07014 2. £13. i Volume 41 No. 1 The Downtonian ostracoderm Sclerodus Agassiz (Osteostraci: Tremataspididae), P.L. Forey. 1987. Pp. 1-30. 11 figs. 0 565070150. £5.51 No. 2 Lower Turonian (Cretaceous) ammonites from south-east Nigeria. P.M.P. Zaborski. 1987. Pp. 31-66. 46 figs. 0 565 07016 9. £6. No. 3 The Arenig Series in South Wales: Stratigraphy and Palaeontology. I. The Arenig Series in South Wales. R.A. Fortey & R.M. Owens. II. Appendix. Acritarchs and Chitinozoa from the Arenig Series of South-west Wales. S.G. Molyneux. 1987. Pp. 67-364. 289 figs. 0 565 07017 Te £59, No. 4 Miocene geology and palaeontology of Ad Dabtiyah, Saudi Arabia. Compiled by P.J. Whybrow. 1987. Pp. 365-457. 54 — figs. 0 565 07019 3. £18: | No. 1 No. 4 | No. 2 | No.2 | No. 1 No. 2 Volume 42 No. 1 Cenomanian and Lower Turonian Echinoderms from Wilmington, south-east Devon. A.B. SMith, C.R.C. Paul, A.S. Gale & S.K. Donovan. 1988. 244 pp. 80 figs. 50 pls. 0 565 07018 5. £46.50 Volume 43 No. | A Global Analysis of the Ordovician-Silurian boundary. Edited by L.R.M. Cocks & R.B. Rickards. 1988. 394 pp., figs. 0 565 07020 7. £70.00 Volume 44 Miscellanea: Palaeocene wood from Mali—Chapelcorner fish bed—Heterotheca coprolites—Mesozoic Neuroptera and Raphidioptera. 1988. Pp. 1-63.0565070215. £12.00 Cenomanian brachiopods from the Lower Chalk of Britain and northern Europe. E.F. Owen. 1988. Pp. 65-175. 0565 07022 3. £21.00 The ammonite zonal sequence and ammonite taxonomy in the Douvilleiceras mammillatum Superzone (Lower Albian) in Europe. H.G. Owen. 1988. Pp. 177-231. 0 565 07023 i £10.30 Cassiopidae (Cretaceous Mesogastropoda): taxonomy and ecology. R.J. Cleevely & N.J. Morris. 1988. Pp. 233-291. 0565 07024 X. £11.00 No. 2 No. 3 Volume 45 No. 1 Arenig trilobites—Devonian brachiopods—Triassic demosponges—Larval shells of Jurassic bivalves—Carboniferous marattialean fern—Classification of Plectambonitacea. 1989. Pp. 1-163. 0 565 07025 8. £40.00 A review of the Tertiary non-marine molluscan faunas of the Pebasian and other inland basins of north-western South America. C.P. Nuttall. 1990. Pp. 165-371. 456 figs. 0 565 07026 6. £52.00 Volume 46 No. 1 Mid-Cretaceous Ammonites of Nigeria—new amphisbaenians from Kenya—English Wealden Equisetales—Faringdon Sponge Gravel Bryozoa. 1990. Pp. 1-152. 0 565 070274. £45.00 Carboniferous pteridosperm frond Neuropteris heterophylla—Tertiary Ostracoda from Tanzania: 1991. Pp. 153-270. 0565 07028 2. £30.00 Volume 47 Neogene crabs from Brunei, Sabah & Sarawak—New pseudosciurids from the English Late Eocene—Upper Palaeozoic Anomalodesmatan Bivalvia. 1991. 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Pp. 1-80. £37.50 Mobility and fixation of a variety of elements, in particular, during the metasomatic development of adinoles at Dinas Head, Cornwall—Productellid and Plicatiferid (Productoid) Brachiopods from the Lower Carboniferous of the Craven Reef Belt, North Yorkshire—The spores of Leclercqia and the dispersed spore morphon Acinosporites lindlarensis Riegel: a case of gradualistic evolution. 1993. Pp. 81-155. No. 2 £37.50 Volume 50 No. | Systematics of the melicerititid cyclostome bryozoans; introduction and the genera Elea, Semielea and Reptomultelea. 1994. Pp. 1-104. No. 2 The brachiopods of the Duncannon Group (Middle-Upper Ordovician) of southeast Ireland. 1994. Pp. 105-175. Volume 51 No. 1 A synopsis of neuropteroid foliage from the Carboniferous and Lower Permian of Europe—The Upper Cretaceous ammonite Pseudaspidoceras Hyatt, 1903, in north-eastern Nigeria—The pterodactyloids from the Purbeck Limestone Formation of Dorset. 1995. 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Pp. 89-305. £37.50 No. 2 51 61 Bulletin of The Natural History Museum . 3 GEOLOGY SERIES Vol. 52, No. 1, June 1996 CONTENTS Zirconlite: a review of localities worldwide, and a compilation of its chemical compositions C.T. Williams and R. Gieré A review of the stratigraphy of Eastern Paratethys (Oligocene—Holocene) R.W. Jones and M.D. Simmons A new protorichthofenioid brachiopod (Productida) from the Upper Carboniferous of the Urals, Russia C.H.C. Brunton The Upper Cretaceous ammonite Vascoceras Choffat, 1898 in north-eastern Nigeria P.M.P. Zaborski S186 A | Geology Series NZ —w NATU RAL HISTORY MUSEUM VOLUME 52 NUMBER2 28 NOVEMBER 1996 The Bulletin of The Natural History Museum (formerly: Bulletin of the British Museum (Natural History) ), instituted in 1949, is issued in four scientific series, Botany, Entomology, Geology (incorporating Mineralogy) and Zoology. The Geology Series is edited in the Museum’s Department of Palaeontology Keeper of Palaeontology: Dr L.R.M. Cocks Editor of Bulletin: Dr M.K. Howarth Assistant Editor: Mr C. Jones Papers in the Bulletin are primarily the results of research carried out on the unique and ever- growing collections of the Museum, both by the scientific staff and by specialists from elsewhere who make use of the Museum’s resources. Many of the papers are works of reference that will remain indispensable for years to come. All papers submitted for publication are subjected to external peer review for acceptance. A volume contains about 160 pages, made up by two numbers, published in the Spring and Autumn. Subscriptions may be placed for one or more of the series on an annual basis. Individual numbers and back numbers can be purchased and a Bulletin catalogue, by series, is available. Orders and enquiries should be sent to: Intercept Ltd. P.O. Box 716 Andover Hampshire SP10 1YG | Telephone: (01264) 334748 | Fax: (01264) 334058 Claims for non-receipt of issues of the Bulletin will be met free of charge if received by the Publisher within 6 | months for the UK, and 9 months for the rest of the world. World List abbreviation: Bull. nat. Hist. Mus. Lond. (Geol.) © The Natural History Museum, 1996 ISSN 0968-0462 Vol. 52, No. 2, pp. 91-173 The Natural History Museum Cromwell Road London SW7 5BD Issued 28 November 1996 Typeset by Ann Buchan (Typesetters), Middlesex Printed in Great Britain by Henry Ling Ltd., at the Dorset Press, Dorchester, Dorset | | | | - | | | Bull. nat. Hist. Mus. Lond. (Geol.) 52(2): 91-102 Issued 28 November 1996 Jurassic bryozoans from Baltéw, Holy Cross Mountains, Poland URSZULA HARA Panstwowy Instytut Geologiczny, Rakowiecka 4, 00-975 Warszawa, Poland i oo PAUL D. TAYLOR Department of Palaeontology, The Natural History Museum, Cromwell Road, London SW7 5BD 1 1 fy . \| ‘ay ~ j FALAEUH SYNOPSIS. Very few Jurassic bryozoans have been recorded from central or eastern Europe. This paper uses scanning electron microscopy to describe a well-preserved Polish Oxfordian bryozoan fauna consisting of five cyclostome species: Oncousoecia sp., Reptomultisparsa norberti sp. nov., Hyporosopora baltovensis sp. nov., Mecynoecia suprabajocina sp. nov. and Apsendesia cristata Lamouroux, 1821. Most abundant are specimens of H. baltovensis with tubular colonies which grew around perished cylindrical substrates. These arboreal colonies may have been epiphytes of algae or alternatively epizoans of sessile animals such as chaetopterid polychaetes, whose organic tubes can be similarly encrusted by bryozoans at the present-day. H. baltovensis is compared with other Jurassic species that have colonies with transversely ridged surfaces. [INTRODUCTION The overwhelming majority of Jurassic bryozoans described in the iterature are from western Europe, particularly England, France and Germany (see Walter, 1970). Knowledge of Jurassic bryozoans from *Isewhere in the world is extremely limited, therefore, and any aunas from outside western Europe are worthy of attention. This is »specially true if, as in the Polish fauna described here, the bryozoans ire well-preserved and include new species. The only previous paper concerning Jurassic bryozoans from oland is that of Reuss (1867) which described 18 species (not neluding Neuropora raristellata Reuss; Neuropora is now regarded is belonging to the ‘sclerosponges’, see Kazmierczak & Hillmer, 974) from the Upper Bathonian or Lower Callovian of Balin, near eracow. Although Reuss’s work is in need of revision, the Balin auna is clearly different from that described here from the Oxfordian if Baltow. A few other species of Upper Jurassic bryozoans have yeen mentioned from the vicinity of Cmiel6w and Sobk6w on the \orth-western margins of the Holy Cross Mts (Malinowska, 1965; *ugaczewska, 1970). _ Our purpose in this short paper is to describe the bryozoan fauna rom Baltéw, utilizing scanning electron microscopy (SEM), and to compare the species present with established species, especially fae the better known, diverse faunas of the western Europe Middle lurassic. faterial and methods. All studied specimens from Poland, which vere collected during 1986-88, are deposited in the palaeontologi- al collections of the Geological Museum of the Polish Geological astitute (Panstwowy Instytut Geologiczny) in Warszawa (MUZ ‘IG numbers). Comparative material from western Europe is in the ollections of The Natural History Museum (BMNH register num- ers). Specimens were studied and micrographs prepared using ISI OA and ABT-55 scanning electron microscopes equipped with 2nvironmental chambers’ for’ back-scattered electron imaging of jncaated material (see Taylor, 1986). The Natural History Museum, 1996 LOCALITY AND GEOLOGICAL SETTING The Upper Jurassic of Baltéw yields one of the most abundant and well-preserved cyclostome bryozoan faunas known from Poland. Baltow is situated approximately 15 km north-west of Ostrowiec Swietokrzyski on the north-eastern margin of the Holy Cross Mts (Fig. 1). The bryozoans come from coral-bearing deposits best exposed in the ravined part of the Kamienna River valley at Baltow. A rich shallow water biota occurs at Balt6w (Karczewski, 1960; Liszkowski, 1962, 1976; Roniewicz, 1966, 1968, 1976; Malinowska, 1967; Barczyk 1968, 1969, 1970; Roniewicz & Roniewicz, 1971; Hurcewicz, 1973; Brochwicz-Lewinski & R6zak, 1976; Maryanska & Kobylinska, 1980, 1984; Gutowski, 1992), including Foraminifera, bivalves, corals, brachiopods, sponges, ammonites, bryozoans, echinoids, crinoids, crabs, serpulid worms, shark teeth and algae (Rhodophycophyta of the genera Solenopora, Parachaetetes and Polygonella). A complex of coral-rich and other limestones is underlain by micritic, platy limestones containing: the bivalves Gryphaea, Isognomon, Nanogyra, Trigonia and Pholadomya; terebratulid and rhynchonellid brachiopods; ammonites; decapod claws; crinoids; and plant debris. The maximum total thickness reaches 100 metres (see Gutowski, 1992). The coralliferous lime- stone complex, approximately 15 metres thick, consists of unbedded or indistinctly bedded limestones overlain by bedded limestones (Roniewicz & Roniewicz, 1971). The former vary in thickness from 4 to 10 metres and comprise coral and pelitic limestones. According to Roniewicz & Roniewicz (1971), spaces between the foliaceous and submassive coral colonies are filled by pelitic or chalky lime- stones with a variable content of organic detritus. The bryozoans described here were collected from these weathered chalky lime- stones. Ammonites have not been found in the bryozoan-bearing beds. However, thick-bedded micritic limestones underlying the coralliferous limestone complex contain ammonites of the Gregoryceras transversarium Zone, and oncolitic limestones over- lying the complex contain ammonites of the Perispkinctes bifurcatus 92 HOLY CROSS MTS Kamienna Fig. 1 Location of Balté6w. A, map of the Holy Cross Mountains with the Upper Jurassic outcrop stippled (based on Roniewicz & Roniewicz, 1971). B, map of the Baltéw area with the outcrop of the coralliferous limestone complex containing the bryozoan fauna stippled (based on Liszkowski, 1976). Zone (J. Gutowski pers comm. to U.H., 1991). Therefore, the bryozoans belong to either the Transversarium or Bifurcatus Zone of the Middle Upper Oxfordian in the Submediterranean ammonite zonal scheme (Cariou et al., 1971; Gutowski, 1992). SYSTEMATIC DESCRIPTIONS Order CYCLOSTOMATA Busk, 1852 Suborder TUBULIPORINA Milne-Edwards, 1838 Family ONCOUSOECIIDAE Canu, 1918 Genus ONCOUSOECIA Canu, 1918 TYPESPECIES. Tubulipora lobulata Hincks, 1880 (=Alecto dilatans Johnston, 1847; see Hayward & Ryland, 1985), Recent. U. HARA AND P.D. TAYLOR nies composed of narrow ramifying branches in which the zooids are arranged multiserially. Gonozooids are small to moderately large, and ovoidal or pyriform in shape. They are not pierced by autozooidal REMARKS. Species assigned to Oncousoecia have encrusting colo- | apertures. Oncousoecia sp. Figs 2-3 9 MATERIAL. MUZ PIG 1601/11/8. / DESCRIPTION. Single colony comprising two coalescing branches detached from their original substrate. } Autozooids have frontal walls about 0.90-1.00 mm long by 0.25—_ 0.35 mm wide, slightly convex distally but more immersed proximally. Apertures circular, about 0.11—0.15 mm in diameter, with short preserved peristomes tapering markedly distally. Pseudopores large, closely-spaced, often distally pointed. Faint transverse wrinkles on autozooid frontal walls continuous across colony surface. Two gonozooids present, both asymmetrical as a result of distor- tion following branch coalescense (Fig. 2). Proximal frontal wall long and indistinguishable from that of an autozooid, raised strongly at its well-marked boundary with the dilated distal part of the frontal © wall. Distal frontal wall roughly pyriform in outline, 1.10 mm long by 1.00 mm wide, slightly inflated in height, relatively smooth- surfaced and possessing a higher density of pseudopores than the autozooids. Ooeciopore subterminal (i.e. within the area of the dilated frontal wall), transversely elliptical, 0.07 mm long by 0.15_ I mm wide, about the same size as an autozooidal aperture. Ooeciostome short, slightly reflexed, bearing very few pseudopores _ (Fig. 3). i eh ee REMARKS. Walter (1970), in his major revision of Jurassic j¥, cyclostomes, described five species of Oncousoecia. The species | \ from Balt6w most closely resembles O. elegantula (d’Orbigny, i 1850) from the Upper Bajocian of Port-en-Bessin, Normandy, France. I However, O. elegantula has slightly narrower autozooids, a differ- t ence which might be significant given that frontal wall width is one ~ a of the more useful morphometric characters for distinguishing )" between species of Jurassic cyclostomes. In view of the sparse — material available from Baltow and the need for SEM study of the” type specimens of O. elegantula and other Jurassic species of Oncousoecia, specific determination of the Baltow specimen is” deferred. ' Family MULTISPARSIDAE Bassler, 1935 (= MACROECIIDAE Canu, 1918) Genus REPTOMULTISPARSA @ Orbigny, 1853 TYPESPECIES. Diastopora incrustans d’ Orbigny, 1850, Bathonian. REMARKS. Nomenclatural problems concerning the type spec-’ ies of Reptomultisparsa, misidentified when selected by Gregory (1896b), were discussed by Taylor (1984) and resolved by ICZN) Opinion 1392 (1986) which designated Diastopora incrustans) d’Orbigny as the valid type species. An earlier concept of Reptomultisparsa encompassing almost all multilamellar tubuli-j porine species has now been superseded by a concept based on the) morphology of the gonozooids, which are longitudinally elongate and have large subterminal ooeciopores (see Taylor and Sequeiros 1982). Some, but not all species of Reptomultisparsa are multilamellar. JURASSIC BRYOZOANS FROM BALTOW, POLAND eptomultisparsa norberti sp. nov. Figs 4-8 OLOTYPE. MUZ PIG 1601/II/1 (Figs 6-7). ARATYPES. MUZ PIG 1601/II/2 (3 specimens). AME. In recognition of the contributions to bryozoology of the ustrian palaeontologist Dr Norbert Vavra. ESCRIPTION. Colony multiserial, sheet-like, unilamellar, either lanar (Fig. 4) or tubular (Fig. 5) in shape. Viewed from the growing dge the colony is thin, generally only one zooid in depth. Early rowth stages unknown. Original substrates not preserved. Autozooids immersed, zooidal boundaries indistinct, frontal walls lat for most of their length though slightly convex distally, about .10-1.50 mm long by 0.25-0.35 mm wide. Apertures widely- paced, circular or a little wider than long, about 0.12 mm in iameter, occasionally closed by terminal diaphragms located at bout the level of the frontal wall. Preserved peristomes moderately hort, tapering distally. Pseudopores longitudinally elongate, slit- ike when unworn (Fig. 8) but elliptical when worn. Gonozooids apparently infrequent, only a single example having een found (Fig. 6). Distal frontal wall almost flat, longitudinally longate, 1.50 mm long by 0.65 mm wide. Ooeciopore (Fig. 7) ubterminal, larger than an autozooidal aperture, transversely elon- ate, 0.10 mm long by 0.25 mm wide. EMARKS. Walter (1970) assigned seven Jurassic species to eptomultisparsa, and Taylor (1980) added one further new species. he autozooids in R. cobergonensis Walter, 1970, R.? margopuncta aagen, 1867), R. cricopora (Vine, 1881), R. oolitica (Vine, 1881) id R. tumida Taylor, 1980, have distinctly convex frontal walls, nlike the rather flat frontal walls of R. norberti. Reptomultisparsa crustans (d’Orbigny, 1850) differs from R. norberti in its much arger gonozooid, as well as its multilamellar colonies invariably ‘ncrusting gastropod shells once occupied by hermit crabs (Buge & ischer, 1970; Taylor, 1994). Reptomultisparsa ventricosa (Vine, 881), a species characteristic of the Aalenian and Bajocian of igs 2-3 Oncousoecia sp., MUZ PIG 1601/II/8, Oxfordian, Balt6w, Poland. Scanning electron micrographs of uncoated specimen. 2, distorted gonozooid and autozooids at coalescence of colony branches, x 45. 3, ooeciopore, x 167. England, is more similar to R. norberti except for its inflated gonozooids with smaller ooeciopores, and subcircular pseudopores. The unilamellar, often tubular colonies of R. norberti prompt comparison with Diastopora, notably the type species D. foliacea Lamouroux, 1821, from the Bathonian of Normandy. There are, however, striking differences in the form of the gonozooid, and in the morphology of the pseudopores as revealed by SEM. In D. foliacea, the gonozooid is transversely elongate and has lateral lobes which extend distally of the ooeciopore (Walter, 1970, pl. 8, fig. 1), whereas in R. norberti it is longitudinally elongate (Fig. 6). Pseudopores in frontal walls of D. foliacea zooids are gull-shaped (PDT unpublished), whereas those in R. norberti are long and slit- like (Fig. 8). These differences underscore the dual importance in cyclostome identification of specimens with gonozooids and of detailed SEM studies of pseudopore morphology. Without access to these two characters it is often difficult to make confident species determinations. Family PLAGIOECIIDAE Canu, 1918 (= DIASTOPORIDAE Gregory, 1899) Genus HYPOROSOPORA Canu & Bassler, 1929 TYPE SPECIES. Bathonian. Hyporosopora typica Canu & Bassler, 1929, REMARKS. Although Hyporosopora was considered to be a jun- ior synonym of Plagioecia Canu, 1918, by Walter (1970), differences exist between the two genera in the morphology of their gonozooids. Colonies of the extant type species of Plagioecia, Berenicea patina Lamouroux, 1816, have gonozooids with exceed- ingly broad, arcuate frontal walls which are profusely pierced by autozooidal apertures (see Hayward & Ryland, 1985). Hyporosopora gonozooids are considerably smaller, typically subtriangular in outline, and are not pierced by autozooidal aper- tures (see Taylor & Sequieros, 1982). 94 Figs 4-8 Reptomultisparsa norberti sp. nov., Oxfordian, Baltow, Poland. Scanning electron micrographs of uncoated specimens. 4, MUZ PIG 1601/11/72) (2), paratype, lamellar colony, x 14. 5, MUZ PIG 1601/II/2 (1), paratype, tubular colony, x 21. 6-7, MUZ PIG 1601/II/1, holotype; 6, autozooids and gonozooids of the fertile colony (ooeciopore just above calcite vein), x 15; 7, detail of ooeciopore, x 138. 8, MUZ PIG 1601/II/2 (1), paratype, slit-like pseudopores on autozooidal frontal wall, x 360. U. HARA AND P.D. TAYLOR | JURASSIC BRYOZOANS FROM BALTOW, POLAND 1] Figs 9-12 Hyporosopora baltovensis sp. nov., Oxfordian, Balt6w, Poland. Scanning electron micrographs of uncoated specimens. 9-10, MUZ PIG 1601/ II/13; 9, lamellar colony, x 18; 10, transverse ridges and autozooids, some with terminal diaphragms, x 64. 11-12, MUZ PIG 1601/II/3, holotype; 11, gonozooid, x 65; 12, ooeciopore (triangular hole beneath is a breakage in the gonozooid frontal wall), x 225. Hyporosopora baltovensis sp. nov. Figs 9-18 HOLOTYPE. MUZ PIG 1601/II/3 (Figs 11-12). Paratypes. MUZ PIG 1601/II/4 and 5, 13-18. AME. After Baltow, the type locality. DESCRIPTION. Colony multiserial, sheet-like, bereniciform, com- only unilamellar but occasionally multilamellar, either planar Fig. 9) or tubular (Figs 13-18) in shape. Tubular colonies possess a istinct suture formed by coalescence of lobes of the growing edge Qn opposite sides of the colony (Fig. 15). Distal fringe of basal amina protrudes beyond budding zone at growing edge (Fig. 14) where two or three generations of zooidal buds, some with mural ustules, are visible. Colony surface ornamented by discontinuous eeimerse ridges, irregularly-spaced 0.02—0.08 mm apart, of low orofile and sometimes indistinct, deflected proximally where they meet apertures (Fig. 10); ridges absent over dilated frontal walls of gonozooids (Fig. 11). Original substrates of encrustation not pre- served. Autozooids (Fig. 10) small, immersed, their frontal walls without well-defined boundaries, short, 0.25—0.50 mm in length by 0.11— 0.17 mm in width. Apertures small, usually longitudinally elongate but sometimes transversely elongate or circular (especially near the edge of colonies), 0.08-0.17 mm long by 0.06-0.14 mm wide, arranged roughly in quincunx, closely-spaced and often crowded close to the colony perimeter where they are larger in diameter. Terminal diaphragms sporadic in distribution, positioned level with or a little beneath the frontal wall (Fig. 10), and also observed occluding immature buds at the growing edge (Fig. 13). Preserved peristomes moderately long, tapering distally. Pseudopores approxi- mately circular in shape, widely spaced, absent on transverse ridges. Gonozooids (Fig. 11) subcircular to subtriangular in outline, wider than long, averaging 0.60 mm in length by 0.90 mm in width, 96 15 16 Figs 13-18 Hyporosopora baltovensis sp. nov., Oxfordian, Baltow, Poland. Scanning electron micrographs of uncoated, tubular specimens. 13, MUZ PIG{_ 1601/11/14, end view showing hollow cavity originally occupied by a cylindrical substrate, x 50. 14-15, MUZ PIG 1601/11/15; 14, wide distal fringe of basal lamina, x 38; 15, side view showing suture formed by anastomosis of opposite colony edges, x 23. 16, MUZ PIG 1601/11/16, x 28. 17, MUZ PIG 1601/11/17, x 20. 18, MUZ PIG 1601/11/18, x 12. inflated in height, lacking transverse ridges, indented by marginal autozooidal apertures. Floor of gonozooid pustulose, corrugated distally by the convexities of the underlying autozooids. Ooeciopore (Fig. 12) terminal, situated just beyond the inflated part of the gonozooid, small, transversely elongate, averaging 0.06 mm long by 0.09 mm wide, subcircular when the ooeciostome is preserved. U. HARA AND P.D. TAYLOR WZ. REMARKS. The main distinguishing feature of this new species ii) | the presence of transverse ridges crossing the colony surface. Trans JURASSIC BRYOZOANS FROM BALTOW, POLAND 97 igs 19-22 Jurassic bereniciform cyclostomes with transversely-ridged colonies similar to Hyporosopora baltovensis sp. nov. 19-20, Hyporosopora enstonensis (Pitt & Thomas, 1969), BMNH D51451, holotype, Bathonian, Hampen Marly Beds, Enstone, Oxfordshire, England; 19, x 13; 20, gonozooid, x 80. 21-22, Plagioecia rugosa (d’ Orbigny, 1853), BMNH BZ51, Lower Kimmeridgian, St Jean des Sables, near La Rochelle, Charente Maritime, France; 21, small colony with prominent transverse ridges, x 14; 22, gonozooid in a larger colony encrusting the same substrate, x 30. 981), H. enstonensis (Pitt & Thomas, 1969) from the Bathonian of xfordshire (Figs 19-20), and Mesenteripora undulata (Michelin, 845) from the Bathonian of Normandy (revised by Walter, 1970). A ew post-Jurassic cyclostomes also possess transversely ridged colo- ies (e.g. Plagioecia plicata (Canu) from the Eocene of France, see uge, 1979a; Berenicea undata Canu & Bassler, 1920 from Eocene f the USA), but this morphology seems to be proportionally less ommon than in the Jurassic. Compared with Hyporosopora baltovensis, the gonozooid in lagioecia rugosa is broader and is penetrated by autozooidal pertures (Fig. 22). In other respects, however, the two species are ery similar, although the transverse ridges tend to be more strongly eveloped in P. rugosa (Fig. 21). Walter (1970: 218) considered P. ugosa to be a junior synonym of Cellepora orbiculata Goldfuss, 826 from the Oxfordian of Streitburg in Germany. The syntypes Universitat Bonn, Goldfuss Collection 104) of C. orbiculata have been studied but are poorly-preserved, lack diagnostic gonozooids, and probably represent more than one species. C. orbiculata is probably better discarded. Hyporosopora portlandica has narrower gonozooids than H. baltovensis, and autozooidal apertures which are more widely- spaced and frequently transversely elongate. Multilamellar growth, rarely seen in H. baltovensis, is very common in H. portlandica, resulting from either spiral overgrowth or eruptive budding of subcolonies onto the colony surface. (Note that the considerable discrepancy in autozooidal size between the holotype of H. portlandica and many other specimens previously assigned to this species, for example by Taylor (1981), suggests that more than one species may be present). Gonozooid morphology is similar in H. enstonensis and H. baltovensis, but the former species has smaller autozooids and more prominent transverse ridges (Figs 19-20). M. undulata has consid- 98 erably larger autozooids, and some colonies apparently develop erect growth (Walter, 1970, pl.10, figs 3-8). In view of the paucity of potential characters for grouping Jurassic bereniciform cyclostome species into genera, consideration must be given to the possibility of using the presence of transverse ridges as a generic character. All of the transversely ridged species seem to be closely-related and are classified within the Family Plagioeciidae. However, they are currently distributed between two or three differ- ent genera. Practical problems are associated with the recognition of transverse ridges because, although the ridges are sharp and clearly- defined in some species (e.g. H. enstonensis), in others (e.g. H. baltovensis) they are gradational with irregular growth checks of the sort which can be found in a wide range of bereniciform cyclostomes. A more complete analysis of character distributions is recom- mended before any attempt is made to group transversely-ridged species into one genus. Genus MECYNOECIA Canu, 1918 TYPE SPECIES. Recent. Entalophora proboscidea Milne-Edwards, 1838, REMARKS. Although Canu (1918) named E. proboscidea Milne- Edwards, 1838, as the type species of Mecynoecia, Canu & Bassler (1922: 11) attempted to change the type species to Pustulopora delicatula Busk, 1875, stating: “The widespread and abundant spe- cies Entalophora proboscidea Milne-Edwards, 1838, was cited as the type of the genus by Canu in 1918, but we have changed the genotype for the reason that several species with different kinds of ovicells are undoubtedly included under this name and it is perhaps impossible at present to determine which one Milne-Edwards de- scribed’. This amendment is inadmissable under the International Rules of Zoological Nomenclature and therefore E. proboscidea stands as the valid type species of Mecynoecia (see also discussion in Buge 1979b; Walter 1987). The probable type specimen of E. proboscidea (Museum Nationale d’ Histoire Naturelle, Paris, Risso Collection 5110) has been exam- ined by PDT. Although its colony-form corresponds with the general usage of Mecynoecia (e.g. by Harmelin 1976), the specimen lacks gonozooids, thus making precise characterization difficult, a prob- lem considered beyond the scope of the current paper which accepts the generic concept as customarily applied. Mecynoecia suprabajocina sp. nov. Figs 23-26, 28 HOLotTyPeE. MUZ PIG 1601/II/11 (Figs 23-25). PARATYPES. MUZ PIG 1601/II/9 and 10. NAME. Indicating its similarity with M. bajocina (d’Orbigny, 1850) and the higher stratigraphical occurrence. DESCRIPTION. Colony erect, branches cylindrical (vinculariiform) and narrow (Fig. 23), 1.0—1.2 mm in diameter, ramifying dichoto- mously. Growth tips low cones in profile, with a radial, spoke-like arrangement of interzooidal walls, visible when viewed from above (Fig. 28). Zooidal budding apparently centred on branch axis. Pseudopores densely-packed and subcircular, absent from broad bands at the zooidal boundaries (Fig. 24). Autozooids with elongate frontal walls, about 1.0 mm long by 0.21—0.30 mm wide, slightly convex distally but sunken beneath the level of the zooidal boundary wall proximally. Apertures widely- spaced, circular or a little longitudinally elongate, about 0.12— 0.14 mm in diameter, sometimes closed by a terminal diaphragm U. HARA AND P.D. TAYLOR Fig. 23 Mecynoecia suprabajocina sp. nov., MUZ PIG 1601/I/11, holotype. Oxfordian, Baltow, Poland. Scanning electron micrographs of uncoated branch with a broken gonozooid close to the distal end, x 21. | | | | . located either atop a short peristome or inclined and positioned proximally to the peristomial rim. Gonozooids (Fig. 26) with globular distal dilated frontal wall, subcircular or transversely elliptical in outline and well inflated. Ooeciopore (Fig. 26) located terminally, more or less semicircular, distal edge markedly convex relative to the almost straight proximal edge, wider than long, about 0.10 by 0.17 mm. Preserved ooeciostomes short. REMARKS. This new species resembles Mecynoecia bajocina from the Upper Bajocian White Sponge Oolite of the Port-en-Bessin area of Normandy, and the contemporaneous Microzoa Beds (a facies : y the Burton Limestone) of Shipton Gorge, Dorset. However, it differs from M. bajocina in the structure of the ooeciopore which is almost semicircular (Fig. 25) compared to the ooeciopore of M. bajocina which is very strongly compressed medially (Fig. 27). In addition, the zooidal boundary areas devoid of pseudopores are substantially wider in M. suprabajocina than in M. bajocina. Genetic studies of living ctenostome and cheilostome bryozoans have shown clearly’ that subtle morphological differences signify different species (€.g.. | i EES OS See ee, Thorpe & Ryland, 1979; Jackson & Cheetham, 1990). Although comparable studies have yet to be made on Recent cyclostomes, it 1s considered reasonable to favour the taxonomic splitting of cyclostomes on the basis of small but consistent differences in skeletal morphology (e.g. McKinney & Jackson, 1989). | il ‘JURASSIC BRYOZOANS FROM BALTOW, POLAND Family THEONOIDAE Busk, 1859 Genus APSENDESIA Lamouroux, 1821 TYPE SPECIES. Apsendesia cristata Lamouroux, 1821, Bathonian, Normandy. REMARKS. Only two species of this genus are recognized: the Jurassic type species and A. neocomiensis d’Orbigny from the | Hauterivian. 99 Figs 24-27 Jurassic Mecynoecia; scanning electron micrographs of uncoated specimens. 24-26, Mecynoecia suprabajocina sp. noy., Oxfordian, Baltéw, Poland; 24-25, MUZ PIG 1601/II/11, holotype; 24, autozooids with broad areas devoid of pseudopores at zooidal boundaries, x 85; 25, ooeciopore, x 200; 26, MUZ PIG 1601/11/10, paratype, gonozooid with intact frontal wall, x 65. 27, Mecynoecia bajocina (d’ Orbigny, 1850), BMNH D59492, Upper Bajocian, Shipton Gorge, Dorset, England, gonozooid with typically compressed ooeciopore, x 55. Apsendesia cristata Lamouroux, 1821 Figs 29-32 1821 Pelagia clypeata Lamouroux: 78, pl. 79, figs S—7. 1821 Apsendesia cristata Lamouroux: 82, pl. 80, figs 12-14. 1854 Apsendesia cristata Lamouroux; Haime: 201, pl. 7, fig. 6 a-k. 1854 Apsendesia clypeata (Lamouroux); Haime: 202, pl. 7, fig. 7 a-d. 1896c Apsendesia cristata Lamouroux; Gregory: 167, pl. 9, figs 4-5. Fig. 28 Mecynoecia suprabajocina sp. nov., MUZ PIG 1601/11/9. Oxfordian, Baltow, Poland. Scanning electron micrograph of branch growing tip showing spoke-like arrangement of zooecial walls, x 85. 1953 21967 Apsendesia cristata Lamouroux; Bassler: 56, fig. 23,4. Apsendesia cristata Lamouroux; Walter: 46, pl. 10, figs 1— Dp Apsendesia cristata Lamouroux; Walter: 202, pl. 20, figs 6— 11. 1970 MATERIAL. MUZ PIG 1601/II/6-7. DESCRIPTION. Colony erect, fungiform, a narrow stalk, about 0.8 mm long, supporting an expanded, cup-shaped head, subcircular in plan view and averaging 4 mm in diameter (Figs 29-30). Autozooids grouped into fascicles which open around the circumference of the head and increase in number through bifurcation. Frontal side of head marked radially by ridge-like fascicles and convex frontal walls of autozooids. Underside of head an exterior wall with pseudopores and concentric growth lines, sometimes giving rise to downward- growing processes or struts each composed of about 3-6 zooids (Fig. Si): Autozooids long, lacking frontal walls and with polygonal aper- tures when situated in the centres of a fascicle, but possessing pseudoporous frontal walls and apertures with a curved external edge when situated at the border of a fascicle. Apertures about 0.15 mm in diameter. Gonozooids not observed in specimens from Baltow (see below). REMARKS. This is one of the most distinctive of all Jurassic bryozoan species. At Baltéw only small colonies, resembling speci- mens from the French Bathonian described as Pelagia clypeata by Lamouroux (1821), have been found. Large colonies depart from a simple cup-shape and become complexly corrugated, like the speci- men described as Apsendesia cristata by Lamouroux (1821). The finding of A. cristata at Baltow in the Oxfordian extends its range upward from the Lower Callovian (Walter, 1970: 204). An apparent occurrence in the Upper Bajocian of Shipton Gorge (Walter, 1967) is questionable: specimens from Shipton Gorge are extremely small, consisting of little more than a stalk and lacking the character- istic head and fascicles. The peculiar gonozooid of A. cristata, not found in specimens from Balt6w, is illustrated here using SEM for the first time (Fig. 32). It develops within a cleft in a fascicle and has a small ooeciopore U. HARA AND P.D. TAYLOR situated in the centre of a bulbous frontal wall. Another unusual feature of A. cristata are the processes which may develop from the undersides of colonies (Fig. 31). These are multizooidal, originate at the growing edge (i.e. not by resorption of the exterior wall in more proximal sites), and may extend down to the substratum to form secondary supports for the colony. Voigt (1993) has described similar structures from a variety of other cyclostomes and cheilostomes. The Balt6w bryozoan fauna consists entirely of small, delicate colonies; there is a total absence of the larger, more robust cyclostomes, notably cerioporines, which characterize many other Jurassic bryofaunas. In this respect, the fauna resembles that found in the Upper Bajocian of Shipton Gorge (Walford, 1889, 1894; Walter, 1967), although known species diversity at Shipton Gorge is considerably greater (ca 19 spp., vs. 5 spp. at Baltow). Other — similarities between the two faunas include the abundance of bereniciform colonies with narrow axial canals, and the shared | presence of closely-related species of Mecynoecia (and possibly also Apsendesia cristata, but see note above). Walter (1967) inter- preted Shipton Gorge as a low energy, shallow water environmentin | which many of the bryozoans grew attached to thin algal filaments. A similar environmental interpretation probably also applies to Baltéw, as suggested by the bryozoans (see Maryanska & Koblinska, 1980, 1984), corals (see Roniewicz & Roniewicz, 1971) and brachiopods (e.g. Barezyk, 1968, 1969, 1970). The presence of calcareous algae certainly implies deposition within the photic zone. Most specimens of H. baltovensis from Balt6w have tubular colonies with narrow axial canals (Figs 13-18). This colony-form is not, however, a specific character of H. baltovensis as some colonies are flat (Fig. 9). The presence of axial canals is clearly a result of encrustation of cylindrical substrates which were soft bodied and did not fossilize. Indeed, axial canals identical in size to those of H. | baltovensis can be found in sponges from Balt6w, which presum- ably lived attached to the same substrates as the bryozoans. Occasionally, the axial canals bifurcate, indicating that the organ- isms concerned had Y-shaped branches. Unfortunately, no informative |) bioimmurations of the surface details of the perished substrates are |) present on the basal laminae of the bryozoans. Therefore, the |} identity of the substrate is equivocal. Recent bryozoans can grow around a variety of cylindrical substrates of both plant and animal | _ origin. For example, Alvarez (1992: fig. 15) illustrated small colo- nies of the Recent cyclostome Disporella sp. growing around) un-named cylindrical substrates and leaving axial canals very simi- lar to those found in the Balt6w specimens. Bone & James (1993: ) fig. 7b) figured several different species of bryozoans attached to the cylindrical stems of sea-grasses from shallow water environments of |} . the Lacepede Shelf, southern Australia. Larger diameter axial canals)| _ result from growth of the cheilostomeSchizoporella floridana around) rhizomes of seagrass (see McKinney & Jackson 1989: fig. 7.12). Among living animals, the polychaete Phyllochaetopterus sociali living at depths beneath the photic zone on the Otago Shelf of New Zealand constructs long horny tubes about 1 mm in diameter. These tubes are fouled by a diversity of bryozoans, including encrustin cheilostomes and cyclostomes, as well as erect colonies of th cyclostomes Telopora and Hornera (P.D.T. unpublished). Colonies wrap around the circumference of the polychaete tubes. Tube decay would leave an axial canal in the centre of the colony. Some of the tubes divide, possibly as a result of asexual fission of the worms} | | | | | PALAEOECOLOGY | | io i JURASSIC BRYOZOANS FROM BALTOW, POLAND 101 32 PIG 1601/11/6, colony upper surface, x 15; 30, MUZ PIG 1601/II/7, colony underside, x 15. 31-32, BMNH D2327 (1), Bathonian, Ranville, Normandy, France; 31, struts on the underside of a colony, x 15; 32, gonozooid between fascicles of autozooidal apertures, x 35. hes 29-32 Apsendesia cristata Lamouroux, 1821. Scanning electron micrographs of uncoated specimens. 29-30, Oxfordian, Baltéw, Poland; 29, MUZ | herefore, caution should be exercised when inferring that fossil ryozoans with axial tubes grew attached to plants and therefore ndicate palaeoenvironments within the photic zone. CKNOWLEDGEMENTS. The British Council is thanked for providing U.H. ith the opportunity to visit and work at the NHM in London. Ivarez, J. A. 1992. 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Warszawa. { / | t Roniewicz, E. 1966. Les Madréporaires du Jurassique supérieur de la bordure des monts de Sainte-Croix, Pologne. Acta Palaeontologica Polonica, Warszawa, 11: 157-256. 1968. Actinaraeopsis, un nouveau genre de Madreporaire Jurassique de Pologne. Acta Palaeontologica Polonica, Warszawa, 13: 305-308. 1976. Baltéw — litologia i ekologia rafy koralowej i koralowo-glonowej oksfordu srodkowego; profil na Zarzeczu. In Liszkowski, J., Problem IB — Rozw6j litofacjalny i paleogeograficzny jury gornej pdlnocno-wschodniej czesci mezozoicznego obrzezenia Gor Swietokrzyskich. Materialy konferencji terenowych. Przewodnik XLVIII Zjazdu PTG Starachowice, Warszawa: 121-127. —— & Roniewicz, P. 1971. Upper Jurassic coral assemblages of the Central Polish Uplands. Acta Geologica Polonica, Warszawa, 21: 399-423. Taylor, P. D. 1980. Two new Jurassic Bryozoa from southern England. Palaeontology, London, 23: 699-706. 1981. Bryozoa from the Jurassic Portland Beds of England. Palaeontology, London, 24: 863-875. 1984. Reptomultisparsa d Orbigny, 1853 (Bryozoa, Cyclostomata): request for the designation of a type species. Z.N. (S.) 2400. Bulletin of Zoological Nomencla- ture, London, 41: 77-79. 1986. Scanning electron microscopy of uncoated fossils. Palaeontology, London, 29: 685-690. 1994. Evolutionary palaeoecology of symbioses between bryozoans and hermit crabs. Historical Biology, 9: 157-205. & Sequeiros, L. 1982. Toarcian bryozoans from Belchite in north-east Spain. Bulletin of the British Museum (Natural History),(Geology Series), London, 36: 117-129. Thorpe, J. P. & Ryland, J. S. 1979. Cryptic speciation detected by biochemical genetics in three ecologically important intertidal bryozoans. Estuarine and Coastal Marine Science, London, 8: 395-398. Vine, G. R. 1881. Further notes on the Family Diastoporidae Busk. Species from the Lias and Oolite. Quarterly Journal of the Geological Society, London, 37: 381-390. Voigt, E. 1993. Stiitz-, Anker- und Haftorgane bei rezenten und fossilen Bryozoen (Cyclostomata und Cheilostomata). Verhandlungen des Naturwissenschaftlichen Vereins in Hamburg, (N.F.) 33 [for 1992]: 121-130. Waagen, W. 1867. Uber die zone des Ammonites sowerbyii. Geognostisch- ' paldontologische Beitrége von Dr. E. W. Benecke, Munich, 1: 507-668, pls XXI-XXXIV. Walford, E. A. 1889. On some Bryozoa from the Inferior Oolite of Shipton Gorge, Dorset. Part I. Quarterly Journal of the Geological Society, London, 45: 561-574. 1894. On some Bryozoa from the Inferior Oolite of Shipton Gorge, Dorset. Part Il. Quarterly Journal of the Geological Society, London, 50: 72-78. Walter, B. 1967. Révision de la faune de Bryozoaires du Bajocien supérieur de Shipton Gorge (Dorset, Grande-Bretagne). Travaux des Laboratoires de Géologie de la Faculté des Sciences de Lyon, Lyon, (n.s.) 14: 43-52. 1970. Les Bryozoaires Jurassiques en France. Documents des Laboratoires de Géologie de la Faculté des Sciences de Lyon, Lyon, 35 (for 1969): 1-328. 1987. Les Bryozoaires Cyclostomes Neocomiens de forme ‘Entalophora et ‘Spiropora’. Revue de Paléobiologie, Geneva, 6: 29-53. EO ————SE s SS “ = ————Ere : | ) | | | | | | Bull. nat. Hist. Mus. Lond. (Geol.) 52(2): 103-107 Issued 28 November 1996 A new deep-water spatangoid echinoid from the Cretaceous of British Columbia, Canada ANDREW B. SMITH Department of Palaeontology, The Natural History Museum, Cromwell Road, London SW7 SBD, U.K. ALAN McGUGAN 1157 Rolmar CR RR 2, Cobble Hill, Vancouver Island, British Columbia, VOR 1L0, Canada. Synopsis. A new species of spatangoid echinoid, Plesiaster vancouverensis, is described from continental slope debris flow deposits of latest Santonian to early Campanian age on Vancouver Island, British Columbia. INTRODUCTION | The fossil record of echinoids is overwhelmingly dominated by species of the continental shelf. Although some of these lived in ‘elatively deep water settings, such as the faunas of the North-West Suropean Upper Chalk facies (Smith & Wright 1989, 1990, 1993), or he shelf marginal faunas of Spain (Neraudeau & Floquet 1991) or Tunisia (Zhagbib-Turki 1989), records of continental slope and basin ‘aunas remain extremely rare. Bather (1934) described the holasteroid Chelonechinus from the Miocene deep-water Suva Formation of Java, and more recently adiverse bathy] echinoid fauna has been discovered nthe early Miocene Morozaki Group of Japan (Mizuno 1991) and the Middle Miocene Tatsukuroiso Mudstone of NE Honshu, Japan ‘Kikuchi & Nikaido 1985). Pliocene deposits of California have also yielded what appear to be bathy] echinoids (Woodring 1938). Here we jescribe a new spatangoid from late Cretaceous outer shelf to upper sontinental slope deposits of western Canada. | The new species described here comes from a single horizon within the Upper Cretaceous Nanaimo Group. The Nanaimo Group ‘epresents a major sedimentary sequence within the Georgia Basin pf south-western British Columbia and was deposited in a fore-arc Jasin along the western margin of the Canadian continental margin |England 1989). Depositional environments represented within this sroup range from alluvial to continental slope facies, with the echinoids coming from within a series of debris flows interpreted as ipper continental slope facies. | The echinoid horizon was discovered in May 1993 by Dr A. McGugan and Dr. T. England during reconnaisance of the French creek area. The echinoids were found in a stream section adjoining dildegard Farm on French Creek, ca. 1 km upstream from Highway 4 bridge at Coombs, which crosses French Creek approximately seven km due south of Qualicum Beach, Vancouver Island, British Columbia (Fig. 1). The succession at this locality begins with ca. 6 n of bedded shales, siltstones and mudstones of the Haslam Forma- ion (= lower part of the Trent River Formation of England, 1989; sig. 2). These are overlain by a chaotic conglomeratic debris flow ‘McGugan, 1992). The matrix is a silty mudstone and included debbles are a mixture of undated meta-volcanics, argillites, Creta- 2e0us calcareous concretions (some with plant remains), sandstone ind siltstone clasts, and many clasts of bedded shale, some up to 4m n length and showing plastic deformation and slump-roll-type eading edges. The fauna of these beds include foraminifera, ammo- nites and inoceramid and other bivalves. The echinoids, all of which belong to the same species, came from 2 The Natural History Museum, 1996 a single bedding plane near the top of the bedded shale series. They are preserved in life orientation and some at least retain associated spines. These echinoids were thus not transported, but lived and died within the environment of deposition represented by the shales. Since the bedded shales are overlain by debris flows, the environ- ment of deposition is taken to be upper continental slope. The dating of the echinoid level is based on the benthic foraminifera, which are indicative of the Inoceramus (Spheno- ceramus) schmidti Zone, latest Santonian to Lower Campanian. The associated macrofauna also support a late Santonian — early Campanian age. A molluscan fauna (Table 1) is associated with the echinoids and in the beds immediately overlying, and has been identified by Dr. J.W. Haggart (Geological Survey of Canada, Vancouver). This includes /. (S.) schmidti, which is known to range from the latest Santonian to early Campanian in British Columbia and Northern California (Haggart, 1984). Table 1. Fossil molluscs found in association with Plesiaster vancouverensis sp. noy. in the Haslam Formation of French Creek, Vancouver Island. Identifications by J.W. Haggart (Canadian Geological Survey, Vancouver). Bivalves: Inoceramus (Sphenoceramus) schmidti Michael I. ex. gr. subundatus Meek Acila (Truncacila) demessa Finlay Ammonite: Canadoceras yokoyamai (Jimbo) SYSTEMATIC DESCRIPTION Class ECHINOIDEA Leske, 1778 Order SPATANGOIDA Claus, 1876 Family MICRASTERIDAE Lambert, 1920 Genus PLESIASTER Pomel, 1883 TYPE SPECIES. Micraster peini Coquand, 1862. OTHER SPECIES INCLUDED. P. cotteaui Gauthier. OCCURRENCE. Late Coniacian to early Campanian of North Af- rica and North America. 104 Area of enlargement Qualicum Beach 3 Ss Parksville Fossil echinoid locality Fig. 1. Map showing the location of the echinoid bed, with insert showing its position on the island of Vancouver. Plesiaster vancouverensis sp. nov. Text-figs 3—5 Types. Holotype, BMNH EE 5078 (Figs 3A, 4A, 4B, 5A); paratypes, BMNH EE5076 (Fig. 3C), EES5077, EES079-83 (Figs 3B, 3D, 3E, 4C-F, 5B). OCCURRENCE. Shales towards the top of the Haslam Formation (= lower part of the Trent River Formation), /. (S.) schmidti Zone, uppermost Santonian to lowermost Campanian, exposed in river bank of French Creek behind Hildegard Farm, ca. 1 km upstream from the highway 4 bridge over French Creek at Coombs, and 7 km due south of Qualicum Beach, south-eastern Vancouver Island, British Columbia, Canada. DIAGNOSIS. A species of Plesiaster with a well-developed peripetalous fasciole around the posterior part of the test, a broad and strongly petaloid anterior ambulacrum, and with lateral and poste- rior petals broad and open, extending most of the distance to the Pachydiscus suciaensis Campanian Bostrychoceras elongatum Santonian England 1989). Zone and Subzone ree tao Se Maastrichtian Nostoceras hornbyense Metaplacenticeras cf. pacificum Hoplitoplacenticeras vancouverense Inoceramus schmidti Haslam Pachydiscus haradai Inoceramus naumanni Fig. 2. Stratigraphic column for the Upper Cretaceous Nanaimo Group showing the level of the echinoid bed (*) (taken from Muller & Jeletzky 1970, and | A.B. SMITH AND A. McGUGAN ambitus. The contact between the labral plate and sternal plates is strongly offset towards the left-hand side. DESCRIPTION. All tests are crushed so accurate dimensions cannot be given. The largest and best-preserved specimen is approximately 63 mm in length and 58 mm in width. Test height and shape in profile are unknown, but the test appears to have been relatively low and gently domed. The test is oviform in outline with the posterior slightly truncated. The widest point on the test lies a little behind the anterior petals at approximately mid-length. The apical disc is ethmophract with all four genital plates similar in size and each bearing a gonopore (Fig. 3B). The posterior genital plates are not separated by the madreporite. The apical disc lies anterior of centre, 39% of test length from the anterior border. All five ambulacra are strongly petaloid adapically. Petals are broad and sunken and remain open distally. The anterior petal extends 80% of the distance to the ambitus. It shallows and more or less disappears towards the ambitus so that the anterior is hardly , notched. There are ca. 45 pore-pairs in a column at 63 mm test length. The pore-pairs are conjugate, with individual pores widely separated (Fig. 3A). The perradial zone is smooth and approxi- mately as broad as a single pore-zone (Fig. 5B). The lateral petals : extend 90% of the distance to the ambitus and are strongly petaloid, — tapering slightly towards the ambitus. They diverge from one an- | | other at an angle of 135°. There are ca. 55 pore-pairs in a column at ———— LS 63 mm test length. Individual pores are strongly elongate and conjugate, with successive pore-pairs separated by single rows of miliary tubercles. The posterior petals are similar, but extend only 75% of the distance to the ambitus. They diverge from one another) | at an angle of 50°. Pores below the petals are single. Around the peristome there are three to five enlarged phyllode pores. Interambulacra are slightly raised adapically but do not form sharp keels. They remain biserial to the apex. On the oral surface the: primibasal plates in interambulacra 1 and 4 are occluded from the: peristome border by proximal ambulacral plates (Fig. 3B). The labral plate is elongate and narrow, more than twice as long as broad. The sternal plates are clearly differentiated, but unequal in size; that towards ambulacrum I being smaller and either just in contact with the labral plate (Fig. 3E) or separated from it (Fig. 3D). The peristome is small and D-shaped; the labral plate does not project over the mouth and there is no distinct lip. The front of the peristome lies 19% of the test length from the anterior border. The periproct is crushed in all specimens. It lies on the posterior surface: and is probably hidden in both apical and oral views. Formation Cedar District Extension Protection Comox H ECHINOID FROM THE CRETACEOUS OF BRITISH COLOMBIA | } Tuberculation is relatively coarse on the aboral surface, with ;cattered primary tubercles (crenulate and perforate) up to 0.6 mm liameter set amongst a fine granulation of miliary tuberculation Fig. 5B). A well-developed peripetalous fasciole, with up to 12 ines of densely spaced (hexagonally packed) miliary tubercles, is leveloped around at least the posterior of the test. It runs immedi- tely beneath the ends of the petals and is indented in the osterio-lateral interambulacra. Unfortunately, most specimens are reserved as internal moulds so it is impossible to tell whether the asciole continues around the anterior of the test. It is also impossible 0 tell whether there was a subanal fasciole present. Some specimens have scattered primary spines preserved flat- ened against the test surface. XEMARKS. The new species is placed in the genus Plesiaster on ccount of its peripetalous fasciole and petaloid anterior ambulacrum. nly a few spatangoid genera have petaloid anterior ambulacra, amely Douvillaster Lambert, 1917, Plesiaster Pomel 1883, Hetero- ampas Cotteau 1862, Isomicraster Lambert 1901 and Barnumia 105 \s 3. Camera lucida drawings of Plesiaster vancouverensis sp. nov. uppermost Santonian; Lower Campanian, / (S.) schmidti Zone, French Creek, Vancouver Island, British Columbia. A, BMNH EE5078; B, BMNH EE5083; C, BMNH EE5076, D, BMNH EE5079: E, BMNH EE5081. Cooke 1953. Neither Douvillaster nor Isomicraster have fascioles and thus differ significantly from the Canadian species. Barnumia can also be dismissed from consideration since its fasciole is not peripetalous but lateral, extending around the ambitus and well- separated from the base of the petals. The type species of Heterolampas, H.maresi Cotteau, has a narrow but continuous peripetalous fasciole and petaloid ambulacra. However, it differs significantly from the Canadian species in its labral and apical disc structure. The apical disc of H. maresi, though ethmophract, is composed of an enlarged madreporite plate which occupies the centre of the disc and separates the posterior two genital plates. Its labral plate is large and broadly triangular, firmly abutting both sternal plates. Finally, the peristome of H. maresi lies considerably further back from the anterior than that of the new species. Plesiaster Pomel, 1883, was established for Micraster-like forms with a partial peripetalous fasciole developed at the base of the petals. Zhagbib-Turki (1987) has given a recent analyses of the North African species and concluded that Plesiaster should be placed in synonymy with Micraster. However, the differentiation of . SMITH AND A. McGUGAN } -ECHINOID FROM THE CRETACEOUS OF BRITISH COLOMBIA 107 Fig. 5. Plesiaster vancouverensis sp. nov. A, BMNH EE5076, adapical view of test showing tuberculation style, x1. B, BMNH EES083, latex cast of external mould showing apical disc and tuberculation style, x2. a partial peripetalous fasciole represents an apomorphy for the genus and the distinction is maintained here. The European species that have been assigned to this genus in the past, P. bucardium Goldfuss, P. coravium Schliiter, P. parvistella Schliiter, and P. minor Schliiter (e.g. Ernst 1972), differ significantly from the North African type, P. peini Coquand. The European species all have short petals, non- petaloid anterior ambulacra, and complete peripetalous and subanal fascioles. They are placed in the genus Diplodetus Schliiter 1900. The two North African species, P. peini and P. cotteaui have much longer petals and incomplete fascioles. The Canadian species comes close to ‘Micraster’ americanus Stephenson in form, but differs from that species in having a much more strongly petaloid anterior ambulacrum. In M. americanus the dores in the anterior ambulacrum are small and separated by a raised nterporal granule, whereas those pores in P. vancouverensis are “ater and widely separated from one another in each pair. “urthermore, the petals of P. vancouverensis are larger, more open and extend closer to the ambitus than they do in M. americanus. F. vancouverensis differs from the North African species of Plesiaster in having a well-developed peripetalous fasciole around {he posterior of the test. P peini and P. cotteaui both have very mpersistent fascioles that are really only present at the base of the detals and do not form a continuous band around the posterior. REFERENCES Sather, F.A. 1934. Chelonechinus n.g., a Neogene urechinid. Bulletin of the Geological Society of America, 45: 799-876. Sooke, W.B. 1953. American Upper Cretaceous Echinoidea. United States Geological Survey Professional Paper, 254—A: 44pp. 16 pls. Ungland, T.D.J. 1989. Lithostratigraphy of the Nanaimo Group, Georgia Basin, southwestern British Columbia. Geological Survey of Canada Paper, 89-1E: 197- 206. Ernst, G. 1972. Grundfragen der Stammesgeschichte bei irregularen Echiniden der nordwesteuropaischen Oberkreide. Geologishes Jahrbuch, A4: 63-175. Haggart, J.W. 1984. Upper Cretaceous (Santonian-Campanian) ammonite and inoceramid biostratigraphy of the Chico Formation, California. Cretaceous Re- search, 5: 225-241. Kikuchi, Y. & Nikaido, A. 1985. The first occurrence of abyssal echinoid Pourtalesia from the Middle Miocene Tatsukuroiso Mudstone in Ibaraki Prefecture, northeastern Honshu, Japan. Annual Report of the Institute of Geosciences, the University of Tsukuba, 11: 32-34. McGugan, A. 1992. Cretaceous submarine debris flow outcrops. Geological Society of America Today, 2: 57 only. Mizuno, Y. 1991. Fossil echinoderms from the early Miocene Morozaki Group in the Chita peninsula, central Japan. /n Yanagisawa, T., Yasumasu, I, Oguro, C, Suzuki, N. & Motokawa, T. (eds), Biology of Echinodermata: Proceedings of the seventh international echinoderm conference, Atami/9-14 September, 1990, p. 582 only. A.A. Balkema, Rotterdam. Muller, J.E. & Jeletzky, J.A. 1970. Geology of the Upper Cretaceous Nanaimo Group, Vancouver Island and Gulf Islands, British Columbia. Geological Survey of Canada Paper, 69-25: 1-77. Neraudeau, D. & Floquet, M. 1991. Les échinides Hemiasteridae: marqueurs ecologiques de la plate-forme castillane et navarro-cantabre (Espagne) au Cretacé superieur. Palaeogeography, Palaeoclimatology, Palaeoecology, 88: 265-281. Pomel, A. 1883. Classification methodique et genera des echinides vivants et fossiles. Alger, Paris, 120 pp. Smith, A.B. & Wright, C.W. 1989. British Cretaceous echinoids. Part I, General introduction and Cidaroidea. Monographs of the Palaeontographical Society, Lon- don:\—101, pls 1-32 (publication number 578, part of volume 141 for 1987). & 1990. British Cretaceous echinoids. Part 2, Echinothurioida, Diadematoida and Stirodonta (1, Calycina). Monographs of the Palaeontographical Society, Lon- don: 101-198, pls 33-72 (publication number 583, part of volume 143 for 1990). — & 1993. British Cretaceous echinoids. Part 3, Stirodonta, part 2 (Hemicidaroida, Arbacioida and Phymosomatoida, part 1). Monographs of the Palaeontographical Society, London: 199-267, pls 73-92 (publication number 593, part of volume 146 for 1993). Woodring, W.P. 1938. Lower Pliocene Mollusks and echinoids from the Los Angeles Basin, California. United States Geological Survey Professional Paper, 190: 1-65, pls 1-9. Zhagbib-Turki, D. 1987. Les Echinides du Cretacé de Tunisie. Paléontologie generale: systématique, paléoecologie, paléobiogeographie. Unpublished Ph. D. Thesis, Faculté des Sciences de Tunis. 613 pp, 25 pls. 1989. Les échinides indicateurs des paléoenvironments: un exemple dans le Cénomanien de Tunisie. Annales de Paléontologie, 75: 63-81. ig. 4. Plesiaster vancouverensis sp. nov. uppermost Santonian; Lower Campanian, / (S.) schmidti Zone, French Creek, Vancouver Island, British Columbia. A, B, BMNH EES078 (internal mould), holotype: A, apical view; B, oral view. C,D, BMNH EES081 (internal mould), paratype: C, apical view; D, oral view. E, F, BMNH EE5S079 (internal mould), paratype: E, apical view; F, oral view. All x1. | Bull. nat. Hist. Mus. Lond. (Geol.) 52(2): 109-114 | Issued 28 November 1996 The cranial anatomy of Rhomaleosaurus thorntoni Andrews (Reptilia, Plesiosauria) ARTHUR R. I. CRUICKSHANK Earth Sciences Section, Leicestershire Museums, The Rowans, College Street, Leicester LE2 OJJ; and Department of Geology, University of Leicester, University Road, Leicester LE] 7RH Synopsis. The skull and lower jaw of Rhomaleosaurus thorntoni Andrews, 1922, from the Upper Lias of Northamptonshire, are figured for the first time. New information shows that the external nares are in a perfectly normal position, just in front of the orbits. There is little difference between R. thorntoni, R. zetlandicus and R. cramptoni, the type species of the genus. As they can be considered to be conspecific, Rhomaleosaurus zetlandicus (Phillips, in Anon, 1854) has priority. R. zetlandicus is of more robust construction than the Rhaetian/Hettangian species R. megacephalus (Stutchbury, 1846), with, among other differences, teeth having fewer striae and the internal nares of a different construction. INTRODUCTION The species Rhomaleosaurus thorntoni was proposed by C W ‘Andrews in 1922 for a pliosauroid plesiosaur (Brown 1981) from the Toarcian (Upper Liassic) of Kingsthorpe, Northamptonshire. The ‘ype specimen (BMNH R4853) comprises a partial skull, partial mandible and much of the postcranial skeleton, but lacks the limbs. Andrews (1922) described the skull and postcranial remains of the specimen in some detail, but illustrated only the sacral vertebrae and he limb girdles. No illustration of the skull and jaw material exists and it is the purpose of this paper to remedy this omission as part of A series of papers to improve knowledge of the Liassic plesiosaurs Taylor 1992a, b; Taylor & Cruickshank 1993a; Cruickshank, 1994a, ?). Andrews discussed the characters of his new species, comparing |hem most closely with those of R. cramptoni (Carte & Baily 1863) INMING F8785). As will be shown below, however, the differences he enumerated between R. thorntoni and R. cramptoni cannot now de Sustained; in addition, many of the characters of R. thorntoni are o be found in R. zetlandicus (Phillips, in Anon, 1854) (Taylor 1992a) (YORYM GS503). ‘INSTITUTIONAL ABBREVIATIONS 3MNH Palaeontology Collections, The Natural History Mu- seum, Cromwell Road, London SW7 5BD (formerly -the British Museum (Natural History)) LEICS Leicestershire Museums, Arts and Records Service, The Rowans, College Street, Leicester LE2 O0JJ AANCH The Manchester Museum, Oxford Street, Manchester M13 9PL NMING National Museum of Ireland, Kilare Street, Dublin 2, | Eire VM Whitby Museum, Whitby Literary and Philosophical Society, Pannet Park, Whitby, Yorkshire YO21 1RE ‘YORYM Yorkshire Museum, Museum Gardens, York YO1 2DR. ) The Natural History Museum, 1996 SYSTEMATIC PALAEONTOLOGY Class REPTILIA Subclass SAUROPTERYGIA Owen, 1860 Order PLESIOSAURIA de Blainville, 1835 Superfamily PLIOSAUROIDEA (Seeley, 1874) Welles, 1943 Family PLIOSAURIDAE Seeley, 1874 Genus RHOMALEOSAURUS Seeley, 1874 TYPE SPECIES. Plesiosaurus cramptoni Carte & Baily, 1863 Rhomaleosaurus zetlandicus (Phillips, in Anon, 1854) Figs 1-6 Plesiosaurus zetlandicus Phillips, in Anon: 19. 1863 = Plesiosaurus cramptoni Carte & Baily: 160. Rhomaleosaurus cramptoni (Carte & Baily) Seeley: 448. 1922 Rhomaleosaurus thorntoni Andrews: 413. 1992 Rhomaleosaurus zetlandicus (Anon, in Phillips, 1854); Taylor: 52. DIAGNOsIS. A Rhomaleosaurus with a more robust and relatively shorter and wider skull, and a steeper profile of the lower jaw symphysis when compared with Rhomalosaurus megacephalus (Stutchbury). Tooth ornament coarse, with widely-spaced ridges and reducing in number towards the tip, triangular in section. Palatal foramina and internal nares lie in the same groove, as opposed to the condition in R. megacephalus. The length-width ratio of the snout is 1: | as opposed to 1.25: 1 for R. megacephalus. The specimen described here is BMNH R4853. Andrews (1922: 413) did not formally diagnose R. thorntoni, except by distinguish- ing it from R. cramptoni in several characters. Andrews (1922: 414) also gave an opinion that Plesiosaurus megacephalus Stutchbury, 1846 belonged to the genus Rhomaleosaurus, but gave no reasons (see Taylor & Cruickshank (1989) and Cruickshank (1994a) for discussion). Plesiosaurus cramptoni (NMING F8785) is the type species of the genus Rhomaleosaurus Seeley, 1874, and comes from Alum 110 Shales at Kettleness on the north Yorkshire coast (Benton & Taylor 1984) of Toarcian (Bifrons Zone) age. Other English species that have been referred to this genus include Plesiosaurus megacephalus Stutchbury, 1846 and P. propinquus Phillips, 1854. Only R. megacephalus is represented so far by more than one specimen, and it alone seems to be from the Lower Liassic (Rhaetian/Hettangian) (Cruickshank 1994a). Plesiosaurus propinquus differs from other species in having a marked boss on the hind end of the inner surface of the lower jaw, just in front of the glenoid and in place of the dorso- median trough (Taylor 1992a, b), and thus its position within Rhomaleosaurus must be reconsidered. Table 1 Abbreviations used on Figs 1-6. alv anterior interpterygoid lgr lateral groove vacuity mto mature tooth carina carina on crown of tooth mx maxilla co coronoid no notch cr crown orb orbit d dentary pal palatine dep depression palv primary alveolus dmfo dorsomedian foramen pmx premaxilla ec ectopterygoid po postorbital en external naris pt pterygoid fac facial processes of the ptb pterygoid boss premaxillae ri ridged ornament on crown fan fan-shaped area of tooth fo foramina Tto replacement tooth er groove salyv secondary alveolus in internal naris sof suborbital fenestra info foramina associated with sp splenial internal naris sym symphysis j jugal Vv vomer 1 position of first tooth Oblique lining represents broken or sectioned bone or tooth. Mechanical stipple represents matrix or crushed bone. DESCRIPTION. Skull (Figs 1, 2). The skull and lower jaw have recently been cleaned and conserved, and they alone will be dealt with here. Only the anterior portion of the skull was collected; the clean break surface runs obliquely from a position in front of the left orbit, through the left external naris, to the front edge of the right orbit, and thence through the postorbital bar. Some bone has been lost from the tip of the premaxillae. The right cheek bar is attached to the snout and runs as far as the end of the maxilla. Attached to the cheek bar is a portion of the palate, comprising the right ectopterygoid and a small part of the pterygoid. The base of the postorbital rests on the posterior end of the jugal. Apart from the obvious break, the skull has been damaged by post-mortem effects which have compressed the bone dorso-ventrally and caused the facial processes of the premaxillae to be shortened, so that the midline of the snout has a step, with the posterior part of the premaxillae, as preserved, being pushed under the anterior part and offset to the right. The maxillae may, in addition, have been squeezed together under the facial processes of the premaxillae. All this disruption has obscured the right external naris. In front of, and lateral to, the position of the hidden right external naris, is a deep depression bottomed with crushed bone and an associated wide groove running to the premax- illary edge. The right jugal is partly visible, and is a narrow bone running under the orbit and ending below the postorbital. However, as the bone is heavily pyritized and crushed; and the sutures much closed up, the prefrontal and lacrimal cannot be distinguished. Similarly the detailed structure of the postorbital — jugal area is obscured. There is no reason to believe that this latter region is any different from that described in R. megacephalus (Cruickshank A.R.I. CRUICKSHANK : dmfo See i ea RL Ta dep RS ptb Fig. 1 Rhomaleosaurus thorntoni Andrews; dorsal view of the skull; scale bar = 100mm. For abbreviations on this and the other figures, see Table 1. 1994a), or indeed the Kimmeridgian species Pliosaurus brachy- spondylus (Taylor & Cruickshank 1993). The anterior palatal surface shows much less damage. A very few fully erupted (mature) teeth are still in their sockets, but several replacement teeth are present, both in primary and secondary al-| be distinguished except the premaxillae and maxillae. The vomers are substantial bones, forming a midline bar on the palate. Anteriorly/) they terminate in a horseshoe-shaped structure with several associ] ated foramina. The vomers widen posteriorly, and are here flanked by grooves which run to the internal nares from fan-shaped areas just | behind, and internal to, each diastema, opposite the notches where} the premaxillae meet the maxillae. These fan-shaped areas arel] » = CRANIAL ANATOMY OF RHOMALEOSAURUS THORNTONI ANDREWS fig. 2 Rhomaleosaurus thorntoni Andrews; ventral view of the skull; | scale bar = 100 mm. povered in a radiating set of shallow grooves, suggesting that they ielped anchor the buccal lining. These features of the internal narial region are different from those of Pliosaurus brachyspondylus and R. megacephalus, where the structure is more fully known (Cruickshank et al 1991; Taylor & Sruickshank 1993b). In P. brachyspondylus, the internal nares lie at the end of grooves in the roof of the mouth, with two prominent “oramina lying on the medial faces of the depression in front of each ternal naris, all equally spaced. This is markedly different from R. negacephalus, where the internal nares lie at the ends of grooves, medial to, and parallel with, supplementary grooves which end in oramina. In neither of these species is there a fan-shaped area nedial to the diastema, nor the extra foramina lying within the limits f the internal narial excavation, as illustrated here. As in all plesiosaurs which I have examined, the internal nares lie anterior to external nares, inviting the explanation that the narial system cted as a hydrodynamically driven olfactory system, and was not sed for respiration (Cruickshank ef al 1991). The internal nares in . zetlandicus and R. cramptoni are not visible, being obscured by he rami of the lower jaw (Taylor 1992b), or matrix. | The badly disrupted posterior palatal elements show that there as a prominent pterygoid boss in exactly the same position as in R. = (Taylor 1992a; b), and an ectopterygoid lying between e jugal and pterygoid. Mandible (Figs 3, 4). Parts of the lower jaw preserved include an Imost complete right ramus as far back as the end of the dentary, the ymphysis and the left ramus to just behind the symphysis, plus a =m 111 Fig. 3 Rhomaleosaurus thorntoni Andrews; lower jaw; 3a, dorsal view; 3b, section through symphysis on line a—b; scale bar = 100 mm. portion of the middle of the left ramus and the left articular region (not illustrated). These portions of the lower jaw correspond almost exactly to the remains of the skull and no doubt represent what was saved during collection, from what must have been an almost complete cranium. The symphysis occupies five tooth positions and a further 26 tooth positions can be counted in the right dentary. The general obscurity of sutures makes it difficult to identify individual bones, but as far as can be seen, the structure of this lower jaw is the same as that of R. zetlandicus (Taylor 1992b). There are both mature and replacement teeth present in the lower jaw, with their associated primary and secondary alveoli. On the portion of the left jaw ramus containing the glenoid fossa, there is a large dorso-median trough on the prearticular and articular (Taylor 1992b: fig 7; Cruickshank 1994a: fig 7), which may be one of the determining characters of the genus Rhomaleosaurus (Taylor 1992a; Cruickshank 1994a). Fig.4 Rhomaleosaurus thorntoni Andrews; symphysis of lower jaw; 4a, ventral view; 4b, section through symphysis on line cd; scale bar = 100 mm. Fig.5 Rhomaleosaurus thorntoni Andrews; teeth; 5a, replacement tooth crown, position 9, right maxilla; 5b, replacement tooth crown, position 24, right dentary; 5e, mature tooth, position 12, right dentary; 5d, mature tooth, position 8, right dentary; teeth are oriented with crowns towards top; drawn with an Abbé drawing apparatus on a Wild M3 stereomicroscope; scale bar = 5 mm. A.R.I. CRUICKSHANK Dentition (Fig. 5). The dentition is that of a powerful predator, with a rosette of interlocking, procumbent teeth in the premaxillae and lower jaw symphysis, followed by tooth-rows which, after two small median teeth, have large caniniforms in the upper jaw overlap- | ping somewhat smaller teeth in the lower jaw. The tooth adjacent to the midline in both the upper and lower dentitions is much smaller | than the more mesial teeth. In the lower jaw there is a marked } reduction in size of teeth immediately behind the fifth position, | which continues in a regular manner to the end of the tooth row on | the dentary. In the upper jaw the fifth tooth position is very small, and is followed behind the diastema by another small tooth. Tooth | positions seven, eight, nine and ten are very much larger caniniforms. | Thereafter there is an even more marked reduction in tooth size, | when compared with the lower dentition, until the sockets become difficult to distinguish. This arangement is very similar to that of R. | zetlandicus (Taylor 19925), allowing for the incompleteness of that |) specimen. It is possible to amplify the description of the individual teeth offered by Taylor (1992b), for R. zetlandicus. Those illustrated come | from the 9th position on the right maxilla, showing the buccal | surface (Fig. 5a); lying across the root of the 23rd tooth on the right |) ramus of the lower jaw (Fig. 5b); the 12th position of the right ramus of the lower jaw (Fig. 5c); and the 8th position of the right ramus of the lower jaw (Fig. 5d). Figs 5a and 5b are replacement teeth, whereas Figs 5c and 5d are erupted, mature teeth. The crowns are covered in a coarse ornament, which reduces in | number of ridges towards the tooth-tip, but which all seem to have carinae on mesial and distal surfaces. The ornament on these teeth is identical with those illustrated by Taylor (1992b: fig. 9), but quite different from the tooth illustrated by Cruickshank (1994a: fig. 10) for R. megacephalus, where the ornament is much finer and more - closely spaced. The ridges are triangular in section, and some start slightly below the crown-root boundary. DISCUSSION Andrews (1922: 413) compared R. thorntoni with R. cramptont, regretting that the shoulder girdle of the latter was not visible and |; that he could not therefore use it for comparative taxonomic pur‘ poses. The skull and vertebral column of each species seemed to be much the same, but he drew attention to the following differences between them. Firstly, he thought that the external nasal openings were much further in front of the eyes in R. thorntoni than in R. cramptoni. Secondly, he recognized differences in the platforms of their cervical neural arches: in R. thorntoni these are nearly horizon- | tal, but in R. cramptoni they are strongly inclined. Thirdly, he}’. pointed out that the humerus in R. thorntoni was relatively larger. with a more expanded distal end. Neither of the external nasal openings are very obvious in R. thorntoni: that on the right side is obscured by the displaced facial processes of the premaxillae, and that on the left is only partly)}) preserved and probably invisible before the skull was recently cleaned properly. However, there is the depression some distance in front of the right orbit which could have been mistaken for af external naris prior to full cleaning of the specimen, and this woul agree with Andrews’ identification of an unusually anteriorly placeq external nasal opening. This depression is floored with crushec bone, and does not penetrate onto the underside of the dermal bone: of the snout. Restoration of the snout region (Fig. 6c) using informa tion now available, shows the external nares to be situated in i normal position relative to the orbits. - ge wF : _CRANIAL ANATOMY OF RHOMALEOSAURUS THORNTONI ANDREWS a b = 100 mm. The differences in orientation of the zygapophyses in the cervical vertebrae of Plesiosauria depend on their relative position in the neck. In general the zygapophyses of the anterior cervical vertebrae are horizontally oriented, becoming inclined after the first ten or so. For instance in MANCH LL8004, a specimen of Macroplata longirostris (Blake) (Broadhurst & Duffy 1971), there are about 32 cervical vertebrae, of which the first ten have horizontal zygapophy- ses, while the remainder have zygapophyses angled at about 45° to the horizontal. Liassic plesiosaurs in general seem to have between 28 and 32 cervical vertebrae. Even in the posteriormost cervicals, the rib articulations are placed close to the lower rim of the centra (Taylor & Cruickshank 1993a), and therefore could still appear to be from a more anterior position. Therefore, it is not always obvious from which part of the neck any single vertebra might come, and hence to draw conclusions about zygapophyseal orientation is pre- mature. The question of the characters of the humeri may well depend on the state of preservation of each. The skull and skeleton of R. cramptoni are very much less damaged than those of R. thorntont, and it seems unwise to make strict taxonomic statements on this character without knowing more about individual variation within the genus Rhomaleosaurus. L Therefore, the principal points of difference between the two species can be interpreted as being due to either their relative state of preservation, their size, or to an unreliable character, as in the case of ‘he neck vertebrae. On the basis of the foregoing discussion, both R. _eramptoni and R. thorntoni are seen to belong to the same species. In ddition they come from approximately the same horizon, in the i= stage of the Liassic (Lower Jurassic) of England. One other similar pliosauroid is known from the Yorkshire (Eng- and) Toarcian, R. zetlandicus (Phillips, in Anon, 1854) (Taylor 19920) Reconstructions of part of the skulls of R. thorntoni, R. -elandicus and R. cramptoni are shown for comparison (Figs 6a-c). The relevant differences lie in the overall size of each and in the \pparent width of the postorbital bar; in R. thorntoni it is relatively ider than in R. zetlandicus and R. cramptoni, but as all specimens re variously damaged in that area, no firm conclusions can be eached on this character. All specimens have the same short, broad nout, which contrasts with the more slender, relatively longer snout 113 Fig.6 Outline reconstructions of the anterior portion of skulls; 6a, Rhomaleosaurus cramptoni (Carte & Baily, 1863), from a photograph of the type NMING F8785; 6b, Rhomaleosaurus zetlandicus (Phillips, in Anon, 1854), after Taylor 1992b; 6c, Rhomaleosaurus thorntoni Andrews, 1922; scale bars of the Hettangian R. megacephalus (LEICS G221.1851) (Cruickshank 1994a). The Toarcian specimens have similar denti- tion, possessing sparsely ridged teeth, which also contrasts with those of R. megacephalus. Taking all three Toarcian species together (Fig. 6), it is probable that they represent only size variants of the same species. They are conspecific and should be referred to the single species Rhomaleosaurus zetlandicus (Phillips, in Anon, 1854), which has date priority. In Fig. 6, which compares that part of the skull preserved in R4853 with the other two types, it will be noted that the premaxillaries of R4853 are apparently narrower than those of the other two specimens. The reconstruction was effected using the most con- servative measurements, and perhaps this is reflected in a false narrowing of the premaxillary facial processes. It is not likely that, for instance, any conclusions can be drawn from such a reconstruc- tion concerning growth rates, or sexual dimorphism. SUMMARY AND CONCLUSIONS 1 The skull of the type specimen of Rhomaleosaurus thorntoni Andrews, 1922, from the Toarcian of Northamptonshire, is illus- trated for the first time. Additional information concerning details of its external nares, and reassessment of other characters dis- cussed in the original description, make it difficult to sustain its supposed differences from R. cramptoni (Carte & Baily, 1863) from the Toarcian of Yorkshire. Comparisons with the type of R. zetlandicus (Phillips, in Anon, 1854), also from the Toarcian of Yorkshire, indicate that R. thorntoni is merely a larger specimen of R. zetlandicus. 3 Since all three specimens are shown here to belong to the same species, the correct name for it is Rhomaleosaurus zetlandicus (Phillips, in Anon, 1854). 4 Rhomaleosaurus zetlandicus was the top predator in the Upper Lias of England. R. megacephalus from the Rhaetian or Hettangian (Lower Lias) has a longer, more slender snout, and different dentition. tO 114 ACKNOWLEDGEMENTS. The type of Rhomaleosaurus thorntoni was cleaned at the Natural History Museum, London, under the direction of Dr Angela Milner, at the request of Dr Michael Taylor. Iam grateful to Dr Milner for the loan of this specimen, and for access to the mounted cast of the type of R. cramptoni (BMNH R34) in the Natural History Museum, and to Dr Taylor for discussion and access to his collection of reference slides. Mr John Martin, Keeper of Earth Sciences, and the Director of the Leicestershire Museums Service provided facilities for the work to be undertaken. The assistance of Mrs Anne Montgomery of the Geology Department, University of Leicester is gratefully acknowledged. Mr Nigel Monaghan (NMING) cheerfully supplied photographs of the type of R. cramptoni. John Martin, David Brown, Angela Milner and Michael Taylor read drafts of the manu- script, to its great improvement. The work was done during the tenure of a Leverhulme Fellowship awarded to Dr M A Taylor. REFERENCES Andrews, C. W. 1922. Notes on the skeleton of a large plesiosaur (Rhomaleosaurus thorntoni sp.n) from the Upper Lias of Northamptonshire. Annals and Magazine of Natural History, 9, 10; 407-415. Anon 1854. Report of the Council of the Yorkshire Philosophical Society. Annual Report of the Yorkshire Philosophical Society for 1853: 7, 8. Benton, M. J. & Taylor, M.A. 1984. Marine reptiles from the Upper Lias (Lower Toarcian, Lower Jurassic) of the Yorkshire coast. Proceedings of the Yorkshire Geological Society, 44: 399-429. Broadhurst, F. M. & Duffy, L. 1971. A plesiosaur in the Geology Department, University of Manchester. Museums Journal, 70: 30-31. Brown, D. S. 1981. The English Upper Jurassic Plesiosauroidea (Reptilia) and a review A.R.I. CRUICKSHANK of the phylogeny and classification of the Plesiosauria Bulletin of the British Museum (Natural History), (Geology Series), 35: 253-347. Carte, A. & Baily, W. H. 1863. Description of a new species of Plesiosaurus from the Lias, near Whitby, Yorkshire. Journal of the Royal Dublin Society, 4: 160-170. Cruickshank, A. R. I. 1994a. Cranial anatomy of the Lower Jurassic pliosaur Rhomaleosaurus megacephalus (Stutchbury) (Reptilia; Plesiosauria). Philosophical Transactions of the Royal Society of London, (B) 343: 247-260. 1994b. A juvenile plesiosaur (Plesiosauria: Reptilia) from the Lower Lias (Hettangian: Lower Jurassic) of Lyme Regis, England: a pliosauroid — plesiosauroid intermediate? Zoological Journal of the Linnean Society of London, 112: 151-178. , Small, P. G. & Taylor, M.A. 1991. Dorsal nostrils and hydrodynamically driven underwater olfaction. Nature, London, 352: 62-64. Phillips, J. 1854. On a new Plesiosaurus in the York Museum. Annual Report of the British Association for the Advancement of Science for 1853, 54. | Stutchbury, S. 1846. Description of a new species of Plesiosaurus in the Museum of t the Bristol Institution. Quarterly Journal of the Geological Society of London, 2: 411-417. Taylor, M. A. 1992a. Taxonomy and taphonomy of Rhomaleosaurus zetlandicus (Plesiosauria; Reptilia) from the Toarcian (Lower Jurassic) of the Yorkshire coast. Proceedings of the Yorkshire Geological Society, 49: 49-55. 1992b. Functional anatomy of the head of the large aquatic predator | Rhomaleosaurus zetlandicus (Plesiosauria; Reptilia) from the Toarcian (Lower 1 Jurassic) of Yorkshire. Philosophical Transactions of the Royal Society of London., (B) 335: 247-280. — & Cruickshank, A. R. I. 1989. The Barrow Kipper, ‘Plesiosaurus’megacephalus © (Plesiosauria; Reptilia) from the Lower Lias (Lower Jurassic) of Barrow upon Soar, Leicestershire. Transactions of the Leicester Literary & Philosophical Society, 83: ) 20-24. —& 1993a. A plesiosaurian reptile from the Linksfield erratic (Rhaetian; | Upper Triassic) of Morayshire. Scottish Journal of Geology, 29: 191-196. & 1993b Cranial anatomy and functional morphology of Pliosaurus | brachyspondylus (Reptilia; Plesiosauria) from the Upper Jurassic of Westbury, Wilt- ( shire. Philosophical Transactions of the Royal Society of London, (B) 341: 399418. | | Bull. nat. Hist. Mus. Lond. (Geol.) 52(2): 115-118 Issued 28 November 1996 The first known femur of Hylaeosaurus armatus and re-identification of ornithopod material in The Natural History Museum, London PAUL M. BARRETT Department of Earth Sciences, University of Cambridge, Downing Street, Cambridge CB2 3EQ SYNOPSIS. The first known femur of the British Lower Cretaceous ornithopod dinosaur Hylaeosaurus armatus Mantell, 1833 is described. The status of Camptosaurus valdensis (Lydekker, 1889) is reviewed, and it is suggested that it might be senior synonym of Valdosaurus canaliculatus (Galton 1975). | | INTRODUCTION |The purpose of this brief paper is to report and describe the first recognised femur of the nodosaurid ankylosaur Hylaeosaurus _armatus Mantell 1833, and to clarify the taxonomic position of |several ornithopod specimens.The status of Camptosaurus valdensis ‘(Lydekker 1889) is also reviewed and it is provisionally placed in synonymy with Valdosaurus canaliculatus (Galton 1975). Unless |otherwise stated, all of the specimens described are of Wealden (Lower Cretaceous) age and are from the Isle of Wight, England. They belong to the Fox Collection housed in the Department of Palaeontology, The Natural History Museum, London (NHM; regis- ‘ter numbers prefixed BMNH in this paper). SYSTEMATIC PALAEONTOLOGY | DINOSAURIA Owen 1841 ORNITHISCHIA Seeley 1887 THYREOPHORA Nopsca 1915, sensu Sereno 1986 ANKYLOSAURIA Osborn 1923 Family NODOSAURIDAE Marsh 1890 Genus HYLAEOSAURUS Mantell 1833a TYPE AND ONLY SPECIES. Hylaeosaurus armatus Mantell 1833b; Lower Cretaceous (Upper Valanginian), East and West Sussex, Southern England. Hylaeosaurus armatus Mantell 1833 Fig. 1 1833b Hylaeosaurus armatus Mantell: 328. HOLOTYPE. BMNH R3775, the anterior part of a skeleton embed- ded in a block of matrix. HORIZON AND LOCALITY. Hastings Beds (Lower Wealden), Upper Valanginian, Lower Cretaceous (Rawson et al 1978). Tilgate Forest, Cuckfield, West Sussex, England. REFERRED MATERIAL. As listed by Pereda-Suberbiola (1993) and MNH R604k (now registered as BMNH R12555), the distal ortion of a right femur. 9° The Natural History Museum, 1996 DESCRIPTION AND COMPARISON. The distal portion of a large right femur (Fig. la), BMNH R604k (Dawson Collection), from the Wealden, near Hastings, East Sussex, may represent the first recog- nised femur of Hylaeosaurus armatus. This specimen was originally identified as /guanodon sp.' but was later referred to as a large individual of Hypsilophodon foxii (Huxley) by Molnar & Galton (1986). However, the femur is much more massive than that of H. foxii, and the absence of a marked anterior intercondylar groove suggests that it is not referable to Jguanodon. The absence of ridges running up the femoral shaft from the condyles supports the view that R604k is not referable to either of these genera. There is a marked posterior intercondylar groove and the lateral and medial condyles extend equal distances posteriorly from the femoral shaft. The lateral condyle is much more massive than the medial one, and there is a distinct ‘step’ between the medial condyle and the medial extremity of the femur (Fig. 1b). The femur bears a strong resemblance to that of the Oxfordian nodosaur Cryptodraco eumerus (Galton 1983) which also displays a ‘step’ between the medial condyle and the medial border of the femur. BMNH R604k can be distinguished from the femora of the other known Wealden nodosaur Polacanthus foxii (BMNH R175) by a number of features. For example, Polacanthus shows no ‘step’ medial to the medial condyle, and the medial condyle is more massive in Polacanthus than in BMNH R604k. It seems unlikely, therefore, that BMNH R604k is referable to Polacanthus. It is suggested that BMNH R604k is not ornithopod as previously supposed but thyreophoran, and the locality and the horizon from which the specimen was recovered suggests that it is referable to Hylaeosaurus. The rela- tively small size of the femur (only 83mm across the distal end) suggests that it belonged to a juvenile (W. Blows, pers. comm.). Unfortunately, due to the fragmentary nature of the specimen, this assignment can only be tentative. Nonetheless, the femur provides valuable information on the alleged synonymy of Hylaeosaurus and Polacanthus (Coombs 1971, Blows 1987, Coombs & Maryanska 1990, Pereda-Suberbiola 1991, 1993). Coombs (1971) and Coombs & Maryanska (1990) suggested that 'Lydekker (1888) listed a number of specimens as BMNH R604a-e. These registered numbers include a number of theropod, ornithopod and ankylosaur remains, all of which come from the Wadhurst Clay near Hastings and are part of the Dawson Collection. The femur in question is labelled BMNH R604k. There is no record of this number in either Lydekker (1888) or in the accessions catalogues of the Natural History Museum (S. Chapman pers. comm.). The specimen label identified the femur as Iguanodon sp. from the Wadhurst Clay near Hastings. 116 la Fig. 1 Hylaeosaurus armatus Mantell. Right femur, distal end; BMNH R12555; la, posterior view, x 0.7; 1b, distal view (note the ‘step’ medial to the medial condyle), x 0.8. Polacanthus is a junior synonym of Hylaeosaurus. However, these authors claimed that no homologous elements are present in the holotype material of Hylaeosaurus and Polacanthus so the proposed synonymy was based largely on the similar geographical and stratigraphical distributions of these genera. In contrast, Blows (1987) suggested that the two genera are stratigraphically distinct, with Hylaeosaurus originating from the Hastings Beds (Valanginian) of East Sussex, whilst Polacanthus remains have been recovered from the Wessex Formation (Barremian) in the Isle of Wight. He also suggested that the arrangement of the dermal armour differs be- tween the two genera. Recently it has been proposed that the North American nodosaur Hoplitosaurus is a junior synonym of Polacanthus (Pereda- Suberbiola 1991, 1993). A specific separation, based on femoral characters, is retained to distinguish the British form (P. foxii) from the North American form (PR marshi Lucas, 1901). If these two genera are genuinely synonymous then a number of elements re- ferred to P. (=Hoplitosaurus) marshi that were previously unknown in the holotype of P. foxii (e.g. humerus and scapula) can be used for comparison with Hylaeosaurus. Comparisons of the pectoral girdle, forelimb, hindlimb and dermal armour led Pereda-Suberbiola (1991, 1993) to the conclusion that Hylaeosaurus and Polacanthus were distinct genera. If BMNH R604k does belong to Hylaeosaurus, this would pro- vide another point of comparison between these two genera. The P.M. BARRETT characters listed above suggest that Polacanthus and Hylaeosaurus | can be distinguished on femoral characters, adding weight to the arguments of Blows (1987) and Pereda-Suberbiola (1993). This | specimen has been re-registered as BMNH R12555 in order to avoid confusion with other specimens allotted to BMNH R604, which contains a variety of other specimens from the same locality includ- ing [guanodon remains (Lydekker 1888). ORNITHOPODA Marsh 1881 Family DRYOSAURIDAE Milner & Norman 1984 Genus VALDOSAURUS Galton 1977 TYPE SPECIES. Dryosaurus canaliculatus Galton 1975; Lower Cretaceous (Valanginian—Barremian), West Sussex and Isle of Wight, southern England. Valdosaurus canaliculatus (Galton 1975) Fig. 2 1975 Dryosaurus? canaliculatus Galton: 747. 1977. Valdosaurus canaliculatus (Galton); Galton: 231. 1982 Valdosaurus canaliculatus (Galton); Galton & Taquet: 147. 2b Fig. 2 Valdosaurus canaliculatus (Galton). Left femur, proximal end, BMNH R12440; note the deep cleft and wide separation between the lesser and greater trochanters. 2a, posteromedial view; 2b, proximal view; x |. | REFERRED MATERIAL. _ & Taquet (1982), BMNH R167 an incomplete left femur (see below) HOLOTYPE. BMNH R184 and BMNH R185, associated left and right femora. FEMUR OF HYLAEOSAURUS ARMATUS HORIZON AND LOCALITY. Hastings Beds, Upper Valanginian, Lower Cretaceous (Rawson ef al. 1978). Tilgate Forest, Cuckfield, | West Sussex, England. Wealden Shales, Barremian, Lower Creta- | ceous (Rawson et al. 1978), Isle of Wight, England. Specimens listed by Galton (1975), Galton _ and BMNH R170 (now registered as BMNH R12440) the proximal | portion of a right femur (see below). DESCRIPTION AND COMPARISON Lydekker (1888) referred a number of unassociated limb bone fragments (BMNH R170) to Hypsilophodon foxii (Huxley). Galton (1975) noted that two of the femoral fragments within BMNH R170 differed significantly from the majority of femora attributed to this _ genus. InH. foxiithe distal end of the femur has a moderate posterior intercondylar groove and lacks an appreciable anterior intercondylar groove. Proximally, the lesser trochanter is separated from the _ greater trochanter by a narrow cleft (Galton 1974). Galton (1975) showed that the two specimens in question possess a distinct anterior Jintercondylar groove and he referred these specimens (now regis- tered as BMNH R8420 and BMNH R8421) to Valdosaurus \(=Dryosaurus) canaliculatus, a dryosaur from the Wealden of the Isle of Wight (Galton 1977, Galton & Taquet 1982). Examination of the specimens remaining within BMNH R170 has yielded a proxi- mal femoral fragment (Fig. 2a) in which the lesser trochanter is separated from the greater trochanter by a deep cleft (Fig. 2b). This jfeature is characteristic of the Dryosauridae (sensu Sues & Norman 1990) and the locality and horizon from which the specimen comes ‘suggests that it is referable to Valdosaurus canaliculatus, BMNH R12440. Several limb bone fragments attributable to H. foxii remain as BMNH R170. Galton (1975) suggested that a small right tibia (BMNH R124), previously listed as [guanodon sp. (Lydekker 1888) was also refer- 124 appears to be more similar to that of /guanodon than to that of H. foxii. In Iguanodon the cnemial crest is longer than in H. foxii. In ddition, the cnemial crest of /guanodon swings laterally near its anterior margin (see Fig. 3). On the basis of these characters BMNH R124 is referred to/. cf. atherfieldensis Hooley. The small size of the specimen suggests that it belonged to a juvenile. Rn to H. foxii. However, the form of the proximal end of BMNH cnemial crest a b Cc fig. 3 Proximal right tibiae; 3a, b, /guanodon atherfieldensis Hooley, 1924: 3a, after Norman 1986, 3b, BMNH R124; 3e, Hypsilophodon foxii (Huxley), after Galton 1974; anterior is towards the top; drawings not to scale. 117 ON THE STATUS OF CAMPTOSAURUS VALDENSIS (Lydekker, 1889) The femur BMNH R167 (Fig. 4) has proved something of an enigma. Lydekker (1888) listed it as H. foxii, but suggested that it may represent a distinct species of Hypsilophodon due to its greater size. Later, Lydekker (1889) suggested that BMNH R167 shared a number of similarities with the femur of his new species Camptosaurus leedsi, and he designated BMNH R167 the holotype of another new species, Camptosaurus valdensis. However, Gilmore (1909) noted several differences between C. valdensis and the North American Camptosaurus dispar Marsh, 1879 (the type species of the genus). For example, the fourth trochanter of C. dispar is situated on the distal half of the femoral shaft, whilst in C. valdensis the fourth trochanter is more proximally placed. Galton (1974) sug- gested that this and several other features indicated that BMNH R167 was not referable to Camptosaurus, but was in fact a large specimen of H. foxii. The use of C. leedsi as a representative specimen for Camptosaurus was also in error. Galton (1980) showed that the holotype of C. /eedsi (a femur, BMNH R1993) differs from North American Camptosaurus in lacking a deep anterior inter- condylar groove and by having a proximally placed fourth trochanter. Due to these and other features, Galton (1980) made Camptosaurus leedsi the type species of a new genus Callovosaurus, which is now regarded as Camptosauridaenomen dubium (Norman & Weishampel 1990). Placing Camptosaurus valdensis is difficult as the areas that provide most of the features used in distinguishing between ornithopod femora, the form of the distal end and the shape and position of the lesser trochanter, are either missing or badly dam- aged. Sues & Norman (1990), in their recent review of the Hypsilophodontidae, regard BMNH R167 as Hypsilophodontidae nomen dubium. It is suggested here that Camptosaurus valdensis Fig. 4 Camptosaurus valdensis (Lydekker, 1889). BMNH R167, anterior view, showing the anterior intercondylar groove; abbreviations: ant.gr. = anterior intercondylar groove, les.tr. = position of lesser trochanter, 4th tr. = fourth trochanter; scale bar = 20 mm. 118 (Lydekker, 1889) may be a senior synonym of the dryosaur Valdosaurus canaliculatus (Galton, 1975), as the beginnings of a marked intercondylar groove can be seen on the anterior face of the distal end of the femoral shaft (contra Galton 1974). This assign- ment is tentative as many diagnostic features are lacking on the specimen. ACKNOWLEDGEMENTS. My sincere thanks to Dr. Angela Milner and Sandra Chapman of the Natural History Museum, London for their continual help throughout the course of this work. Dr. Milner read several earlier drafts of this paper and made many useful comments. Dr. David Norman and Mr. William Blows provided useful discussion and an anonymous reviewer made constructive comments on a draft of this paper. Photographs were provided by the Photographic Unit of the Natural History Museum. Much needed artistic and photographic advice was provided by Hilary Alberti and Dudley Simmons. This paper results from work undertaken during a Natural History Museum vacation studentship, and was written during the tenure of a NERC studentship (GT4/93/128/G) at the University of Cambridge (Department of Earth Sciences). REFERENCES Blows, W.T. 1987. The armoured dinosaur Polacanthus foxii from the Lower Creta- ceous of the Isle of Wight. Palaeontology, 30 (3): 557-580. Coombs, W.P. 1971. The Ankylosauria. Ph.D. thesis (unpublished), University of Columbia, 487pp. & Maryanska, T. 1990. Ankylosauria Jn Weishampel, D.B., Dodson, P. & Osmolska, H. (Eds.) The Dinosauria. University of California Press, Berkeley: 456— 483. Galton, P.M. 1974. The ornithischian dinosaur Hypsilophodon from the Wealden of the Isle of Wight. Bulletin of the British Museum (Natural History), Geology, 25: \-152. 1975. English hypsilophodontid dinosaurs (Reptilia; Ornithischia). Palaeontol- ogy, 18: 741-752. P.M. BARRETT 1977. The ornithopod dinosaur Dryosaurus and a Laurasia — Gondwanaland connection in the Upper Jurassic. Nature, 268: 230-232. 1980. European Jurassic ornithopod dinosaurs of the families Hypsilophodontidae and Camptosauridae. Neues Jahrbuch fur Geologie und Palaontologie. Stuttgart. Abhandlungen, 160: 73-95. — 1983. Armoured dinosaurs (Ornithischia; Ankylosauria) from the Middle and Upper Jurassic of Europe. Palaeontographica A, 182: 1-25. & Taquet, P. 1982. Valdosaurus, a hypsilophodontid dinosaur from the Lower Cretaceous of Europe and Africa. Geobios, 15: 147-159. Gilmore, C.W. 1909. Osteology of the Jurassic reptile Camptosaurus, with a revision of the species of the genus, and a description of two new species. Proceedings of the United States National Museum, 36; 197-332. i | Lydekker, R. 1888. Catalogue of the Fossil Reptilia and Amphibia in the British Museum; Part I. British Museum (Natural History), London. 309pp. — 1889. On the remains and affinities of five genera of Mesozoic reptiles. Quarterly Journal of the Geological Society of London, 45: 41-59. Mantell, G.A. 1833a. Observations on the remains of the /guanodon, and other fossil reptiles, of the strata of Tilgate Forest in Sussex. Proceedings of the Geological Society of London, 1: 410-411 1833b. The Geology of the South-East of England. Longman, London. xix + - | 376pp. Molnar, R.E. & Galton, P.M. 1986. Hypsilophodontid dinosaurs from Lightning — Ridge, New South Wales, Australia. Geobios, 19: 231-239. Norman, D.B. 1986. On the anatomy of /guanodon atherfieldensis (Omithischilil ! Ornithopoda). Bulletin de l'Institute Royal des Sciences naturelles de Belgique: Sciences de la Terre, 56: 281-372. & Weishampel, D.B. 1990. Iguanodontidae and related ornithopods. In! Weishampel, D.B., Dodson, P. & Osmolska, H. (Eds.) The Dinosauria. University of California Press, Berkeley: 510-533. Pereda-Suberbiola, J. 1991. Nouvelle evidence d'une connexion terrestre entre’ Europe et Amerique du Nord au Cretace inferieur: Hoplitosaurus, synonyme de Polacanthus (Ornithischia: Ankylosauria). Comptes Rendus de l'’Academie de Sci- ences de Paris (Series 2), 313: 971-976. 1993. Hylaeosaurus, Polacanthus, and the systematics and stratigraphy of { Wealden armoured dinosaurs. Geological Magazine, 130: 767-781. | Rawson, P.F., Curry, D., Dilley, F.C., Hancock, J.M., Kennedy, W.J., Neale, J.W.,. Wood, C.J. & Worssam, B.C. 1978. A correlation of Cretaceous rocks in the British Isles. Geological Society of London, Special Report, 9: \—70. Sues, H-D. & Norman, D.B. 1990. Hypsilophodontidae, Tenontosaurus, Dryosauridae. In Weishampel, D.B., Dodson, P. & Osmolska, H. (Eds.) The Dinosauria. Universtiy of California Press, Berkeley: 498-509. , | Bull. nat. Hist. Mus. Lond. (Geol.) 52(2): 119-171 Issued 28 November 1996 e ° 7 | Bryozoa from the Lower Carboniferous (Viséan) of | | County Fermanagh, Ireland | PATRICK N. WYSE JACKSON | Department of Geology, Trinity College, Dublin 2, Ireland CONTENTS / WoetOC Ct Oneeeemereer ses eesrscee tee Seamer sever steerer treet eee ee RE TA ieee ed a rte 120 TEV IOUSRWOLKerametne sees scececreee er ere a ee ee or en eee’ I nar) 120 IMIate nial Mews mreteemrr te cectrrce tose timetable ap Meh Caer i eileen mbanamvalbes. 120 Sy stematicialacontol a py ren-tecs eae ee ee eke rh RS IS SS) lS ee hes, coca ai 122 Orden GRP MOSTO MATA See eee Se eee aeeee eee ene a, Seen, 0) Nn ee) Lae farts ee) Morea oe 122 Cenls ladies STUERANGIO Gil), JOSS), cpocccecscecoocecnacacecutsnococuo broontecseenonacenacezese cea ebnncrosencssehsciavecscecie oecebeectiacosaroeeenecbbe 122 PLEX ILESSDOTAAONUS! SP) HOV.) ate steer te nn aman ee Ais De ote eth ee 9 123 GenusiNematopora Ulrich MSS Sate net wet eas ee ee ce ant Een Ine EN tet) eee teats 124 INCTIGLODOLCHIDETMICAISD NOV: sussver cee teeter Te ee ee ee ee ee eee ee 124 GenUSVENCUAOneMatopOra Balakint MOWAm Mentin Oe. 2 Seen ne Noten Aan oe arene ee antes een See See Se 125 FESCUE ONEIALOD OL AND | GNGTUSHS pM OV eer en ete ee eee ae ee ee ee 126 Ceitis /Ralaclaraanon MONDE Ge MOWME, ISTE scecccconosesesseceaonccen coc daeudee seen cotaotheo.aconsebe dence nenesiatoscoxocooeoseb/ Soncoceecoroeoocuaecoscece 128 habdomesonprogracile Wysevackson\ <1 Bancroit. 1995s ee 128 IAGO TONON AAO MAH OLiae (WPAN ES., 1131S) ecoacccoccreoccceeeonenosoecodetosteroscopobconcbsonocnaocoace seFenceaoboocacobcacesontesageneceadbscotonaccs 128 Gems OmiPOPO/GINICCK MSH Direc csc te coc sc ue ar ce ee ee ee AR II oi cece R et csR ices shescacies 129 IR LOMBO DOL Cy LTC ri CCSD) AL OV sass PD RO ig ty 7s Mee. ese cot en 129 IR OMB OD OF GILELAR ONGISD MOV snc sonezcuet erase tot varesieotnes ea eR oN eee eR ER. Big Seo ceed tener digest 132 CenMs Siaaloinyag (Ulm e MIMS.) Witney NEY cores ocnncncponconcaosecors see oreaceanoncdFoce cusee-nopeesn se oneeepencn coc caprosndosebeecee-ne steer ehorcean: 135 SireDlalgy pal (|S) IPECtinataOwenrgl 96 Gemeente rac eae cae ae eee Pe ees es 136 Genusi Glaser pal BasSlerglO 2 OF mene te setts g tesco cs ae ee ere RR IER NO cea ume ties tenes ae 137 (GIGUS Cy DENTGIOS Gl (Wena lOH/3) KOLO yams eee ener ee 137 (iraleie VBINIS SIN LUE VERS eae Pr ee AS die cle Seer iced oa eee ee CE ee ee ee ee 139 | GenUsBGGVIOPOTaIW ySENlACKSOn IS 88 mer eeters secre tree wos ects oeee eek pe RS asc ects Sone ESS eR ee 139 / BAGUIO DOTGNME ASTON (NI @ OYA S AA) Pamir tenets enc fc cce cece cocccese se ccse cat vae vot oca dass eueveps fui gnesesreen eeeinves evans patie 139 Genus DiploporariGaNickles: SHB ASS ler plO OO be. eaeeset ccs cceeecer cree eeces meee ease ores eens ee eee as a eee 140 Diplonoraniainarciialin NOUN Eg SavOUNEmIS/5) Reece nee ete enter ee ee 140 Diploporaniaitenc lIGAW ySCHAGKSONMIOS Sirs. eee eececsocecse ots cecc esses caeectteec se eceoosere soccer erect ecitaens ese ne 141 | GenusiiGhinvoraGnisiMaGoyal Raven. 2e wees ARE GEE cn. cccclrtcse tern es eet eee nee 142 lichihyoraanisinew ennamia Nis Cova SAA sete cert cre eee eee ee 142 GE MUS WTA TEES CLESPISAN PSS Oath ase AAU acacSciscec ee ee ee 143 MAT AISGUSKCOICL NN SEM ACKSON Bll 8 8 pects. ce cnaes ses vaee vest ens cacao coe AGS ase Ears 143 GenustRRomPocladidiROPers OOo... ocn sites camer cL ee ee ee 143 | Riombogladialdichooma (Ni Coy 9844) combsnoy.. 2. et ee ee ee eee 144 Or ers © SMO NAT A Cs re cases cas coe isa cet vaces spuecei sive vasa se eet ee Te Ee 146 GenusvheroclemawU TiC NS 82g ces crac estate ncntet tarsi Zoe A ON eee eee Es, een ys eee 146 HEC LOGIE TMAMIN AGNI ALGISID MON sc saree xt oversee es ears coe s torres so. 5s (sR Nes OE ER, Sais Raa leh te er ctea tei, leer er 146 Genus Dyseritell ai Girty eNO a csees cress ccsss sere Oeecer tates AU hxc ae oo ee OP res Me ceentiedi, Eoecal weet. wun Nee sd aaeieh 148 DS GRUCICMTILIATIGN INICHOIS ONS Oils) erecta tere eres cases sacree ese Ree eee ce eee ea A tras race ee eee 148 (GETS TIGA QOS OUI NRSC ces dee en eae ce a SE CERF rere a EE 150 GD UL OF CRLTIL (TLETIUN CMOS) metres eR anne meen re ee eee eee ee Tn cere 150 Habuliporanowsit (NICHOSOU PUSS) eetettet crs ee 151 TABUMPONCHNINIMG LEC AO Die ee ee ea oc ces Sos eens ceree eee eo eee eee ee 153 GENUS SICHODNTARINIGIUMB ASS CL ML OS OeReeee ENR neve hee ee ee eae ee ee 154 | ISLE OD ECR INIA Sp 8 menteertas cate erate: oe eset ees o soak se Tea ee 155 | Order CUGMOLOR ATA 2 te cir ee eee, A... eaten ates a ave hel ere, clebecgenladege 155 GenustisiZliporaiNidCoy ml SAO ese eae er a te... Meee, Oe C8 ree, cl recon te eS 155 | Histuliporamiicnas tansy (bil lips WSS 6) ieee seeeseee eee as. «<<. cae Seavcsesnuch ee sctraveaeeneess eae Some theses nS ee eee vanes 155 Genuspyilenrerenora dé Orbip nyse B49 As. peeeee ees saceccsustek ete cee ne aes: Me 157 Sulaoretcporalparallelak (Dilip sx 6) ie ecoecascesea leds: sc sss co ee ones eicPa seep eae ee Sad caaae Sena ee a as 158 GemusyGonigaladia\Bthend Se 5518 7 Ocaveaactr gS as tase rscceas cs s.nss izes ede Seneca sone Stes casas a acca ece estas oa cB Sea ie aes atte se onRee eae ea se 159 (Gonroaadiaicelluliferai(EthermG sea 81/5) res aes ate ose x reece ra oe ae aac ar eke as eastside sees ee ee 159 Falaececolopysouthe County bermanas hibryOZOan faim ales .u.ccseceesessasvaraiess---ce sadeedeasscat fas sacasass asorsdnavacissssessseasndeveesatoenesesies 162 ) The Natural History Museum, 1996 P.N. WYSE JACKSON (Comparnisoniwithiothe mAs b tampa im asieecrerere-eeeereneenest ee tceeerare sc eccaya sate neniasense sense snties cmedteter nese cnen = reeenrenaneteernernetenes te recrese eer eeteenee 164 RattemStoib yy OZOznlZzOanlame pl aGementn by Vgs Gal wee eter eee neers ee cee teetering er ees 164 Keystoithendentificationtofisomel Garb onitenous) By OZOAncersrre.- trea aececte ste ere tes caret tect te tare nace meets eee ees ae eee a 165 Acknowled semmenits}t.c:..2-esss te reet cen AIR Raion ea areas ns eas ene nce sce Uaans ivan disenaets ts seca Serco ents nremaere. » Sc05 ssn see nero 168 REfEremCe pars ss tec ceivesssiconnssturtamensevdeqsaorvet feestestectet avenue ctatwaes eae ..- 168 Synopsis. A systematic appraisal of the partially silicified Lower Carboniferous bryozoan fauna of County Fermanagh has demonstrated a rich and diverse bryozoan fauna of which the fenestrate portion has been largely described earlier by other authors. This paper describes the remaining cryptostome, trepostome, and cystoporate elements of the fauna, as well as a few previously ignored fenestrate taxa. 24 species are described (9 cryptostome species; 6 fenestrate species; 6 trepostome species; and 3 cystoporate species) of which 6 are new species and 2 new combinations. The new species are the arthrostylid cryptostomes Hexites paradoxus, Nematopora hibernica, and Pseudonematopora planatus, the rhomboporid cryptostomes Rhombopora cylindrica, and Rhombopora hexagona, and the trepostome Leioclema indentata. The new combinations are Clausotrypa ramosa (Owen), and Rhombocladia dichotoma (M‘Coy). For completeness brief descriptions are given of the following taxa, which have been described more fully elsewhere: Rhabdomeson progracile Wyse Jackson & Bancroft, Baculopora megastoma (M‘Coy), Diploporaria tenella Wyse Jackson, Thamniscus colei Wyse Jackson, and Fistulipora incrustans (Phillips, 1836). The genus Leioclema and the species Streblotrypa pectinata Owen, Diploporaria marginalis (Young & Young), Dyscritella miliaria (Nicholson), Tabulipora howsii (Nicholson), and Tabulipora minima Lee are reported from Ireland for the first time. The following genera are reported from the British Isles for the first time: Hexites, Pseudonematopora and Clausotrypa. Lectotypes are designated for Diploporaria marginalis (Young & Young), Ichthyorachis newenhami M‘Coy, Rhombocladia dichotoma (M‘Coy), Dyscritella miliaria (Nicholson), Tabulipora howsii (Nicholson), and Tabulipora minima Lee. Nomenclature problems for several species have been clarified. A tabular and dichotomous key is given for the complete fauna (including taxa described earlier by other authors). Patterns of silicification show that replacement of calcified bryozoan zoaria by silica was delayed. INTRODUCTION Bryozoans comprise a significant component of Lower Carbonifer- ous faunal assemblages in Ireland. However, they are often fragmentary in nature which has made them difficult to study. Nevertheless, Lower Carboniferous bryozoans have been the subject of research since the mid-1800s when M‘Coy (1844) described many species. In the last thirty years recent studies (Miller 1961a, 1961b, 1962a, 1962b, 1963, Owen 1973, Tavener-Smith 1973, Olaloye 1974, Bancroft 1985, 1986b, Bancroft & Wyse Jackson 1995, Wyse Jackson 1988, Wyse Jackson & Bancroft 1995a) have resulted in the description of new taxa, the redescription of previ- ously described taxa, and give detailed quantitative and statistical analysis of these taxa. While these studies have increased the biostratigraphical value of Carboniferous bryozoans from Ireland, there is still considerable work to be carried out to assess faunas of particular Brigantian stages. This present study adds to the taxonomic diversity of bryozoans described from Ireland, and shows some similarities at generic level, to Lower Carboniferous faunas of the Russian Platform. This paper describes 24 bryozoan species of Lower Carboniferous (Viséan, Asbian) age from County Fermanagh, Ireland. An unusual nodular trepostome and a species of the cystoporate genus Goniocladia, that exhibits atypical branching patterns will be described elsewhere. PREVIOUS WORK The largely silicified fauna from County Fermanagh, dominated by bryozoans and brachiopods, has been the subject of several papers: Tavener-Smith (1965a) erected the genus Prilofenestella, described a species of Minilya (1965b, 1981), noted the occurrence of ovicells in Fenestella (1966), described a new species of Polypora (1971), and monographed 32 species from eight genera of which three were new (1973). Olaloye (1974) examined the acanthocladiid element, describing nine species of Penniretepora, five being new. Three new fenestrate taxa that were discovered in the Carrick Lough fauna during the present study have been described elsewhere (Wyse Jackson 1988), and two species of the cryptostome genus / Rhabdomeson and the cystoporate taxon Fistulipora incrustans (Phillips, 1836) are described more fully by Wyse Jackson & | Bancroft (1995a), and Bancroft & Wyse Jackson (1995). MATERIAL The bryozoans described in this study were collected at two locali- ties, Carrick Lough and Sillees River (Fig. 1) from thin beds of pale grey and muddy limestones that have been assigned to the upper part | of the Glencar Limestone (Brunton & Mason 1979, George er al. | 1976) (Fig. 2). Nearly 50 kg of rock from Carrick Lough containing | both silicified and calcified bryozoan zoaria was processed, and additionally many thousands of unsorted etched silicified specimens | (from the Brunton and Tavener-Smith collections in the Natural | History Museum) and a small number of limestone blocks (from the } Mason collection in the Ulster Museum) were examined. A small | from drift deposits close to Lough Gara, County Roscommon were | also included in this study. | Silicified bryozoan colonies were acid-etched from the surround-} ing limestones. Calcified bryozoan zoaria were extracted from thei} the bryozoans. Type and other material from the Griffith Collection in they National Museum of Ireland, the Owen Collections in the Manches- ter (prefix LL) and Ulster Museums, the Vine Collection in the National Museum of Wales, Cardiff (prefix NMW), the Nicholson} Collection in Aberdeen University (prefix AUGD), the National} Museum of Scotland, Edinburgh (prefix RSM), the Whidborne and} LOWER CARBONIFEROUS BRYOZOA IRELAND LOWER LOUGH ERNE Cc DERRYGONNELLY@® —s ENNISKILLEN @ eCastle Hill Carrick Lough — * Fossil Locality Fault {-—1Dartry Limestone E==Glencar Limestone tudy material. A number of taxa from these collections have been assigned and redescribed in the light of taxonomic work carried ut on the County Fermanagh fauna. Specimens have been deposited in a number of museums. The gest collection is lodged in the Natural History Museum, London refix BMNH PD). A voucher collection which includes some aratype material has been lodged in the Geological Museum, trinity College Dublin (prefix TCD. ). In addition some paratypes of | number of fenestrates (see Wyse Jackson 1988) have been depos- _ted in the National Museum of Ireland (prefix NMING:F) and the Jister Museum (prefix BELUM K). | Morphometric measurements for every taxon were taken on at past 12 specimens, if available, and up to ten measurements were nade on each parameter on each specimen, using a Leitz binocular -ficroscope fitted with a linear graticule at magnifications of be- ‘ig. 1 Location map showing the collecting localities at Carrick Lough and Sillees River, County Fermanagh, Ireland. tween x40 and x100.The mean, standard deviation, and intracolonial and intercolonial coefficients of variation were computed and are tabulated within the description of each taxon. Abbreviations used in these tables are: CV = Coefficient of Variation; CV w = Intra (within) Colonial Variation; CVb = Inter (between) Colonial Variation; NM = Number of measurements; Mn = Minimum value recorded; Mx = Maximum value recorded; x = Mean value; N = Number of speci- mens measured; explanations for other abbreviations used are given in Figs 3, 18, 42, 59 and 84. The measurement scheme for cryptostomes is modified from Newton (1971) (Figs 3, 18); for fenestrates from Tavener-Smith (1973) (Fig. 42); for trepostomes from Boardman (1960) and Cuffey (1967) (Fig. 59); and for cystoporates from Warner & Cuffey (1973) (Fig. 84). 122 DERGVONE SHALE CARRAUN SHALE BELLAVALLY FMN GLENADE SANDSTONE MEENEYMORE FMN DARTRY LIMESTONE (including reef facies) BRIGANTIAN GLENCAR LIMESTONE BENBULBEN SHALE HOLKERIAN MULLAGHMORE SANDSTONE BUNDORAN SHALE (UPPER) DOWRA SANDSTONE BUNDORAN SHALE (LOWER) BALLYSHANNON LIMESTONE CHADIAN COURCEY AN BASAL CLASTICS LIMESTONES SHALES SANDSTONES Fe=9>1 COARSE CLASTICS Fig. 2 Stratigraphical succession in west County Fermanagh. Bryozoan horizon at the top of the Glencar Limestone arrowed. SYSTEMATIC PALAEONTOLOGY During the course of this study three cases of misidentification of taxa in earlier described collections came to light. All cases have a bearing on the systematics of the Asbian bryozoan fauna described below. The first two cases of misidentification were those of P.N. WYSE JACKSON | M‘Coy (1844) who identified two Lower Carboniferous bryozoans | as being conspecific with two Devonian taxa (Millepora gracilis | and Millepora similis) described three years earlier by Phillips (1841). The Carboniferous taxa have been redescribed and named, below and elsewhere (Wyse Jackson & Bancroft, 1995a) as | Rhombopora cylindrica sp. nov. and Rhabdomeson progracile re- spectively. Millepora gracilis as originally described by Phillips (1841) contains specimens herein considered to belong to two \ genera — Rhabdomeson and Rhombopora. The third case of misidentification is that of Owen (1966). Rhombopora radialis ) Owen, 1966 is synonomised with Pseudonematopora turkestanica © (Nikiforova, 1948). More detailed discussion of the contrasting | } taxa is given in the discussion section of Pseudonematopora planatus sp. nov. and Rhombopora cylindrica sp. nov., and in Wyse Jackson & Bancroft (1995a) For completeness brief descriptions are given of the following = taxa which have been described more fully elsewhere: Rhabdo- - meson progracile Wyse Jackson & Bancroft 1995, Rhabdomeson | rhombiferum (Phillips, 1841), Baculopora megastoma (M‘Coy, | 1844), Diploporaria tenella Wyse Jackson, 1988, Thamniscus } colei Wyse Jackson, 1988, and Fistulipora incrustans (Phillips, _ 1836). Phylum BRYOZOA Ehrenberg, 1831 Class STENOLAEMATA Borg, 1926 Order CRYPTOSTOMATA Vine, 1884a | Suborder RHABDOMESINA Astrova & Morozova, 1956 | Family ARTHROSTYLIDAE Ulrich, 1882 Genus HEXITES Shulga-Nesterenko, 1955 TYPE SPECIES. Hexites triangularis Shulga-Nesterenko, 1955 by monotypy from the Lower Carboniferous of Chekhurskiv in the Russian Platform. EMENDED DIAGNOSIS. Arthrostylid with dendroid, erect zoaria | composed of small delicate branches, with polygonal cross-sec- tions. Perpendicular lateral branches occasionally developed. Jointing unknown. Autozooecial apertures oval to elliptical in shape, ar ranged in five to eight longitudinal rows and separated by a distinct) ridge. Autozooecial chambers triangular to sub-triangular in cross-) section. Zooecia seven to ten times longer than wide, diverging atal}) low angle with slightly inflated bases and sublinear chambers.) Hemisepta and diaphragms not present. Small acanthostyles fre-\) quently developed on ridges. DISCUSSION. The genus Hexites Shulga-Nesterenko 1955 was erected for distinctive small dendroid six-sided arthrostylids. Late Dunaeva (1974: 93) included the eight-sided variety Hexites quadrangularis. The taxon from County Fermanagh described here has very similar morphological features to that of Hexites triangula\) ris but differs from it in that as many as eight longitudinal rows ma} | be developed. It might be acceptable to erect a new genus tt} incorporate H. quadrangularis and the Irish form but is probably}. taxonomically unnecessary; rather the diagnosis of Hexites has beet} emended here to include all three taxa. STRATIGRAPHICAL RANGE. Lower Carboniferous (Viséan). DISTRIBUTION. Soviet Union). Only known from Ireland and the CIS (forme LOWER CARBONIFEROUS BRYOZOA lexites paradoxus sp. nov. Figs 3a, 4-5, 8 {OLOTYPE. BMNH PD9410; Upper part of the Glencar Lime- fone (Viséan, Asbian), Carrick Lough, County Fermanagh. ARATYPES. BMNH PD9411-9429; TCD.34012-34014, 34125, 4127, 34164, 34167, 42593c, all from the same locality and /orizon as the holotype. TCD.42514-42515, Upper part of the Glen- Limestone (Viséan, Asbian), Sillees River, County Fermanagh. ERIVATION OF TRIVIAL NAME. This Hexites species has between e and eight rows of autozooecial apertures, unlike the type species thich has six, so therefore is a paradox. IAGNOSIS. Hexites with a delicate dendroid zoarium. Branches e straight to gently curved with a polygonal to sub-polygonal _foss-section. Lateral branches diverge nearly perpendicular to the ain stem and are infrequently developed. Autozooecia developed . five to eight longitudinal rows around the complete branch. The “verse surface is seen only as a very thin groove. Autozooecial ertures are small, oval to elliptical in shape. Apertural rows are vided by sharp distinct longitudinal ridges. Stylets developed on _ dges and occasionally between successive autozooecial apertures. JESCRIPTION. Colonies are small, delicate, and dendroid. Branches “fe straight and of constant diameter along their length. Branch fOss-sections are polygonal to rarely sub-polygonal. Lateral branches ~a similar diameter diverge at unknown intervals perpendicular to ie main branch. There is a very slight increase in branch width at mifications. The largest fragment examined measured 9.8 mm in ngth. Autozooecia are developed in five to eight longitudinal rows ound most of the branch. In some specimens the reverse surface is ig. 3. Measurements taken on arthrostylid cryptostomes in this study. a, Hexies paradoxus sp. nov.; b, Nematopora hibernica Sp. NOV.; ¢, Pseudonematopora planatus sp. nov. BW = Branch diameter; AD1 = Autozooecia apertural diameter measured parallel to growth direction; AD2 = Autozooecia apertural diameter measured perpendicular to growth direction; AS1 = Autozooecia apertural spacing measured parallel to growth direction; AS2 = Autozooecia apertural spacing measured perpendicular to growth direction; AR = Number of longitudinal apertural rows around zoarium; Z2 = | Number of autozooecial apertures contained within a 2mm line measured parallel to growth direction. represented by a thin groove (Fig. 5). Autozooecia arise from a thin central axis and diverge from it a low angle. Chamber bases are slightly inflated. Chambers are sublinear in shape, 0.60 to 0.71 mm in length and at least ten times as long as wide. In cross-section they are triangular to polygonal in shape. The vestibule, which shallows distally, is orientated at an angle of between 45° and 60° to the zoarial surface. The distal wall is thin with slight thickening of the proximal frontal wall. Hemisepta and diaphragms are not present. Autozooecial apertures (0.28 x 0.14mm) occur in longitudinal rows that are separated by a sharp to rounded ridge 0.08 mm wide. They are oval to elliptical in shape, and occasionally narrow distally. They are regularly spaced one diameter apart within rows and one to two diameters apart between rows. Autozooecial aperture size is marginally greater and apertural spacing slightly less in those rows closest to the groove on the reverse of branches. One to two rows of small short acanthostyles 0.02mm wide occur on the crest of ridges. Interapertural areas may be smooth or be decorated with up to twelve acanthostyles in three rows. Table 1 Measurements of Hexites paradoxus (in mm). N=18. NM Xx Mn Mx CVw CVb BW 136 0.59 0.37 0.72 31,715) 8.43 ADI 152 0.28 0.20 0.43 10.55 7.00 AD2 tS7 0.14 0.08 0.22 10.88 6.64 AS1 142 0.22 0.09 0.42 14.08 4.78 AS2 158 0.26 0.13 0.40 14.81 7.31 DISCUSSION. Hexites is easily recognised by the arrangement of autozooecia in well-developed longitudinal rows, with strong interapertural ridges. 124 P.N. WYSE JACKSON Table 2 Quantitative comparison between Carboniferous Hexites species (dimensions in mm). AR BW H. paradoxus sp. nov. 5-8 0.37-0.72 H. triangularis Shulga-Nesterenko, 1955 6 0.18-0.38 H. quadrangularis Dunaeva, 1974 8 0.52 i Data from original sources. | | | AD1 AD2 AS1 AS2 0.20-0.43 0.08—-0.22 0.09—-0.42 0.13-0.40 0.17 0.08 Genus NEMATOPORA Ulrich, 1888a | H. paradoxus is only the third Hexites species to be described and | first outside of the CIS (former Soviet Union). It differs from the / type species H. triangularis Shulga-Nesterenko 1955 in having a- larger branch width, a variable number of autozooecial rows (five to eight and not the consistent six of the latter), no peristomes, andy acanthostyles in interapertural areas. It bears a close resemblance to { H. quadrangularis Dunaeva 1974, which has 8 rows of autozooecia. } However, H. paradoxus shows some morphological differences: branches are often thicker, autozooecial apertures are larger, ellipti-i cal to oval in shape, and are spaced considerably further apart. On i the basis of these morphological differences H. paradoxus is erected | as a new species (Table 2). | 0.16 0.12 0.18 0.15 STRATIGRAPHICAL RANGE. Lower Carboniferous (Viséan—Asbian). | DISTRIBUTION. Carrick Lough and Sillees River, County Ferman) agh, Ireland. TYPE SPECIES. Trematopora minuta Hall, 1876 by original desig- nation, from the middle Silurian of Waldron, Indiana, U.S.A. REVISED DIAGNOSIS. Arthrostylid with delicate, erect, dichoto- mously branching zoarium. Branches straight, circular to sub-circulat in cross-section. Autozooecia arranged in four to ten longitudinal, rows, either completely around branches or concentrated on one side of branch. Interapertural areas smooth with acanthostyles developed along ridges. Autozooecial apertures are oval to rhombic, dorsally, flared. —— i} STRATIGRAPHICAL RANGE. Middle Ordovician—Lower Permian. DISTRIBUTION. British Isles, Europe, North America, the CIf (former Soviet Union), Asia. Nematopora hibernica sp. nov. Figs 3b, 6—7,8) HOLoTYPE. BMNH PD9430; Upper part of the Glencar Lime) stone (Viséan, Asbian), Carrick Lough, County Fermanagh. | | Figs 4—5 Hexites paradoxus sp. nov. Upper part of the Glencar Limestone (Viséan, Asbian), Carrick Lough, County Fermanagh. 4, BMNH PD9410 (holotype); 4a, colony fragment comprising a thin slender octagonal to circular-shaped branch; autozooecia are arranged 1 distinct longitudinal rows divided by strong flexous ridges, and their apertures are oval in shape, x30; 4b, detail of 4a showing autozooecial | apertures and intervening ridges showing the disposition of small stylet: on the crest of ridges, x150. 5, BMNH PD9414 (paratype), reverse surface showing longitudinal groove, x12. Figs 6,7 Nematopora hibernica sp. nov. Upper part of the Glencar Limestone (Viséan, Asbian), Carrick Lough, County Fermanagh. 6, BMNH PD9430 (holotype); 6a, small colony fragment showing bifurcation of branches, and regular arrangement of autozooecial apertures in offset rows on obverse surface, x20; 6b, detail showing distal growing tip of branch and pyriform autozooecial apertures separated by thin interapertural walls patterned by a single row of smal’ stylets, x120. 7, BMNH PD9442 (paratype), reverse surface showing longitudinal rows of small nodes, x14. LOWER CARBONIFEROUS BRYOZOA Fig. 8 Hexites paradoxus sp. nov. Line drawing of external features of | BMNH PD9410; scale bar = 1 mm. fig.9 Nematopora hibernica sp. nov. Line drawing of external features of BMNH PD9430); scale bar = 1 mm. PARATYPES. BMNH PD943 1-9449: TCD.34015-34017; BELUM _ all from the same locality and horizon as the holotype. ERIVATION OF TRIVIAL NAME. reland. fable 3 Measurements of Nematopora hibernica (in mm), N=19. NM x Mn Mx CVw CVb 3W From the Latin hibernica meaning 3 176 0.59 0.41 0.76 TS) 9.81 \DI 176 0.33 0.21 0.44 8.63 9.84 \D2 172 0.12 0.08 0.20 10.34 6.89 Asi 165 0.27 0.11 0.52 22.04 6.04 $2 170 0.12 0.08 0.21 17.54 8.29 IAGNOSIS. Nematopora with delicate dendroid zoarium. Branches ichotomise irregularly, are straight in outline and subcircular in ‘Toss-section. Autozooecia are developed in four to five longitudinal Ows on one surface only. Autozooecial apertures are oval and jurrounded by small acanthostyles. Reverse surface barren, with our to five longitudinal rows of faint pustules. ESCRIPTION. Colonies are small, delicate, erect with irregularly ichotomising branches. The largest fragment examined measures BW able 4 Quantitative comparison between Carboniferous species of Nematopora (in mm). 125 16.4mm in length. Branches are straight (range of diameter from 0.41mm to 0.76mm), sub-circular in cross-section. Branch width increases slightly prior to bifurcation. Interapertural areas may bear one to two rows of small acanthostyles which surround autozooecial apertures. The reverse surface is barren, either smooth or with small acanthostyles occurring in four longitudinal rows (Fig. 7). Autozooecia are arranged quincuncially in four to five longitudi- nal rows, irregularly spaced within and between rows. Autozooecial apertures are oval to ellipical in shape, often nar- rower distally. Vestibules are steep-sided and shallow distally. DISCUSSION. From the study area only 22 fragments of Nematopora hibernica were found. All are hollow silicified fragments in which only the surface has been replaced, and consequently details of internal morphology are unknown. Externally Nematopora is very distinctive. The rhombic shape of the autozooecial apertures, and the abundance of acanthostyles resembles that of Rhabdomeson rhombiferum (Phillips), but unlike the latter autozooecial apertures do not occur all the way around the branch. N. hibernica is only the second species of Nematopora to be described from the British Isles Nematopora hexagona having been described from the Silurian (Wenlock) of Shropshire (Owen, 1962). Only 27 species of Nematopora have been described worldwide throughout its stratigraphic range (Goryunova 1985). Of these only seven, found in the U.S.S.R., Afganistan, and Japan, occur in the Carboniferous: N. afgana Termier & Termier, 1971; N. donbassica, Dunaeva, 1961; N. kusbasensis Trizna, 1958; N. ivanovi Shulga- Nesterenko, 1955; N. parvula Shulga-Nesterenko, 1955; N. tulensis Morozova, 1955; and N. sp. indet. Sakagami, 1962. STRATIGRAPHICAL RANGE. Lower Carboniferous (Asbian). DISTRIBUTION. Carrick Lough, County Fermanagh, Ireland. Genus PSEUDONEMATOPORA Balakin, 1974 TYPE SPECIES. Nematopora? turkestanica Nikiforova, 1948 by original designation from the Lower Carboniferous of the CIS (former Soviet Union). EMENDED DIAGNOSIS. Arthrostylid with slender dendroid zoarium, with occasional dichotomising branches. Branches are of constant width and are circular to semicircular in cross-section. Autozooecia occur in 6 to 16 longitudinal rows, and are budded in an annular manner. Autozooecial apertures are circular to oval in shape, with proximal peristomes. Autozooecia originate from a central axis. Skeletal cysts may be present in the exozone. Terminal diaphragms developed in some species. Acanthostyles are absent. STRATIGRAPHICAL RANGE. naisian—lower Viséan). Lower Carboniferous (lower Tour- . hibernica sp. nov. 0.41-0.76 /. afgana Termier & Termier, 1971 - 0.6-1.5 . donbassica Dunaeva, 1961 - 0.89-1.20 /. kusbasensis Trizna, 1958 10 0.96—1.00 t: ivanovi Shulga-Nesterenko, 1955 - 0.70-0.90 V. parvula Shulga-Nesterenko, 1955 - 0.25-0.30 . tulensis Morozova, 1955 0.30-0.40 . Sp. indet. Sakagami, 1962 8-10 1.00-1.10 bata from original sources. ADI AD2 AS1 AS2 0.21-0.44 0.08—0.20 0.11—0.52 0.08-0.21 0.24-0.32 0.12-0.14 0.29-0.36 0.12 0.18-0.22 0.08-0.12 - - 0.40-0.45 0.15 - - 0.30-0.35 0.07-0.10 — - 0.20-0.22 0.10 = ~ 0.24 0.13-0.16 = - P.N. WYSE JACKSON DISTRIBUTION. Ireland, England, Belgium, and the CIS (former Soviet Union). Pseudonematopora planatus sp. nov. Figs 3c, 10-15 | t HOLOTYPE. BMNH PD9450; Upper part of the Glencar Lime- | stone (Viséan, Asbian), Carrick Lough, County Fermanagh. PARATYPES. BMNH PD9451-9472, 9741; TCD.34018-34025, 34137, 34160, 34161, 42561, 42607; BELUM K2222; all from) same locality and horizon as above. TCD.42516-42519, Upper part | of the Glencar Limestone (Viséan, Asbian), Sillees River, County Fermanagh | DERIVATION OF TRIVIAL NAME. From the Latin planus meaning plain and unornamented. | DIAGNOSIS. Pseudonematopora with slender dendroid noon Branches dichotomise infrequently and at a high angle. They are’ circular to sub-circular in cross-section and are a constant width’ throughout their length. Autozooecial apertures occur in 6 to 8) longitudinal rows. They are circular in shape and have an arcuate, proximally located, peristomial rim. A faint longitudinal ridge oc- curs in the smooth interapertural area. Autozooecial budding is/ annular from a central axis. Chambers diverge at a low angle in the) endozone before bending in the exozone to become orientated at an’ angle of 60° to 70° to the zoarial surface. Zooecial walls are thin ini the endozone and do not thicken in the exozone. Terminal dia~ phragms may be developed. Skeletal cysts are lacking. | DESCRIPTION. Colonies are small, delicate and have irregularly dichotomising straight branches, which are circular in cross-section and undulatory in outline. The largest fragment examined is 13.2 mm in length. On no specimen were two dichotomies seen. Autozooecia occur in 6 to 8 longitudinal rows around the circum ference of the zoarium except for a thin barren area on the reverse They are budded from a distinct central axis in an annular pattern) Zooecial chambers diverge from the median wall at an angle of 10° to 15° in the endozone. The exozone is reached when the chambery bend fairly abruptly through 60° to 70°. The living chambers art orientated nearly perpendicular to the zoarial surface. The complet chamber is nearly four times as long as it is wide. Interzooecial wall are very thin in the endozone but thicken considerably in thy exozone. Basal diaphragms are not developed and the zooecia chambers are simple tubular structures. Interapertural areas are smooth with a single faint longitudine Figs 10-14 Pseudonematopora planatus sp. nov. Upper part of the Glencar Limestone (Viséan, Asbian), Carrick Lough, County Fermanagh; 10, BMNH PD9450 (holotype); 10a, colony fragment | showing cylindrical branch shape, with circular autozooecial apertures | developed in irregular longitudinal rows; apertures surrounded by proximal peristomes that extend beyond the branch margin giving an | uneyen outline, x12; 10b, detail of 10a showing the smooth interapertural areas, x80. 11, BMNH PD9452 (paratype), reverse surface showing longitudinal sinuous series of small nodes, x12. 12, BMNH PD9470 (paratype), tangential section showing the central axia region with a row of autozooecial either side of it, and the marginal protrusion of the proximal peristmes, x35. 13, BMNH PD9471 (paratype); 13a, longitudinal section showing thin axial region from which are budded autozooecial chambers and the thickened exozone, x20; 13b, detail of 13a showing brown bodies in autozooecial chambey trapped behind a thin linear terminal diaphragm (arrowed), x35. 14, BMNH PD9452 (paratype), transverse section showing radial arrangement of seven autozooecial chambers around the central axis, x35. LOWER CARBONIFEROUS BRYOZOA Fig. 15 Pseudonematopora planatus sp. noy. Line drawing of external features of BMNH PD9450; scale bar = | mm. Lidge developed between adjacent autozooecial rows. On the reverse surface the interapertural areas are slightly wider than those on the ybverse surface. A strong ridge may be developed there. Autozooecial apertures are small and circular. The apertures of he autozooecia adjacent to the reverse surface are divergent from it Fig. 11) and are marginally larger than those in other rows on the pbverse surface. Peristomes, situated proximally, are commonly : around apertures. Thin terminal diaphragms close off ome autozooecial apertures, behind which small circular brown bodies are found in chambers (Fig. 13b). These brown bodies, which fe similar in morphology to those reported by Morrison & Anstey 1979) in some Ordovician trepostomes, represent the degenerated _ of the polypide soft tissues. able 5 Measurements of Pseudonematopora planatus (in mm), N=13. NM x Mn Mx CVw CVb W 130 0.80 0.61 1.20 7.80 7.96 D1 130 0.18 0.10 0.26 10.84 8.60 D2 130 0.13 0.10 0.23 14.74 6.08 \S1 130 0.39 0.24 0.63 15.32 6.69 \S2 130 0.18 0.10 0.41 26.97 5.65 L2 130 3.6 3 5 11.31 13.50 ISCUSSION. Pseudonematopora is reported from outside the CIS ormer Soviet Union) for the first time. In the County Fermanagh una P. planatus is quite common. Only three other species have reviously been recorded, all from Lower Carboniferous strata: P. etchorensis Gorjunova, 1985, the type species P. turkestanica 227 (Nikiforova, 1948) [Balakin, 1974], and P. balakini Gorjunova, 1988. P. planatus differs from these three species in a number of respects. Zoarial width is narrower in P. turkestanica and the number of autozooecial rows is less. More importantly the autozooecial apertures in P. planatus are at least half the size as those of the other three species. Skeletal cysts are absent in P. planatus but may be developed in the other species. Terminal diaphragms have been reported from both P. turkestanica (Owen, 1966) and P. balakini (Gorjunova, 1988), and are present in P. planatus. Balakin (1974) noted that variation in zoarial width and fluctua- tion in the number of autozooecial rows in P. turkestanica are both large. P. planatus does not show such variation. Coefficients of variation for all measured parameters are low (Table 5). Variation within colonies is greater than variation between colonies in all features except zoarial width (ZW) and the number of autozooecia in a 2mm line (Z2). In these two cases variation within and between colonies is virtually identical. Rhombopora radialis Owen, 1966 is herein considered to be conspecific with Pseudonematopora turkestanica (Nikiforova, 1948). Comparison of Owen’s type material (LL.2984 holotype; LL.2985- 2989 paratypes; Upper Viséan; Treak Cliff, Castleton, Derbyshire) with illustrations of Pseudonematopora turkestanica from the former Soviet Union (Balakin 1974) shows these taxa to have a similar morphology. Pseudonematopora is characterised by autozooecia budded from a central axis in an annular fashion, with short cham- bers and terminal diaphragms often developed, circular apertures with proximal peristomes, and a lack of acanthostyles and metapores. Conversely, Rhombopora zoaria are dendroid, with long autozooecia containing hemisepta, and with oval zooecial apertures, many acanthostyles, and occasionally metapores. Pseudonematoporais avery distinct genus witha straight, occasion- ally branching zoarium, autozooecia budded from a central axis, aper- tures with proximal peristomes, and occasional terminal diaphragms. Externally the taxon resembles the cystoporate Cheilotrypa Ulrich 1884. However, internal structures and budding patterns in the two are quite different: inCheilotrypa diaphragms are present and autozooecia are budded from a central hollow axial tube (Utgaard 1983). Nematopora has been regarded as ancestral to Pseudonematopora (Balakin 1974) because the two taxa display a similar colony shape, autozooecial chamber shape, aperture size and shape, and budding pattern. However, in a computer-based phenetic study on the Rhabdomesina using cluster analysis of 44 features, Blake & Snyder (1987) suggested that Pseudonematopora was more closely related to Osburnostylus (88% similarity) than to Nematopora (78% simi- larity). Externally, however, Osburnostylus with rapid thickening of the zoarium at the level of the autozooecia apertures appears more different from Pseudonematopora than is Nematopora. Resolution of this phylogenetic problem may be achieved through finds of Pseudonematopora, extending its range, both geological and geo- graphical, and by more work on Palaeozoic bryozoans. STRATIGRAPHICAL RANGE. Lower Carboniferous (Viséan, Asbian). DISTRIBUTION. agh, Ireland. Carrick Lough and Sillees River, County Ferman- able 6 Quantitative comparison between Pseudonematopora species (dimensions in mm). | BW AR AD1 AD2 AS1 AS2 Z2 _—_ sp. nov. 0.61—1.20 6-8 0.10-0.26 0.10-0.23 0.24-0.63 0.10-0.41 3-5 ) balakini Gorjunova, 1988 0.88-1.10 - 0.35-0.45 0.22-0.26 = - 4 petchorensis 0.72-1.08 ~ 0.33-0.36 0.15-0.19 0.33-0.36 0.20-0.25 sy) | turkestanica (Nikiforova, 1948) 0.80—2.80 8-16 0.25-0.37 0.17-0.22 0.25-0.37 0.15-0.25 3-4 ata from original sources. 128 Family RHABDOMESIDAE Vine, 1884a Genus RHABDOMESON Young & Young, 1874 TYPE SPECIES. Millepora gracilis Phillips, 1841, by monotypy, from the Devonian of north Devon, England (for discussion relating to the problems with this type species see Wyse Jackson & Bancroft 1995a, 1995b). Rhabdomeson progracile Wyse Jackson & Bancroft, 1995 Fig. 16 Fig. 16 Rhabdomeson progracile Wyse Jackson & Bancroft 1995. Upper part of the Glencar Limestone (Viséan, Asbian), Carrick Lough, County Fermanagh; BMNH PD9473 (paratype); 16a, typical zoarial fragment showing cylindrical shape of branch, with the spiral arrangement of autozooecia in curved interlocking rows; autozooecial apertures are oval to elliptical in shape; one large acanthostyle is placed distally of apertures, x25; 16b, detail of 16a, x130. Fig. 17 Rhabdomeson rhombiferum (Phillips, 1836). Upper part of the Glencar Limestone (Viséan, Asbian), Carrick Lough, County Fermanagh; BMNH PD9485; 17a, growing tip of branch showing cylindrical colony form; autozooecia are arranged in longitudinal and obliquely intersecting rows; apertures are oval in shape, and narrow slightly distally; short blunt stylets surround each autozooecial aperture, x50; 17b, detail of 17a showing autozooecial aperture surrounded by stylets, x130. P.N. WYSE JACKSON | MATERIAL. BMNH PD9473-9484, TCD.34026-34028, BELUM | K3095, Upper part of the Glencar Limestone (Viséan, Asbian), Carrick Lough, County Fermanagh, Ireland. TCD.42520, Upper | part of the Glencar Limestone (Viséan, Asbian), Sillees River, } } { County Fermanagh. DESCRIPTION. Zoariaare dendroid, with cylindrical branches rang- ing in diameter from 0.61 to 1.07mm. There may be some increase in branch diameter prior to, or subsequent to, lateral branch develop- ment. Bifurcation is rare. The longest zoarial fragment examined | measures 8.1mm in length. Autozooecia are budded from a straight hollow cylindrical axis _ 0.14 to 0.29mm in diameter, in an annular or spiral pattern. In thin section autozooecial chambers are triangular to pentagonal in shape when seen in transverse section. Vestibules are orientated at a high angle to the zoarial surface. Acanthostyles arise as rods of granular calcite in the lower portions of the exozone. Interchamber endozonal walls are 0.1mm in width and are composed of an inner granular layer surrounded by a fine laminated skeleton. Autozooecial apertures are pyriform to oval in shape, and moder - ate to small in size. They are crowded or arranged in quincunx in 14 - to 18 longitudinal rows around the branch. Interapertural spacing is greatest longitudinally where apertures are spaced one diameter | apart and up to 5 in a 2mm line. Transversely adjacent apertures are | spaced less than one diameter apart. Autozooecial apertural dimen- | sions and spacing are approximately constant in each branch! fragment. However, some considerable differences are found be- | tween zoarial fragments. A large acanthostyle, up to 0.12mm in height, is always found i) distal to autozooecial apertures. Rare zoaria bear only this single acanthostyle (Fig. 16b); more frequently one or two smaller: acanthostyles lie proximal to the first in a longitudinal line between } adjacent autozooecial apertures. Acanthostyles are usually abraded, and appear as faint protruberances on the zoarial surface. DISCUSSION. A complete systematic description of R. progracile’ is given in Wyse Jackson & Bancroft (1995a). Rhabdomeson rhombiferum (Phillips, 1836) Fig. 17 MATERIAL. BMNH PD9485-9506; TCD.34029-34036, 42591b, stone (Viséan, Asbian), Carrick Lough, County Fermanagh. Ireland; TCD.42521-42524, Upper part of the Glencar Limestone (Viséan, Asbian), Sillees River, County Fermanagh. DESCRIPTION. The dendroid zoarium is composed of irregularly bifurcating delicate branches, with a polygonal or circular cross-)_ section, that range in diameter from 0.41 to 0.86mm. Branch width i remains approximately constant along their entire lengths. Branch: ing either by bifurcation or development of lateral branches at a high angle of between 68° and 90° from parent branch. There is ne increase in branch diameter prior to or subsequent to branch devel opment. In no specimen was there more than one bifurcation 0 L. lateral branch observed. Autozooecial apertures are moderate to large in size, pyriform tt)’ oval or ellipsoidal in shape, and are arranged in quincunx in eight te} eleven longitudinal rows around branches. Apertural size, shape and spacing is very variable around thi branch. A distinct barren area 0.25mm in width, with four longitudi) nal rows of small acanthostyles, is found on some branches and ca’ be regarded as delineating the branch reverse surface. In all zoanl the autozooecial apertures are long, thin, and oval in shape on thi reverse surface. Towards the obverse surface apertures becom|}) LOWER CARBONIFEROUS BRYOZOA y AH/AW oe wil @ @ e @|MD1 ADI | MD2 — e AD2 | fig. 18 Measurements taken on rhomboporid and hyphasmoporid cryptostomes in this study: ZD = Width of zoarium measured perpendicular to growth direction; MD = Metapore diameter; MD1 = Metapore diameter measured parallel to growth direction; MD2 = Metapore diameter measured perpendicular to growth direction; AH = Acanthostyle height from base to tip; AW = Acanthostyle width measured at its base; AD1 = Autozooecia apertural diameter measured parallel to growth direction; AD2 = Autozooecia apertural diameter measured perpendicular to growth direction; IWT1 = Autozooecia apertural spacing measured parallel to growth direction; [WT2 = Autozooecia apertural spacing measured perpendicular to growth direction. Z1 = Number of autozooecial apertures contained in 1mm/?; Z2 = Number of autozooecial apertures contained within a 2mm line measured parallel to growth direction; AR = Number of autozooecial apertural rows measured around zoarium; ET = Endozone thickness; TE | =Exozone thickness. ncreasingly pyriform and equidimensional. This variation in jpertural size is reflected in apertural spacing; these two parameters apertural size and apertural spacing) are inversely proportional to ach other. Immediately after branching, elongate autozooecial pertures developed around the complete circumference of the aughter branch. Differentiation of apertural dimensions occurs a within two or three generations along the branch. | Interapertural walls are gently sinuous or occasionally straight nd may be raised to produce a ridge between apertural rows. One or wo rows of small short acanthostyles (0.02—0.04mm in diameter) re developed along this ridge. When two rows are present they are 2parated by a distinct furrow. Autozooecial apertures are surrounded by 24 to 30 acanthostyles, various patterns. Commonly they flank only lateral margins and 'p to six acanthostyles may occur proximal to apertures. Less Fensnity acanthostyles are arranged in a rhombic pattern, with nly one acanthostyle proximal to apertures. ISCUSSION. A complete systematic description of Rhabdomeson hombiferum is given in Wyse Jackson & Bancroft (1995a), as well 5 a discussion of budding, branching and other features in nabdomesonids. Family RHOMBOPORIDAE Simpson, 1895 | Genus RHOMBOPORA Meek, 1872 YyPE SPECIES. Rhombopora lepidodendroides Meek, 1872 by 129 original designation, from the Upper Carboniferous of Nebraska City, Nebraska, U.S.A. Rhombopora cylindrica sp. nov. non 1841 = Millepora similis Phillips: 21, fig.32. non 1843 = Millepora similis Phillips; Morris: 42. 1844 Millepora similis Phillips; M‘Coy: 196. non 1854 Ceriopora similis (Phillips); Morris: 121. 1854b Millepora similis Phillips; M‘Coy: 104. 1862 Millepora similis Phillips; Griffith: 196. 1871 Ceriopora similis (Phillips); Young & Armstrong: 33. 1876 Ceriopora similis (Phillips); Armstrong, Young & Robertson: 46. 1877 Ceriopora similis (Phillips); Young & Robertson: 175. 1881 Ceriopora similis (Phillips); Vine: 338. 1885 Rhombopora similis? (Phillips); Vine: 93 pro parte. non 1887 Rhombopora persimilis Ulrich; Vine: 226, pl.1, fig.6. 1887 Rhombopora similis (Phillips); Vine: 226, pl.1, fig.7. 1889 Rhombopora similis (Phillips); Vine: 198. 1987 Rhombopora similis (Phillips); Bancroft: 196. HOLOTYPE. BMNH PD9507; Upper part of the Glencar Lime- stone, Lower Carboniferous (Viséan, Asbian); Carrick Lough, County Fermanagh. Figs 19-25 PARATYPES. BMNHPD9508-9534, 9576, upper part of the Glencar Limestone, Lower Carboniferous (Viséan, Asbian); Carrick Lough, County Fermanagh; Tavener-Smith and Wyse Jackson Collections. BMNH D294 (2 zoaria in a cavity slide of five), D295, Lower Carboniferous, Gayton Boring, Northamptonshire, England; Vine Collection. BMNH D303 (thin section of several zoarial fragments), Shales; Lower Carboniferous; Argyleshire, Scotland; Vine Collec- tion. TCD.28317, 28369, Nant-y-Gamar buildup, Llandudno Pier Dolomite Formation (Viséan, Asbian), near Llandudno, north Wales. TCD.34037-34044, 34122, 34126, 34128, 34165, 42592a, b; BELUM K2175, Upper part of the Glencar Limestone, Lower Carboniferous (Viséan, Asbian); Carrick Lough, County Ferman- agh; Wyse Jackson Collection. TCD.41515, Shales above Main Limestone, Pendleian, Upper Carboniferous, Hurst, near Richmond, Yorkshire, U.K. [NZ044 023], Bancroft Collection. TCD.42525- 42528, Upper part of the Glencar Limestone (Viséan, Asbian), Sillees River, County Fermanagh, Wyse Jackson Collection. DERIVATION OF NAME. branches. From the cylindrical nature of zoarial DIAGNOSIS. Rhombopora with zoaria comprised of irregularly dividing, thin, cylindrical branches. Autozooecia are budded in a spiral manner from a central linear axis. The exozone region is thin. Autozooecial apertures are oval in shape, moderate to large in size and arranged in quincunx in longitudinal rows around branches. Metapores are rare and occur proximal to autozooecial apertures. Stylets are common and structurally varied: characteristically one to two acanthostyles may be situated at junctions of interapertural walls, and many small heterostyles occur in interapertural areas. DESCRIPTION. Colonies are composed of delicate, thin bifurcating branches. The largest fragment examined measures 16.2mm in length. Branches range in diameter from 0.54 to 1.15mm and retain a constant width along their length except prior to lateral branch development when a 25% increase in diameter occurs. Bifurcation is infrequent and irregular; lateral ramifications deviate at high angles of between 75° and 87°. Autozooecia are budded from a straight to undulatory central axis Figs 19-24 Rhombopora cylindrica sp. nov.; 19-23, Upper part of the Glencar Limestone (Viséan, Asbian), Carrick Lough, County Fermanagh; 19, BMNH PD9509 (paratype), colony form showing long straight branches, with interconnected longitudinal and oblique rows of autozooecia, x20; 20, BMNH PD9507 (holotype); 20a, zoarial fragment with distal growing tip, showing regular arrangement of autozooecial apertures around branch; large acanthostyles are situated at the proximal and distal ends of apertures with smaller heterostyles developed on surrounding walls, x20; 20b, detail of 20a showing autozooecial apertures and acanthostyles on interapertural walls, x110; 21, BMNH PD9534 (paratype), transverse section showing radial budding pattern of autozooecia and the differentiation of endozone and exozone; 22, BMNH PD9532 (paratype), tangential section showing oval-shaped autozooecia and heterostyles (C-type stylets) developed on interapertural walls, x100; 23, BMNH PD9531, (paratype), longitudinal section showing autozooecial chamber shape and the thickened exozonal walls, x40; 24; Shales above Main Limestone, Pendleian, Upper Carboniferous, Hurst, near Richmond, Yorkshire, U.K. TCD.41515; 24a, longitudinal section, x25; 24b, detail of 24a showing morphology of acanthostyles, x80. at low angles of 10° to 25° and chambers are eight times as long as their maximum width. The chamber bends through 30° to 40° at the endozone/exozone boundary and vestibules are orientated at an angle of 45° to the zoarial surface. In cross-section chambers are rhombic, pentagonal or subcircular in shape. Chamber walls are thin (0.01—0.03 mm), compound (a very thin granular core covered by laminated skeleton) in the endozone and thicker, with a predomi- nantly laminated skeleton, in the narrow exozone region. The exozone varies in width between 0.1 and 0.2 mm and is approximately one fifth the width of branches. Thin terminal diaphragms may be present. Autozooecial apertures are large to moderate in size, oval to circular in shape, regularly spaced approximately one diameter apart, and spirally arranged in quincunx in 10 to 16 longitudinal rows around branches. 3 to 6 apertures occur longitudinally and 8 taj), 10 diagonally along a 2mm line. Autozooecial apertural size is) constant on a zoarium except at branch nodes. The first autozooecial apertures on new branches are long and thin, particularly on the reverse surface of branches. Uniformity of size is regained 4 to * apertures along branches. Metapores are rare. They are small (0.01- 0.13 x 0.02-0.10mm), irregular in shape and one or occasionally two are found proximal of autozooecial apertures, with other: sparsely distributed elsewhere on interapertural walls. They at usually developed close to branch divisions and zoarial thickening’ They originate within the exozone. Stylets are numerous and structurally varied. They occur in one 0 two rows, between autozooecial apertures. One or rarely twi acanthostyles (up to 0.07mm wide) occur at autozooecial apices 0) in w LOWER CARBONIFEROUS BRYOZOA | Fig. 25 Rhombopora cylindrica: sp. nov. Line drawing of external | features of BMNH PD9507; scale bar = 1 mm. ] some but not all zoaria. 20 to 24 heterostyles (0.01—0.03mm wide) in ne or two rows flank autozooecial apertures on all zoaria A longitudinal groove frequently occurs between heterostyle rows which probably marks the position of the zooecial boundary. Acanthostyles have a thick granular core and develop from the base pf the exozone. Heterostyles have a thinner granular core and grow from within the exozone. Skeletal lamellae are bent around hcanthostyles. {able 7 Measurements of Rhombopora cylindrica (in mm). N=23. xe Mn Mx CVw CVb 0.76 0.54 1.15 4.40 7.11 8.75 8 10 10.94 - v2 155 4.26 3 6 9.56 10.57 ADI 219 0.19 0.10 0.35 12222) 7.21 AD2 219 0.11 0.07 0.18 12.18 8.56 WT! 219 0.25 0.12 0.55 18.87 6.80 WT2 219 0.15 0.09 0.32 24.78 6.55 AD 1 Pil 0.04 0.01 0.13 32.17 2.04 AD2 21 0.04 0.02 0.10‘ 24.85 2.10 \H 34 0.04 0.01 0.10 27.86 1.79 \W 63 0.02 0.01 0.07 20.64 2.01 fE 18 0.15 0.10 0.20 13.53 5.66 DISCUSSION. Rhombopora cylindrica is quite distinctive and may € easily distinguished by the presence of oval-circular autozooecial pertures, a central axis, a thin exozone, and structurally varied canthostyles. Coefficients of variation for both zoarial (ZW) and zooecial AD1, AD2, IWT1, and IWT2) parameters within colonies are low. ‘Vw values for metapore diameter (MD1 and MD2) as well as ee height (AH) and width (AW) are large. They are due to € space-filling function of metapores, abrasion of acanthostyles, nd poor replacement by silica of small skeletal elements. This is eflected by examining autozooecia aperture dimensions which yere more varied in silicified specimens than in calcified specimens. ‘oefficients of variation between colonies are all extremely low. Millepora similis was first described by Phillips (1841) as a pposed coral from the Devonian of south-west England. Phillips 131 collected specimens from two localities: Cannington Park, north Devon, and Hope, near Torquay, south Devon. M‘Coy (1844) noted Millepora similis from the Lower Carboniferous of Ireland (the Courceyan of St. Doulagh’s, County Dublin and the Courceyan/ Chadian of Gort, County Galway). This identification was the first of many that confused two distinct taxa of Devonian and Carbonif- erous age. It is unfortunate that of the two slabs labelled Millepora similis from the Griffith Collection (NMING F7081, 7082) exam- ined by M‘Coy neither contains specimens referable to either taxa; but it is evident that M‘Coy described a taxon that is different from the Devonian Millepora similis of Phillips (M‘Coy, 1844: 196). Subsequently Morris (1854) classified Millepora as a zoophyte and transferred M. similis into the genus Ceriopora, considered then to be a coral, but now known to be a cyclostome bryozoan. Later still, Young & Robertson (1877) described some Carbonif- erous bryozoans from the Carboniferous of Scotland, which they regarded as being conspecific with Ceriopora similis. Vine (1881) followed this description but later (1885) deciding that the former generic assignment was incorrect, placed all Carboniferous mate- rial, as well as Phillips’ Devonian taxon, into the newly erected genus Rhombopora Meek, 1872. Rhombopora similis (Phillips, 1841) sensu Vine 1885 has only been found in strata of Carboniferous age. It is clear that M‘Coy (1844) misassigned a new undescribed Lower Carboniferous bryozoan and that this mistake was compounded and reinforced in later descriptions of Lower Carboniferous material. Phillips’ figured and only extant Millepora similis specimen (GSM 7110, ?Hope’s Nose Limestone, Middle Devonian (Eifelian), Hope, near Torquay, Devon, England) has been examined. It is a poorly preserved specimen which displays both rhomboporid and ptilodictyid affinities. The zoarium is composed of dendroid, mod- erately delicate, flattened lense-shaped straight bifurcating branches 1.35—1.80mm in diameter. Autozooecia are developed in eight to ten longitudinal rows. Autozooecial apertures are moderately large, 0.28 x 0.13mm, distinctly rhombic in shape, and closely packed less than one diameter apart. Interapertural walls are thin and appear to be smooth. A single proximal acanthostyle may be associated with autozooecial apertures. These features contrast with the cylindrical branches and oval to circular-shaped autozooecial apertures devel- oped in R. cylindrica. Vine’s figured material (BMNH D294-5: Vine 1887, pl.1, figs 7— 8) in the collections of the Natural History Museum, London, and some Vine material in National Museum of Wales, Cardiff has been examined, and all specimens are correctly assigned to the genus Rhombopora. They are not conspecific with Phillips’ Devonian taxon. The Carboniferous material represents a new taxon which is described and named here as Rhombopora cylindrica. A new epithet is required; similis of M‘Coy cannot be used on account of original misapplication of the name through misidentification (Article 49 — Code of Zoological Nomenclature, 1985). A holotype for Rhombopora cylindrica sp. nov. is designated from the Lower Carboniferous of Carrick Lough, County Ferman- agh, Ireland. STRATIGRAPHICAL RANGE. Carboniferous (Asbian—Pendleian). The range of Rhombopora cylindrica has been increased down- wards into the Asbian by its discovery in County Fermanagh and Nant-y-Gamar, north Wales, where the taxon is quite uncommon. DISTRIBUTION. Carrick Lough and Sillees River, County Ferman- agh and Nant-y-Gamar, north Wales. Previously recorded and described (see discussion) from the Brigantian of the Midland Valley of Scotland (Young & Armstrong 1871, Young & Robertson 1877) 132 and the Arnsbergian of Northamptonshire (Vine 1887) and Lanca- shire (Vine 1885), and the Pendleian of Yorkshire (Bancroft 1984, Vine 1881). Rhombopora hexagona sp. nov. Figs 26-31 HoLoTyPE. BMNH PD9535, Upper part of the Glencar Lime- stone (Viséan, Asbian), Carrick Lough, County Fermanagh. PARATYPES. BMNH PD9536-9564; TCD.34045-34048, 34121, 34154, 34170, 42591a, 42592c, 42602a; BELUM K2186, from the same locality and horizon as above, TCD.25687, near base of Michelinia Beds, Hook Head Formation (Courceyan), Locality 8 (of Dresser 1960), Lyraun Cove, Hook Head, County Wexford; TCD.25884, Michelinia Beds, Hook Head Formation (Courceyan), Locality 15 (of Dresser 1960), Brecaun Church, Hook Head, County Wexford, TCD.25885, Michelinia Beds, Hook Head Formation (Courceyan), Locality 40 (of Dresser 1960), Patrick’s Bay, Hook Head, County Wexford; TCD.25886, Linoproductus Beds, Hook Head Formation (Courceyan), Locality 92 (of Dresser 1960), Hook Head, County Wexford. DERIVATION OF TRIVIAL NAME. From the hexagonal pattern of heterostyles disposed around autozooecial apertures. DIAGNOSIS. Rhombopora with thin dichotomising cylindrical branches. Autozooecia are budded from a central linear axis in a spiral manner. Hemisepta are common: a robust inferior hemiseptum is present at the base of the vestibule while a thin superior hemiseptum is found on distal chamber walls high in the endozone. Diaphragms are absent. Autozooecial apertures are oval in shape, vary in size around branches, and are arranged in intersecting oblique and longitudinal rows around the zoarium. Small heterostyles are ar- ranged on ridges between autozooecial chambers in an interlocking hexagonal pattern. DESCRIPTION. Zoaria are composed of thin cylindrical dichotomising branches that form delicate erect dendroid colonies. No complete colonies were observed: the largest fragment measured 3.38mm in length. Branching is infrequent and irregular with either dichotomous bifurcation or lateral perpendicular ramification, producing second- ary branches which are slightly narrower than those from which they were derived. All branches retain a constant width along their length. Autozooecia deviate from a central axis in a spiral fashion at low angles of 18° to 27°. Chambers are sub-linear in shape, five to six times long as wide, with a slightly attenuated zooecial base. Cham- bers bend marginally at the exozone and vestibules are orientated at low angles of between 25° and 40° to the zoarial surface. Chamber walls are thin (0.01 mm) in the endozone, with a compound structure of a granular centre surrounded by thin skeletal laminae. The exozone is very thin (0.03—0.15mm). In cross-section chambers are polygonal in shape. Autozooecial apertures are large to moderate in size, oval in shape, and arranged in 9 to 15 longitudinally and diagonally inter- secting rows around branches. The angle of this intersection varies from 45° to 75°. Long, narrow autozooecial apertures occur on reverse surfaces, while more equidimensional apertures are found towards and on obverse surfaces. Interapertural spacing is inversely proportional to apertural size (see Tables 9 and 12, and Figs 26a and 28). Autozooecial apertures are surrounded by as many as 40 small, blunt, circular heterostyles (0.01mm in diameter) which arise from the base of, or from within the exozone. They are arranged in single, or occasionally several (particularly on reverse surfaces), rows along the crests of otherwise smooth interapertural walls, or occasionally Figs 26-30 Rhombopora hexagona sp. nov. Upper part of the Glencar Limestone (Viséan, Asbian), Carrick Lough, County Fermanagh; 26, BMNH PD95335 (holotype); 26a, colony form, x25; 26b, detail of 26a showing oval autozooecial apertures and disposition of heterostyles ina } hexagonal pattern on interapertural walls, x75; 27, BMNH PD9536 (paratype), colony fragment, x40; 28, BMNH PD9537 (paratype), view of ‘reverse’ surface showing long, thin autozooecial apertures, x25; 29, BMNH PD9560 (paratype), longitudinal section showing autozocecial chambers with thin superior (labelled ‘s’) and thick inferior (labelled ‘i’) ” hemisepta, x35; 30, BMNH PD9563 (paratype); 30a, transverse section showing circular branch outline, zooecia budded from a central axial area, with thin walls in the endozone and thicker walls in the exozone, x35: 30b, detail of 30a showing heterostyles on interapertural walls, x75.| LOWER CARBONIFEROUS BRYOZOA Fig.31 Rhombopora hexagona sp. nov. Line drawings of external features; a, colony form; scale bar = 1 mm; b, autozooecial apertures surrounded by heterostyles in hexagonal arrangement, scale bar = 0.1 mm. 0.30 | ria ee | 0.10 i aan : O ADI | ¢ AD2 mm 0.00 : ie2eomd 5) 6 7) 6584) 3) 21 Reverse Obverse Reverse Autozooecial rows ja 0.40 : 0.30 0.20 | Oo \wil ¢ IWwt2 mm 0!0 ———< ——— Se 1072 Sa 4s) 6) 7) wan) 4e5e2.) Reverse Obverse Reverse Autozooecial rows b ig. 32 Rhombopora hexagona sp. nov. Graphs of mean values of | aperture size and spacing; a, apertural size; b, apertural spacing. (For explanation of AD1, AD2, IWT1 and IWT2 see Fig. 18). 133 in an interlocking hexagonal to pentagonal pattern. These hexagons range in size from 0.66 x 0.25mm to 0.23 x 0.20mm, with the greatest dimensions occurring on reverse surfaces. Autozooecial apertures are generally situated distally within these areas. Hemisepta are common and are of two types.A prominent superior hemiseptum occurs within the endozone four-fifths along the cham- ber on distal walls (Fig. 29). They are thin, short (0.04—0.06mm) and have a similar skeletal structure to chamber walls. At the exozone the proximal chamber walls bend through 30° and thicken rapidly to form robust inferior hemisepta 0.13 long by 0.04mm thick. These have a sharp pointed distal extremity and bend marginally into the vestibules. They are composed of laminated skeleton in which lamellae are orientated parallel to the zoarial surface. Table 8 Measurements of Rhombopora hexagona (in mm). N=19. NM x Mn Mx CVw CVb Jé\D) 135 0.63 0.48 0.92 4.62 Uh Z2 25 5.23 4 6 9.93 WS AD1 145 0.16 0.10 0.26 20.94 8.32 AD2 146 0.09 0.04 0.18 15.57 5.13 IWT1 139 0.26 0.12 0.66 30.34 3.74 IWT2 136 0.17 0.07 0.39 27.71 4.37 AH 14 0.01 0.01 0.01 50.00 - AW 31 0.01 0.01 0.02 16.04 - ET 21 0.43 0.32 0.50 3.1/2 8.75 TE 42 0.08 0.03 0.15 17.37 2.96 DISCUSSION. Rhombopora hexagona 1s only the second species of the taxon, after R. cylindrica, to be recorded from Carboniferous strata of the British Isles and has previously been noted from Courceyan strata of Hook Head, County Wexford (Dresser 1960 MS).A previously recorded species R. radialis Owen, 1966 from the Viséan of Derbyshire is regarded as being an arthrostylid rather than a rhomboporid and is reassigned to the genus Pseudonematopora. Rhombopora hexagona is easily recognised externally from its cylindrical branches on which autozooecial apertures of varying dimensions (which is unusual) are surrounded by a hexagonal pattern of small heterostyles, and internally by the possession of two hemisepta of different sizes and a thin exozone. The relationship between autozooecial apertural diameter and apertural spacing is illustrated graphically in Fig. 32. Where aper- tures are long (high AD1) and thin (low AD2) longitudinal autozooecial spacing is moderate (low-high WT 1), and autozooecial spacing between adjacent rows is great (high IWT2). Where aper- tures are short and fat (low ADI values; high AD2 values), autozooecial spacing tends to be moderate and narrow (moderate IWT1 values; low IWT2 values). There is an inverse correlation between autozooecial apertural diameters AD1 and AD2 (Fig. 32a) and a moderately positive correlation between autozooecial apertural spacing IWT1 and IWT2 (Fig. 32b). In one specimen (C on Tables 9-12) where 14 autozooecial rows, as against the mean of 10, are developed, these correlations are not good. Dimensions of 56 other Carboniferous Rhombopora taxa are tabulated below. R. hexagona differs sufficiently from them, both morphologically and dimensionally, to justify its erection as a new species. It most closely resembles R. attenuata Ulrich 1890, in which two acanthostyle types are present, and R. gracilis Ulrich 1890, in which acanthostyles are developed at interapertural wall junctions only, but differs in acanthostyle development as well as in the size and spacing of autozooecial apertures. STRATIGRAPHICAL RANGE. Asbian). Lower Carboniferous (Courceyan— 134 P.N. WYSE JACKSON Table 9 Measurements of autozooecial aperture length (AD1) of Rhombopora hexagona around the zoarium from reverse to obverse surface (in mm). N=9 (A-I). SS eee ee ee SS Reverse Obverse Reverse | ROW 1 2 3 4 5 6 7 7 6 5 4 3 2 l | A - - OnW 0.15 0.15 0.13 0.15 0.15 0.14 0.13 0.18 0.18 - - | B - - 0.13 0.21 0.17 0.13 0.16 0.13 0.11 0.12 0.10 0.17 ~ ~ | € 0.22 0.18 0.19 0.18 0.16 0.13 0.13 0.12 0.12 0.13 0.13 0.17 0.20 0.20 { D - - 0.23 0.13 0.23 0.17 0.15 0.13 0.13 0.14 0.15 0.22 — - } E ~ - 0.19 0.23 0.17 0.17 0.20 0.10 0.10 0.14 0.14 0.16 - - 8 0.17 0.12 0.18 0.22 0.20 0.16 0.21 0.22 - - - G - - = 0.18 0.19 0.25 0.18 0.20 0.22 0.30 0.22 0.22 ~ - H - - 0.25 - 0.21 0.20 0.18 0.16 0.14 0.15 0.18 - = - | I - | 0.24 0.26 0.23 0.17 0.16 0.12 0.14 0.17 0.15 0.21 0.20 - - X 0.22 0.21 0.20 0.18 0.17 0.16 0.16 0.14 0.14 0.16 0.17 0.18 0.20 0.20. | | Table 10 Measurements of autozooecial aperture width (AD2) of Rhombopora hexagona around the zoarium from reverse to obverse surface (in mm). N50 (ASD. Ne SEE EEE EE eee Reverse Obverse Reverse | ROW 1 2 3 4 5 6 i] 7 6 5 4 3 2 1 A - - 0.08 0.07 0.08 0.08 0.09 0.08 0.08 0.07 0.09 0.07 - - B - - 0.09 0.07 0.12 0.10 0.10 0.07 0.08 0.08 0.07 0.06 - - C 0.08 0.08 0.10 0.06 0.09 0.10 0.10 0.08 0.10 0.09 0.09 0.08 0.09 0.06 D - - 0.12 0.08 0.09. 0.12 0.18 0.08 0.08 0.09 0.08 0.12 ~ - 18} - - 0.09 0.08 0.07 0.07 0.06 0.07 0.08 0.06 0.07 0.07 - - F - - - 0.16 0.13 0.14 0.14 0.11 0.11 0.13 0.13 - - - G - - ~ 0.11 0.11 0.10 0.11 0.10 0.09 0.12 0.15 0.13 - - H - - - 0.12 0.10 0.12 0.11 0.11 0.10 0.10 0.10 - - - | I ~ 0.04 0.06 0.08 0.07 0.09 0.08 0.08 0.09 0.11 0.08 0.07 - - Xx 0.08 0.06 0.09 0.09 0.09 0.10 0.10 0.08 0.09 0.09 0.09 0.08 0.09 0.06 Table 11 Measurements of interapertural wall thickness (IWT1) of Rhombopora hexagona around the zoarium from reverse to obyerse surface (in mm). N=9 (A-I). Reverse Obverse Reverse ROW 1 2 3 4 5) 6 7 i 6 5 4 3 2 1 A ~ - 0.19 0.18 0.35 0.31 0.13 0.15 0.20 0.18 0.18 0.21 - - B ~ - 0.42 0.29 0.54 0.28 0.25 0.23 0.35 0.30 0.26 0.30 ~ =.§ ce 0.19 0.22 0.20 0.17 0.24 0.25 0.28 0.26 0.30 0.22 0.20 0.20 0.22 0.36 D - - - 0.21 ORS 0.21 0.12 0.16 0.13 0.15 ~ - - ~ E - - 0.24 0.26 0.21 0.23 0.34 0.15 0.16 0.16 0.35 0.32 - - F - - - 0.43 0.35 0.32 0.18 0.23 0.13 0.12 0.14 - - - G - - 0.64 0.48 0.30 0.37 0.34 0.37 0.39 0.24 0.28 - - - H - ~ ~ 0.52 0.40 0.37 0.34 0.30 0.33 0.32 0.16 - - - I - 0.51 0.24 0.43 0.40 0.24 0.30 0.33 0.35 0.36 0.40 0.30 - - Xx 0.19 0.36 0.32 0.33 0.32 0.28 0.25 0.24 0.26 0.22 0.24 0.26 0.22 0.36 Table 12 Measurements of interapertural wall thickness (IWT2) of Rhombopora hexagona around the zoarium from reverse to obverse surface (in mm). N=9 (A-I). a Reverse Obverse Reverse ROW l 2, 3 4 5 6 7 7 6 5 4 3 2 1 A - - 0.24 0.23 0.18 0.17 0.16 0.21 0.22 0.22 0.13 0.26 - - B — - 0.17 0.31 0.39 0.17 0.20 0.21 0.17 0.23 0.22 0.29 - - E 0.32 0.35 0.30 0.21 0.29 0.15 0.13 0.11 0.12 0.13 0.13 0.18 0.17 0.18 D = - 0.17 0.10 0.21 0.22 0.20 0.23 0.19 - E - = 0.32 0.24 0.25 0.23 0.25 0.23 0.23 0.25 0.26 0.29 - - F _ = - 0.36 0.17 0.13 0.16 0.13 0.15 0.23 0.18 = - - G - - 0.25 0.25 0.14 0.13 0.07 0.06 0.12 0.18 0.23 — - - H 0.13 - 0.10 0.10 0.08 0.09 0.07 0.11 — - - I - 0.26 - 0.20 0.17 0.12 0.12 0.12 0.13 0.17 0.13 0.25 = - 0.25 0.17 0.18 Pad S ae) ns) S wo So So we n S i uD S iV S a S a S a S ex S a So 3 LOWER CARBONIFEROUS BRYOZOA 135 : Table 13. Quantitative comparison of Rhombopora cylindrica sp. noy. and Rhombopora hexagona sp. nov. with some other Carboniferous Rhombopora species (dimensions in mm). | AR : _R. cylindrica sp. nov. 10-16 _R. hexagona sp. nov. 9-15 +R. Ybifurcata Campbell, 1961 = R. nova Ceretti, 1963 ee 'R. multipora Foerste, 1887 20 R. prompta Gorjunova, 1988 = | R. johnsvalleyensis Harlton, 1933 - _R. nitidula Harlton, 1933 'R. millepunctata McFarlan, 1942 - *R. lepidodendroides Meek, 1872 - R. ampla Moore, 1929 = R communis Moore, 1929 - R. constans Moore, 1929 17 R. cortica Moore, 1929 = _R. fovata Moore, 1929 = -R. munda Moore, 1929 26 'R. muralis Moore, 1929 = R. pilula Moore, 1929 17 *R. tersiensis Nekhoroshev, 1926 - R. binodata Trizna, 1958 - R. floriformis Trizna, 1958 - |R. insinuata Trizna, 1958 = 'R. novitia Trizna, 1958 - R. perpera Trizna, 1958 - R. sarcinulata Trizna, 1958 - \R. simplex Trizna, 1958 - IR. charasensis Sakagami, 1972 - R. murthyi Sakagami, 1972 - R. diaphragmata Shulga—Nesterenko, 1955 - R. riasanensis Shulga-Nesterenko, 1955 = JR. variaxis Shulga-Nesterenko, 1955 - _’R. armata Ulrich, 1884 - 1R. crassa Ulrich, 1884 ~ JR. elegantula Ulrich, 1884 - . pulchella Ulrich, 1884 = IR. incrassata Ulrich, 1888b - R. ohioensis Ulrich, 1888b = R. angustata Ulrich, 1890 6? R. ?asperula Ulrich, 1890 ~ R attenuata Ulrich, 1890 = AR decipiens Ulrich, 1890 - dichotoma Ulrich, 1890 - R . exigua Ulrich, 1890 - . gracilis Ulrich, 1890 = R. minor Ulrich, 1890 = R. nickesi Ulrich, 1890 = R. simulatrix Ulrich, 1890 - . 2spiralis Ulrich, 1890 =- R. tabulata Ulrich, 1890 - . tenuirama Ulrich, 1890 10 -R. transversalis Ulrich, 1890 = . varians Ulrich, 1890 - R. pseudonovita Yang & Lu, 1962 = . Staffordotaxiformis Yang et al., 1988 ~ ZD Z2 AD1 AD2 0.50-1.10 4 0.10-0.35 0.07-0.18 0.40-0.90 5 0.10-0.26 0.04—0.18 1.00—-1.60 - 0.20-0.23 0.10-0.13 1.08-1.25 3 0.38 0.16—0.20 1.40 7] 0.15 0.09 2.52-0.27 = 0.30 0.20-0.22 0.60-0.80 5 0.19 0.10 0.4 5 0.29 0.10 0.6 - 0.14 0.06 1.00-3.60 4 0.16—0.29 0.09-0.21 1.00 = 0.31 0.17 1.00 3 0.28 0.14 1.00 - 0.29 0.14 1.80—2.70 4 0.28-0.29 0.16—-0.17 1.00-1.15 = 0.37 0.29 125) 3 0.33 0.18 1.00 = 0.28-0.34 0.20-0.25 1.50—1.70 4 0.43 0.26 1.50—2.40 5 0.16-0.25 0.08-0.15 1.69-2.00 4 0.50 0.4 1.60-1.70 7 0.50 0.45 1.20-1.40 6 0.35 0.30 1.15—1.30 5 0.30 0.25 2.00 3 0.60 0.50 1.60 5 0.14-0.15 0.08 1.10-1.50 6 0.15 0.13 2.50-3.80 5 0.19-0.26 0.12-0.18 iNe7/ J 0.21-0.3 0.10-0.14 2.00-3.00 5 = = 0.8 4 - - 2.00—2.50 5 0.25 0.15 1.00-1.10 - - - 2.50-4.50 5 = = 2.50 4 - - 0.88 4 - - 1.00-1.10 6 0.35 0.30 1.00-1.30 - 0.11-0.25 0.07-0.13 0.40—0.50 4 0.17 0.08 1.00—1.60 - - - 0.70-1.00 6 0.15 0.10 1.50-3.00 1] 0.15 0.10 3.00 4 0.12 0.12 0.60—0.80 - 0.11 - 1.30 7 0.10 = 0.50—0.90 ~ 0.12 - 0.40-0.90 - 0.17 - 1.00-2.10 6 0.12 - 1.50—2.00 5 0.21 - 1.00-1.50 5 0.18 0.12 0.40-0.50 5 0.11 0.08 2.50-4.00 5) 0.12 - 2.00—-4.00 7 0.12-0.13 = 1.10-1.70 6 0.12-0.20 0.07-0.11 1.14-1.53 4 0.16-0.21 0.07-0.10 Dimensions condensed from primary sources except where indicated: : Sabattini, 1972; *: Huffman, 1970; *: Sakagami, 1972; *: Ulrich, 1890; ': Lu, 1989. ISTRIBUTION. Carrick Lough, County Fermanagh, and Hook dead, County Wexford. Family HYPHASMOPORIDAE Vine, 1885 Genus STREBLOTRYFPA (Ulrich MS.) Vine 1884b YPE SPECIES. Streblotrypa nicklesi (Ulrich MS.) Vine, 1884b, by onotypy, from the Lower Carboniferous of Hurst, Yorkshire, Eng- and and Illinois, U.S.A. TYPES. Duncan (1949) has discussed the problem of Vine’s lost material and the implications for the type specimens. She designated a suite of specimens collected by Ulrich in North America (USNM 43311) as (neo)types. Indeed, she should have only designated one neotype. Hageman (1993) has designed one of Ulrich’s syntypes (USNM 114392) as the neotype. DISCUSSION. Confusion has existed over the authorship and the concept of the type species of the genus. J.M. Nickles of the U.S. Geological Survey sent specimens of a bryozoan from the Carbon- 136 iferous of Kaskasia, Illinois to Vine and Ulrich in 1883. Ulrich, in an 1884 manuscript, named them Streblotrypa nicklesi and this de- scription subsequently appeared in print (Ulrich 1890: 667). However, before the appearence of Ulrich’s paper, Vine (1884b: 391) pub- lished a description of the American specimens together with some specimens from Yorkshire and named them as‘. nicklesi, stating that this was the name used by Ulrich in his manuscript which as such ‘had no validity’. Yet in his subsequent papers Vine (1885, 1889) credits the genus and species, which he consistently names as nicklesi, to Ulrich. As was clear later Ulrich had in fact named the taxon S. nicklesi, and Hageman (1993) has corrected Vine’s error and quotes Ulrich’s spelling as the correct name of the type species, and credits Ulrich in Vine as the author. S. minuta, described as a new variety by Vine in 1885, was later considered by him to be a variety of S. ‘nicklesi’ (Vine, 1889). Unfortunately, Vine’s original specimens are lost, and it is possi- ble that the American and English forms are not conspecific. Therefore, what is now in question is the original concept of the species and the validity of Hageman’s designation of the neotype: should the concept be based upon the American specimens (as is generally agreed (Blake 1983)) or on the now missing British specimen? If it is shown that the original material belonged to two separate taxa then perhaps either the American or British material needs renaming. New collecting is needed at Vine’s original locality at Hurst, north Yorkshire, which may yield specimens of Streblotrypa, so that comparison with the American material can be made. Subgenus STREBLOTRYPA (STREBLOTRYPA) (Ulrich MS.) Vine 1884b DISCUSSION. Blake (1983: 590) recognized two subgenera in Streblotrypa: S. (Streblotrypa) and S. (Streblascopora) Bassler, 1952. This differentiation is based on the number of axial zooecia contained in the endozone, the location of metapores between autozooecial apertures, and the presence or absence of hemisepta. Species of S. (Streblascopora) display a distinct axial area with more than 10 axial zooecia. Metapores are frequently found beyond the lateral margins of autozooecia, and hemisepta are rare or absent. The opposite holds for S. (Streblotrypa). Streblotrypa (S.) pectinata Owen, 1966 Figs 33-36 v1966 Streblotrypa pectinata Owen: 144, pl.10, figs A-C. MATERIAL. BMNH PD9565-9579; TCD.34049, 34124, 34129, 34130, 34140: BELUM K3239, Upper part of the Glencar Lime- stone (Viséan, Asbian), Carrick Lough, County Fermanagh. TCD.42529-42530, Upper part of the Glencar Limestone (Viséan, Asbian), Sillees River, County Fermanagh. DESCRIPTION. Zoarium ramose and composed of cylindrical branches 0.67 to 1.04mm in diameter with a circular cross-section. Figs 33-35 Streblotrypa pectinata Owen, 1966; Upper part of the Glencar Limestone (Viséan, Asbian), Carrick Lough, County Fermanagh. 33, BMNH PD9565; 33a, zoarial fragment showing dendroid colony form, oval-shaped autozooecial apertures arranged in longitudinal rows, with three to four rows of metapores developed between, x35; 33b, detail of 33a, x150. 34, BMNH PD9578; 34a, tangential section showing autozooecial apertures surrounded by small metapores (arrowed), x100; 34b, detail of 34a, x100. 35, BMNH PD9579; 35a, longitudinal section showing thin exozone pierced by acanthostyles, x30; 35b, detail of 35a showing acanthostyles in exozonal wall — the calcite rods clearly deflect the skeletal laminae of the outer wall, x100. LOWER CARBONIFEROUS BRYOZOA Fig. 36 Line drawing of external features of Streblotrypa pectinata Owen, 1966 (BMNH PD9565); a, colony form, scale bar = | mm; b, detail of autozooecial apertures surrounded by metapores, scale bar = 0.1 mm. No complete colonies were observed; the largest fragment examined measured 10.2mm in length. Branches bifurcate at irregular inter- vals, and lateral ramifications diverge at a high angle of between 83° nd 90°. Branches retain a constant width between ramifications and there is only a slight increase just prior to and after branching. Autozooecial apertures are moderate in size (0.08-0.19 by 0.05— ).10mm), oval in shape, evenly spaced 2 to 2.5 diameters apart, and pecur in 12 to 20 longitudinal rows around the entire branch. Within ‘any one Zoarial fragment apertures are of approximately constant dimension. Small oval to occasionally circular-shaped metapores € abundant; twelve to twenty occur in 3 to 4 longitudinal rows petween the distal and proximal extremities of adjacent autozooecial ‘Apertures, and beyond the lateral margins either a single or double ow of metapores is present. In cross-section metapores have a thin eck and flare towards the endozone. They are approximately -).09mm deep; only a small proportion of metapores penetrate to the ase of the exozone. | Autozooecia are budded from an axial region in which axial fooecia are not present. Chambers are initally recumbent in the ‘tndozone and diverge from the branch axis at a low angle of less than '}5°. At the exozone they bend through 65° to become orientated early perpendicular to the zoarial surface. From this surface the yroximal wall of the vestibule slopes at a moderate angle, while the jistal wall is perpendicular (Fig. 35a). In cross-section chambers are entagonal and slightly inflated laterally. Chamber walls in the jndozone are thin (0.01mm) and composed of a granular core overed with very thin laminated layers. The walls thicken rapidly trazooidaly (up to 0.4mm) in the exozone; much of this expansion due to metapore development. Acanthostyles are small (0.02-0.05mm in diameter) and blunt nd are randomly distributed in interapertural areas where they lie at € proximal end of metapores. Acanthostyles arise at the endozone/ ozone boundary, thicken slightly laterally and have solid cores mposed of granular skeleton, around which is bent laminated eleton. They form dark granular circles on the zoarial surface. ble 14. Measurements of Streblotrypa pectinata in mm. N=13. 137 DISCUSSION. Streblotrypa pectinata is very rare in the Lower Carboniferous of the British Isles. From the limestones of County Fermanagh less than 20 zoarial fragments and a small number of specimens in section were found. The presence of metapores, small acanthostyles, and a thin exozone make this bryozoan very distinctive. Only three other species of Streb- lotrypa have been described from strata of Carboniferous age in the British Isles: Streblotrypacortacea(Owen 1966), Streblotrypaminuta (Vine 1889), and Streblotrypa nicklesi (Ulrich in Vine 1884b).S. pect- inatadiffers fromS. cortacea which possesses athick exozone and few metapores; S. minuta, in which sharp longitudinal ridges and a small number (6 to 8) of metapores are developed; and. nicklesi, which has 9 to 15 metapores and 12 longitudinal rows of autozooecial apertures. STRATIGRAPHICAL RANGE. Lower Carboniferous (Asbian). DISTRIBUTION. Apart from the occurrences at Carrick Lough and Sillees River, County Fermanagh Streblotrypa pectinata has previ- ously only been recorded from Castleton, Derbyshire (the type locality). Genus CLAUSOTRYFPA Bassler, 1929 TYPE SPECIES. Clausotrypa separata Bassler, 1929 by original designation from the Permian of Timor. DISCUSSION. The taxon has both trepostome and cryptostome features. Of the former the long autozooecial chambers, moderately thick exozone, many acanthostyles particularly associated with autozooecial apertures. The dendroid zoarial form, arrangement of autozooecial apertures and the presence of metapores strongly Suggest cryptostome affinities. I consider Clausotrypa to have stronger cryptostome than trepostome affinities. Bassler (1929) assigned the genus to the Order Cryptostomata, family Rhabdomesidae. Many Soviet authors have placed it in the suborder Rhabdomesina Astrova & Morozova, 1956 (Romantchuk 1970, Morozova 1970, 1981), while others assign the genus to the suborder Streblotrypina Gorjunova, 1985 (Gorjunova 1985). Blake (1983: 592) does not consider Clausotrypa to be a rhabdomesonid, while Gorjunova (1985) does. Blake & Snyder (1987) show, based on a cluster analysis of 44 characters, that Clausotrypa is rather unusual. It does not easily fall into any familial grouping. Recognising the obvious need for fuller taxonomic and compara- tive studies the genus is tentatively placed here in the family Hyphasmoporidae, Vine 1885. Clausotrypa ramosa (Owen, 1973) comb. noy. Figs 37-41 v1973 Sulcoretepora? ramosa Owen: 304, pl. 9a—c. HOLOTYPE. The holotype of Sulcoretepora? ramosa Owen, 1973 is represented by a zoarial fragment and three thin sections cut from it, collected from shales below the Rossmore Mudstone (upper Viséan), Tullaghoge, County Tyrone, in the collections of the Ulster Museum (BELUM K1830). MATERIAL. BMNH PD9577; 9627-9637; 9730-9739:TCD.34067- 34078, 34136, 34163, Upper part of the Glencar Limestone (Viséan, Asbian), Carrick Lough, County Fermanagh. TCD.42513, 42531- 42534, Upper part of the Glencar Limestone (Viséan, Asbian), Sillees River, County Fermanagh. DIAGNOSIS. Clausotrypa forming semi-robust erect cylindrical dichotomising zoaria. Autozooecia more frequent on one side of branch than the other. Metapores, closed to the surface, are devel- oped in interapertural areas. Autozooecial apertures are circular, moderate in size, widely spaced, and are surrounded by six to eight P.N. WYSE JACKSON acanthostyles. Strong undulating ridges and short acanthostyles are common in interapertural areas. DESCRIPTION. Zoaria form dendroid expansions of unknown maxi- mum height and are composed of cylindrical dichotomising or bifurcating branches 0.46 mm to 1.12 mm in diameter. Autozooecia are arranged in poorly defined longitudinal rows and are developed throughout the zoarium. They are budded from a | central undulating axis in an annular or irregular pattern. Autozooecial chambers have a sub-linear shape, are eight times as long as wide, and diverge distally from the axis at low angles of between 10° and 20°. They bend slightly at the exozone and vestibules are orientated |) at a high angle to the zoarial surface. In cross-section chambers are sub-rounded in shape. Endozonal walls are thin, undulatory, and | retain a constant width along their length. Chamber walls are © thickened in the exozone to a maximum width of 0.52 mm. The ) exozone averages 0.07 mm in width and is approximately one sixth of the branch diameter on either margin. Metapores are developed at the top of the endozone and the base |) of the exozone. They are small, circular to oval structures, usually , closed at the zoarial surface. One or two are disposed between |) autozooecia. Autozooecial apertures are moderate in size, oval in shape, widely | spaced approximately four to five diameters apart, and are arranged :| in longitudinal rows around branches. On most zoaria apertures are | more abundant on one side of branches than on the other. Acanthostyles are common. Six to ten surround autozooecial | apertures, often resembling a peristome, and they also occur ran- | domly and widely scattered in interapertural areas. They develop i) from the base of the exozone only. Strong longitudinal ridges also) decorate interapertural areas. DISCUSSION. Clausotrypa ramosa was first described as Sulcor-)) etepora? ramosa by Owen (1973: 305). He suggested that the taxon | is either a sulcoreteporid or arhabdomesonid depending on which of Owen), or the ramose zoarial form, is considered to be of stronger) generic importance. He assigned the taxon to the former, but ignored the diagnostic features of the genus Sulcoretepora, namely the bifoliate zoarial habit and the arrangement of autozooecia in longi- tudinal rows, budded from a plicated median carina. Several Claustotrypa species have been previously described. Of | Figs 37-40 Clausotrypa ramosa (Owen, 1973) comb. nov.; 37-39; Upper part of the Glencar Limestone (Viséan, Asbian), Carrick Lough, County Fermanagh. 37, BMNH PD9627; 37a, dendroid zoarium showing aborted lateral branch development, oval-shaped autozooecial apertures arranged in crude longitudinal rows, with striated ridged interapertural areas, x25; 37b, detail of 37a, showing oval-shaped autozooecial aperture surrounded by seven large acanthostyles. Acanthostyles are also developed in interapertural areas, x200. 38, BMNH PD9636, longitudinal section showing sublinear autozooecial chambers, budded from a poorly defined axis. Circular metapores are present at the endozone/exozone boundary, x25. 39, BMNH PD9637, transverse section showing pyriform autozooecial chamber cross- sections, circular/polygonal metapores, and acanthostyles in exozone, x25. 40, Limestone and Shales below Rossmore Mudstone (Upper Viséan), Tullaghoge, County Tyrone; Owen Collection. BELUM K.183¢ (lectotype); 40a, transverse section showing pyriform autozooecial chamber cross-sections, and circular/polygonal metapores (figured as Sulcoretepora ramosa by Owen, 1973, pl. 9c, x25); 40b, longitudinal section showing autozooecial chambers budded from an irregular axis, and small circular/polygonal metapores developed at the base of the thi! exozone region (figured as Sulcoretepora ramosa by Owen, 1973, pl. 9b, x25). LOWER CARBONIFEROUS BRYOZOA ‘TE 10 Fig. 41 Clausotrypa ramosa (Owen, 1973) comb. nov. Line drawing of external features of BMNH PD9627; a, colony form, scale bar = 1 mm; b, detail of autozooecial apertures; scale bar = 0.1 mm. ‘Table 15 Measurements on Clausotrypa ramosa (in mm). N=17. MN x Mn Mx (CW wCVb 2D 102 0.73 0.46 TEND) 7.56 5.97 AD1 107 0.11 0.06 0.16 13.06 9.68 AD2 106 0.08 0.05 0.14 14.87 7.67 IWT1 Si7/ 0.53 0.28 0.85 12.85 4.14 ~TWT2 64 0.25 0.15 0.42 17.38 4.38 Z2 13 3.4 2, 5 16.05 3.9 ET 6 0.42 0.31 0.52 10.60 4.34 0.07 0.03 0.17 14.54 1.33 these only one, C. limpida Gorjunova, 1988, is from the Carbonifer- ous, while all the rest occur in Permian strata. Comparison of C. 139 ramosa with these species shows it to be distinct from them all (see Table 16). It is morphologically most similar to C. monticula (Eich- wald, 1860) but differs significantly by having thicker branches and larger autozooecial apertures. Bassler (1929) regards Rhombopora? spiralis Ulrich 1890 from the Carboniferous of Kentucky as belong- ing to Clausotrypa. However, after examination of the original description and figures I consider the taxon to be correctly identified by Ulrich. STRATIGRAPHICAL RANGE. Lower Carboniferous (Asbian). DISTRIBUTION. Carrick Lough and Sillees River, County Ferman- agh and Tullaghoge, County Tyrone, Ireland. Order FENESTRATA Elias & Condra, 1957 Family ACANTHOCLADIIDAE Zittel, 1880 Genus BACULOPORA Wyse Jackson, 1988 TYPE SPECIES. Vincularia megastoma M‘Coy, 1844 by original designation, from the Lower Carboniferous (Viséan, Brigantian) of Killymeal, Dungannon, County Tyrone, Ireland. Baculopora megastoma (M‘Coy, 1844) Fig. 43 MATERIAL. BMNH PD8109-PD8127, TCD.29284-29303, TCD.34124, 34131, 34156, 34162; NMI: F19501-F19520; BELUM K3137, K3436, K12088-K12107, Upper part of the Glencar Lime- stone, Viséan (Asbian), Carrick Lough, County Fermanagh. BMNH PD8128-PD8132; TCD.42535-42538, Upper part of the Glencar Limestone (Viséan, Asbian), Sillees River, County Fermanagh. fig.42 Measurements taken on fenestrates in this study. a, Diploporaria tenella; b, Ichthyorachis newenhami:; ¢, Rhombocladia dichotoma. AD = Autozooecia apertural diameter; AD1 = Autozooecia apertural diameter measured parallel to growth direction; AD2 = Autozooecia apertural diameter measured perpindicular to growth direction; AR = Number of longitudinal autozooecial rows; AS = Autozooecia apertural spacing: minimum distance between two adjacent autozooecial apertures, measured from their centres; AS] = Autozooecia apertural spacing measured parallel to growth direction; AS2 = Autozooecia apertural spacing measured perpendicular to growth direction; AWT = Autozooecial chamber wall thickness; BW = Branch width: ET = Endozone thickness; HL = Hemiseptum length; LBW = Lateral branch width measured perpendicular to growth direction; LBS = Spacing between the centres of two successive lateral branches; MBW = Main branch width measured perpendicular to growth direction; NS = Nodal spacing: distance between two adjacent carinal nodes; TE = Exozone thickness; ZD1 = Length of autozooecial chamber; ZD2 = Width of autozooecial chamber; ZL2 = Number of autozooecial apertures contained in a 2mm line drawn parallel to growth direction; ZT = Zoarial thickness; ZT 1 = Number of autozooecial apertures contained in a 1mm line drawn perpendicular to growth direction; ZW = Zoarial width. 140 Table 16 Comparison of Clausotrypa species (dimensions in mm). P.N. WYSE JACKSON a ZD ADI AD2 IWT1 IWT2 C. ramosa (Owen, 1973) comb. nov. 0.46—1.12 0.06—0.16 0.05—0.14 0.28-0.85 0.15—0.42 C. limpida Gorjunova, 1988 0.6-0.9 0.25 0.13 -- C. clara Krutchinina, 1986 2.5-3.0 0.4-0.5 0.2-0.25 0.2 0.12 C. conferata Bassler, 1929 2.8 0.32 0.25 0.2 0.1-0.15 C. costata Romantchuk, 1981 4.2-4.3 0.21 0.16-0.2 0.3-0.6 0.43-0.64 C. exillis Sakagami, 1961 1.3-1.4 0.1-0.13 = = = C. minor Bassler, 1929 1S 0.2-0.25 0.15—-0.20 1.0 C. monstruosa Morozova, 1970 4.04.5 0.16-0.2 - - C. monticola (Eichwald, 1860) 1.0-2.5 0.17-0.2 0.12-0.14 - C. petaloides Romantchuk, 1970 4.8-5.0 0.24-0.25 - - - C. separata Bassler, 1929 0.15-2.6 0.3 0.17 0.4 0.3 C. spinosa Fritz; 1932 1.0-1.5 0.26-0.27 0.13-0.14 0.35 0.3 a Data from original sources. DESCRIPTION. Colonies consist of very slender branches that di- vide at irregularly spaced intervals, with branches bifurcating at low angles, and with lateral ramifications also occurring. Branches are straight or gently flexous, and are circular in cross-section. No complete colonies have been discovered, and the largest fragment examined was 1.53mm in length. Bifurcations and lateral branches appear to be widely spaced, as two laterals have not been observed on the same colony. Distal branches are thinner than proximal branches, with a range in diameter from 0.26mm to 0.36mm ob- served in one colony fragment. Branch width decreases slightly after bifurcation but soon increases to equal the width of the preceeding link. Lateral branches are thinner than main branches. The obverse surface bears faint undulating striae, occasionally with rows of small circular stylets (weathering to small pits) along the crests of striae. Autozooecial apertures are regularly arranged in quincunx, in four to seven longitudinal rows. They are small, circular, lack peristomes and are evenly spaced along the length of the branch. Some apertures are surrounded by six small pustules, giving them a stellate appearance. The reverse surface is smooth or faintly striated. Autozooecial chambers are rectangular in profile with pentagonal bases. Internally the skeletal arrangement is tripartite; a primary granu- lar layer is surrounded by an inner laminated layer lining zooecial chambers, and an outer laminated layer. Small stylets composed of granular skeleton penetrate through the outer laminated skeleton where they appear as pustules or weathered pits. DISCUSSION. A complete systematic description of the genus Baculopora and the species B. megastoma is given in Wyse Jackson (1988). Genus DIPLOPORARIA Nickles & Bassler, 1900 TYPE SPECIES. Glauconome (Diplopora) marginalis Young & Young, 1875, by original designation from the Upper Limestone Shales (Lower Carboniferous) of the British Isles (cited localities: Hairmyres, East Kilbride; Beghill, near Hamilton; Gillfoot, near Carluke; Hook Head, County Wexford). Diploporaria marginalis (Young & Young, 1875) Figs 44, 49 1875 Glauconome (Diplopora) marginalis Young & Young: 326, pl.3, figs 14-21. 1877. Glauconome (Diplopora) marginalis Young & Young; Vine, fig. 207. 1881 | Glauconome (Diplopora) marginalis Young & Young; Vine: B39) | | | 1885 Diplopora marginalis Young & Young; Vine: 83. 1900 Diploporaria marginalis (Young & Young); Nickles & Bassler: 233. 1953 Diploporaria marginalis (Young & Young); Bassler: G127, figs 87 — 6a, 6b. 1975 Diploporaria marginalis (Young & Young); Graham: 9, pl. 4, figs 6, 6a, 6b. 1987 Diploporaria marginalis (Young & Young); Bancroft: 196. 1 1991 Diploporaria marginalis (Young & Young); Billing: 41. i . LECTOTYPE. Graham (1975) designated a lectotype but cited al cavity slide that contained several zoaria. Consequently Bancroft! (1984-ms) designated specimen number 14 in cavity slide HM D144 (Hunterian Museum) as lectotype. This designation is formalised herein. MATERIAL. Three zoarial fragments, BMNH PD9580;TCD.34050- 34051, Upper part of the Glencar Limestone (Viséan, Asbian), Carrick Lough, County Fermanagh. DESCRIPTION. Zoaria are small non-pinnate expansions, composed) of delicate straight branches 0.21 to 0.34 mm in diameter with sub-) circular cross-sections. Lateral branches were not developed in the specimen examined. Autozooecia are arranged in two longitudinal rows along the length of the branch. Autozooecial apertures are small (0.07—0.09 mm in diameter), circular, and are surrounded by a complete peristome. They are regularly spaced two to two and a half diameters, apart either side of a median carina. The lateral margin and up to hall i the apertural diameter protrudes beyond the margin of the branch This produces a sharp serrated branch outline. A faint median carina carries regularly spaced oval to circulal nodes 0.02 mm in diameter. Two equally faint longitudinal ridges li¢ either side of the median carina inside the inner margins 0}; autozooecial apertures. Interapertural areas are smooth. The reversé surface is gently rounded and smooth. Internal features were no seen. Table 17 Measurements of Diploporaria marginalis (in mm). N=1. | NM Xx Mn Mx CVw CVby BW 10 0.27 0.21 0.34 4.40 Z| AD 10 0.08 0.07 0.09 10.52 = AS 11 0.29 0.28 0.33 28.85 = NS 6 0.30 0.28 0.32 22.69 = DIscussION. Diploporaria marginalis is very rare in the Asbian q County Fermanagh. In the present study only three zoarial frag 1" LOWER CARBONIFEROUS BRYOZOA 141 Fig. 43 Baculopora megastoma (M ‘Coy, 1844); Upper part of the Glencar Limestone (Viséan, Asbian), Carrick Lough, County Fermanagh. BMNH PD8109, obverse surface detail with five longitudinal rows of autozooecia and bifurcation of zoarium, x22. \Fig. 44 Diploporaria marginalis (Young & Young, 1875); Upper part of the Glencar Limestone (Viséan, Asbian), Carrick Lough, County Fermanagh. __ BMNH PD9580; 44a, obverse surface of branch fragment showing disposition of autozooecia in two longitudinal rows, one either side of a strong | median carina. The carina consists of adjacent longitudinal ridges, the central one of which bears distinct nodes. Autozooecial apertures are circular and | their edges protrude far beyond the branch margin, x40; 44b, lateral view showing elevated carinal nodes, x40. | Fig. 45 Diploporaria tenella Wyse Jackson, 1988; Upper part of the Glencar Limestone (Viséan, Asbian), Carrick Lough, County Fermanagh. BMNH | PD8138 (holotype), obverse surface detail of a slender zoarium with one row of autozooecial apertures developed either side of a central carinal ridge, x25. bad 46-47 = Ichthyorachis newenhami M‘Coy, 1844; 46, Upper part of the Glencar Limestone (Viséan, Asbian), Carrick Lough, County Fermanagh. BMNH PD9581, obverse surface showing a strong mainstem and broken lateral branches. Four longitudinal rows of autozooecia are developed on the former, fewer rows on the latter. Autozooecial apertures are small and circular in shape, x20. 47, Carboniferous Limestone (Dinantian), Kilmallock, | County Limerick. NMING:F6044 (lectotype), large colony fragment consisting of a straight mainstem with straight lateral branches diverging at moderate angles. Preservation of the specimen is poor and autozooecia of the mainstem are not seen; some lateral branches carry four autozooecial rows. Figured by M‘Coy, 1844, pl. 29, fig. 8, x0.8. apertures surrounded by complete peristomes, x20. ments were recovered. D. marginalis is a distinct species which can easily be recognised from its delicate zoarium with strongly serrated nargins caused by autozooecial apertures that project laterally. It was first described as a Glauconome species by Young & Young 1875). They noted the presence of small orifices proximal to iutozooecial apertures which were divided from them by a thin septum. Abrasion of this septum produced a pyriform aperture. Such ipertures and ‘orifices’ have not been observed by subsequent vorkers. They probably result from the abrasion of the zoarial urface and the revealing of the superior hemiseptum (Ulrich 1890, 3ancroft 1984). _ Vine (1881, 1885) added nothing to the original description of the pecies. He refers to a more robust form found in Scotland. However, is he does not illustrate these forms, and as his specimens are lost it Sy impossible to substantiate these records. While the two species of Diploporaria found in the British Isles D. marginalis and D. tenella) show morphological similarities there re a number of important differences between them. The jutozooecial apertures in D. marginalis possess a prominent outer eristomial rim which extends markedly beyond branch margins jiving branches a strongly serrated outline. In D. tenella peristomial ms are absent and autozooecial apertures hardly protrude beyond i branch margins giving them a smooth sinuous outline. Apertures € generally spaced further apart in D. tenella. Carinal nodes in D. Fig. 48 Thamniscus colei Wyse Jackson, 1988; Upper part of the Glencar Limestone (Viséan, Asbian), Carrick Lough, County Fermanagh. BMNH PD8959 (holotype), obverse surface detail showing circular bifurcating branches, autozooecia developed in three to four irregular rows, and circular tenella are more evenly and closely spaced than in D. marginalis. STRATIGRAPHICAL RANGE. Lower Carboniferous (Asbian- Brigantian). The range has been extended downwards into the Asbian for the first time. DISTRIBUTION. This is the first record of this species outside Great Britain where itis common in the Midland Valley of Scotland andrarer in Yorkshire and Lancashire. It is very rare in County Fermanagh. Diploporaria tenella Wyse Jackson, 1988. Figs 42a, 45 MATERIAL. BMNHPD8138-PD8149, PD8950-8958, TCD.29303- 29313, TCD.34132; NMI:F19521-F19530, BELUM K12108- K12117, Upper part of the Glencar Limestone, Viséan (Asbian), Carrick Lough, County Fermanagh. TCD.42539-42542, Upper part of the Glencar Limestone (Viséan, Asbian), Sillees River, County Fermanagh. DESCRIPTION. Colonies are very small and branches dichotomise irregularly. The largest fragment examined measured 5.9mm in length. Branches are slender, gently flexuous, and have a sub- circular cross-section. Lateral branches diverge at angles of between 70° to 80° from the main stem and slight flaring of lateral branch bases accompanies their development. Branch surfaces are smooth 142 to faintly pustulose. A narrow but prominent median carina is developed on the mainstem and lateral branches, and distinct nodes are regularly spaced on the carina at distances equal to the interapertural spacing. Autozooecial apertures are small, circular, and lack peristomial rims. They are regularly spaced (about twice their diameter apart) and are usually alternately arranged in two longitudinal rows on either side of the median carina, but may occasionally may be paired across the carina. The outer margins of autozooecial apertures protrude slightly beyond the lateral margin of branches, producing a gently sinuous branch outline. Internally the chambers are pentago- nal in transverse section and longitudinally rectangular. Hemisepta are not developed. DISCUSSION. A complete systematic description of D. tenella is given in Wyse Jackson, 1988. Genus ICHTHYORACHIS M ‘Coy, 1844 TYPE SPECIES. Ichthyorachis newenhami M‘Coy, 1844, by monotypy, from the Lower Carboniferous (Viséan, Chadian?) of Killmallock, County Limerick, Ireland. M‘coy’S ORIGINAL DIAGNOSIS. ‘Coral plumose, composed of a straight, central stem or midrib, having on either side a row of short, simple branches or pinnae, all in the same plane; obverse both of the midrib and lateral branches rounded, without keel, and each bearing several rows of small, prominent, oval pores, arranged in quincunx; reverse rounded, smooth or finely striated.’ EMENDED DiAGNosis. Acanthocladiid with pinnate zoarium composed of a mainstem and regularly-spaced, co-planar lateral branches which diverge from the mainstem at a high angle. Dissepiments are absent. Branches are circular to sub-circular in cross-section. Interapertural areas and the branch reverse surface are smooth or faintly striated. Autozooecia are arranged in 4 to 6 longitudinal rows on the mainstem, and in 3 to 4 rows on lateral branches. Autozooecial apertures are small, circular to oval in shape, regularly-spaced, and occur on the obverse surface only. STRATIGRAPHICAL RANGE. Lower Devonian—Lower Carbonifer- ous. DISTRIBUTION. British Isles, Europe, United States. Ichthyorachis newenhami M‘Coy, 1844 Figs 42b, 46-47, 51 v1844 Ichthyorachis newenhami M‘Coy: 205, pl.29, fig.8. 1854b Ichthyorachis newenhami M‘Coy; M*Coy: 104. 1857. Ichthyorachis newenhami M‘Coy; Jukes: 454. 1862 Icthyorachis [sic] newenhami M‘Coy; M ‘Coy, pl.29, fig.8. 1883 Ichthyorachis sp. Vine: 171. 1884a Ichthyorachis newenhami M‘Coy; Vine: 196. 1886 Ichthyorhachis [sic|nevenhami[sic] M‘Coy; Hoernes: 230, fig. 233. 1953 Ichthyorachis newenhami M‘Coy; Bassler: G128, fig. 88 (3a-c). 1966v Penniretepora triserialis Owen: 141, pl.9, figs A-C pro parte. LECTOTYPE. Herein designated NMING:F6044; Kilmallock, County Limerick (Viséan, Chadian?); C.B. Newenham Collection; figured M‘Coy 1844, pl.29, fig.8. This is the only extant specimen of Ichthyorachis from the collection on which M‘Coy based his de- P.N. WYSE JACKSON scription. Newenham also collected specimens from County Cork (M‘Coy 1844: 206). MATERIAL. BMNH PD9581-9590, TCD.34052, Upper part of the Glencar Limestone (Viséan, Asbian), Carrick Lough, County Fer- managh. } M‘COY’S ORIGINAL DIAGNOSIS. ‘Stem and lateral branches with} five rows of oval, prominent pores, closely arranged in quincunx; - reverse flattened, slightly convex, divided by a deep groove along}: the middle; obsoletely striated longitudinally; lateral branches half} the thickness of the midrib, space between them equal to the)}) diameter of the midrib.’ EMENDED DIAGNOSIS. Ichthyorachis with small delicate pinnate). zoaria. Two sets of straight lateral branches diverge from either side} of a thin, straight central main stem at a high angle. Lateral branches are regularly spaced and may be offset from each other either side of, the main stem, but more frequently occur paired. Branches are }_ circular to subcircular in cross-section. The obverse surface is} smooth and rounded, with faint longitudinal striae in interapertural) areas. The reverse surface is barren, rounded or slightly flattened: smooth, longitudinally striated, or with a central groove occurring }_ down the centre. Autozooecia are arranged in longitudinal rows 0 branches with 4 to 5 on the main stem and 3 to 4 on laterals. Autozooecial apertures are small, circular to oval in shape, lack peristomes, and are regularly arranged in quincunx. | DESCRIPTION. Zoaria form small pinnate expansions, consisting }/ of a main stem and lateral branches. The largest fragment in the } County Fermanagh assemblage examined is 17.3 mm in length (the | lectotype, which was collected in County Limerick, is larger, anc measures 53 mm in length (Fig. 47)). The main stem is thin, straighi or gently flexuous, and circular in cross-section. A small increase it main stem diameter precedes lateral branch development. Lateral}: branches lie in the same plane as the mainstem, and branch from i)}¥ at angles of between 50° and 60°. They are regularly spaced, abou! j= 2 diameters apart, and usually paired either side of the main stem 7 fe than the mainstem with a straight or undulatory margin, and / circular cross-section. The longest lateral branch observed is only 1) mm in length: most are broken at their bases. The obverse surface 1 }= rounded and faint longitudinal striae are developed along its lengt }_ The reverse surface is also rounded, and may bear indistinct longit ; dinal ridges or be smooth. 7 .- = eters apart. Apertural spacing becomes fractionally closer towards’ branch node. Details of internal features are unknown as only silicified fraj ments have been recovered from Carrick Lough. Discussion. Although only 10 colonies of Ichthyorachis newe }\, hami were measured it was found that they showed very lit| }, | LOWER CARBONIFEROUS BRYOZOA | morphological variation. Main stem width, lateral branch width, _ lateral branch spacing, and autozooecial apertural diameter all dis- play low coefficients of variation, both within and between colonies. | A slightly larger variation occurs in autozooecial apertural spacing. j Ichthyorachis newenhami is a distinctive but uncommon fenestrate _ form found in the Carboniferous of the British Isles. It has previ- i ously been reported only from Counties Cork and Limerick (M‘Coy | 1844), Hook Head in County Wexford (Courceyan) (Dresser 1960), and Castleton in Derbyshire (Vine 1883, 1884a). Examination of some of the type specimens of Penniretepora triserialis Owen, 1966 from the Upper Viséan of Treak Cliff, Castleton, Derbyshire (holotype: LL.2978; paratype: LL.2980) shows that autozooecia are arranged in three rows on mainstems and in two Ichthyorachis, and conceptually cannot be attributed to Penniretepora. The gross size of the specimens shows them to lie within the range exhibited by /. newenhami. The remaining type material of P. triserialis (paratypes: LL.2979, LL.2981-2983), also from the Upper Visean of Treak Cliff, Castleton, Derbyshire, has been examined and the ‘third row of autozooecia’ on mainstems are found to be the cores of abraided carinal nodes. These specimens belong to an indeterminate Penniretepora species and P. triserialis Owen, 1966 is herein regarded as a species inquirenda. | Lower Devonian species of Ichthyorachis are also known. I. Vreis occurs in the Helderbergian of New York (Hall 1874), and Ichtyorachis [sic] sp. has been found in the Emsian of France (Rondeau 1890, Bigey 1973). King (1849, 1850) referred Ichthyorachis to his genera Acanthocladia and Thamniscus owing to the similarities of the branching pattern with the former, and autozooecial arrangement with the latter. Comparison of the three taxa shows King’s reasoning 0 be incorrect. Ichthyorachis is very distinct from the other two ‘taxa. Branches in both Acanthocladia and Thamniscus are more robust and irregular than in Ichthyorachis, and distinct peristomial e are developed around autozooecial apertures in Thamniscus. 1 Ichthyorachis does, however, resemble Baculopora Wyse Jackson 1988 in the development of 4 or more rows of autozooecia on branches. However, the two taxa are generically distinct in that Ichthyorachis bears regular lateral branches on both sides of the ‘main stem and Baculopora does not. ' STRATIGRAPHICAL RANGE. Asbian). | _ DISTRIBUTION. | | Family FENESTELLIDAE King, 1850 | Genus THAMNISCUS King, 1849 YPE SPECIES. Keratophytes dubius Schlotheim, 1820 by original esignation from the Permian of Germany. Lower Carboniferous (Courceyan— British Isles. Thamniscus colei Wyse Jackson, 1988 Fig. 48 ATERIAL. BMNH PD8960-8982; TCD.29314-29323, CD.34152, 42606b; NMI:F19531-F19540; BELUM K2154, | <2157, K2166, K2223, K12118-K12127, Upper part of the Glencar i mestone, Viséan (Asbian), Carrick Lough, County Fermanagh. _ }MNH PD8959, 8983-8985; TCD.42543-42546, Upper part of the - blencar Limestone (Viséan, Asbian), Sillees River, County Ferman- gh. Boers The zoarium is small and develops from a flared _[asal holdfast to an open basket 7mm wide. The heavily calcified rows on lateral branches. This arrangement is characteristic of ; 143 holdfast is barren of apertures, whereas apertures open on the outer side of the basket. Branches are very robust (1.40 mm maximum width), circular in cross-section and bifurcate at irregular intervals at a high angle. Branches maintain a constant width along their length, and there is no increase in branch width prior to or following bifurcation. Branch surfaces are smooth. Autozooecial apertures are arranged in two to five sub-linear rows in an irregular pattern on the obverse branch surface. The outer peristomial rims of the lateral rows of apertures protrude slightly beyond normal branch margins giving them an undulatory appearance. Autozooecial apertures are circular, large and are surrounded by prominent thin peristomes. Autozooecial chambers are elongate, circular in cross-section, and narrow proximally. Discussion. A complete systematic description of ZT. colei is given in Wyse Jackson, 1988. Suborder PHYLLOPORININA Lavrentjeva, 1979 Family CHAINODICTYONIDAE Nickles & Bassler, 1900 Genus RHOMBOCLADIA Rogers, 1900 TYPE SPECIES. Rhombocladia delicatula Rogers, 1900 by original designation from the Upper Carboniferous of Kansas, U.S.A. REVISED DIAGNOSIS. Chainodictyonid with unilaminate, ramose zoarium, with dichotomous branches which are oval to elliptical in cross-section. Autozooecia are arranged in 4 to 12 longitudinal rows on obverse surfaces only. Superior hemisepta well developed; basal diaphragms rare. Autozooecial apertures oval. Reverse surface coy- ered with thin semi-circular ridges. STRATIGRAPHICAL RANGE. Lower Carboniferous—Lower Permian. DISTRIBUTION. British Isles, Europe, North America, the CIS (former Soviet Union), China. 49 50 Fig. 49 Diploporaria marginalis (Young & Young, 1875a). Line drawing of external features of BMNH PD9580; scale bar = 1 mm. Fig. 50 Ichthyorachis newenhami M‘Coy, 1844. Line drawing of external features of BMNH PD9581; scale bar = 1 mm. 144 P.N. WYSE JACKSON Figs 51-57 Rhombocladia dichotoma (M‘Coy, 1844) comb. nov.; 51-56, Upper part of the Glencar Limestone (Viséan, Asbian), Carrick Lough, County Fermanagh; 51, BMNH PD9591; 51a, obverse surface details of a growing tip; autozooecia are arranged in eight longitudinal and obliquely intersecting rows; autozooecial apertures are rhombic in shape, x40; 51b, detail of autozooecial apertures together with that of the hemiseptum developed from proximal walls; note the position of a single acanthostyle distal of autozooecia, x150; 52, BMNH PD9592, obverse surface of a branch with 11—12 rows of autozooecia, x20; 53, BMNH PD9593, reverse surface detail showing characteristic semi-circular pattern of basal laminae, x40; 54, BMNH PD9593, broken zoarial fragment showng typical cross-section shape of branches, x80; 55, BMNH PD9618, longitudinal section showing autozooecial chamber | shape, thin chamber walls, and long hemisepta, x50; 56, BMNH PD9614, shallow tangential section showing longitudinal arrangement of autozooecia, with cross-cut hemisepta, and heterostyles developed on interapertural walls, x40; 57, NMI.F6030, (lectotype), Carboniferous Limestone, Dinantian (Viséan); locality uncertain; Griffith Collection, reverse surface of branched zoarium showing characteristic semi-circular pattern of basal laminae, and | longitudinal lines where autozooecial walls meet basal wall; figured by M‘Coy, 1844, pl. 27, fig. 15 as Vincularia dichotoma; x2. Rhombocladia dichotoma (M ‘Coy, 1844) comb. nov. fig. 15. NMING:F7056-F7057, F7059-F7060; Black Lion, ne Figs 42c, 51-58 Enniskillen, County Cavan (Viséan, Asbian); Griffith Collectior NMING:F7061; Millicent, Clane, County Kildare (Viséan, Chadian Griffith Collection. NMING:F7486-F7489; Kildare, County Kij dare (Viséan); Griffith Collection. SMC:E5188; Howth, Coun Dublin (Courceyan/Chadian, Dinantian); Griffith Collectioy SMC:E5189/a-b; Killymeal, Dungannon, County Tyrone (Viséa Brigantian); Griffith Collection. SMC:E5190; Kildare, County Ki dare (Viséan); Griffith Collection. LecToryPE. NMING:F7058, here designated; Black Lion, near fareRIAL. BMNH D2564; ?Ireland (?Carboniferous); Shrubso| c Enniskillen, County Cavan (Viséan, Asbian); Griffith Collection. Collection. BMNH PD9591-9620: TCD.34053-34064, 3415} PARALECTOTYPES. NMING:F6030, here designated; no locality 34164, 34167, 42563a, b, 42565-42567, 42594, 42604c, 42606) given (?Viséan); Griffith Collection; figured M*Coy 1844, pl. 27, 42609; BELUM K2159, Upper part of the Glencar Limestot) v 1844 Vincularia dichotoma M‘Coy: 198, pl. 27, fig. 15. non1850_ -Vincularia dichotoma @ Orbigny, p. 195, pl. 682, figs 7— } 1854b Vincularia dichotoma M‘Coy; M‘Coy: 104. 1857 Vincularia dichotoma M‘Coy; Jukes: 454. 1862 Vincularia dichotoma M ‘Coy; Griffith: 198, 235. LOWER CARBONIFEROUS BRYOZOA | Fig. 58 Rhombocladia dichotoma (M‘Coy, 1844) comb. noy. Line drawing of external features; a, obverse surface (BMNH PD9591): b, reverse surface (BMNH PD9593); scale bar = | mm. |(Viséan, Asbian), Carrick Lough, County Fermanagh. TCD.42547- 42549, Upper part of the Glencar Limestone (Viséan, Asbian), Sillees River, County Fermanagh. )M‘Coy’s ORIGINAL DIAGNOSIS. ‘Dichotomous; obverse rounded, jwith about six equal, parallel, slender, longitudinal ridges, in the \concave furrows, between which are five rows of oval, prominent \cells, the marginal furrow on each side free of cells: reverse flat, with jnumerous, semicircular, scale-like wrinkles, and about six longitu- ‘dinal striae.’ JEMENDED DIAGNOSIS. Rhombocladia with a ramose zoarium of dichotomising branches oval to elliptical in cross-section. Seven to twelve longitudinal rows of autozooecial apertures open onto the obverse surface. A single superior hemiseptum is developed on the proximal side of apertures and basal diaphragms are rare. Inter- apertural areas smooth, or with small pustules. Autozooecial apertures oval to rhombic in shape; a single large acanthostyle occurs proxi- ally at each autozooecial aperture. Reverse surface barren, with parallel semi-circular ridges along the length of branches. DESCRIPTION. Zoaria are ramose with dichotomously dividing attened branches. In cross-section branches are oval to elliptical in shape, and convex frontally. The largest fragment examined meas- ures 30.4mm in length. Autozooecial apertures open on the obverse surface only, and are wranged into 7 to 12 longitudinal rows. The number of rows along branch is usually stable until a bifurcation point is reached, where he shape and size of apertures becomes variable and the regularity wf their organization is lost. Uniformity returns distally beyond ifurcations. Narrow, barren marginal zones on the obverse surface ire common in most zoaria. Autozooecial apertures are commonly val to rhombic, rarely an acute hexagonal shape. Oval apertures are ost common along branch margins, with rhombic shapes predomi- ating towards the branch centre. Interapertural walls are thin and smoothly rounded. They are mooth or bear faint pustules, and a single large acanthostyle, up to .18mm in length and 0.03—-0.08mm in width, is situated proximally f each aperture. In most zoaria the large acanthostyles have been braded down to the zoarial surface and are evident only as small eas of coarser skeleton. The reverse surface, formed by the colony basal wall, is undula- ory, very thin (0.1mm), and bears thin, parallel, semicircular lines ong the entire length of branches (apparently marking former ositions of the growing tips of branches). Where the basal wall is “braded a series of up to 10 longitudinal rows, representing the _roximal portions of autozooecial chamber walls, is visible. _| Autozooecial chambers originate on the basal wall and distally 145 Table 19 Measurements of Rhombocladia dichotoma (in mm), N=18. ————— ee ee eae NM Xx Mn Mx CVw CVb ZW 71 1.79 0.80 33,113} 7.95 4.39 ZT 23 0.47 0.30 0.56 8.16 8.97 AR 41 10 7 12 6.66 8.60 ZL2 134 4.89 4 ff] 7.26 8.19 ZT1 97 3.87 3 6 10.72 8.89 ADI 160 0.27 0.19 0.42 9.14 6.97 AD2 160 0.12 0.08 0.18 METS 6.64 AS1 160 0.09 0.07 0.32 19.81 4.50 AS2 160 0.07 0.04 0.13 14.13 4.63 AW 87 0.04 0.03 0.08 15.26 4.13 AH 21 0.12 0.02 0.18 17.29 0.65 HL 6 0.15 0.10 0.19 9.82 7533) ET 9 0.33 0.30 0.38 6.01 15.79 TE 8 0.13 0.09 0.17 10.86 3.81 AWT 7 0.01 0.01 0.02 16.67 9.19 —_—_—_—_———-_———————————— curve upwards at a low angle. At the junction of the endozone and exozone the chamber bends abruptly upwards. Here a prominent superior hemiseptum (average length 0.15mm) is developed, and is a little reflexed distally. Large acanthostyles originate at the base of the exozone. Endozonal walls are thin (0.01mm). Diaphragms are thin and are found only in the basal areas of the endozone. DISCUSSION. M‘Coy (1844) described and figured Vincularia dichotoma from the Carboniferous of Ireland. On the basis of M‘Coy’s types and conspecific specimens from the Viséan of County Fermanagh, this species is here reassigned to the genus Rhombocladia Rogers, 1900. Of the eleven specimens examined by M ‘Coy and still in existence, none show the obverse surface. M‘Coy must therefore have had additional specimens available which are presumed lost. The lateral margin of specimen NMING:F7058 is slightly worn and shows some detail of internal structure, enabling it to be compared to material from Carrick Lough, examined in the present study. All the material is conspecific and NMING:F7058 is here selected as the lectotype for the species Rhombocladia dichotoma. In many cases early workers assigned cylindrical Bryozoa to the genus Vincularia which is in fact a cheilostome genus. 16 species of Rhombocladia have been previously described. Of these 13 occur in the Carboniferous and have been recorded from from the United States (McKinney 1972, Rogers 1900), the CIS (former Soviet Union) (Dunaeva 1961, Gorjunova 1988, Shulga-Nesterenko 1955), the Carnian Alps (Ceretti 1963, 1964), and China (Lu 1989). Three species have been recorded from the Permian: R. aktashensis from the CIS (former Soviet Union) (Lavrentjeva 1985) and R. minor and R. spinulifera from Western Australia (Crockford 1944). Table 20 shows morphological measurements for all species of Rhombocladia. R. dichotoma differs significantly from all species except R. delicatula Rogers which has a similar zoarial thickness, and number of autozooecial rows. However, in R. delicatula promi- nent superior hemisepta are not present as they are in R. dichotoma. Rhombocladia dichotoma displays the greatest zoarial width of any of the taxa (maximum observed value = 3.13 mm) and the largest number of autozooecial rows. Apertural diameters within the genus are relatively constant. This explains the large width of branches observed in R. dichotoma. Zoarial thickness in all species is similar, with the exception of R. aktashensis whose branches are sub-circular in cross-section. The ratio of zoarial width to zoarial thickness in Carboniferous species is approximately 3:1, compared to 3:2 for the Permian species. McKinney (1972: 60) postulated an erect growth habit for Rhombocladia on the basis of the occurrence of a species of Hederella encrusting the reverse surface. This interpretation is questionable, 146 EXTERNAL Z2 IMS INTERNAL Fig. 59 Measurements taken on trepostomes in this study: ZD = Zoarial diameter measured perpendicular to growth direction; IMS = Inter- monticule spacing; AD = Autozooecia apertural diameter: maximum value seen in aperture; [WT = Interzooecial wall thickness; ED = Exilazooecia apertural diameter: maximum value seen in aperture; MD = Mesozooecial apertural diameter: maximum value seen in aperture; Z1 = Number of complete autozooecial apertures enclosed in 1mm*; Z2 = Number of complete autozooecial apertures measured along a 2mm line; AR = Axial ratio: this measures the ratio of the exozone to the zoarial diameter [AR = 2(TE)/ZD; the method employed follows that of Boardman (1960: 21) rather than that of Cuffey (1967: 31) who multiplied the ratio by 100, which makes the obtained values unnecessarily large]; ET = Endozone thickness; TE = Exozone thickness; FD = Diameter of ring septum foramen. and Hederella may have grown over adead skeleton ofRhombocladia lying face downwards on the sea-bed. Rhombocladia may have hadan encrusting habit as some Carrick Lough specimens show that the zoarium grew around and engulfed some adjacent pinnate bryozoan colonies. The perpendicular attitude of these pinnate colonies, which grew erect, relative to colonies of Rhombocladia, suggests that Rhombocladia grew horizontally on or just above the sea-bed. Table 20 Quantitative measurements for all described species of Rhombocladia (in mm). P.N. WYSE JACKSON _ STRATIGRAPHICAL RANGE. Lower Carboniferous (Tournasian | [Courceyan]—Viséan [Brigantian]). DISTRIBUTION. British Isles (Counties Cavan, Fermanagh, Kil-| dare, and Tyrone, Ireland; Mill Gill, North Yorkshire, England). | Order TREPOSTOMATA Ulrich, 1882 Suborder HALLOPOROIDEA Astrova, 1965 Family HETEROTRYPIDAE Ulrich, 1890 Genus LEIOCLEMA Ulrich, 1882 TYPE SPECIES. Callopora punctata Hall, 1858, by original desig- nation from the Lower Carboniferous of Iowa, U.S.A. Leioclema indentata sp.nov. Figs 60-62 HOLOTYPE. BMNH PD9621, Upper part of the Glencar Lime- stone (Viséan, Asbian), Carrick Lough, County Fermanagh. PARATYPES. BMNH PD9564, 9622-9626; TCD.34065-34066 34123, Upper part of the Glencar Limestone (Viséan, Asbian)| Carrick Lough, County Fermanagh. TCD.42550, Upper part of the Glencar Limestone (Viséan, Asbian), Sillees River, County Ferman: agh. TCD.28152, Base of reef, Lower Carboniferous (Viséan Asbian), Shanvaus Cross, County Leitrim, G.D. Sevastopulo Col} lection. DERIVATION OF TRIVIAL NAME. From the indentation of the autozooecial apertures by acanthostyles. DIAGNOSIS. Leioclema forming ramose erect cylindrical, irregu_ larly dichotomising zoaria. Zooecial walls considerably thickene¢ in exozone. Diaphragms present at zooecial bend. Mesozooeci: common between chambers in exozone, closed to the surface Autozooecial apertures indented by several large acanthostyles. DESCRIPTION. Zoaria are robust, composed of straight, cylindricé I branches 1.15 to 2.13 mm in diameter. Branches are circular 1 cross-section and bifurcate at irregular intervals. There is a sligh increase in branch diameter prior to division; subsequent branche are slightly narrower. The largest fragment examined measures 24.) |. mm long. ZW ZT AR AD1 AD2 AS R. dichotoma (M‘Coy,1844) 1.79 0.47 7-12 0.27 0.12 0.07-0.14 R. aktashensis Laurentjeva, 1985 1.50-1.80 1.00-1.1 6-8 0.22-0.24 0.12 0.10-0.15 R. borissiaki Shulga-Nest., 1955 1.00-1.35 0.45-0.50 7-8 0.26—-0.34 0.12-0.14 - R. carnica Ceretti, 1964 0.60 - - 0.30 0.13 0.09-0.10 R. coronata Shulga-Nest., 1955 1.10 0.45 6-8 0.19-0.20 0.08-0.15 0.20-0.27 R. delicatula Rogers, 1900 1.50 0.52 10 0.26 0.15 0.10 R. johnseni Ceretti, 1964 1.07 0.50 7 0.20 0.21 0.09 R. kasimovensis Shulga-N., 1955 0.80-1.20 0.45-0.70 5-7 0.15-0.22 0.08-0.12 - R. minor Crockford, 1944 0.46-0.57 - 2-3 0.24-0.29 0.10-0.15 = R. multispinosa McKinney, 1972 1.4 <1.00 9 0.30-0.42 0.12-0.19 - R. ninae Shulga-Nest., 1955 1.70-2.30 0.70-0.80 8-9 0.20-0.22 0.10-0.12 0.22-0.28 R. orientalis Lu, 1989 - 0.30-0.50 - 0.22 0.16-0.18 - R. punctata Dunaeva, 1961 1.00-1.08 0.48 - 0.24—0.27 0.12 - R. ramosa Gorjunova, 1988 1.10-1.32 0.86-0.88 6-7 0.18-0.22 - - R. septata Shulga-Nesterenko, 1955 1.25 0.45-0.52 il 0.27-0.32 0.12-0.14 0.15-0.20 R. spinulifera Crockford, 1944 0.70-0.95 - 4-6 0.19-0.24 0.16 - b R. tenuata Shulga-Nesterenko, 1955 0.80 0.50 7 0.18—0.20 0.10-0.12 - Lal Data from original sources. LOWER CARBONIFEROUS BRYOZOA 147 Figs 60,61 Leioclema indentata sp. noy.; 60, Upper part of the Glencar Limestone (Viséan, Asbian), Carrick Lough, County Fermanagh; BMNH PD9621 | (holotype); 60a, ramose colony with circular autozooecial apertures arranged in crude longitudinal rows, surrounded by large acanthostyles, with ridged interapertural areas, x15; 60b, detail of 60a, showing oval-shaped autozooecial apertures surrounded by six to eight large acanthostyles; smaller very thick exozone region, x30; 61, base of reef, Lower Carboniferous (Viséan, Asbian), Shanvaus Cross, County Leitrim, TCD.28152; 61a, ramose | acanthostyles are developed in interapertural areas, x85; 60c, cross-sectional view showing central axial region of thin walled autozooecial chambers and . colony., x20; 61b, detail of 61a showing small circular mesozooecial apertures between autozooecial apertures, x100. | Autozooecia are developed throughout the zoarium. The exact shape of the zooecial chambers is unknown. It appears that the cham- bers diverge from the centre of the branch at a low angle of 25° to 30°. Zooecia bend through 50° at the endozone/exozone boundary so that vestibules are orientated perpendicular to the zoarial surface. Cham- ber walls are thin (0.01—0.02 mm) in the endozone and thicken rapidly at the exozone which reaches a maximum thickness of 1.10 mm.Thin diaphragms are common at the base of the vestibule. In cross-section chambers are circular, pentagonal or polygonal in shape. 1 ig.62 Leioclema indentata sp. nov. Line drawing of external features of _ | BMNH PD9621; a, colony form, scale bar = 1 mm; b, Detail of surface | features, scale bar = 0.1 mm. Mesozooecia are common in the exozone, where they are dis- posed in three to five irregular rows between autozooecial chambers, but are not present in the endozone. They are circular, polygonal or irregular in shape, 0.03 to 0.11 mm in diameter, thin walled, hollow, and closed to the surface. No internal diaphragms were observed, perhaps due to the nature of the preservation. Autozooecial apertures are oval in shape, moderately large 0.12 to 0.20 mm in diameter, and fairly regularly spaced one to two diam- eters apart. They are arranged in 16 to 20 poorly defined longitudinal rows. Each aperture is surrounded by six to seven large acanthostyles (0.05 to 0.08 mm in width and up to 0.1 mm in length), which indent the aperture margin. Smaller ramdomly orientated acanthostyles, and thin wavy longi- tudinal grooves are found in interapertural areas which are slightly Table 21 Measurements of Leioclema indentata (in mm). N=6. NM x Mn Mx CVw CVb ZD 61 1.66 i315) DAB! 6.92 7.38 AD 60 0.15 0.12 0.20 9.40 9.88 IWT 60 0.18 0.06 0.26 17.33 9.22 Zi 60 6.23 4 8 10.48 11.11 Z2 60 3.60 3) 5 13.48 9.11 ET 9 0.84 0.68 1.10 2.69 S1117/ TE 18 0.38 0.28 0.51 8.89 6.68 148 Table 22 Quantitative comparision of Leioclema indentata with other Carboniferous Leioclema species (dimensions in mm). P.N. WYSE JACKSON ZD AD IWT Z2 ET TE L. indentata sp. nov. 1.15-2.18 0.12-0.20 0.06—0.26 4-8 0.68-1.10 0.28-0.51 L. porosum Crockford, 1947 20 0.2—0.25 0.08 - - - L. shumaradi Girty, 1908 3F5 0.15-0.20 0.20 - - = L. punctatum (Hall, 1858) 2.00—5.00 0.15—0.20 = 5-6 = = L. pushmatahensis Harlton, 1933 0.73=1.02 - - - - L. avonense Lee, 1912 22.00 ~ - 6 - = °L. hirsutum Moore, 1929 1.00-1.40 0.16—0.20 0.1 - 0.9 0.40-0.45 L. tubulosa Nekhoroshey, 1956 3.00-5.00 0.13-0.17 - 5.5-6.5 - = L. bifoliata Nikiforova, 1927 4.0 0.20-0.25 0.1 = 12 0.8 ?L. kobayashii Sakagami, 1962 2.60-3.60 0.21-0.29 0.32-0.40 = ~ - ?L. uzuraense Sakagami, 1964 = 0.16—0.22 - 10 - - L. cassis Trizna,-1958 1.80 0.18—0.20 - 10 - = L. echinata Trizna, 1958 1.60 0.16—0.18 - 7-10 = = L. podunskense Trizna, 1958 2.20-3.20 0,15-0.18 = 5-7 - - L. rojkiensis Trizna, 1958 2.00 0.20-0.22 - 6-7 - - L. semetra Trizna, 1958 2.75-3.00 0.14-0.16 = 11-12 - - L. textila Trizna, 1958 0.80-0.90 0.10-0.16 - 6 = - L. clara Trizna, 1962 3.30 0.16—0.20 - 8-9 ~ - L. gracillimum Ulrich, 1888b 1.00-1.50 0.10-0.15 - 8-9 - — L.araneum Ulrich, 1890 Adnate 1.00 0.2 - 9-10 - - L.foliatum Ulrich, 1890 Adnate 1.00—1.50 0.2 - 6 - - L. subglobosum Ulrich, 1890 - 0.15—0.20 ~ 8-9 - - L. washsmuthi Ulrich, 1890 Adnate <1.00 0.2 - 6 - - Data from original sources. elevated above the autozooecial apertures. Monticules are not devel- Dyscritella miliaria (Nicholson, 1881) Figs 63-66 ) OSes v1881 Monticulipora tumida var. miliaria Nicholson: 123, pl.3, DISCUSSION. Leioclema indentata is very rare in the Viséan of fig.2. ; ¥ . a ‘ | County Fermanagh. Only ten zoarial fragments were found; all of 1884c Monticulipora tumida var. miliaria Nicholson; Vine: 101. | these are silicified and consequently internal structures are not well 1912 Dyscritella miliaria (Nicholson); Lee: 178, pl.16, figs 7 known. However, some fine skeletal features such as autozooecial 1950 Dyscritella miliaria (Nicholson); Termier & Termier, p.15, diaphragms are preserved, and are seen in broken fragments. pl.5s, figs 4-6, 9, pl.60, fig.7, pl.65, figs 7-9. } L. indentata can easily be recognised by the indentation of the 1987 Dyscritella miliaria (Nicholson), Bancroft: 196. [ autozooecial apertures by six to seven large acanthostyles, and from LECTOTYPE. Herein designated AUGD 10135a, Carboniferous surface ornamentation. Coefficients of variation for measured pa- rameters are all low, including that for interapertural wall thickness (IWT). 25 other Leioclema species have been described from the Carbon- iferous (McKinney 1973, Astrova 1978, and herein), of which only L. avonense Lee, 1912, from the Avon area, and Leioclema sp. from North Wales (Bancroft, Somerville & Strank, 1988), occur in the British Isles. All are compared with L. indentata in Table 22 above. L. avonense is unusual in that the zoarium is very thick, exozone walls are not thickened, and that mesozooecia are few. Thus, its taxonomic position 1s debateable, and may only be resolved by examining Lee’s original material. L. indentata has a gross morphology similar to the North Ameri- can speciesL. gracillimum Ulrich, 1888b. However, apertural spacing and acanthostyle diameter are considerably less in the latter. STRATIGRAPHICALRANGE. Lower Carboniferous ( Viséan, Asbian). DISTRIBUTION. Counties Fermanagh and Leitrim, Ireland. Suborder AMPLEXOPORINA Astrova, 1965 Family DYSCRITELLIDAE Dunaeva and Morozova, 1967 Genus DYSCRITELLA Girty, 1911 TYPE SPECIES. Dyscritella robusta Girty, 1911, by original desig- nation from the Lower Carboniferous of Arkansas, U.S.A. shales; Redesdale, Northumberland, England; Nicholson Collec- tion. PARALECTOTYPES. Herein designated AUGD 10135b (2 speci- mens in cavity slide); Carboniferous shales; Redesdale,, Northumberland, England; Nicholson Collection. RSM 1967.66.384, |” RSM 1967.66.387; Carboniferous shales; Redesdale, Northumber-~ land, England; Nicholson Collection. 1 MATERIAL. BELUM K3442, K3608 Upper part of the Glenc Limestone (Viséan, Asbian), Sillees River, County Fermanagh. DIAGNOsIS. Dyscritella with robust dendroid zoarium and irregu- larly dichotomising branches. Autozooecia are developed throughou the zoarium. Zooecial walls are thin in the endozone but are thick | * and of constant width in the exozone. Basal diaphragms are rare an indistinct. Autozooecial apertures are circular and of moderate size Interapertural areas are of variable width, and exilazooecia ar abundant. Maculae are quite common. Prominent, large acanthostyle: occur at interzooecial intersections interspersed with smalle acanthostyles between. DESCRIPTION. The dendroid zoarium is robust, the longest frag) | ment examined attaining a length of 30.6mm. Bifurcation is rare bul | where it occurs no increase in branch width accompanies it. Autozooecia originate in the endozone by interzooecial budding) }, From the endozone they diverge at a low angle, but in the exozon| |» they turn through 80° to lie perpendicular to the zoarial surface. I} the endozone zooecial walls are thin and undulatory. Walls thicke} LOWER CARBONIFEROUS BRYOZOA 149 ‘rapidly at the endozone/exozone boundary, and wall thickness re- ‘mains constant (mean thickness 0.07mm) throughout most of the ‘exozone. Thin basal diaphragms are infrequently developed in the endozone, whereas very thin diaphragms are occasionally devel- oped in the exozone. 4 Autozooecial apertures are of moderate size, circular in shape, ‘and are closely packed, approximately their own diameter apart. Interapertural areas are irregular in width. Exilazooecia are very bundant between autozooecia and vary greatly in shape and size. ‘They are usually circular to oval, but in interapertural angles small bentagonal forms occur (0.02—0.1mm in diameter). Frequently, one pr two exilazooecia are located between autozooecia. Exilazooecia inate within the exozone where they form short, narrow tubes. “Maculae, 0.4mm in diameter, occur occasionally and comprise Seistcrs of exilazooecia. Acanthostyles are abundant in interapertural walls. Relatively arge acanthostyles (mean diameter 0.03mm) persist at interapertural ngles around autozooecia while smaller acanthostyles (mean ‘Table 23 Measurements of Dyscritella miliaria (in mm). N=5. 'D 19 4.59 3.17 6.15 9.65 5.06 4 21 16.06 9 DD) 28.53 451 2 25 8.35 7 10 10.16 147.80 AD 30 0.12 0.09 0.22 12.08 5.34 WT 30 0.08 0.03 0.28 38.77 2.30 oD 30 0.04 0.02 0.10 31.90 4.04 NT 9 2.64 1.91 3.52 13.47 5.50 18 0.97 0.57 1.38 8.00 1.24 Figs 63-65 Dyscritella miliaria (Nicholson, 1881); 63, Upper part of the Glencar Limestone (Viséan, Asbian), Sillees River, County Fermanagh; BELUM ) K3442; 63a, large zoarial fragments showing ramose growth-form and large circular autozooecial apertures surrounded by irregularly-placed exilazooecia, x1; 63b, detail of 63a, x15; 63c, cross-section showing autozooecial chambers with thickened exozonal walls, x50: 63d, detail of 63c, | showing exilazooecium developed in exozone between adjacent autozooecial apertures, x120; 64-65, Carboniferous shales (probably Redesdale Ironstone Shale (Asbian), Lower Limestone Group), Redesdale, Northumberland; 64, AUGD.10135a (lectotype), ramose branching zoarial fragment with crowded arrangement of autozooecial apertures surrounded by much smaller circular or irregularly-shaped exilazooecial apertures, x5; 65, GSM 1967.66.384 (paralectotype), tangential section showing oval autozooecial apertures surrounded by numerous exilazooecia, x15. diameter 0.01mm) are developed in a line on interapertural walls between both autozooecia and exilazooecia. DISCUSSION. This is the first reported occurrence of the trepostome Dyscritella miliaria from the Carboniferous of Ireland. The abun- dance of exilazooecia (“interstitial tubuli’ of Nicholson, 1881) makes the taxon very distinctive. It was first described, as Monticulipora tumida var. miliaria, from the Lower Carboniferous of England (Nicholson 1881). Subsequent specimens from the Midland Valley of Scotland were discovered in the Young Collection at the Hunterian Museum (Bancroft 1984). D. miliaria also occurs at Llangollen, North Wales, in strata of Asbian age (A.J. Bancroft, pers. comm., April 1988). Dyscritella miliaria is rare in the Lower Carboniferous of the Fig. 66 Dyscritella miliaria (Nicholson, 1881). Line drawing of external features of BELUM K3442; scale bar = 0.1 mm. 150 British Isles. In the large bryozoan sample examined in this study only 4 specimens were found. In his original description Nicholson failed to notice basal dia- phragms in the endozone. Lee (1912) redescribed the earlier material in which he found ill-defined ‘tabulae’ (basal diaphragms). Conse- quently, he correctly placed the specimens in the genus Dyscritella and elevated miliaria from variety to specific rank. Coefficient of variation values in Table 23 illustrate a number of features. Zoarial width (ZW) and autozooecial aperture diameter (AD) do not vary greatly either within or between colonies. The within-colony ranges of exilazooecial diameter (ED), autozooecial spacing (AS), and the number of autozooecia in an area of 1mm (Z1) are all large. However, between-colony CVs for these parameters are small because all colonies are of a similar size. There is little within- colony variation, but large between-colony variation in the number of autozooecia in a 2mm line (Z2). This large CV value may be a sampling error arising from the small sample size. STRATIGRAPHICAL RANGE. Lower Carboniferous (Asbian— ?Brigantian). DISTRIBUTION. Ireland, northern England, North Wales, Midland Valley of Scotland, Morocco. Family STENOPORIDAE Waagen & Wentzel, 1886 Genus TABULIPORA Young, 1883a TYPE SPECIES. the Lower Carboniferous of East Kilbride, Scotland. | Tabulipora urii (Fleming, 1828) Figs 67—70, 71a 1828 v1883a v1883b v1883 1884b 1912 1912 1935 1953 1961 1969 1970 1973 1977 non v 1986 1987 Cellepora urii Fleming, 1828 by monotypy from Cellepora urii Fleming: 532. Tabulipora urii (Fleming); Young: 154. | Tabulipora urii Young; Young: 264. | Tabulipora urii Young [sic]; Nicholson: 295. Tabulipora urii Young [sic]; Vine: 380, pl. 20, figs 3a— | . / P.N. WYSE JACKSON | | b, 4. | Tabulipora urei Young [sic]; Lee: 150. Tabulipora scotica Lee: 162, pl.14, figs 4a—d, pl. 15, figs 12, 13, 17, 18. Tabulipora scotica Lee, Anderson & Lamont: 216, fig. 6.13. Tabulipora scotica Lee; Bassler: G105, fig. 70 — la, . Ib. 1 Tabulipora scotica Lee; Wilson: 91. Tabulipora scotica Lee; Owen: 262, pl. 22, figs d-e. | | Tabulipora urii (Fleming); Gautier: 19, pl. 7, fig. 1; pl. 8, figs 1-2. | Tabulipora scotica? Lee; Owen: 302. | Tabulipora urii (Fleming); McKinney: 313, pl. 1, fig. 3 Tabulipora scotica Lee; Kora & Jux: 91, figs 3, g1-4. ~ Tabulipora urii (Fleming); Bancroft: 196. lq Figs 67-70 Tabulipora urii (Fleming, 1828); Upper part of the Glencar Limestone (Viséan, Asbian), Sillees River, County Fermanagh; 67, BELUM | K 15200, ramose colony fragment; autozooecia with circular to oval shaped apertures arranged over the whole surface, except on monticules; mesozooecia are developed between autozooecia, x3; 68, BMNH PD9638, small encrusting colony on 2echinoid spine, consisting of one layer of | | | | | autozooecia with circular apertures with polygonal-shaped mesozooecia situated between, x40; 69, BMNH PD9472, shallow tangential section showing | autozooecial and mesozooecial apertures, and interapertural walls with large acanthostyles at junctions and smaller stylets in between, x35; 70, BELUM) K15207; 70a, longitudinal section through thin-walled endozonal area and thickened exozone region; the initial portions of autozooecial chambers lie at) |) a high angle to the zoarial surface, but bend at the exozone and become nearly perpendicular to it, x12; 70b, longitudinal section showing exozone and autozooecial chambers; ring septae develop at the top of the endozone with eight per autozooecium; septal necks are inflated and deflected interiorly, | x20; 70c, longitudinal section showing exozone and autozooecial chambers, and zone of regeneration where the endozone is characteristically thin, x20/ | | i | LOWER CARBONIFEROUS BRYOZOA 1988 Tabulipora urii (Fleming); Yang, Hu & Xia: 69, pl. 22, figs 5-8. MATERIAL. BM(NH) PD9472, 9638-9642, 9704; TCD.34079, 42593b, 42604b, Upper part of the Glencar Limestone (Viséan, Asbian), Carrick Lough, County Fermanagh; BELUM K15200- 15208, Upper part of the Glencar Limestone (Viséan, Asbian), Sillees River, County Fermanagh. DESCRIPTION. Zoaria are erect, ramose expansions of bifurcating cylindrical branches reaching 8.20mm in diameter. Autozooecial chambers are budded interzooecially from the branch centres. Chambers diverge from the centre of branches at a low but constant angle of 30° in the endozone. They bend abruptly at the exozone; vestibules are orientated at angles of between 70° and 80° to the zoarial surface. Autozooecial chambers are ten to twelve times longer than wide. Chamber walls are very thin (0.01mm) in the endozone, but thicken considerably (to 0.05mm) in the exozone, where in some specimens as many as five monilae may occur. They are pear to oval-shaped thickenings of the chamber wall, and may be separated by lengths of thin chamber wall. Skeletal laminae within the wall are deflected away from the zoarial surface from a central dark zone at autozooecial boundaries. In cross section and deep tangential section chambers are polygonal in shape. Ring septa are developed in autozooecial chambers towards the top of the endozone and throughout the exozone. A solitary or pair of thin, widely spaced endozonal ring septa contrast with up to seven thicker and more closely spaced ring septae developed in the exozone. Foramen are circular or oval in shape and are placed either centrally or slightly laterally. The central walls of ring septa constitute a thickened ring that is bent posteriorly. Autozooecial apertures are large, circular to oval in shape, and closely spaced at less than one diameter apart. They are irregularly arranged over the zoarial surface. They are : ; 3 : Pod. J ‘s ig.71 Line drawing of external features of the Tabulipora species described in this study. a, Tabulipora urii (Fleming, 1828) (BELUM K15200); b, Tabulipora howsii (Nicholson, 1881) (BMNH PD9644); ¢, | Tabulipora minima Lee, 1912 (BMNH PD9650); scale bar = 0.1 mm. | | | | 151 poorly developed on monticules where they are marginally larger. The long axes of oval shaped apertures radiate out from monticules and maculae. Interapertural walls are thin. Exilazooecia are very common, and are disposed in one or two rows between autozooecial apertures, or in radiating maculate clusters of up to forty individuals. They are small, thin walled, and circular to polygonal in shape. Stylets are common and structurally variable, and developed on the thin interapertural walls. Large acanthostyles, 0.04mm in diam- eter, are found at interapertural junctions. These have a distinctive dark grey core developed from the base of the exozone. Smaller heterostyles (0.01—0.02mm in diameter) are found in one or two rows between acanthostyles. They arise from within the exozone. Table 24 Measurements of Tabulipora urii (in mm), N=8. NM X Mn Mx CVw CVb ZD 69 59) 3.79 8.20 3.84 5.34 ZI 80 10 7 14 IHES9 10.48 Z2 80 5.85 4 a 10.73 10.48 AD 80 0.19 0.11 0.29 15.55 9.71 IWT 80 0.11 0.06 0.17 18.08 13.33 ED 80 0.07 0.05 0.14 22.98 6.95 ET 17 LST 0.70 2.76 5.99 2M TE 33 0.97 0.47 1.98 Sl PAU IMS 8 4.72 4.10 5.94 17.47 11.32 DISCUSSION. Tabulipora urii was first described and figured by Ure (1793) as Millepore from the Carboniferous of Kilbride, West Scotland. Subsequently, Fleming (1828) cited Ure’s material as the type species of his species Cellepora urii. This species was later described by Young (1883a) who noted ring septa in the autozooecial chambers and on the strength of this erected the subgenus Tabulipora. Cellepora urii is the type species of Tabulipora by monotypy. Lee (1912) revised the British Trepostomata. He examined Young’s material and erected a new species, Tabulipora scotica, designating it as the type species of Tabulipora. He demoted T. urii (misspelt ‘urei’) because he felt that Young had not really proposed it as a new specific name, and because his material contained several species, none of which had been figured. Gautier (1970) stated that 7. urii is the type species by monotypy, and thus T. scotica is invalid. T. scotica is regarded as a junior synonym of T. urit (Bancroft 1984: 372). The figured specimens of T. scotica described from Egypt (Kora & Jux, 1986) have been examined. Up to nine hemiphragmas are developed in chambers indicating that the specimens are referable to the genus Stenophragmidium. STRATIGRAPHICAL RANGE. Brigantian). Lower Carboniferous (Tournaisian— DISTRIBUTION. ?China. County Fermanagh, Midland Valley of Scotland, Tabulipora howsii (Nicholson, 1881) v1881 v1883 Figs 71b, 72-78 Stenopora howsii Nicholson: 83, fig. 12. Stenopora howsii Nicholson; Nicholson: 285, pl. 10, figs 1-10, text-figs la—c. 1886 Stenopora howsii Nicholson; Nicholson & Etheridge jun.: ae Stenopora howsii Nicholson; Nicholson & Lydekker: 356, sia, 2B Vd: 1891 Stenopora howsii Nicholson; Etheridge jun.: 48. 1912 Tabulipora howsei |sic] (Nicholson); Lee: 166, pl. 14, figs 9a—c; pl.15, figs 22-24. v1889 Figs 72-78 Tabulipora howsii (Nicholson, 1881); 72-75, Upper part of the Glencar Limestone (Viséan, Asbian), Carrick Lough, County Fermanagh; 72, BMNH PD9643; 72a, adnate zoarium showing regular arrangement of autozooecia with circular apertures, x12; 72b, detail of 72a, showing circular autozooecial apertures with ring septa and circular-oval foramen, and acanthostyles positioned on interapertural wall junctions, x40; 73, BMNH PD9645, adnate zoarium, x20: 74, BMNH PD9644, as 72a, including ring septa with circular-oval foramen, x50; 75, BMNH PD9561, longitudinal section { showing thin autozooecial chamber walls with acanthostyles developed on interapertural wall junctions, x100; 76-78, probably Redesdale Ironstone | shale (Asbian), Lower Limestone Group, Redesdale, Northumberland; 76, AUGD.10134a (paralectotype), detail of zoarial surface, x20, 77, ' AUGD.10134b (lectotype), shallow tangential section showing arrangement of polygonal autozooecial chambers, centrally placed circular to oval | | foramen in some autozooecial chambers, rare exilazooecia, and stylelets of two sizes on thin interapertural walls — acanthostyles at wall junctions and heterostyles in between, x40; 78, AUGD.10134c (paralectotype), transverse section through branch showing inner endozone with thin-walled polygonal | autozooecial chambers, and outer exozone with moniliform walls and 7-8 ring septa developed per chamber, x20. | 1913 Tabulipora howsei [sic] (Nicholson); Wright et al.: 72. 1925 Tabulipora howsei [sic] (Nicholson) var. nov. Smyth: 150. 1987 Tabulipora howsii (Nicholson); Bancroft: 196. 1991 Tabulipora ct. howseii (sic) (Nicholson); Billing: 41. LECTOTYPE. Herein designated AUGD 10134b (fig. 12a of Nicholson 1881) PARALECTOTYPES. Herein designated AUGD 10134, 10134a, and 10134c (fig. 12b of Nicholson 1881). AUGD 10132 and 10141 may be part of the original Nicholson material and so may also be paralectotypes (Benton & Trewin 1978, Benton 1979). MATERIAL. BMNH PD9561, 9643-9649, 9653, 9722-9729, TCD.34080-34087, 34121, 34158, 34164, 42564, ; BELUM K2177, P.N. WYSE JACKSON K3234, Upper part of the Glencar Limestone (Viséan, asbidlll Carrick Lough, County Fermanagh. TCD.42551-42554, Upper p' | of the Glencar Limestone (Viséan, Asbian), Sillees River, County Fermanagh. DESCRIPTION. Only small encrusting zoaria were examined; ne " solid branches were recovered. | Adnate zoaria are thin, up to 1.30mm thick, and form smal! | circular expansions up to 6.2mm in diameter. They encrust fenes: i trate bryozoans, and crinoidal material. Sheets are only one |” autozooecial chamber high; young autozooecia do not encrust olde ; autozooecia (as in Fistulipora incrustans). Autozooecia are budded from a thin basal wall 0.02—0.03mm) })j thick and diverge distally in the narrow endozone. At the exozont|}); LOWER CARBONIFEROUS BRYOZOA they bend slightly and vestibules are orientated perpendicular to the zoarial surface. Chamber walls are only 0.01—0.02mm thick in the endozone, but are considerably thicker, ranging from 0.15—0.18mm, in the exozone. These endozonal walls are straight, and rarely moniliform. Ring septa are developed within the exozone where generally three to four are found. (A greater number have been reported from _ solid ramose zoaria where the exozone is thicker (Lee 1912, Bancroft | 1984)). The inner margins of the ring septa are unthickened, thin and not deflected posteriorly. Foramen are 0.08 to 0.12mm in diameter, centrally placed, and circular to occasionally reniform in shape. Autozooecial apertures are large, and polygonal to infrequently circular in shape and are very closely spaced at less than one | diameter apart. Exilazooecia are small, with circular apertures. They are uncommon, and usually occur as isolated individuals between autozooecia at interapertural angles. Interapertural walls are very thin and angular or rounded with stylets developed along their crests. Large acanthostyles are found at interzooecial junctions, while as many as 20 smaller heterostyles may occur between them, around autozooecia. Acanthostyles reach 0.05mm in thickness and 0.09mm in length. Table 25 Measurements of Tabulipora howsii (in mm). N=14. NM x Mn Mx CVw CVb MTZ 41 0.45 0.24 1.30 9.10 33) 1Z1 72 9 6 1] 8.24 11.02 Z2 74 6.41 5 8 7.74 10.48 \AD 130 0.27 0.20 0.35 9.16 TY IWT 126 0.03 0.01 0.08 24.2 13.08 FD 35 0.09 0.06 0.12 11.19 WI ED 4 0.09 0.05 0.13 - 2.67 MTZ = Maximum thickness of adnate zoarium from basal wall to external surface. | “Discussion. Tabulipora howsii is easily recognised by its thick /ramose zoaria which may reach a diameter of 20mm, its moniliform een wall, the presence of numerous ring septa, and from the ! | often polygonal to angular autozooecial apertures. However, no ramose specimens were found; this is unusual as jthey have previously been recorded in large numbers (Lee 1912, Bancroft 1984). Adnate colonies of 7: howsii have been reported : rom Scotland (Bancroft 1984) but these have not been examined in the present study. _ The number of exilazooecia may vary greatly from zoaria to Zoaria. In the County Fermanagh specimens exilazooecia are gener- ally few in number. However, Bancroft (1984: 376) describes them “As common, occurring as either scattered individuals or in maculae, “im ramose zoaria from the Midland Valley of Scotland, which Be ccsts that the Fermanagh specimens were young individuals hich had not developed ramose branches characteristic of older ions Smyth (1925: 150) noted a variety of the taxon from the Asbian of north Wales, and stated that it conformed in every aspect with Lee’s iliagnosis except in two features: 30 not 40 autozooecial apertures “were contained in a Icm line, and the foramen were oval not om Within the Scottish and County Fermanagh T. howsii populations there is variation in these characters and they are not _ponsidered herein to merit varietal status. Examination of Smyth’s inaterial (TCD.R11a—g, TCD.21545a, b) shows it to lie within the _ range of T. howsii as given by Bancroft (1984). ; : ; : : TRATIGRAPHICAL RANGE. Lower Carboniferous (Asbian— 8rigantian). 153 DISTRIBUTION. Rare, but recorded from Counties Fermanagh and Donegal (Wright ef al. 1913), the Midland Valley of Scotland, northern England, and North Wales. Tabulipora minima Lee, 1912 Figs 71c, 79 1912 Tabulipora minima Lee: 164, pl.15, fig.21. 1987 Tabulipora minima Lee; Bancroft: 196. LECTOTYPE. Herein designated GAGM 01-53 BYC (longitudinal section in thin section, figured by Lee, 1912). MATERIAL. Three zoarial fragments, BMNH PD9650, BELUM K2182. Upper part of the Glencar Limestone (Viséan, Asbian), Carrick Lough, County Fermanagh; K3474 Upper part of the Glencar Limestone (Viséan, Asbian), Sillees River, County Fermanagh. DESCRIPTION. Zoaria are erect, ramose, and composed of moder- ately robust sub-circular branches. Only two zoaria were examined; the larger measures 14.7mm in length. The nature of branch division is not known. Autozooecia are budded from within the endozone. Autozooecial chambers are long, reaching a maximum length of 4.3mm in length. They diverge at low angles of less than 25° in the endozone, and hardly bend posteriorly at the exozone. Endozonal walls are very thin, undulating or straight. The major- ity of the autozooecial tube is contained within the endozonal area. Consequently the axial ratio is high. The exozone is only 0.11— 0.21mm wide, where chamber walls are of regular thickness. Three to occasionally four ring septa are found per autozooecium. They are thin, with a central circular foramen, and are found towards the top of the endozone and within the exozone. Autozooecial apertures are large, oval to circular in shape and closely spaced at less than one diameter apart. Interapertural walls are thin and smooth. Exilazooecia are small, and range in diameter from 0.06 to 0.13mm. They are circular to oval in shape and occur as scattered individuals or in small clusters between autozooecia. Stylets are of one structural type only. Between 12 and 15 large acanthostyles 0.06mm in diameter surround autozooecial apertures. They are arranged on interapertural walls, with one consistently positioned at interapertural junctions. Table 26 Measurements of Tabulipora minima (in mm). N=2. NM X Mn Mx CVw CVb ZD 11 DES 1.80 3.36 4.16 2.71 ZI 16 12.95 10 16 ETS 9.63 Z2 16 4.71 4 5 9.83 28.99 AD 20 0.23 0.15 0.30 13.43 4.74 IWT 19 0.08 0.05 0.12 22.92 12.02 ED 7 0.09 0.06 0.13 19.52 3.18 ET >) 1.99 1.38 2.58 1.46 2.38 TE 12 0.17 0.11 0.21 13.33 8.24 DISCUSSION. Tabulipora minima is easily recognised by its mod- erately robust and often flattened branches, by the presence of a very thin exozone, and by the occurrence of a small number of ring septa in each autozooecium. It is rare in the British Isles. Only Lee (1912) and Bancroft (1984) have previously recorded it. At Carrick Lough three specimens were recovered. Although one of these was silicified a considerable amount of internal detail, including ring septa, is preserved. STRATIGRAPHICAL RANGE. Carboniferous (Viséan—Arnsbergian). | P.N. WYSE JACKSON DISTRIBUTION. Only recorded from County Fermanagh, York- | shire, and the Midland Valley of Scotland. | | TYPE SPECIES. Stenophragma lobatum Munro, 1912, by original | designation from the Lower Carboniferous of Ravenstonedale, Cum- ; bria (formerly Westmoreland), England. | Genus STENOPHRAGMIDIUM Bassler, 1952 REVISED DIAGNOSIS. Stenoporid with encrusting, rarely ramose zoaria. Encrusting zoaria commonly form flat adnate colonies or’ hollow erect dichotomising expansions on which monticules are regularly developed. \ Autozooecial chambers have thickened walls in the exozone, | where they are of uniform width or occasionally moniliform. | Autozooecial chambers diverge distally at a low angle in the thin endozone, becoming perpendicular to the zoarial surface in the wider exozone. Up to five hemiphragms are developed on proximal | walls, at the top of the endozone and in the exozone, where they extend halfway across chambers. Autozooecial apertures are of moderate size, circular to oval in shape, and closely spaced. Exilazooecia are rare. Large acanthostyles may be situated at zooecial wall junctions, and heterostyles may be disposed between them. [ DISCUSSION. This genus, which is restricted to the Carboniferoni was first described from Northern England (Munro 1912). To date) nineteen species have been reported; four species have been de- scribed from the Carboniferous of the British Isles. The British species are Stenophragmidium grandyense (Munro 1912), S. lobatum, (Munro 1912) [the type species], S. incrustans Owen 1973 and S. ramosum Owen 1969. To these, five Tabulipora species of Le (1912) may be added, as well as Tabulipora serrata Smyth, 1922 (from the Lower Carboniferous (Brigantian) of Ballycastle, County 7 Antrim). They possess ‘tabulae’ which ‘extend only partly, leaving |_ an untabulated space, ... always situated on the distal side ...’ (Le i. 1912: 171). The ‘tabulae’ are clearly hemiphragms. Examination and revision of Lee’s and Smyth’s material may reveal that their six species are referable to Stenophragmidium. STRATIGRAPHICAL RANGE. Lower—Upper Carboniferous DISTRIBUTION. British Isles, Belgium, the CIS (former Soviel Union), North America, Asia. | | Fig. 79 Tabulipora minima Lee, 1912; Upper part of the Glencar | Limestone (Viséan, Asbian), Carrick Lough, County Fermanagh; BMNH PD9650; 79a, portion of ramose colony showing arrangement 0} autozooecia with oval apertures, x20; 79b, detail of 79a, showing oval autozooecial apertures divided by thick interapertural walls on which are developed large acanthostyles, x100. Figs 80-82 Stenophragmidium sp. Upper part of the Glencar Limeston (Viséan, Asbian), Sillees River, County Fermanagh; 80, BELUM K15209, ramose hollow zoarial fragment, with regular disposition of monticules and autozooecia; small exilazooecial apertures are found | between autozooecia, particularly on monticules; autozooecia adjacent | to monticules are larger than those in inter-monticular areas, x3; 81, BELUM K15210, zoarial fragment, details as for 80, x2.5; 82, BEL K15214; 82a, longitudinal section through colony showing thin undulatory basal wall, low angle of divergence of autozooecia in endozone, increasing in exozone region; chamber walls thin in endozone, thicken rapidly in exozone; a single hemiphragm is situated on proximal chamber walls, bends inwards, and seals half the vestibule, x25; 82b, detail of 82a, x75. LOWER CARBONIFEROUS BRYOZOA Stenophragmidium sp. Figs 80-83 MATERIAL. BELUM K3436, K15209-K15215 (5 zoarial frag- ments and 2 longitudinal sections), Upper part of the Glencar Limestone (Viséan, Asbian), Sillees River, County Fermanagh. DESCRIPTION. Zoaria consist of large erect hollow expansions up to 11.17mm in width, or thin encrusting adnate sheets 0.28—0.30mm thick. Autozooecia are budded from a very thin undulating basal wall. Autozooecial chambers are recumbent in the thin endozonal region (0.15 mm) and straight in the wider exozone (0.28 mm). The vestibule lies at a high angle of 80° to 85° to the zoarial surface, and is moderately wide (0.19-0.20 mm). Autozooecial walls are 0.03—0.04 mm thick in the endozone, but expand rapidly in the exozone to between 0.15 and 0.18 mm. They are usually of constant width, occasionally undulatory, and rarely -moniliform. Walls consist of a well defined central hyaline zone (0.02 mm wide) in which laminae are orientated parallel to the growth direction. Thin lateral laminae are derived from this central zone and bend sharply proximally. This is typical leioclemid-type wall structure (after Boardman 1960). Up to three hemiphragms are developed within and at the base of the exozone, where they arise from proximal walls. They are thin, often straight and occasionally bend slightly proximally. They are |short (0.04—0.07 mm in length), usually extend less than half-way ‘into chambers, and have rounded bulbous tips. Autozooecial apertures are polygonal or irregular in shape, occa- \sionally circular and are closely spaced at less than one diameter apart Apertures decrease in diameter away from monticules. Interapertural walls are rounded and moderately thick, with as many as 16 stylets occurring along their crests. Large acanthostyles (0.05—0.07 mm) are usually found at autozooecial wall junctions, with smaller stylets (0.02—0.03 mm) between. Exilazooecia are rare; they are small, with circular to angular apertures. They occur as isolated individuals between autozooecia, or infrequently in groups of 10 to 12 in monticule- centred maculae which are widely spaced at up to 5.53 mm apart. DISCUSSION. This taxon is known from 14 specimens from which details of both external and internal structure are known. It does not resemble previously described British species, but may be synonomous with Tabulipora crassimuralis Lee, 1912. The genus ieeds revision, which will be undertaken at a future date and until hen I prefer to leave this taxon unassigned to any species. The hollow portions of the zoaria examined all had a posthumous ig. 83 Stenophragmidium sp. Line drawing of external features of BELUM K15209; a, Portion of hollow ramose colony, scale bar = | mm; b, detail of autozooecia and exilazooecia and disposition of acanthostyles, scale bar = 0.1 mm. 155 Table 27 Measurements of Stenophragmidium sp. (in mm). N=6. NM Xx Mn Mx CVw CVb ZD 39 7.92 4.81 11.17 11.28 4.38 Zl 60 WB) 5 10 12.08 13.44 Z2 60 5):9) 4 8 13739) 6.16 AD 60 0.24 0.15 0.32 10.75 10.84 IWT 60 0.07 0.04 0.12 20.86 7.96 ED 60 0.08 0.06 0.15 21.26 125) It 4 0.64 0.25 1.07 10.60 1.22 WB 4 1.64 0.90 2.76 16.76 1.68 IMS 6 4.45 3.38 5.5) 18.34 14.30 infill of micritic mud which was also found in autozooecial cham- bers. Zoaria possibly encrusted soft algal branches which have now disappeared, or may simply have been hollow. STRATIGRAPHICAL RANGE. Lower Carboniferous (Viséan, Asbian). DISTRIBUTION. Ireland, North Wales. Order CYSTOPORATA Astrova, 1964 Suborder FISTULIPORINA Astrova, 1964 Family FISTULIPORIDAE Ulrich, 1882 Genus FISTULIPORA M ‘Coy, 1849 TYPE SPECIES. Fistulipora minor M‘Coy, 1849 by subsequent designation (Milne-Edwards & Haime 1850: lix) from the Lower Carboniferous of the British Isles. Fistulipora incrustans (Phillips, 1836) Figs 84a, 85-88, 105 A complete systematic description, full synomony and discussion of the type specimens is given in Bancroft & Wyse Jackson (1995). MATERIAL. BMNH PD9651-9676, 9740; TCD.34088-34103, 34138-34139, 34146, 34157, 34159, 34166, 42590, 42610; BELUM K2151,K2153, K2193, Upper part of the Glencar Limestone (Viséan, Asbian), Carrick Lough, County Fermanagh. TCD.42511, Upper part of the Glencar Limestone (Viséan, Asbian), Sillees River, County Fermanagh. DESCRIPTION. Zoaria form thin unilaminate button-like discs up to 1.3cm in diameter; encrusting unilaminate or multilaminate sheets up to 1.58mm thick, or small chaetetiform nodular expansions 30mm wide by 17mm high. Colonies often encrust crinoid stems, fenestellid Bryozoa, and occasionally brachiopod valves. Autozooecia are budded from a very thin basal wall (0.015—0.025 mm thick). They are often initially slightly recumbent, but subse- quently become erect and orientated at 80° to 90° to the zoarial surface. Autozooecia are straight, tubular and thin walled. Thin diaphragms are commonly developed at the base of chambers and are less common and irregularly dispersed throughout the remaining portions of chambers. Autozooecia are arranged in straight to curved rows which produces an offset pattern on the zoarial surface. Autozooecial apertures are large (0.17mm to 0.40mm); usually circular to oval, occasionally D-shaped, rarely sub-polygonal in shape and spaced 1—2.5 diameters apart. Lunaria are indistinct and not consistently present. They are small — only one fifth the circum- ference of apertures, crescent-shaped, marginally elevated above the zoarial surface, and discretely indent apertures (by as much as 0.04mm). Lunaria are found on the proximal sides of autozooecia closest to monticules. In a few zoaria lunaria completely encircle autozooecial apertures, forming low peristome-like collars. Rarely, 156 P.N. WYSE JACKSON Fig. 84 Measurements taken on cystoporates in this study; a, Fistulipora incrustans (Phillips, 1836); b, Sulcoretepora parallela (Phillips, 1836); ¢, Goniocladia cellulifera (Etheridge, 1873b); ZT = Zoarial thickness from basal wall to upper zoarial surface; BW1 = Branch width measured parallel to | median wall: BW2 = Branch width measured perpendicular to median wall; MS = Distance between adjacent monticule measured from centre to centre; V1 = Number of vesicles contained along a 1mm line; FL = Fenestrule length; FW = Fenestrule width; AD1 = Autozooecia apertural diameter measured) parallel to growth direction; AD2 = Autozooecia apertural diameter measured perpendicular to growth direction; AS = Autozooecia apertural spacing: minimum distance between two adjacent autozooecial apertures, measured from their centres; LAD = Autozooecia apertural length measured from lunaria to opposite side; TAD = Autozooecia apertural width measured perpendicular to LAD; LID = Autozooecia apertural spacing in same longitudinal row: TID = Autozooecia apertural spacing between adjacent rows; Z1 = Number of complete autozooecial apertures contained in 1 mm?; Z2 = Number | of complete autozooecial apertures contained in a 2mm line; LW = Lunarium width; LT = Lunarium thickness; LD = Lunarium depth; V2 = Number of _ vesicles seen in transverse section contained along a Imm line; D1 = Number of diaphragms contained within 1mm. large hood- or cowl-like lunaria are developed. These are 0.1 mm interzooecial vesicular tissue, and by the regular monticular a high and cover approximately half the aperture. They are similar to rangement. those illustrated by Warner & Cuffey (1973) in F. incrustans Moore and F: carbonaria. Table 28 Measurements of Fistulipora incrustans (in mm). N=15. Monticules are regularly arranged between 2 and Smm apart and are conspicuously elevated above the general surface of zoaria. Small maculae (sensu Anstey 1981) are found at the apex of each ZT 4] 0.67 0.20 1.58 18.10 1 monticule. Autozooecia develop in intersecting semi-circular rows AD 119 0.27 0.18 0.40 10.73 6 away from monticules. LAD 20 0.24 0.20 0.32 10.10 1Lg Vesicles are arranged in 3 to 4 vertical stacks between autozooecia, we av Jay wad sale Us if with 5-8 contained within 1mm per stack. They are small, rectangu- BS ae oes Gao i. 2 z . ‘ ; : i LID 20 0.41 0.22 0.66 22.42 2 lar, polygonal or inverted cup-like structures. They become TID 20 0.27 0.20 0.45 14.59 4 increasingly thinner towards the zoarial surface. Between ten and Zl il 35 D) 6 16.88 6 fifteen vescicles are found in each vertical stack. Z2 109 3.9 3 6 MAS 7 In some large colonies five to six encrusting cycleshave beenseen ——-V! 16 4.6 3 6 16.04 5: (this is not unusual — many more such multilaminate sheets have V2 ZY Sif 2 : 16.60 7 been observed in massive colonies from other localities) ws 8 zh eee cae ee : i re : e4 LW 18 0.20 0.17 0.23 7.84 20. . . * : : : : LD 12 0.01 30.0 10.02 29.25 3 DISCUSSION. Fistulipora incrustans is easily recognised by its LT 12 0.03 0.02 0.07 3312 3 encrusting habit, its large circular autozooecial apertures, its stacked exe ewe ee LOWER CARBONIFEROUS BRYOZOA _ It is the only species of Fistulipora described from the British (sles. Several previously reported taxa are considered synonymous with it. These are Fistulipora minor M‘Coy, 1849 (see Owen 1969), and Berenicea megastoma M‘Coy, 1844, which Young (1882) con- sidered to be an immature F. incrustans colony (Bancroft & Wyse lackson 1995). F. excelens Ulrich, 1884, has a similar form and morphometric measurements to F incrustans (Phillips, 1836) and might be ynonymus. F. incrustans Moore (1929) (a secondary homonym) fom Texas has significantly larger elliptical to oval-shaped utozooecial apertures, which are surrounded by complete eristomes, and is not conspecific. On the other hand, the specimens lescribed as F. incrustans Moore by Warner & Cuffey (1973) from ne Lower Permian of Kansas are similar in most respects, and are robably conspecific with F. incrustans (Phillips, 1836). Some morphological variation is seen both within and between olonies although this is not as considerable as the variation re- orded by Warner & Cuffey (1973). They measured parameters in yhich high variation would be expected (eg. lunaria width and yapore dimensions). In this study a smaller number of parameters /ere measured: these are associated with the autozooecia (eg. oecial chamber width and spacing) and variation in them is Figs 85-88 Fistulipora incrustans (Phillips, 1836); Upper part of the Glencar Limestone (Viséan, Asbian), Carrick Lough, County Fermanagh; 85, BMNH PD9651, small dome-shaped adnate zoarium (encrusting Baculopora megastoma fragment) showing regular arrangement of autozooecia in curved intersecting rows; lunaria are situated on the upper edges of apertures, x10; 86, BMNH PD9652, chaetetiform colony showing monticular arrangement, x3.5; 87, BMNH PD9676;: 87a, transverse view of circular autozooecial apertures separated by vesicular tissue, x20; 87b, detail of 87a, | x60; 88, BMNH PD9675; 88a, longitudinal section showing three growth increments, simple tubular autozooecial chambers, and vesicular tissue _ comprising box-like vesicles with domed upper surfaces, x20; 88b, detail of 88a, x60. thought to be indicative of minor environmental changes (Farmer & Rowell 1973). STRATIGRAPHICAL RANGE. Lower Carboniferous (Courceyan)— Lower Permian. DISTRIBUTION. Fistulipora incrustans is acommon species with a wide geographical range. It is frequent in the Lower Carboniferous of the British Isles, and has been recorded from the CIS (former Soviet Union), and North America. Family CYSTODICTYONIDAE Ulrich, 1884 Genus SULCORETEPORA @ Orbigny, 1849 TYPE SPECIES. Flustra? parallela Phillips, 1836 by original desig- nation from the Lower Carboniferous of Whitewell, Yorkshire, England. REVISED DIAGNOSIS. Cystodictyonid with erect zoaria composed of dichotomising bifoliate branches, elliptical to oval in cross- section. Branches retain a constant width along their length. Autozooecia are budded from a straight or zig-zag median wall 158 which is composed of a dark central granular skeleton surrounded by laminated skeleton. Autozooecia are arranged in longitudinal rows. Zooecial chambers are long and narrow in the endozone and bend sharply in the exozone. In tangential section, chambers are rectangu- lar in shape. Vesicles are found in the outer endozone and inner exozone, where they occur as irregular to circular cavities. They become less abundant towards the zoarial margin. Indistinct lunaria are found on the proximal edge of the large oval-shaped autozooecial apertures. Interapertural areas are smooth with a single longitudinal ridge developed between adjacent autozooecial rows. DISCUSSION. Taxonomically Sulcoretepora has presented many problems and prompted much argument. At ordinal level it was first regarded as a cryptostome (Vine 1884a), and this view was main- tained until recently (Cuffey 1973). It is now recognised as a cystoporate (Morozova 1970, Utgaard 1983). At generic level Sulcoretepora was first described by d’Orbigny (1849), who desig- nated Phillip’s species Flustra? parallela as type species. Subsequently, Ulrich (1882) erected the genus Cystodictya (type species, C. ocellata from the Mississippian of Somerset, Kentucky, U.S.A.) and placed Sulcoretepora in synonymy with it on account of the shared presence of a median wall. Many authors have followed this opinion (eg. Young 1887, Vine 1888). Close examination of the two genera shows that there is a difference in the shape and nature of the median wall. It is always straight in Cystodictya, but is undula- tory or sharply folded inSulcoretepora. Mstainia Shulga-Nesterenko, 1955, has a plicated median wall and is regarded a junior subjective synonym of Sulcoretepora (Elias 1964). STRATIGRAPHICAL RANGE. Devonian—Permian. DISTRIBUTION. British Isles, Europe, the CIS (former Soviet Un- ion), United States, Asia. Sulcoretepora parallela (Phillips, 1836) Figs 84b, 89-93 1836 = Flustra? parallela Phillips: 200, pl.1, figs 47, 48. 1843 = Flustra? parallela Phillips; Morris: 37. v1844 ~~ ~Vincularia parallela (Phillips); M*Coy: 198, pl. 27, fig. 14. 1849 Sulcoretepora parallela (d’Orbigny); d’Orbigny: 152. 1854 Sulcoretepora parallela (d’ Orbigny); Morris: 105. 1862 — Vincularia parallela (Phillips); Griffith: 227. 1877 — Sulcoretepora parallela (Phillips); Vine: 273. 1880c Ptilodictya? parallela (Phillips); Vine: 508. 1884a Arcanopora parallela (Phillips); Vine: 204. 1885 Cystodictya parallela (Phillips); Vine: 95. 1887 Cystodictya parallela (Phillips); Young: 461. 1888 Cystodictya parallela (Phillips); Vine: 74. 1953 Sulcoretepora parallela (Phillips); Bassler: 142, fig. 103. 1964 Sulcoretepora parallela (Phillips); Elias: 380, pl. 5, figs 3-6. 1969 = Sulcoretepora parallela (Phillips); Owen: 265, pl. 23, figs E-F. 1983 Sulcoretepora parallela (Phillips); Utgaard: 429, fig. 210, la-f. 1986a Sulcoretepora parallela (Phillips); Bancroft: 23. 1987 — Sulcoretepora parallela (Phillips); Bancroft: 196. 199] Sulcoretepora parallela (Phillips); Billing: 41. MATERIAL. BMNH PD9563, 9619, 9677-9700; TCD.34104- 34111, 34138, 34142, 34146, 34148-34153, 34157-34158, 34172, 42596, 42600b, 42605; BELUM K2158. Upper part of the Glencar Limestone (Viséan, Asbian), Carrick Lough, County Fermanagh. P.N. WYSE JACKSON Figs 89-92 Sulcoretepora parallela (Phillips, 1836); Upper part of the Glencar Limestone (Viséan, Asbian), Carrick Lough, County Fermanagh; 89, BMNH PD9677, branch fragment showing arrangement of autozooecia in longitudinal rows separated by longitudinal ridges, x15; 90, BMNH PD9678, as 89; lunaria are more pronounced proximally of autozooecial apertures; the size and spacing of autozooecial apertures increases in rows from left to right, x15; 91, BMNH PD9619, transverse section showing scalloped branch margins, atypical straight median wall, and elongate polygonal autozooecial chambers with rounded lateral margins, x50; 92, BMNH PD9563, transverse section with more typical plicated median wall, x20. TCD.42555-42558, Upper part of the Glencar Limestone (Viséan Asbian), Sillees River, County Fermanagh. DESCRIPTION. Zoaria form quite large expansions of dorso ventrally flattened branches that are elliptical or oval in cross-section Bifurcation of branches is more common than the development 0 lateral branches, which are thinner than the main branch. The larges fragment measured is 38.8mm in length and no lateral branche LOWER CARBONIFEROUS BRYOZOA Fig. 93 Sulcoretepora parallela (Phillips, 1836). Line drawing of external features of BMNH PD9678; scale bar = 1 mm. } were developed. Mature branches are usually of constant width along their length, but a small amount of thinning distally can occur. _ The ratio of branch thickness (BW2) to branch width (BW1) ranges from 1:4 to 1:2. _ Autozooecia are arranged in 4 to 6 longitudinal rows on the lateral sides of branches. The number of longitudinal rows remains con- stant along the length of a branch. Autozooecial apertures are of medium to large size and circular to oval in shape. A thin proximal lunarium is present around each autozooecial aperture. Apertures lare spaced two and a half to three diameters apart. This spacing increases slightly in the rows adjacent to the median ridge. Here three autozooecial apertures generally occur in a 2mm line, while | four occur in the same length in rows away from the median ridge. Between adjacent rows of autozooecial apertures a strong narrow jlongitudinal ridge is developed. A smaller, fainter ridge is often found either side of the main ridge. This second ridge is found between the distal and proximal ends of two autozooecial apertures in the same row. The interapertural areas are smooth except for the _ longitudinal ridges. | Branches are internally divided by a thin plicated median wall, from either side of which are budded autozooecia. The median wall _is composed of pale laminated skeletal material. | Autozooecial chambers are elongate, narrow, and bend distally in _ the endozone region. The exozone is extremely thin, being about one sixth the thickness of the branch. Autozooecial chamber walls are ; thin and straight. In shallow tangential section chambers are tear- Shaped and narrower proximally. Deeper sectioning shows the | chambers to have a rectangular shape. In cross-section chambers are hexagonal to pentagonal in outline. _ Small vesicles, 0.03 mm in diameter, are commonly found be- Ween autozooecial chambers and the branch margin. They are most _ frequently developed in the outer endozone and inner exozone. They "are thin-walled, circular to irregular in shape, and may be infilled with stereom in the exozone. _ Table 29 Measurements of Sulcoretepora parallela (in mm). N=21. NM x Mn Mx CVw CVb 3W1 2, 1.02 0.68 1.43 4.39 5.34 ~3W2 38 0.49 0.31 0.75 7.92 3.14 14 63 ~ 6 8 8.46 19.38 2) 170 - 3 4 11.57 23.28 ADI 200 0.19 0.11 0.29 9.90 6.70 *AD2 201 0.12 0.08 0.20 11.49 6.64 PAS 201 0.56 0.38 0.82 10.72 lili EEE eee DISCUSSION. Sulcoretepora parallela is unmistakable in appear- _ jce due to its strap-like branches with a regular arrangement of 159 autozooecial apertures which are divided by longitudinal ridges. This bryozoan displays very little variation either within or between colonies in a population. Computed coefficients of variation for branch width (BW1), branch thickness (BW2), and autozooecial apertural diameter (AD1 and AD2) are all very low. The values for the other parameters, those that are a measure of autozooecial spacing (Z1, Z2, and AS), are higher, but are still regarded as low when compared with other bryozoan taxa. Sulcoretepora parallela has a wide distribution in the Carbonifer- ous of the British Isles. It is common in the Carrick Lough/Sillees River assemblage. Three other species of Sulcoretepora have been described from the Carboniferous of the British Isles: S$. ramosa Owen 1973, S. raricosta (M‘Coy, 1844), and S. robertsoni (Young & Young, 1877). S. ramosa is nota sulcoreteporid but is a hyphasmoporid cryptostome. Branches are circular and not bifoliate, it lacks a median wall and lunaria, and acanthostyles are common. It occurs in County Ferman- agh and is redescribed herein as Clausotrypa ramosa (Owen, 1973). The other species of Sulcoretepora differ from S. parallela in a number of respects. S. raricosta has autozooecia of similar dimen- sions to those in S. parallela but has more autozooecia developed on one side of the branch than the other. Branches of S. robertsoni are nearly circular in cross-section, autozooecial apertures are larger, and interapertural areas are pitted and more ornate (Young & Young 1877). STRATIGRAPHICAL RANGE. Carboniferous (Holkerian—Pendleian). DISTRIBUTION. British Isles. Family GONIOCLADIIDAE Waagen & Pichl, 1885 Genus GONIOCLADIA Etheridge, 1876 TYPE SPECIES. Carinella cellulifera Etheridge, 1873, by original designation from the Lower Carboniferous of Carluke, Scotland. Goniocladia cellulifera (Etheridge, 1873) Figs 84c, 94-102 1873a_ Carinella cellulifera Etheridge: 433. 1873b Carinella cellulifera Etheridge; Etheridge: 101. 1876 Goniocladia cellulifera (Etheridge); Etheridge: 522. 1880b Goniocladia cellulifera (Etheridge); Vine: 81. 1880c Goniocladia cellulifera (Etheridge); Vine: 507. 1885. Goniocladia cellulifera (Etheridge); Waagen & Pichl: 804. 1887 Goniocladia cellulifera (Etheridge); Young: 463. 1888 Goniocladia cellulifera (Etheridge); Vine: 77. 1888 Goniocladia cellulifera (Etheridge) var. robusta Vine: 78. 1953. Goniocladia cellulifera (Etheridge); Bassler: 89, fig. 54. 1983 Goniocladia cellulifera (Etheridge); Utgaard: 434, figs 213, la-h. 1986a Goniocladia cellulifera (Etheridge); Bancroft: 23. 1987 Goniocladia cellulifera (Etheridge); Bancroft: 196. MATERIAL. BMNH PD9563, 9701, 9703-9721; TCD.34112- 34120, 34135, 34146-34147, 34150-34154, 34157, 42589, 42600a, 42602b, 42604a, 42606c; BELUM K2162-5, K12003, Upper part of the Glencar Limestone (Viséan, Asbian), Carrick Lough, County Fermanagh. TCD.42512, Upper part of the Glencar Limestone (Viséan, Asbian), Sillees River, County Fermanagh. EMENDED DIAGNOSIS. Goniocladia with large reticulate or occa- sionally adnate zoaria composed of bifoliate straight to gently curved branches. Branches anastomose at regular intervals to form 160 P.N. WYSE JACKSON Figs 94-100 Goniocladia cellulifera (Etheridge, 1873b); Upper part of the Glencar Limestone (Viséan, Asbian), Carrick Lough, County Fermanagh; 94, BMNH PD9719, colony fragment with typical branching pattern at a triple point, x9; 95, BMNH PD9704, laterally flattened branch with sharp median ridge on left and rounded carinal ridge on right; autozooecia are arranged in longitudinal rows with circular apertures: lunaria are indistinct; cystopores open to the surface and are seen as small circular ‘pits’ in interapertural areas, x50; 96, BMNH PD9718, close-up of branch with circular autozooecial apertures and small cystopore-openings, x45; 97, BELUM K2164, portion of reticulate colony. Branches divided perpendicular to the plane of the median wall; they coalesce to form large open rhombic, polygonal to irregularly-shaped fenestrules, x6; 98, BELUM K2163, robust holdfast of colony with portion of branch reticulation, x6; 99, BMNH PD9721; 99a, longitudinal section. Autozooecia diverge from a planar thin median wall at low angles; chambers are separated by vesicular tissue which consists of irregular to spherical vesicles, x20; 99b, transverse section; the carinal ridge is on the left hand side while the sharper median ridge forms a sharp keel on the right. Autozooecia are polygonal, squat, and the median wall is straight, x40; 100, BMNH PD9563, transverse section through a triple point reflected by the division of the median wall, x20. large pentagonal or polygonal fenestrules. In cross-section branches are pyriform. They are bisected by a compound median wall which protrudes as a faint carina on the rounded barren surface and as a strong keel on the obverse surface. Autozooecia are large, circular and arranged in quincunx in four to seven longitudinal rows either side of the median wall, from which they are budded. Low indistinct lunaria are frequently devel- oped around autozooecial apertures. Interapertural areas are smooth. Autozooecial chambers are narrow, recumbent, and closely packed in the endozone. They diverge away from adjacent chambers in the exozone, and vestibules are orientated at a high angle to the zoarial surface. Autozooecial chamber walls are thin. Basal diaphragms are rare. DESCRIPTION. Zoaria form large planar reticulate colonies of un- known maximum dimensions. The largest reticular fragment | LOWER CARBONIFEROUS BRYOZOA Fig. 101 Goniocladia cellulifera (Etheridge, 1873b). Line drawing of external features of BMNH PD9704; scale bar = 1 mm. Fig. 102 Goniocladia cellulifera (Etheridge, 1873b). Line drawing of | protion of a reticulate colony of BELUM K2164; scale bar = 1 mm. examined measured 19 x 12mm. Branches are bifoliate, laterally ‘flattened, straight, and maintain a constant width along their length except prior to division when a small increase in width occurs. Branches divide, at angles of 30° to 70°, at short regular intervals. -Coalescing of branches produces a regular pattern of pentagonal, “nexagonal, or polygonal fenestrules upto 4.9mm long by 3.4mm wide. Branch division frequently produces three branches 60° apart. _ Dissipiments are absent. Branch cross-sections are narrow with a pyriform to rhombic outline. The barren reverse surface is well rounded with a faint to “distinct longitudinal carina while the celluliferous frontal surface is divided by a strong narrow angular median keel. These ridges are the _pxternal manifestations of the internal median wall. | Autozooecial apertures are arranged in quincunx in four to seven ongitudinal rows, either side of the median ridge. Autozooecial "ipertures are large, circular to rarely oval in shape. Proximal lunaria "ire developed around most apertures. The size and thickness of the _junaria decreases towards the median keel. Interapertural areas are smooth and featureless. Autozooecial apertures are regularly spaced within longitudinal rows. Interapertural spacing decreases towards he median keel from five to two diameters apart (Table 31 and Fig. 103). 161 Colonies arise from stout holdfasts up to 13mm in width. Initial colony growth is encrusting where autozooecial apertures are large, circular in shape and closely spaced. From this adnate portion three to six erect branches arise, which either remain isolated as erect eschariform colonies or anastomose to form reticulate colonies. Internally branches are divided by a thin, straight compound median wall. It is composed of a dark coarse central layer sur- rounded by a thin pale laminated layer. Autozooecia are budded from this wall. In the endozone autozooecial chambers are long, narrow, recumbent, and the thin chamber walls are shared. In transverse section they are semi-circular to pentagonal in shape. Autozooecial chambers curve distally from the median wall at angles of between 60° and 20°, and they diverge away from each other so that in the exozone they are isolated. The thickness of the exozone is greatest at the widest portion of the branch, where vestibules are oriented at a high angle to the zoarial surface, and decreases in the autozooecial rows towards the median keel, where vestibules lie at a low angle to the zoarial surface. Small hemispherical vesicles 0.01 to 0.03mm in diameter are commonly found between autozooecial chambers. They are thin- walled, irregularly shaped and may be infilled with stereom in the endozone. Table 30 Measurements of Goniocladia cellulifera in mm. N=18. NM x Mn Mx CVw CVb BWI 137 1.47 0.83 2.30 7.38 5.00 BW2 44 0.69 0.33 0.98 9.03 6.21 Zl 62 6.30 4 9 13.06 12.01 Z2 161 3.90 2 6 SA 9.41 AD 180 0.12 0.10 0.20 11.73 10.28 AS 178 0.57 0.32 1.12 24.90 7.35 FL Wf 4.91 3.69 6.56 21.30 -l FW 7 3.40 2.56 4.40 17.89 = Table 31 Differences in apertural spacing in longitudinal rows in Goniocladia cellulifera (in mm). Carinal ridge -------------------------------------------- >Median keel (barren surface) (obverse surface) 1 2 3 4 5 6 7 NM 13 13 13 12 12 10 5 Mn 0.73 0.52 0.43 0.42 0.39 0.33 0.47 Mx 1.12 1.04 1.00 1.02 1.00 0.73 0.75 X 0.86 0.71 0.64 0.61 0.59 0.55 0.53 DISCUSSION. Goniocladia cellulifera is quite common in the Lower Carboniferous of the British Isles, where it has been described from the Midland Valley of Scotland, Cumbria, Northumberland, and Yorkshire. It is very distinct with an unusual laterally flattened branch bearing a number of rows of autozooecial apertures and a sharp median ridge. It was first described as Carinella cellulifera by Etheridge (1873a), but he later realised that that generic name was pre-occupied by a nemertean worm, and so proposed the name Goniocladia (Etheridge 1876). He suggested (Etheridge 1873a, 1873b), on the evidence of external features, that Goniocladia was an intermediate between Polypora M‘Coy and Fenestella Lonsdale. While Goniocladia does possess a median keel (as in Fenestella) and has more than two rows of autozooecial apertures (as in Polypora), internally it resembles neither. In Goniocladia the zoarium is divided into two by a straight 162 Interapertural spacing (in mm) 0 y 2 4 6 8 inal Rid Median Ri sane e Autozooecial rows oe ols Fig. 103 Goniocladia cellulifera (Etheridge, 1873b). Graph of apertural spacing. vertical median wall from which autozooecia are budded; Fenestella and Polypora have no such median wall. Autozooecia in these two genera are budded from a basal wall. STRATIGRAPHICAL RANGE. Lower Carboniferous. DISTRIBUTION. British Isles. PALAEOECOLOGY OF THE COUNTY FERMANAGH BRYOZOAN FAUNA The silicified Viséan fauna found towards the top of the Glencar Limestone has proved to be very diverse. It has been the subject of a number of research papers: the bryozoan element of the fauna has been systematically described by Bancroft & Wyse Jackson (1995), Olaloye (1974), Tavener-Smith (1965a, 1965b, 1971, 1973), Wyse Jackson (1988) and Wyse Jackson & Bancroft (1995a); the brachiopods by Brunton (1966a, 1966b, 1968, 1984); and the sponges by Reid (1970). These workers have recorded a large number of taxa (Tables 32 and 33). Other palaeontological work includes that of Gardiner & Mason (1974) who reported the occurrence of palaeoniscid fish from strata just overlying the Glencar Limestone, at a nearby locality west of Carrick Lough. The fauna in County Fermanagh is essentially a bryozoan- brachiopod assemblage with rare taxa of other groups, such as trilobites, gastropods, bivalves, sponges and corals. This community is similar to that developed on the slopes of Asbian reefs of the Cracoe area, Yorkshire (Mundy 1978). Brunton (1987) tabulated the diverse and abundant fauna from Carrick Lough and Sillees River (Table 33). The number of bryozoan species described has been increased in this paper, and Bryozoa are now the largest element (numerically by species) in the community. The bryozoan fauna is dominated by fenestrates, which in turn are dominated by fenestellids. However, the delicate cryptostomes are also quite common (Table 32). During this study one blastoid species and one ostracod species not recorded by Brunton (1987) have been found. Three specimens of the blastoid Monoschizoblastus rofei (TCD.9605), common in Asbian strata (Waters & Sevastopulo 1984a, 1984b), and one speci- men of the ostracod Polytylites (TCD.9606) complete with both left and right valves were found. P.N. WYSE JACKSON Table 32 List of bryozoans from the Lower Carboniferous (Viséan, Asbian) of Carrick Lough and Sillees River (Bancroft & Wyse Jackson 1995; Olaloye 1974; Tavener-Smith 1965a, 1965b, 1971, 1973; Wyse Jackson 1988 and herein; Wyse Jackson & Bancroft 1995a). Order CRYPTOSTOMATA (9)* *Hexites paradoxus sp. NOV. *Nematopora hibernica sp. nov. *Pseudonematopora planatus sp. nov. *Rhabdomeson progracile Wyse Jackson & Bancroft *R. rhombiferum (Phillips) *Rhombopora cylindrica sp. nov. *R. hexagona sp. nov. *Streblotrypa pectinata Owen *Clausotrypa ramosa (Owen) comb. nov. Order TREPOSTOMATA (7) *Leioclema indentata sp. nov. *Dyscritella miliaria (Nicholson) *Tabulipora urii (Fleming) *T, howsii (Nicholson) *T, minima Lee *Stenophragmidium sp. Nodular trepostome Order CYSTOPORATA (4) *Fistulipora incrustans (Phillips) *Goniocladia cellulifera (Etheridge, jun.) +Goniocladia sp. *Sulcoretepora parallela (Phillips) Order FENESTRATA (49) *Baculopora megastoma (M‘Coy) *Diploporaria marginalis Young & Young *D. tenella Wyse Jackson *[chthyorachis newenhami M‘Coy *Thamniscus colei Wyse Jackson *Rhombocladia dichotoma (M‘Coy) comb. nov. Penniretepora pluma (Phillips) P. gracilis (M*Coy) P. frondiformis Olaloye P. normalis Olaloye P. cucullea Olaloye P. cf. flexicarinata Young & Young P. sinuosa (Hall) P. elegantula (Etheridge, jun.) P. rotunda Olaloye P. tortuosa Olaloye Fenestella frutex M‘Coy EF. ivanovi Shulga-Nesterenko F. multispinosa Ulrich F. modesta Ulrich F, hemispherica M*Coy F. parallela Hall F. rudis Ulrich multinodosa Tavener-Smith F, plebeia M‘Coy FE. cf. arthritica (Phillips) F. praemagna Shulga-Nesterenko F. fanata Whidborne carrickensis Tavener-Smith F. cf.spinacrisata Moore F. cf.funicula Ulrich F. cf filistriata Ulrich E. subspeciosa Shulga-Nesterenko EF. pseudovirgosa Nikiforova F. cf. albida Hall F. oblongata Koenig F. cf. delicatula Ulrich F, polyporata (Phillips) F. irregularis Nekhoroshev Hemitrypa hibernica M‘Coy Levifenestella undecimalis Shulga-Nesterenko Minilya plummerae (Moore) M. binodata (Condra) M. oculata (M‘Coy) Polypora dendroides M*Coy P. verrucosa M‘Coy P. stenostoma Tavener-Smith Ptilofenestella carrickensis Tavener-Smith Ptiloporella varicosa (M*Coy) Ptylopora pluma M‘Coy parva Tavener-Smith Septopora hibernica Tavener-Smith * Described in this study; + to be described elsewhere. | | | LOWER CARBONIFEROUS BRYOZOA Table 33 Abundance and diversity of the fauna at Carrick Lough and Sillees River. Element Number of genera* Number of species* Bryozoans 30 69 Brachiopods 47 56 Arthropods Trilobites 6 7 Ostracods 1 1 Echinoderms Crinoids 2+ 2+ Blastoids 1 1 Molluscs Gastropods 4 4/5 Bivalves 2 22 Sponges ? 210 Corals 3 24 Annelids 1 ! *Modified from Brunton (1987) Although the fauna from County Fermanagh is very well-known, its palaeoecology has been a matter of some debate. The succession at Carrick Lough consists of thin alternating beds of dark argillaceous muddy limestones and paler grey to yellowish bioclastic limestones, while at Sillees River the fauna was etched from biomicritic lime- Stones. The lithology of the bryozoan-bearing strata is that of a distal reef or off-reef deeper water facies. __ Extensive reef development occurred in the Asbian of north west Ireland (George et al. 1976); one such reef lay just south of the DINANTIAN [GR [CH [AR [HO|AS [ER [PE_ ORDER Hexites paradoxus Nematopora hibernica Pseudonematopora planatus Rhabdomeson progracile Rhabdomeson rhombiferum Rhombopora cylindrica Rhombopora hexagona Cryptostomata Streblotrypa pectinata Clausotrypa ramosa Baculopora megastoma Diploporaria marginalis Diploporaria tenella Ichthyorachis newenhami |s 1& \2 | Thamniscus colei Rhombocladia dichotoma Leioclema indentata Dyscritella miliaria Tabulipora urii Tabulipora howsii Tabulipora minima Trepostomata Stenophragmidium sp. Fistulipora incrustans Sulcoretepora parallela Goniocladia cellulifera Cystoporata lAbbreviations: SIL: Silesian; CR: Courceyan; CH: Chadian; AR: Arundian; HO: Holkerian; AS: Asbian; BR: Brigantian; PE: Pendleian. Fig. 104 Range chart of bryozoans described in this study. | 163 collecting localities which are in proximal reef or off-reef facies (Tavener-Smith 1973). Bryozoans favour clean sediment-free water in which suitable substrates are available (Schopf 1969, Cuffey 1970).The depositional environment at Carrick Lough and Sillees River did not favour extensive bryozoan colonisation because the water was too muddy, as was the substrate. Fistulipora incrustans and Tabulipora howsii colonised substrates such as brachiopod shells, spines and other bryozoans. They developed small button-like zoaria, which were size-controlled because the adjacent muddy sediment could not be extensively encrusted by bryozoans. Similar evidence for the soft nature of the substrate is suggested by brachiopod attachment styles. The 56 brachiopod species discussed by Brunton, 1987) are more or less equally divided into those that are pedicle supported, quasi- infaunal, and spine supported. Only 3 species were found to be permanently cemented, probably to other brachiopod shells. It is suggested that bryozoans found permanent points of attachment to small shelly substrates from which erect or adnate colonies devel- oped, and that they were not attached directly to any lithic substrate. Bryozoans are abundant at Carrick Lough and Sillees River, and calcified, decalcified and silicified zoaria have been obtained. Silicification seems to have occurred in the pale limestones and not in the argillaceous and muddy limestones. It is impossible to quan- tify the abundance of bryozoan colonies in pure numerical values. This is due to the very fragmented nature of the zoaria. Almost without exception all colonies are broken or disarticulated but not heavily abraded nor shredded into fine hash. Bryozoan colonies are found lying on bedding planes and are stacked one upon another. The fauna comprised bryozoans of four of the five stenolaemate orders (cyclostomes were absent), with a numerical and taxonomic weighting towards the fenestrates. The cryptostome taxa all formed small delicate erect expansions and were quite abundant. In contrast, while the trepostomes are taxonomically moderately diverse they occurred in small numbers. The cystoporates which comprised three taxa were reasonably abundant. The bryozoan community was dominated by large erect planar and conical fenestrate zoaria, which exploited the seawater up to 20 cm above the substrate. Between Fenestella s.l. zoaria grew the smaller fenestellids such as Baculopora, Diploporaria, Ichthyorachis, Penniretepora and Thamniscus, as well as the delicate cryptostomes, two Tabulipora species and the cystoporates Sulcoretepora and Goniocladia. Colonising the sea floor were Rhombocladia, Tabulipora, Stenophragmidium and Fistulipora. In addition Stenophragmidium grew epiphytically ona soft cylindrically shaped substrate (possibly algae), and Tabulipora and Fistulipora encrusted brachiopod spines and crinoid ossicles. Few holdfasts were recovered from this fauna, which is in keeping with other fenestrate-rich faunas (F.K. McKinney pers. comm.). Of those that were found, the majority belong to the cystoporates Goniocladia cellulifera and Sulcoretepora parallela, with a single holdfast of Thamniscus colei being present in the sample. The attitude and prevalence of fenestellid fronds on bedding planes, the limited abrasion, and lack of holdfasts indicates that the fauna has been translocated. This movement has taken place downslope off reef slopes into deeper water. However, the move- ment distance cannot have been great as fragemnts display little abraision of fine surface skeletal detail, such as carinal nodes and the superstructure of Hemitrypa hibernica. If, as Brunton (1987) postulates, the Carrick Lough and Sillees River fauna is an in situ assemblage then bryozoans preserved in growth position would be expected. As outlined above they are not thus preserved. Tavener-Smith (1973) considers that the Carrick Lough fauna is an accumulated assemblage, that is it has not been 164 fossilized in situ. From the sedimentological and bryozoan evidence this latter interpretation is preferred. However, the possibility that the fauna represents an in situ brachipod community into which bryozoans have been washed from adjacent reef slopes cannot be excluded. i COMPARISON WITH OTHER ASBIAN FAUNAS. ee Asbian faunas in the British Isles have received little attention, and many species recorded from County Fermanagh have not been recorded elsewhere. Bancroft (1984) and Wyse Jackson, Bancroft & Somerville (1991) document the faunas of several sites in Britain: Ashfell Road Cutting, Cumbria [1 cryptostome species / 7-8 fenestrate species / several trepostome species / 2 cystoporate spe- cies]. Odin Fissure, Treak Cliff, Derbyshire [1/8/1/1]. Penruddock, Cumbria [1/12/several/2]. Redesdale, Northumberland [1/1 1/3/1]. Nant-y-Gamar, near Great Ormes Head, north Wales [1/3/5/1]. This fauna is somewhat unusual as it is trepostome-dominated. Carrick Lough and Sillees River, County Fermanagh [9/49/7/4]. These faunas are generally dominated by fenestrates and fenestellids in particular, with taxa of other bryozoan orders, par- ticularly cryptostomes, being conspicious by their scarcity or apparent absence. This may be due to poor preservation, which makes identi- fication of delicate cryptostome species difficult. The fauna in County Fermanagh encompasses all species re- ported from the above localities with the exception of five: Fenestella matheri, Septopora cestriensis, Batostomella sp., an undescribed species of Leioclema, and Stenodiscus tumida. Fenestella plebeia is common to all six localities; Fistulipora incrustans and Hemitrypa hibernica occur in five while Fenestella bicellulata occurs in three. The importance of the Fermanagh fauna lies in its rich taxonomic diversity and reasonable fossil preservation which will allow for future comparison with other Asbian faunas. Such investigations may reveal that the British and Irish faunas are more diverse than hitherto appreciated. PATTERNS OF BRYOZOAN ZOARIA REPLACEMENT BY SILICA Replacement of calcified bryozoan zoaria by silica has allowed for easy extraction from their carbonate matrix by acid digestion. The large number of specimens obtained in this way allows for detailed Figs 105, 106 Patterns of silica replacement in bryozoan zoaria from the upper part of the Glencar Limestone (Viséan, Asbian), Carrick Lough, County Fermanagh, as illustrated by examples of Fistulipora incrustans colonies; 105, BMNH PD9740; 105a, euhedral and sub-hedral microquartz replacement of calcite autozooecial skeletal structure, producing a thin exterior rim that is usually fuzzy in appearance, megaquartz crystals infill the autozooecial chambers, x25; 105b, grading of euhedral and sub-hedral microquartz rim into spherulitic chalcedony which replaces the remainder of the skeleton (Pattern 2), x25; 105¢e, total replacement of skeletal walls and autozooecial chamber megaquartz infill by pervasive chalcedony and obliteration of original skeletal structure, x25; 106, BMNH PD9741, replacement of crinoid ossicle stereom by radial spherulitic chalcedony; typical Pattern 2 to 3 replacement, x25. P.N. WYSE JACKSON | LOWER CARBONIFEROUS BRYOZOA quantitative taxonomic, ontogenetic and palaeoecological studies. However, it has been argued that the silicification process is selective and does not necessarily preserve all elements of a fauna (Whittington & Evitt 1954), and secondly, that preservation is poor and only retains the external features of the skeleton (Cooper & Grant 1972— 1975, Tavener-Smith 1973, Taylor & Curry 1985). Both calcified and silicified bryozoan fragments are found at ~ Carrick Lough and Sillees River, and comparison of both fractions : indicates that more taxa are represented silicified than calcified. This : probably reflects the ease by which silicified specimens are re- _ trieved. Some authors report excellent preservation by silica of _ invertebrate skeletal microstructure (Brunton 1976, Holdaway & Clayton 1982), and of plant vessels (Stein 1982), while Schmitt & Boyd (1981) noted that relict ultrastructure is commonly observed in _ megaquartz crystals. There is no doubt that the process of silicification can produce both excellent and poor preservation. Schmitt & Boyd _ (1981) noted that the quality of preservation is related to the timing of replacement: poor preservation is due to delayed replacement, | whereas excellently preserved fine detail is produced by immediate | replacement of the calcite skeleton by silica. They sub-divided silica | patterns into five major types, of which patterns | to 4 represent delayed replacement and pattern 5 immediate replacement. A number of randomly orientated thin sections were cut from silicified blocks collected from Carrick Lough. In general bryozoan | autozooecial chambers are infilled with megaquartz, whereas the | skeleton is replaced by a mixture of spherulitic chalcedony which is | radial in appearance, tiny sub-hedral to euhedral microquartz crys- tals (<20um), and occasionally some megaquartz. Silicification may occur in descrete steps, which will produce different silica styles. Initially the calcite bryozoan zoarium is quite faithfully replaced by euhedral and sub-hedral microquartz crystals, | producing a thin exterior rim that is usually fuzzy in appearance (Fig. 105a). As the intensity of silicification increases autozooecial _and vesicular chamber walls are totally replaced by spherulitic | chalcedony (Pattern 2 of Schmitt & Boyd 1981) (Fig. 105b). Finally, chalcedony pervades both the walls and the megaquartz crystals that infilled autozooecial chambers obliterating all original skeletal struc- ture (Fig. 105c). Overgrowing this ubiquitious chalcedony are found small rhombic dolomite crystals. In some zoaria silicification has / not been totally pervasive. Where this occurs the outer laminated cE: (eg. in Fenestella s.l.) is replaced, with the inner granular tissue left unaltered. These silica patterns indicate that replacement of the bryozoan colonies in County Fermanagh was delayed. In many cases replace- ment is incomplete: colonies may be lightly silicified on their outer surfaces and not silicified internally, or else portions of branches may be silicified and others not. In these situations colonies are very delicate and easily fragmented. Rarely are zoaria completely '\silicified. However, where this has occurred, considerable skeletal detail is retained, which allows for reasonable taxonomic determinations and morphological descriptions. In these cases the silica style is that of Pattern 2. In total 29 genera, containing 68 species have been described from the Viséan of County Fermanagh. This key is an aid to their identi- fication. It has been necessary to construct the key in two styles: a multi-element dichotomous-trichotomous portion and a tabular por- ion. Where possible taxa are keyed out in the former, but where ‘ 165 there are a large number of species in a genus (as with Fenestella s.1.) the latter has been used. In some cases several routes through the key will lead to the same taxon. No attempt has been made to designate the various Fenestella species to the genera erected by Morozova (1974). The key to Fenestella s.l. species and other fenestrate taxa is based on the findings of Bancroft (1984), Olaloye (1974), and Tavener- Smith (1965a, 1965b, 1971, 1973). No attempt has been made to check their taxonomic determinations. Key 1 Autozooecia developed on obverse surface only, tripartite skeleton GeV eloped iraascacrceseierer ct reer reie ee cere ce ee ee ci 2 Autozooecia developed throughout ZoariuM ...........c:ccccccseceeeeeeeeeees 26 Dey Oaniate th CUlale meets. tect rese a eee ee ea 3 Zoaria small pinnate or non-pinnate expansions Zoaria basket or cup-shaped ............ccecssereeseeeseenee Se LOWS Ol atozOOeCialOnlbranches ae ee ee eee 4 More than two rows of autozooecia developed ..............cccccccceeereee 11 fame btaniches:allthicrsalne SiZcwemnacmter se ret tte are nee enn 5 Branches of two sizes: a few primary and many secondary ............... CRG Seta See sae Tracsn uni ered teem ace gaaersesiauers Ptiloporella varicosa See Mediantcanina presen trcreeree re rs eee ee 6 Median carina absent ...........cccccccceeceeeeees Levifenestella undecimalis 6 Obverse surface protected by a honeycomb-like superstructure ......... Rois REE PSE ET acon ems vds aaa eee acne ev ona Sees Hemitrypa hibernica Obverse surface not covered by a superstructure ............0.0cccccceeeee0-2e q 7 Carinal nodes arranged in a straight lineFenestella s.1. (go to Table 34) Carinal nodes in two offset Zigzag COWS ..........cccccccceceecseseeseeeeeeseseees 8 8 Fenestrules quadrate, hourglass-shaped, serving four autozooecial ap- (GIRS ccceacdeoosed 370200262 econsaceacboce-Ehaneonesraasaaceacesncaisedtaadie6 oasesncoacedasancas6%0 9 Fenestrules elongate, serving 8 autozooecial apertures .................. 10 9 Peristomes large, median carina well developed, reverse surface often MOGUILOSE 22 A aoeres eget fetaa pense ab oscsze se Bogen vosac eines Minilya plummerae Peristomes slight, median carina weak, reverse surface smooth to mareimallygestht Aled mewsees meee ee. ayaa eee eee Minilya nodulosa 10 Branch margins slightly indented by autozooecial apertures, median carina weak, one carinal node situated at branch/dissepiment junction OSCE COSE CLOSE CCU BCOCE PER ERECT TE ORS onerer cee ern eee Minilya binodata Branch margins straight, median carina moderate, arrangement of carinal nodes somewhat irregular ................:c0c0eesee0e0s Minilya oculata 11 3 rows of autozooecia, keyhole-shaped apertures ..................00c00000000- Seeee smoot aan Saeidegs ouaee seas ace cate Seer a oe ena cee ook Polypora stenostomata 4—5 rows of autozooecia, oval apertures, fenestrules oval 2.50mm long Bec CCRS SOE GOARCCE OO AEB eer ER ECRELOE EHR CRS CICeNSSORE Polypora dendroides 4 rows of autozooecia, peristomes around apertures, fenestrules elon- gates 30-47 Omi on oer ere eee: Polypora verrucosa WD ZO ATIUITANP IM ALC cerns ne ree aeccserseae eters rien eect vat ceases see eee 13 ZO ATAU TIAN OM P UUM ALE eee ec oavc saa cee ee ec tessce ee az 21 13 Lateral branches lack dissepiments .................... 14 Lateral branches connected with dissepiments .............:ccccccccc:cceeeeees Seti ae Oeste cic teee vas coe eat ae ES tae Sess Heese Wa Sean Ptylopora pluma parva Wateral branches|coalescelmesss es ssesssee reese Septopora hibernica 14) 2rOWSiOtautozooeciaioni branches |pesecsse-ceescsecessesescessenceseesee senses IS) More than two rows of autozooecia ......... Ichthyorachis newenhami 15 1 autozooecial aperture between lateral branches ................cc00000000- 16 2 autozooecial apertures between lateral branches ... ae Site) >5 autozooecial apertures between lateral branches .............c:0000e000+- Teste Nate cette sist ones easeeataraerrnentinscuvetetetcoeecotaee Penniretepora tortuosa 166 26 OT 28 30 31 Lateral branches straight ............:ccccscccscssecereereeceesessersseressenenssensssae® 17 Lateral branches sinuous (margins broken by protruding autozooecial ADEMUUKES) ee wiseesscureeaetane-seeeteeseen ceaeese renee cee Penniretepora gracilis Median carina strong, straight Penniretepora frondiformis Median carina weak, straight Penniretepora normalis Median carina consists of 3 sinuous pustulose ridges ...........-.-:e0e Penniretepora flexicarinata Mictinistenmis tial oii tpec steerer sescstesnnersers eerecehecreeet eee ccrearrnssnenronswre nen 19 Main stem Sinuous .............c:ccccescereeeereeeeeee Penniretepora elegantula [LAL ATMO NES STNELN? o5-cocrscoscacecqacnncnenertouéadobsossacoaacoossesecnscascesece¢ 20 Lateral branches sinuous ............:::ccceeereeeeees Penniretepora cucullea Peristomes, horseshoe-shaped, developed on distal side of autozooecial BYDSTURELRSS ares cocoreaneniccncabeoouncosce-cecctouaseocacacebaashcnatoe Penniretepora pluma PETIStOMeS ASEM tereeeeeee neeneseeeeesenense ener Penniretepora sinuosa 2 rows of autozooecia ON branches ..............:.ssscseesccsecsseseneseeeeceneeee 2D, More than two rows of autozooecia on branches .............:0:0+ee 23 Lateral margins of branches serrated .......... Diploporaria marginalis Lateral margins flexuous or smooth ................++ Diploporaria tenella Branches delicate, flexuous, oval/circular apertures ............:0:eees SOE rR Do coca COLON EE CY EON DSDOTBOOTONOOECO Baculopora megastoma Branches robust, apertures with peristOMEs ..........:::seecesereeeeereeees 24 Complete peristomes surround apertures .............-. Thamniscus colei Proximal peristomes Only ...........::cccceeceseeeeereee Thamniscus rankin Branches connected by dissepiments ...... Ptilofenestella carrickensis Branches bear no dissepiment ............:.::000:0c008 Thamniscus colei Zoaria ramose or encrusting, with polymorphic zooecia, distinct endozonal and exozonal differentiation .............scccscsesereesereteeeees 27 Zaria POSSESS CYStOPOTES .......-.csceccccsseeceeeeseneneeeesceeneersecsseesestesseees 32 Zoaria dendroid with zooecia budded from central axis ................. 34 Mesozooecia developed .................scsssscsssessssenenscoerenceesrecccurcnsseseters 28 Ext aZOOECLAIGEVE LOPE feeeesseetee cence easnsene tne eneeeeeee neers erect eeeeceeen 7S) Zoaria ramose with thick exozone and autozooecia heavily indented by FISANTU NODES 5s:ccenosecoecaococco Lessee anoaonicsoneasoasosoondes Leioclema indentata Exilazooecia common with many disposed between autozooecia...... DE Oc ee eo Otc coe ne eeE eae nce eSaares Dyscritella miliaria Exilazooecia relatively rare 50) Ring septa developed 31 Hemiphragma developed ...............:0c1eeee: Stenophragmidium sp Zoaria ramose, autozooecial apertures circular, ring septa tips slightly thickened, deflected proximally, exilazooecia in clusters ............-.2++ ESECaRSSaNIEIE eNOS DIOTgTROCODEE Sona A a00 cabs ze RSCG OG SEER Ea BNaGMaCEECERC HCE Tabulipora urii Ww i) 33 40 P.N. WYSE JACKSON Zoaria thin adnate sheets, autozooecial apertures polygonal, ring septa | planar/thin, exilazooecia rare ..........cccceeeeeeeeeeneees Tabulipora howsii Zoaria robust, exozone thin, autozooecial apertures oval, exilazooecia | Tabulipora minima — Zoaria encrusting, adnate, with well developed rectangular (vesicles) | cystopores between tubular autozooecia ......... Fistulipora incrustans Branches bifoliate with spherical CyStOpoOres ............:ccscssceeseeseeeesees 35 Branches strap-like, zoaria budded from a plicated median wall, autozooecial apertures with proximal lumaria ............:.0::ccceeeeeeeeeeeene cares ca TAN Seeds RSE ERE OT ee Rca ane Sulcoretepora parallela | Branches sub-rounded, with a distinct keel and carina, autozooecia budded from a straight median wall .............. Goniocladia cellulifera Branches articulated, very slender (01—25mm), circular or polygonal in cross-section, hemisepta absent ..............c.-cescerceseeseeseeseeseeseeseeeeteeees 35 Branches not articulated, cylindrical (05—60mm), hemisepta usually | JSYTSSTSVOLE HA ORAM) opp: cer eeesaco cocnnanone cee erBeeaBcOnec0a LoGaN: ae Sbo0e cece eS Autozooecial apertures present on all branch surfaces Autozooecial apertures absent from a distinct reverse surface don NE ee soe essen oad nade ee Sica eer ees ay Nematopora hibernica Branch cross-section polygonal, autozooecia budded in annular fash- ion, autozooecial apertures oval, separated from each other by longitudinal nid esis eseeesenessrscttennrerese-eteeesteeenmae Hexites paradoxus Branch cross-section circular, autozooecial budding radial, autozooecial apertures circular, with proximal peristOMES .........-s:eeseeeeetererereseees slssedisteleascaceaieeeninea race. ee tes enema oe eaSe Pseudonematopora planatus Metapores very common, arranged in linear rows in interapertural areas Streblotrypa pectinata | Metapores irregularly disposed in interapertural areas ...........2-.-::0000 dcouddsengstensvost Gees OnBetsUc Es co ge te eeoRRE eee eno st sso een Clausotrypa ramosa Metapores either absent or only 1 per autozooeciuM .........-.2-.-100+- 38) Central axial cylinder present Axial cylinder not present ........2....-c1sssecsccsseseeseecenencecseceensassentecstenses Autozooecial apertures oval, of constant dimensions, with a large acanthostyle situated proximally, smaller stylets may be present ....... BOE Ee ce POOLS ESLER eee econ sce ob ode MEPREEES- Rhabdomeson progracile Autozooecial apertures pyriform, distally flared, of varying dimen sions, over 20 small stylets developed around apertures ra se CR ERE EPEC EN so -coccAREEE Rhabdomeson rhombiferum’ Autozooecial apertures circular, of constant size, with acanthostyles and heterostyles disposed in a circle around them, a single metapore may be situated proximal to autozooecia ..... Rhombopora cylindrica Autozooecial apertures circular to oval, of varying size around zoarium) LOWER CARBONIFEROUS BRYOZOA Table 34 Tabular key for the identification of Fenestella species from the Viséan of County Fermanagh and from other localities in the British Isles [* = not recorded from Carrick Lough by Tavener-Smith (1973)]. CHARACTER 9 12 144 15 16 17 18 1 3 4 5 6 7 0 NEV ESEY ES CA CSESEN CAEN CS ACNE Late [alatel*[a[afef= [a lela] a [a] FUGIEN ES EVEN CIEUCS COUSIN FAESCUCICUCAESES A 11 13 > ie Seas Fenestella bicellulata * A Fenestella ivanovi Fenestella frutex Fenestella multispinosa Fenestella tuberculo-carinata * ii Fenestella plebeia Fenestella papilliata * ASE) AEE) [@| EI] fa - 4 JABS FIRES] [| Fenestella morristi * Fenestella polyporata FEBS SSHAAAe Bog P a E rs QO Q e ra Fenestella quadridecimalis * jelafe[e|o[alala[a [al C A Fenestella modesta pa BE pa Fenestella hemispherica Fenestella parallela Fenestella rudis multinodosa Fenestella cf. arthritica ra] : Tefefelele[stelalotelole[m[o lala [al efefefefefelalelalal+lalelele[=[a [a] Fenestella praemagna Fenesiella fanata carrickensis Fenestella cf. spinacrista Fenestella cf. filistrata Fenestella cf. funicula A Fenestella subspeciosa Fenestella cf. albida Fontes |» fo [ele le) HEE HES al) HE AE ae (a Fi [| fal ey fa] Fz i afejcfela[alaslelalalate|s[alec[s[s | s| cle}o[=lalalelelelelo[e]alelalaa| elelc[s}alelala jatelela[slelela| fe[pe[slale|s|a Jajafalalcle A A jate}alo[ stata fei Tee) Explanation of characterers used in Table 34: 7. Zoarial appearance: 1. Branch shape: A, Rigid; B, Lax; C, High angle cone A, Straight; B, Sinuous; C, Irregular; D, Partly straight partly sinuous 8. Fenestrule shape: |2. Fenestrule length: A, Square; B, Rectangular; C, Hour-glass; E, Various; F, Elongate- A, < 0.5 mm; B, 0.5—1.0 mm; C, 1.0-2.0 mm; D, 2.0-3.0 mm; E, > 3.0 rectangular; G, Elongate-oval mm 9. Fenestrule edges: 3. Fenestrule width: A, Straight; B, Indented (by peristomes), C, Slightly undulating / A, < 0.5 mm; B, 0.5—0.75 mm; C, 0.75—1.0 mm; D, > 1.0 mm 10. Carina: |4. Number of autozooecial apertures per fenestrule: A, Distinct; B, Indistinct; C, Absent A, 2; B, 3; C, 4; D, 5; E, 5-8; F, 2-3; G, 3-4; H, 4-5: 1, >8 11. Inclination of branch sides from carina: 5. Interapertural distance: A, Steep; B, moderate; C, Gentle A, < 0.2 mm; B, 0.2-0.3 mm; C, > 0.3 mm 12. Carinal nodes: 6. Zooecial aperture/branch/dissepiment relationship: A, Large; B, Moderate; C, Small; D, Poorly developed A, Aperture occurs at branch/dissepiment junction; B, Apertures situ- 13. Spacing of carinal nodes: ated anywhere along branch length A. Regular — close < 0.25 mm; B, Regular — moderate 0.25 — 0.50 mm; 168 C, Regular — wide 0.50 — 1.00 mm; D, Regular — very wide > 1.00 mm; E, Irregular 14. Dissepiment depression on obverse surface: A, Well; B, Moderate; C, Slight; D. None (flush with branch surface — indicative of Fenestella parallela) 15. Dissepiment depression on reverse surface: A, Well; B, Moderate; C, Slight; D, None (flush with branch surface) 16. Incipient 3rd row of zooecial chambers developed before bifurcation: A, Present; B, Absent 17. Extra zooecial chamber in angle of bifurcation: A, Present; B, Absent 18. Rib(s) on dissepiment obverse surface: A, Present; B, Absent; C, Present (extending to meet carina); D, Occasionally present ACKNOWLEDGEMENTS. I thank Professor C.H. Holland and Dr George Sevastopulo who supervised the Ph.D. study of which this formed a part. This paper benefitted considerably from a review by Dr F.K. McKinney for which I thank him. I am grateful for the advice of Drs A.J. Bancroft, P.D. Taylor, I.P. Morozova, F.K. McKinney, C.J. Buttler, and all the other bryozoologists who sent me copies of their work. A.J. Bancroft is thanked for permission to examine and figure some of his material from the Upper Carboniferous of Yorkshire. Mr Bill Baird (National Museums of Scotland), Mr Gaynor Boon (Sheffield City Museum), Mr Philip Doughty, Mr John Wilson and Dr Andrew Jeram (Ulster Museum), Mr Stephen Howe and Mr Tom Sharpe (National Museum of Wales), Dr H.C. Ivimey Cook (formerly of the British Geological Survey), Mr Nigel Monaghan (National Museum of Ireland), Dr John Nudds (Manchester Museum), the late Dr David Price (Sedgwick Museum), Dr Ian Rolfe and Dr Neil Clark (Hunterian Museum), Dr Paul Taylor (Natural History Museum, London) and Dr Nigel Trewin (Aberdeen University) all kindly lent material in their care. I am grateful for the help of Mr Jeremy Smith and Dr Phillip Tubbs of the I.C.Z.N. and the late Professor David Webb (TCD) in answering some of my nomenclatural questions. I thank Garry and Bassia Bannister of Dublin who translated some Russian and Ukrainian texts for me at very short notice and in a very short time, also Ide nf Thuama (Royal Irish Academy) and Dr John Moore who located some Soviet and Australian papers for me. This work was carried out while in receipt of grants from Trinity Trust (Dublin) and the Irish Geological Asso- ciation which are gratefully acknowledged. 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Transac- tions and Proceedings of the Palaeontological Society of Japan, (NS) 46: 227-242. 1964. Bryozoa of Akiyoshi. Part 2. Lower Carboniferous Bryozoa from the Uzura Quarry. Transactions and Proceedings of the Palaeontological Society of Japan, (NS) 56: 295-308. 1972. Carboniferous Bryozoa from Bukit Charas, near Kuantan, Pahang, Malaya. In, Kobayashi, T. & Toriyama, R. (eds.), Geology and Palaeontology of Southeast Asia, 10: 35-62. Schlotheim, E.F. von 1820. Die Petrefactenkunde auf ibrem jetzigen Standpunke durch die Beschreibung seiner Sammlung versteinerter und fossiler Uberreste des Thier- und Pflanzenreichs der Vorwelt erlautert von E.F. von Schlotheim. Part 1. Gotha. Schmitt, J.G. & Boyd, D.W. 1981. Patterns of silicification in Permian pelecypods and brachiopods. Journal of Sedimentary Petrology, 51: 1297-1308. Schopf, T.J.M. 1969. Paleoecology of Ectoprocts (Bryozoans). Journal of Paleontology, 43: 234-244. Shulga-Nesterenko, M.I. 1955. Carboniferous bryozoans of the Russian Platform. Akademii Nauk CCCP, 57: 1-207. Simpson, G.B. 1895. A Handbook of the Genera of North American Paleozoic Bryozoa; with an introduction upon the structure of living species. New York State Geology 14th Annual Report: 403-669. Smyth, L.B. 1922. On some new species from the Lower Carboniferous of Ballycastle, County Antrim. Geological Magazine, 59: 21-24. 1925. A contribution to the geology of Great Orme’s Head. Proceedings of the Royal Irish Academy, 18: 141-164. 2; The Mountain P.N. WYSE JACKSON Stein, C.L. 1982. Silica recrystallization in petrified wood. Journal of Sedimentary Petrology, 52: 1277-1282. Tavener-Smith, R. 1965a. A new fenestellid bryozoan from the Lower Carboniferous of County Fermanagh. Palaeontology, 8: 478-491. 1965b. A revision of Retepora nodulosa Phillips, 1836. Geological Magazine, 102: 135-142. — 1966. Ovicells in fenestrate cryptostomes of Viséan age. Journal of Paleontology, 40: 190-198. 1971. Polypora stenostoma: a Carboniferous bryozoan with cheilostomatous | features. Palaeontology, 14: 178-187. 1973. Fenestrate bryozoa from the Viséan of County Fermanagh, Ireland. Bulletin of the British Museum (Natural History), Geology, 23: 389-493. 1981. The neotype of Retepora nodulosa Phillips, 1836. Geological Magazine, 1 118: 565. Taylor, P.D. & Curry, G.B. 1985. The earliest known fenestrate bryozoan, with a short review of Lower Ordovician Bryozoa. Palaeontology, 28: 147-158. Termier, G. & Termier, H. 1950. Paleontologie Marocaine, II. Invertebres de Lere Primaire, pt. 2, Bryozoaires et Brachiopodes. Maroc Service Geologie Notes et Memoires, 77: \—253. & 1971. Bryozoaires du Paléozoique supérieur de I’ Afganistan. Documents | des laboratoires de Géologie de la faculté des sciences de Lyon, 47: 1-52. Trizna, V.B. 1958. Early Carboniferous bryozoans of the Kuznetsk Basin.7rudy Vsesojuznyj nauchno-Issledovatel'skyj geologo-razvedocnyj Instituta, 122: 1436. 1962. Carboniferous bryozoans. Jn, Khalfin, L.L., Paleozoic biostratigraphy of Sayano-Altaiskoi. Trudy Sibiran nauchno-Issledovatel’skyj Instituta geologo- geofizikii Mineralnogo Syr’ya, 21: 55-61, 124-143. Ulrich, E.O. 1882. American Paleozoic Bryozoa. Journal of the Cincinnati Society 0, Natural History, 5: 121-175. 1884. American Paleozoic Bryozoa (continued). Journal of the Cincinnati Society of Natural History, 7: 24-51. 1888a. On Sceptropora a new genus of Bryozoa, with remarks on Helopora Hall, and other genera of that type. American Geology, 1: 228-234. 1888b. A list of the Bryozoa of the Waverly Group in Ohio; with descriptions of new species. Bulletin Scientific Laboratories Denison University, 4: 63-96. 1890. Paleozoic Bryozoa. Bulletin of the Geological Survey of Illinois, 8: 283 688. Utgaard, J. 1983. Paleobiology and Taxonomy of the Order Cystoporata. Jn, Robison, R.A. (ed.), Treatise on Invertebrate Paleontology. (Part G) Bryozoa — Revised 1; 327-439. 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Sorby and Mr. G.Ri Vine, appointed for the purpose of reporting on fossil Bryozoa. Report of the Britis) | Association for the Advancement of Science (Southport, 1883): 161-209. 1884b. Further notes on new species, and other Yorkshire Carboniferous Polyzo: described by Prof. John Phillips. Proceedings of the Yorkshire Geological ane Polytechnic Society, 8: 377-393. 1884c. Micro-palaeontology of the Northern Carboniferous shales. III, The Ostracoda, Monticulipora, and Miscellaneous Forms: Redesdale Shales, Northum) berland. The Naturalist, (NS) 10 (113): 97-103. | 1885. Notes on the Yoredale Polyzoa of North Lancashire. Proceedings of th Yorkshire Geological and Polytechnic Society, 9: 70-98. } 1887. Notes on the Polyzoa and other organisms from the Gayton Boring Northamptonshire. Journal of the Northamptonshire Natural History Society ani Field Club, 4: 255-266. 1888. A Monograph of Yorkshire Carboniferous and Permian Polyzoa. Part |. Proceedings of the Yorkshire Geological and Polytechnic Society, 11: 68-85. 1889. A Monograph of Yorkshire Carboniferous and Permian Polyzoa. Part 2 Proceedings of the Yorkshire Geological and Polytechnic Society, 11: 184-200. Waagen, W. & Pichl, J. 1885. Salt Range Fossils. Palaeontologica Indica, Series 13 1: 771-834. —— & Wentzel, J. 1886. Salt Range Fossils. Palaeontologica Indica, Series 13, 835-924. Warner, D.J. & Cuffey, R.J. 1973. Fistuliporacean bryozoans of the Wrefor Megacyclothem (Lower Permian) of Kansas. University Kansas Paleontologice Contributions, 65: 1-24. | Waters, J.A. & Sevastopulo, G.D. 1984a. The paleobiogeography of Irish and Britis |) LOWER CARBONIFEROUS BRYOZOA Lower Carboniferous blastoids. /n, Keegan, B.F. & O'Connor, B.D.S. (eds.), Echino- dermata: 141-147. Balkema, Rotterdam. — & 1984b. The stratigraphical distribution and palaeoecology of Irish Lower Carboniferous blastoids. Jrish Journal of Earth Sciences, 6: 137-154. j Whittington, H.B. & Evitt, W.R. 1954. Silicified Middle Ordovician trilobites. | Geological Society of America Memoir, 59: 1-137. Wilson, R.B. 1961. Palaeontology of the Archerbeck borehole, Canonbie, Dumfries- \ shire. Bulletin of the Geological Survey of Great Britain, 18: 90-106. Wright, W.B., Carruthers, R.G., Lee, G.W. & Thomas, I. 1913. On the Lower } Carboniferous succession at Bundoran in South Donegal. Proceedings of the Geolo- gists’ Association, 24: 70-77. Wyse Jackson, P.N. 1988. New fenestrate Bryozoa from the Lower Carboniferous of County Fermanagh. /rish Journal of Earth Sciences, 9: 197-208. '—, Bancroft, A.J. & Somerville, ID. 1991. Bryozoan zonation in a trepostome- ) dominated buildup from the Lower Carboniferous of North Wales. In, Bigey, EP. (ed.), Bryozoaires actuels et fossiles: Bryozoa living and fossil. Bulletin de la Société | des Sciences Naturelles de l'Ouest de la France; Mémoire, (HS) 1: 551-559. i—_ & 1995a. Generic revision of the cryptostome bryozoan Rhabdomeson Young & Young, 1874 with descriptions of two species from the Lower Carbonifer- ous of the British Isles. Journal of Paleontology, 69: 28-45. & 1995b. Rhabdomeson Young & Young, 1874 (Bryozoa): proposed designation of Rhabdomeson progracile Wyse Jackson & Bancroft, 1995 as the type species. Bulletin of Zoological Nomenclature Yang Jingzhi & Lu Linhuang 1962. Palaeozoic Bryozoa of Mt. Qilianshan. Memoirs of the Geology of Mt. Qilianshan, 4: 1-114 [In Chinese]. 171 , Hu Zhaoxuh & Xia Fengsheng 1988. Bryozoans from Late Devonian and Early Carboniferous of Central Hunan. Palaeontologica Sinica, 173 (23): 1-197 [In Chinese and English]. Young, J. 1882. On the Identity of Ceramopora (Berenicea) megastoma, M‘Coy, with Fistulipora minor, M*Coy. Annals and Magazine of Natural History, (5) 10: 427-431. —— 1883a. On Ure’s ‘Millepore, Tabulipora (Cellepora) Urii, Flem. Annals and Magazine of Natural History, (5) 12: 154-158. 1883b. Notes on Ure’s ‘Millepore’, Tabulipora Urii, J.Young (Cellepora Urii, Flem.). Transactions of the Geological Society of Glasgow, 7: 264-272. 1887. Note on a new family of the Polyzoa Cystodictyonidae (E.O. Ulrich) — with notice of three Carboniferous species. Transactions of the Edinburgh Geological Society, 5: 461466. —— & Armstrong, J. 1871. On the Carboniferous fossils of the West of Scotland. Glasgow. 103pp. & Robertson, D. 1877. Note on the Polyzoa of the Hairmyres Limestone Shale, East Kilbride. Transactions of the Geological Society of Glasgow, 5: 173-175. Young, J. & Young, J. 1874. On a new Genus of Carboniferous Polyzoa. Annals and Magazine of Natural History, (4) 13: 335-339. & 1875. New species of Glauconome from Carboniferous Limestone strata of the West of Scotland. Proceedings of the Natural History Society of Glasgow, 2: 325-335. —— & —— 1877. Ona new species of Sulcoretepora. Proceedings of the Natural History Society of Glasgow, 3: 166-168. Zittel, K. 1880. Handbuch der Paleontologie; Band 1, Bryozoa: 573-641. Miinchen und Leipzig. = > ‘Ta \ Vv 14 Ah i vT ia ry a) © q's Lm) é o< : 2 =e ‘(ep @ . ce = + th my as Bay 9 aot eae me) a ) tpl area eT pimp Hemi > Camireyrtea ai orc . - oe 1 ee) @ axvlinetn') ‘Cad sped rw meet § P AA ee A er eee 1 Aelia rk umctondl AS Eb eget dl ere ee re ee ST a eS try fA ill ees = é nyntLe- MM ease ove Sty Mea ea — a parremin . qupligd wihiatd) CHE! eo iin} Mr iy. tly ere ce @ creat, ame eee simens> chica am aA eon] seeeti’ mn oe). Shy = iW 4 a=’ eee ork)", Uy Ase a) Oeegyeiiet & nl oat = — oka Le ieameoalill an is gD tacit wien) iited ire os atl? 4904, i ee ie es Eel ee Lites oe nai tee aeminantins inenel pean ot ee ee er ee ee - i Se il a) ie [1a + « ee | ba weet hq phioah) My omnadenetier cee bourt - 2 tae npg , Pe a a Salil tie) aie ‘en tllome mde are a ‘qi tall ad vig e"T > ed ; — wt ——~ hn Rin tak) ee Ai GS Qetbeni yy cA nt »* Was cme ped, om oy stormed F 4 wes) ep 7 : ay ee - —— miguel aa 5 >, Vt, 34 = —" = he = My : i = 2.6 \¢ 4 Pe oe > Atee See a —-——eeom aa a ie eS y 2 : - — ene ee - a hes * ba Ps) ce j _ ee insitlony ij “ill 7 oe - ; Sat» ot; &. iy A ao mm =a ?<* @.fheres @ co a, tite glo ae T= @ at) ee oS je y*- - . =) Cee @ « ven, A ra ; ve. + 2, (> Pie. aa Ps fico - weXipemt. ppd ~ wai per a apa i J $ — . : ll SAG : aes, » a , [pine = j=, Cy i Y Pepe a Sees, Ti ~ ete Pome pil ae lb-~~s Wom Dish ein hee - i eas i 08 ~ oa Ps —a = Zi ee ae : = "om AS ret pan 6. 173 CORRIGENDA Corrections to: Jones, R.W. & Simmons, M.D. 1996. A review of the stratigraphy of Eastern Paratethys (Oligocene-Holocene). Bull. nat. Hist. Mus. Lond. (Geol.) 52 (1): 25-49. 1. InFig. 11 (p. 39) the density of the shading of the land and sea areas has been reversed, due to an error. The figure should show a large area of land (with dark shading) containing the two smaller areas of sea (with pale shading) that cover most of the present-day Black Sea and the southern part of the Caspian Sea. 2. In Figs 7, 8, 10-12 (pp. 34-41) some of the ticks are wrongly shown on the seaward (pale) side of the lines rather than the landward (dark) side. Bulletin of The Natural History Museum Geology Series _ shown, this may also be obtained from the same address. | - Volume 42 | No. | Cenomanian and Lower Turonian Echinoderms from Wilmington, south-east Devon. A.B. SMith, C.R.C. Paul, A.S. Gale & S.K. Donovan. 1988. 244 pp. 80 figs. 50 pls. 0 565 07018 5. £46.50 Volume 43 No. | A Global Analysis of the Ordovician-Silurian boundary. Edited | by L.R.M. 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Barrett Bryozoa from the Lower Carboniferous (Viséan) of County Fermanagh, Ireland PN.Wyse Jackson Corrigenda Vol. 52, No. 2, November 1996 IS r) | THE NATURAL HISTORY MUSEUM | i 21 AUG 1997 PRESENTED | PALAEONTOLOGY LIBRARY Geology Series NZ a NATU RAL HISTORY MUSEUM VOLUME 53 NUMBER1 26 JUNE 1997 The Bulletin of The Natural History Museum (formerly: Bulletin of the British Museum (Natural History) ), instituted in 1949, is issued in four scientific series, Botany, Entomology, Geology (incorporating Mineralogy) and Zoology. The Geology Series is edited in the Museum’s Department of Palaeontology _Keeper of Palaeontology: Dr L.R.M. Cocks Editor of Bulletin: Dr M.K. Howarth Assistant Editor: Mr C. Jones Papers in the Bulletin are primarily the results of research carried out on the unique and ever- growing collections of the Museum, both by the scientific staff and by specialists from elsewhere who make use of the Museum’s resources. 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(Geol.) © The Natural History Museum, 1997 Geology Series ISSN 0968-0462 Vol. 53, No. 1, pp. 1-78 The Natural History Museum Cromwell Road London SW7 5BD Issued 26 June 1997 Typeset by Ann Buchan (Typesetters), Middlesex Printed in Great Britain by Henry Ling Ltd, at the Dorset Press, Dorchester, Dorset Bull. nat. Hist. Mus. Lond. (Geol.) 53(1): 1-9 Issued 26 June 1997 The status of ‘Plesictis’ croizeti, ‘Plesictis’ gracilis and ‘Lutra’ minor: synonyms of the early Miocene viverrid Herpestides SATS MIECZYSLAW WOLSAN Instytut Paleobiologii PAN, ul. Twarda 51/55, 00-818 Warszawa, Poland MICHAEL MORLO Forschungsinstitut Senckenberg, Senckenberganlage 25, 60325 Frankfurt am Main, Germany \ (Mammalia, Carnivora) AUG IDA PRESENTED PALAEONTOLOGY Lt LIBRARY | SYNOPSIS. The reputed musteloid carnivorans *Plesictis’ croizeti Pomel, 1847, ‘Plesictis’ gracilis Pomel, 1853, and ‘Lutra’ minor Lydekker, 1885 are recognized as junior subjective synonyms of the European early Miocene (Agenian) viverrid carnivoran Herpestides antiquus (de Blainville, 1842). The name ‘Plesictis’ gracilis is a junior objective synonym of ‘Plesictis’ croizeti, whose type locality is identified as Langy, of Agenian age, central France. The type locality of ‘Lutra’ minor is Mainz- Mombach, ofAgenian age, western Germany. The taxonomic histories of ‘Plesictis’croizeti, ‘Plesictis’ gracilis, and ‘*Lutra’minor are reviewed, synonymies are provided, and the holotypes described and figured. INTRODUCTION The name-bearing types of the reputed musteloid carnivorans Plesictis’croizeti Pomel, 1847, ‘Plesictis’ gracilis Pomel, 1853, and ‘Lutra’ minor Lydekker, 1885 constitute a part of the unique collec- tions of The Natural History Museum, London. The specimens have not previously been adequately described, and only the holotype of ‘Plesictis’ croizeti Pomel, 1847 has been figured. The taxonomic histories of the specific names given to them are highly confused. | The present paper provides comprehensive descriptions of the holotypes of ‘Plesictis’ croizeti, ‘Plesictis’ gracilis, and ‘Lutra’ minor, and also reports the complicated taxonomic histories of these names, with evidence that they are all junior synonyms of the Buropean early Miocene viverrid carnivoran Herpestides antiquus (de Blainville, 1842). | The following abbreviations are used in this paper: BMNH and NHM, Department of Palaeontology, The Natural History Museum formerly British Museum (Natural History), London; /CZN, Inter- hational Code of Zoological Nomenclature (The International Commission on Zoological Nomenclature 1985); NMB, Natural distory Museum, Basle; SMF, Department of Palaeozoology, The Senckenberg Research Institute and Museum, Frankfurt am Main. PLESICTIS’ CROIZETI AND ‘PLESICTIS’ GRACILIS | “AXONOMIC HISTORY. Pomel (1847) introduced the specific name -lesictis Croizeti to designate the partial mandible illustrated in fig. | of his pl. 4. Although neither description nor definition accompa- jied that name, it has nevertheless been made available by indication haccordance withArticle 12 (b, 7) of the JCZN. Pomel’s fi g.4of pl. |, as well as its reproductions published in Bronn & Roemer (1856: I 60, fig. 14b), Pictet (1857: pl. 4, fig. 8), and Viret (1929: text-fig. 3), represent the reversed mirror image of the original that is stored ) The Natural History Museum, 1997 in the NHM under register number 26702. As noted in the old vertebrate register at the NHM, this name-bearing type of ‘Plesictis’ croizeti was purchased by the British Museum (Natural History) in June 1851 from ‘M. Pomel’ (M.J. Pomel according to Lydekker 1885: xiii). In 1853 (reprinted in 1854) Pomel proposed the new name Plesictis gracilis for the partial mandible figured by him in 1847 under the name Plesictis Croizeti, and he incorrectly applied the latter name to a skull of a mustelid carnivoran. Because both specific names were based on the same type specimen, they are objective synonyms according to Article 61 (c, iv) of the JCZN. The senior synonym, ‘Plesictis’ croizeti Pomel, 1847, is the valid name of the taxon in accordance with the Principle of Priority (Article 23 of the ICZN). The taxonomic status of ‘Plesictis’ croizeti (=‘Plesictis’ gracilis) was further complicated by Filhol (1879a—b) who considered ‘Plesictis’ croizeti and ‘Plesictis’ gracilis to be distinct varieties of Plesictis robusta Pomel, 1853, which is indeed a synonym of the musteloid carnivoran Amphictis antiqua Pomel, 1853. Although the type specimen of both ‘Plesictis’ croizeti and ‘Plesictis’ gracilis had already been figured or briefly described in Pomel (1847, 1853, 1854), Gervais (1852b, 1859), Bronn (1856), Bronn & Roemer (1856) and Pictet (1857), and the British Museum had been explic- itly indicated by Gervais (1852b, 1859) as the institution where the holotype had been kept, Filhol (1879a—b) was, nevertheless, appar- ently unaware of its existence. At any rate, he made no mention of this specimen. Instead, he assigned a mustelid skull to ‘Plesictis’ croizeti and two musteloid partial mandibles to ‘Plesictis’ gracilis, believing that the characters of one of those mandibles, illustrated in fig. 5 of his pl. 22, corresponded to those of the holotype of ‘Plesictis’ gracilis (his p. 128: ‘J'ai trouvé dans la collection du musée de Lyon un maxillaire inférieur possédant des caractéres correspondants a ceux que M. Pomel avait fait connaitre comme devant servir a faire distinguer spécifiquement le Plesictis gracilis’). In addition, in his quotations of Pomel’s (1853, 1854) descriptions of “Plesictis croizeti’ and ‘Plesictis’ lemanensis Pomel, 1853, Filhol (1879a—b) mistakenly reversed the two descriptions, giving Pomel’s The mandibular foramen lies a little below the level of the alveolar border of the body, about 4.5 mm above the ventral border of the ramus and about 13 mm behind the alveolus for M,. The foramen faces posteriad and somewhat laterodorsad. P, and P, are double-rooted, with the posterior root being larger than the anterior one. The base of the crown bears cingula anterobuccally and posteriorly on both teeth. The cingula are stronger on P,; much of the posterior cingulum of P, has been broken away. The posterior cingulum of P, is much better developed than the anterior one, which was also true for P, judging from the preserved base of its crown. The posterior cingular region of the base of the P, crown is little deflected linguad. The crown base of both P, and P, bears three projections arranged one behind the other anteroposteriorly to form a blade compressed buccolingually. The blade slightly curves linguad on the most ante- rior of the projections, the anterior accessory cusp, in both the premolars and on the most posterior projection, the posterior acces- sory cusp, in P,. The middle cusp, the protoconid, culminates slightly anterior to the half of the tooth length and is distinctly largest, whereas the anterior accessory cusp is smallest. The anterior and posterior accessory cusps are stronger and larger relative to the protoconid on P, than they are on P,. The cusps are divided by prominent v-shaped notches on P.. In both teeth, the tips of the anterior and posterior accessory cusps are noticeably worn away exposing the dentine. Wear has also broken through the enamel at the tip of the protoconid on P,,, but over a very small area only. The crown of M, is supported by two strong roots. There is no cingulum on the talonid, but there are two cingula running along the buccal base of the trigonid from the anterior end of the paraconid to the most anterior portion of the protoconid buccally and to that of the metaconid lingually. The buccal cingulum is very strong, whereas the lingual one is poorly developed. The trigonid is notably arched buccad, making its lingual contour concave when viewed occlusally. The carnassial blade comprises the buccal ridge of the paraconid and the anterior ridge of the protoco- nid, which are divided by a deep, slit-shaped carnassial notch. The carnassial blade is rather deeply worn exposing dentine. The shear- ing surface on the buccal side of the paraconid and protoconid is considerably worn. Viewed from the occlusal surface, the carnassial edge of the paraconid abruptly turns anteriorly into a long, trenchant lingual ridge descending towards the metaconid from which it is set off by a valley. The carnassial edge of the protoconid curves posteriorly at an obtuse angle to continue into a sharp, partly damaged ridge that descends obliquely until it meets the metaconid. The anterior and lingual ridges of the protoconid delimit the lingual wall of this cusp, which flanks posterobuccally a deep, spacious valley that sets the protoconid off from the paraconid and metaconid. In addition to the anterior and lingual ridges, the protoconid exhibits a very short ridge, which is mostly worn away, on the base of its posterior wall. This short ridge ascends occlusolinguad from the anterior end of the anterior edge of the hypoconid. There is an extensive wear facet on the posterior surface of the protoconid. The metaconid is stout, well detached from the protoconid, and proportionally short anteroposteriorly. In lingual view, it resembles an isosceles triangle with its anterior and posterior profiles being slightly convex. In posterior view, the lingual contour of the metaco- nid is also slightly convex. A small part of the metaconid projects posteriorly beyond the protoconid so that its posterior edge is visible in buccal view. The slopes of the metaconid are angulated anteriorly, buccally, and posteriorly into ridges of which the buccal ridge is most trenchant or sharpened and the posterior one is most rounded or blunt. The anterior ridge descends towards the lingual ridge of the paraconid, from which it is separated by a valley. The buccal ridge is M. WOLSAN AND M. MORLO united with the lingual ridge of the protoconid at a prominent, V- | shaped notch. The posterior ridge meets the lingual wall of the talonid. The occlusal part of the posterior surface of the metaconid is worn. Viewed occlusally, the posterior wall of the trigonid is almost ~ straight, while the buccal and lingual contours of the crown are — concave at the area where the trigonid meets the talonid. The buccal | concavity is much better marked than the lingual one. ! The talonid is deeply basined. Its buccal wall consists of an | anteroposteriorly elongate hypoconid, which is the largest cusp on the talonid. The hypoconid is buccolingually wider and has its outer surface more inclined than is the case for the lingual wall of the | talonid, making the talonid basin appear to be shifted linguad in | occlusal view. Although wear has breached the enamel along the © hypoconid, it is evident that the tip of this cusp was originally situated within the posterior half of the cusp length. The hypoconid is detached from the posterior wall of the talonid by a distinct V- shaped notch that is continued into an occlusobasal groove on the | outer surface of the talonid. The posterior wall of the talonid is lower than the buccal and_ lingual walls. It is somewhat worn occlusally and produced into — three low, poorly differentiated elevations. ; The lingual wall of the talonid forms two projections separated from each other by a notch. The anterior of these projections, the © entoconulid, is small, whereas the posterior one, the entoconid, is | much larger, being the second largest cusp on the talonid. The tips of — both the cusps are worn, exposing dentine facets. | TYPELOCALITY. Although Pomel (1847) did not indicate the place \ of collection of the name-bearing type of *Plesictis’ croizeti explic- itly, it is obvious from the contents of his article that the specimen had been excavated from Tertiary deposits in the region of Vaumas and Saint-Gérand-le-Puy (Allier, France). Several years later, Pomel (1853: 97, 1854: 61) expressly attributed that fossil to the Tertiary sediments of Langy (‘Terrain tertiaire 4 Langy’), which is a village | situated about 3 km west of Saint-Gérand-le-Puy and some 25 km) southwest of Vaumas. Most subsequent authors listed the locality as) | ‘Saint-Gérand-le-Puy’. It deserves to be noted, however, that the name Saint-Gérand-le-Puy has generally been applied in the litera-/ ture to encompass various fossil sites discovered in several quarries i in the region of the village Saint-Gérand-le-Puy, including the! locality Langy as well (Cheneval, 1983). The only published statements about the type locality of “Plesictis’, croizeti that were significantly different were those of Gervais)” (1859) and Dawkins (1880a—b). According to the former author, the): ‘ holotype of ‘Plesictis’ croizeti was found in calcareous marls of) ~ Miocene age in the environs of Issoire in the department of Puy-dey 4 Dome (his p. 250: ‘Fossile dans les marnes calcaires de étage) 5 miocéne aux environs d’Issoire (Puy-de-D6me)’). Dawkins in C : a tte statements (1880a: 386, ‘Issoire, Volvic (Puy-de-Déme)’; 18805: 505, ‘Issoire, Volvic, Puy-de-dome [sic]’) simply quoted Gervai (1859). However, neither Gervais nor Dawkins presented any sup ¥ porting evidence for their assertions. Gervais’s (1859) referral of the holotype of ‘Plesictis’ croizeti to) ~ the locality Issoire is rather intriguing since that author was appar ently familiar with Pomel’s (1847, 1854) papers as indicated by theil) citations in his work, and since he studied that fossil during his visi to the British Museum (Natural History) shortly after it had beer acquired by that institution, which is evident from footnote 2 on p| ) 11 in Gervais (18525). In addition, one of the two labels on the typ¢ specimen of ‘Plesictis’ croizeti, which lies in its box and refers ittd ‘Herpestes croizeti’ (the only other label on the fossil is its registe number), identifies the holotype as coming from the ‘Uppel )), ~ iL r -HERPESTIDES ANTIQUUS fig. 2 The holotype of ‘Lutra’ minor Lydekker, 1885 (BMNH 25449), a _ partial left dentary with P, and M,; a, dorsal view; b, lateral view; c, medial view. ligocene’ of “Issoire, Puy-de-Dome [sic], France’. It is thus essen- tally consistent with Gervais’s (1859) statement.There is noevidence, 1owever, to support claim that just this label accompanied the iolotype when it was examined by Gervais, or that the data included nit corresponded to those of any original but now lost label known to hat worker. On the contrary, it seems to be more probable that the yerson who wrote the present label simply followed Gervais (1859), specially as both the old vertebrate register at NHM (which recorded Ilier’ as the locality of the holotype) and Lydekker (1885: 185) (who lescribed it as ‘the Lower Miocene of St. Gérand-le-Puy (Allier)’) upport Pomel’s (1853, 1854) statement that Langy was the place vhere the type specimen of ‘Plesictis’ croizeti was collected. Inconclusion, the correct name of the type locality of ‘Plesictis’ roizeti Pomel, 1847 is Langy in the department of Allier, central france. The accurate placement of this fossil site is vague. Its age _ orresponds to theAgenian, early Early Miocene, as indicated by the xclusively Agenian occurrence of many taxa (e.g. Herpestides ntiquus) attributed by Pomel (1853, 1854) to Langy. UTRA’ MINOR AXONOMIC HISTORY. Jame Lutra minor to a ‘[f]}ragment of the right ramus of the mandi- le, containing the last premolar and the carnassial; from the Lower iocene of Mombach, near Mayence’, purchased in ‘1850’ by the jritish Museum (Natural History). He referred that specimen to Lydekker (1885: 195) applied the specific 5) register number 25440. However, as seen from the vertebrate regis- ters at NHM, the number 25440 has never been allocated. Instead, the old vertebrate register records under number 2544¢g, a ‘[f]ragment of lower jaw of S[tephanodon]. minor’ with ‘2 molars in situ’ from “Mayence’, purchased in ‘Aug[ust]. 1850’ from ‘M. Becker’. The specimen BMNH 25449 is accompanied by two labels. One of them, which is glued to the fossil, displays its register number in which the ‘9’ is of blurred appearance (see Fig. 2a—b), which may have been the reason for Lydekker’s mistake. The other label, lying in the specimen’s box, reads as follows: ‘Fragmentary mandibular ra- mus[;] Potamotherium minor, Meyer sp.[;] Form" Lower Miocene[;] Loc’ Mombach, near Mayence[;] Purch* 1850[;] Cat. 1, p. 149[;] Brit. Mus. Geol. Dept. 25449. The reference to p. 149 in the first part of Lydekker’s catalogue (1885) is an error, of course, because this page is actually devoid of any mention of this fossil; instead, the name of the ursoid carnivoran Cephalogale minor Filhol, 1879a is quoted there. Another inconsistency between Lydekker’s (1885) account of “Lutra’ minor and the data available on BMNH 25449 is that the latter represents the left branch of the mandible, and not the right one as indicated by that author. Otherwise, BMNH 25449 fits Lydekker’s description exactly. Moreover, there is no other fossil in the collections of The Natural History Museum in London, which could represent Lydekker’s specimen. Accordingly, we conclude that BMNH 25449 must be the specimen referred by Lydekker (1885) to Lutra minor. Lydekker (1885) treated the name Lutra minor as a new combina- tion for Stephanodon minor, deemed by him to be erected by Hermann von Meyer. However, Lydekker (1885: 195, footnote 1) ‘ha[d] been unable to find a reference to this species’. The old vertebrate register at NHM refers the specific name Stephanodon minor to specimen 25449, but without any relation to von Meyer’s name. Instead, this German palaeontologist is cited in the register in connection with number 25448 (“Stephanodon monbachensis [sic] V. Meyer’ ) attributed to the holotype of Stephanodon mombachensis von Meyer, 1847, which is indeed a junior synonym of the arctoid carnivoran Potamotherium valletoni (Geoffroy Saint-Hilaire, 1833). According to the register, that fossil and three others catalogued under numbers 25450-25452 were purchased in August 1850 from M. Becker as coming from ‘Mayence’, exactly as specimen 25449. In all likelihood, all these fossils had been studied by von Meyer before they were conveyed to the British Museum. It is thus just possible that H. von Meyer gave the name Stephanodon minor to specimen 25449. At any rate, it is very probable that a label stating this name accompanied the specimen originally and was known to Lydekker. Its existence was explicitly stated by Pohle (1920: 17; ‘Das Stiick war von v. Meyer mit dem Namen etikettiert worden’), who, however, provided no evidence to support his statement. Regardless of this, even if von Meyer was really responsible for the name Stephanodon minor, as declared by Lydekker (1885) and followed by Trouessart (1885, 1897, 1904), Schlosser (1888), Pohle (1920), and Haupt (1935), he has not satisfied the criteria of avail- ability of that name and therefore cannot be considered its author according to Article 50 (a) of the JCZN. Instead, Lydekker (1885), who satisfied these criteria through both publishing the name of this taxon and stating in footnote | on his p. 195 that ‘this species [. . .] may be only a smaller form of [Lutra valetoni]’, is the author of the name whose correct original spelling is Lutra minor. The name-bearing type of “Lutra’ minor has never been figured or adequately described in the literature. The only published informa- tion relating to its size and morphological characteristics is that of Lydekker (1885: 195, footnote 1) that ‘Lutra’ minor ‘may be only a smaller form of’ Potamotherium valletoni. The subsequent authors confined themselves to following this assumption. Trouessart (1885, 6 1897, 1904) held ‘Lutra’ minor to be a subspecies of Potamotherium valletoni while Pohle (1920), Haupt (1935), and Savage (1957) simply placed it in the synonymy of that species. Schlosser (1888: 123) expressly denied the specific status of “Lutra’ minor (“Ebenso ist auch Stephanodon minor H. v. Meyer auf keinen Fall als besondere Art zu betrachten’), including it in Potamotherium valletoni, but later (1890) he quoted it as a separate species of Potamotherium. The synonymy list of ‘Lutra’ minor Lydekker, 1885 includes the following names: 1885 Lutra minor [or] [Lutra] minor Lydekker: xxi, 195, 266. 1885 Stephanodon minor, Lydekker: 195, 267. 1885. [Lutra valetoni| minor; Trouessart: 47. 1888 Stephanodon minor; Schlosser: 123. 1890 [Potamotherium] minor; Schlosser: 82. 1897 [Potamotherium Valetoni] minor, Trouessatt: 281. 1904 [Potamotherium valetoni] minor, Trouessart: 212. 1920 Stephanodon minor [or] [Stephanodon] minor; Pohle: 16— Wy 223s 1935 Stephanodon minor; Haupt: 38. 1957 Potamotherium minor; Savage: 155. DESCRIPTION OF THE HOLOTYPE. The holotype, by monotypy, of ‘Lutra’ minor Lydekker, 1885 is BMNH 25449, a fragment of a left dentary with partially eroded P, and M, (Fig. 2, Tables 1-3). The side-walls of the preserved fragment of body in the holotype dentary are convex in cross-section, excepting the ventral part of the medial wall where the surface of the dentary bone is somewhat depressed along the ventral border. The alveoli for P,-M, are arranged one behind the other and closely spaced. P, and M, slightly overlap each other and have pairs of alveoli. Only the anterior part of the M, alveolus is preserved; judging from this preservation, the alveolus was single and anteroposteriorly elongated. The masseteric fossa extends anteriorly to the level of the alveolus for M.,,. The morphological patterns of P, and M, are congruent with those of the corresponding teeth in the type specimen of *Plesictis'croizett, with the exception of the following differences concerning M.;: in the holotype of ‘Lutra’ minor the lingual ridge of the paraconid is shorter; the lingual contour of the metaconid is slightly concave; the metaconid is somewhat deflected posteriad, making its posterior contour slightly concave when viewed from the lingual side; there is no crest on the anterior face of the metaconid, so that the anterior slope of this cusp is widely rounded and blunt; and, finally, no elevation could be detected on the posterior wall of the talonid. The crowns of P, and M, are generally less worn in the holotype of ‘Lutra’ minor than those of the type specimen of *Plesictis’ croizeti. TYPE LOCALITY. The old vertebrate register at NHM reports the holotype of ‘Lutra’ minor as having been collected in ‘Mayence’ (=Mainz). Lydekker (1885: 195) described it as coming ‘from the Lower Miocene of Mombach, near Mayence’ (now Mainz- Mombach), perhaps on the basis of an original, but now missing, specimen label. Lydekker’s attribution is consistent with that on the label accompanying the holotype at present. Schlosser (1890), who knew both Lydekker’s (1885) catalogue and H. von Meyer’s unpub- lished drawings of carnivoran remains from Mainz-Weisenau (as seen from Schlosser 1887: 4, 6), referred ‘Lutra’ minor (his Potamo- therium minor) to ‘Mainz (Weissenau)’ (=Mainz-Weisenau). No evidence exists, however, to suggest that Schlosser’s assignment concerned specimen(s) other than the holotype and, moreover, none of the copies of von Meyer’s drawings preserved in NMB represents the type specimen of ‘Lutra’ minor. Consequently, we conclude that Schlosser’s Potamotherium minor pertained to the holotype of “Lutra’ minor, and hence its referral to the locality Mainz-Weisenau resulted from confusion. M. WOLSAN AND M. MORLO | Table 1 Mandible measurements (in mm) of the holotype of “Plesictis’ croizeti Pomel, 1847 and ‘Plesictis’ gracilis Pomel, 1853 (BMNH 26702), and the holotype of ‘Lutra’ minor Lydekker, 1885 (BMNH 25449). BMNH BMNH 26702 25449 Distance between posterior-most points of C, and M., alveolar rims B19 - Greatest distance between alveolar rims for M, and M, 11.6 - Length of P, alveolus (greatest diameter of P, alveolar rim) 2.0e —- Width of P, alveolus (least diameter of P, alveolar rim) 1.0 - ; | | | | | Length of P, alveoli (greatest distance between rims of anterior and posterior alveoli for P,) 4.6 - ; Width of P, alveoli (least distance from line connecting : lingual-most points of P, alveolar rims to buccal- most point of these rims) ep) = Length of P, alveoli (greatest distance between rims of anterior and posterior alveoli for P,) 60) = Length of P, alveoli (greatest distance between rims of anterior and posterior alveoli for P,) q-3} 6.62 Length of M, alveoli (greatest distance between rims of anterior and posterior alveoli for M,) 8.6 8.5 Length of M, alveolus (greatest diameter of M, alveolar rim). ‘ 2.6e - Width of M, alveolus (least diameter of M, alveolar rim) PIgp - Greatest horizontal distance between lateral and medial walls of dentary below M, perpendicular to long axis of dentary 5.6 5.8 Least distance from alveolar border of dentary between P, and P, to its ventral border, measured on medial side 93 = Least distance from alveolar border of dentary between M_ and M, to its ventral border, measured on medial side 10.5 WE7/ a ‘e’ indicates an estimated value. Table 2 Measurements (in mm) of premolar teeth in the holotype of ‘Plesictis’ croizeti Pomel, 1847 and ‘Plesictis’ gracilis Pomel, 1853 (BMNH 26702), and the holotype of ‘Lutra’ minor Lydekker, 1885 (BMNH 25449). BMNH BMNH 26702 25449 Length of P, (from anterior-most to posterior-most points of crown) 6.2 = Width of P, (greatest distance between buccal and lingual borders of crown perpendicular to antero- posterior length of tooth) 2.8+ = Height of P, (least distance from occlusal-most point of tooth to basal margin of crown, measured on buccal side) 3.8 - Length of P, (from anterior-most to posterior-most points of crown) UA gal Width of P, (greatest distance between buccal and lingual borders of crown perpendicular to antero- posterior length of tooth) 333) 3.4 Height of P, (least distance from occlusal-most point of tooth to basal margin of crown, measured on buccal side) ‘4? indicates a minimum measurement on an incomplete structure, “e’ indicates an estimated value. HERPESTIDES ANTIQUUS Table 3 Measurements (in mm) of M, in the holotype of ‘Plesictis’ croizeti Pomel, 1847 and ‘Plesictis’ gracilis Pomel, 1853 (BMNH 26702), and the holotype of ‘Lutra’ minor Lydekker, 1885 (BMNH 25449). BMNH BMNH 26702 25449 Length (from anterior-most to posterior-most points of crown) 8.7 8.8 Width (least distance from buccal-most point of crown _ to line joining lingual-most points of paraconid wing and talonid) 4.6 4.7 Trigonid length (least distance from anterior-most point of crown to line connecting notch between protoconid and hypoconid with notch posterior to metaconid) 5.9 6.2 Least distance between buccal and lingual borders of crown across carnassial notch 3.8 3.7 Talonid length (least distance from posterior-most point of crown to line connecting notch between protoconid and hypoconid with notch posterior to metaconid) 2.8 2.6 | Talonid width (greatest distance between buccal and | lingual borders of talonid perpendicular to antero- posterior length of tooth) 3a) 3).7/ /Paraconid height (least distance from occlusal-most point of paraconid to basal margin of crown, measured on lingual side) 3.7+ 3.8+ Protoconid height (least distance from occlusal-most point of protoconid to basal margin of crown, measured on buccal side) Salle BO) ‘Metaconid height (least distance from occlusal-most point of metaconid to basal margin of crown, | measured on lingual side) 3h57/ 3.6+ \Hypoconid height (least distance from occlusal-most point of hypoconid to basal margin of crown, measured on buccal side) 234 0 DS Entoconid height (least distance from occlusal-most point of entoconid to basal margin of crown, measured on lingual side) 1.9+ 2.0+ t . . cae . ‘+ indicates a minimum measurement on an incomplete structure. _ To summarize, the correct name of the type locality of ‘Lutra’ Ininor Lydekker, 1885 is Mainz-Mombach in Rhineland-Palatinate, western Germany. The exact location of this fossil site is uncertain at oresent. Tobien (1980) assigned the fossil fauna from Mainz- ombach (his Mombach) to the Hydrobia beds of late Agenian age, arly Early Miocene. SONSPECIFICITY WITH HERPESTIDES \NTIQUUS during their taxonomic history, ‘Plesictis’ croizeti, ‘Plesictis’ graci- _(s, and “Lutra’ minor have been referred to various arctoid caniform enera, including the mustelids Plesictis Pomel, 1846, Lutra srunnich, 1771, and Mustela Linnaeus, 1758 (for ‘Plesictis’croizeti: -g. Gervais 1852b: 12), the amphictid musteloid Amphictis Pomel, 853 (for ‘Plesictis’ gracilis; Viret 1929), and the arctoid otamotherium Geoffroy Saint-Hilaire, 1833 (= Stephanodon von ae 1847; for “Lutra’ minor). However, the fact that they really flows to none of those genera needs no elaboration. Their name- aring types differ from the characteristics of these genera in most pects of their morphology, and more importantly, ‘Plesictis’croizeti ‘Plesictis’ gracilis) and ‘Lutra’ minor do not belong even to the border Caniformia Kretzoi, 1943, which is evidenced below. A ‘Plesictis’ croizeti and ‘Plesictis’ gracilis BS ‘Lutra’ minor 6 O Herpestides antiquus 7 8 9 10 11 Fig. 3 A diagram of length versus width of M,, showing the distribution of individuals in a sample of Herpestides antiquus from the Agenian locality Montaigu-le-Blin in France (collection of NMB), compared to that of the holotypes of ‘Plesictis’ croizeti, ‘Plesictis’ gracilis, and ‘Lutra’ minor. As seen from the descriptions, illustrations (Figs 1-2), and meas- urements (Tables 1—3) presented in this paper, the holotype of ‘Plesictis’ croizeti and ‘Plesictis’ gracilis and that of ‘Lutra’ minor differ only insignificantly from each other, plainly justifying the conclusion that they represent the same species. The available morphological features of that species unanimously point to its affiliation with the viverrid feliform Herpestides de Beaumont, 1967, known from the lower part (Agenian) of the European Lower Miocene. According to de Beaumont (1967), that genus included the single, extensively variable species Herpestides antiquus (de Blainville, 1842). A comparison of the holotypes of ‘Plesictis’ croizeti (and *Plesictis’ gracilis) and ‘Lutra’ minor with the corre- sponding portions of dentary and lower dentition of Herpestides antiquus from the French locality Montaigu-le-Blin, stored in NMB, revealed that both the morphological traits and size of the holotypes are well within the variability range observed in Herpestides antiquus (Fig. 3). To conclude, on the evidence presented above we consider the name-bearing types of ‘Plesictis’ croizeti Pomel, 1847, ‘Plesictis’ gracilis Pomel, 1853, and ‘Lutra’ minor Lydekker, 1885 to represent the species Herpestides antiquus (de Blainville, 1842). Accordingly, we synonymize the first three names with the last one that is the valid name of the species in conformity with the Principle of Priority. ACKNOWLEDGEMENTS. Our sincere thanks go to J.J. Hooker (NHM) and B. Engesser, P. Jung, C. Médden, and F. Wiedenmayer (NMB) for their aid and generous hospitality during our research at those institutions. This study was supported by the State Committee for Scientific Research (KBN) grant 6 P204 051 05, the Swiss National Science Foundation, the Alexander von Humboldt Foundation, and the German Research Association (DFG) grant Ro 143/12-1. REFERENCES Anonymous 1839. Marte fossile. L’Echo du Monde Savant, 5: 42-43. 1849. A. Pomel: Note iiber die im Allier-Dept. entdeckten fossilen Thiere. 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Die Affen, Lemuren, Chiropteren, Insectivoren, Marsupialier Creodonten und Carnivoren des europaischen Tertiars und deren Beziehungen ZU ihren lebenden und fossilen aussereuropdischen Verwandten. Beitrage — - nt) ati @@ ey ety reir na | . aa i ~~! et : > oes a Oe Oe —estett mlipeelinaee tk = | « Pee i a real ivmal 4 —— hee a ee gon toned A eit aoa adam pee! Ot? aa oamiee —- ae J , A ay - ed et — ‘ . onion, Ray ae ell aad me — ala — ee ' a - —_——a isl 7s in " a iis J = e * = at, ore - a - —— - *¢ a } cy ss = i ‘ on : _ - S San = ms ? e- = > ee - ola - - = = “* a CO-.@@e i a ae ia 4 mm ¢ = = a6 _ = oo) ~ a, = — =~ * A Tee -> ee* L - <> a A ; £ = . 7 — a > Pad a, » » z © ce = 7 @ A = —— j Se a 4 - x - e sheay mie. = ‘6 ‘ y he a 6 io : | Bull. nat. Hist. Mus. Lond. (Geol.) 53(1): 11-70 Issued June 1997 Baryonyx walkeri, a fish-eating dinosaur from | the Wealden of Surrey | ALAN J. CHARIG AND ANGELA C. MILNER : Department of Palaeontology, Natural History Museum, Cromwell Road, London SW7 5BD CONTENTS | SS VLG DISS es Sabon Ceca eo NR sh rt cca ce Medan pce eee iar 1] Introduction ..... sob oocentbonerroa06ce-0oeG0deHagH_NCAAoCINS CGD SEIS TO OCOD SEO EERE caSHa-acouodeod aK70"200300qs0OCuO HDA JUAdooH AG BOnEEegoRGcHeR ace Daemactcnotnatensron 13 Sy SLE MALES CLAD CL ONI ra..cosccure se srvass teat sae tastiest fo0 08st ate eee dO ES nh cone OTE 13 OUTTA SECIAUICS) cvccecc. sacs .scess nessa scpateetee ere t= 5- MERE ORs a PETE ALLO ST 13 SI ccaecaac Soe SUSS eS RE aE rea eRe Arcee Se is uae eee Bt tae semis ae ee PN ee ok yee 14 TBO CIA AW bee, «ct aneeneness Sete Neen tare STL coe vs AEA. cee Ree ced REE EL SGN 0) Owe See cede ne ne 23 IISA STOR saceg etc stc08c Sconce eee CEP ee ceca anrnieecchooke enn Mian Giorteetctussaiee ere ai ata eat is etna ea ore 28 Werte bralicolumnmernurnerter sects sss. octek irs eee cassis te eee tr ee OTs toe, A ee ee OL GC ON 30 ISS ssocoocdaconaacttocd codon Sone eer ea erent ee oe OO ec RC 41 Chev ronsh(ilaemapophy Ses) ems.ce cette, oetcastee: sort hes ae Binnie eianecd ee. ae, ane ces We ad Se 0 Denes 41 SS LILLLID] eemeenees eee Meee cere it, OS AL A Ae er em AO erty en on ON ei 42 | Hee Gtotal spit dl ceeceenanres serra cesses fa, i Wet eter Oe UR ng esd eee one as Ge Rie SEWER, tess eed ae 42 BOLUM re arate eee oer cea eve aE Race, ne eee. Cheon Cath ats Wee ln 43 IPGIMS (SARS cs aceccecceayg eA aac teAN oo SeBee bs 8 io rac ne eee REN PRP cy oe et NO 47 IF BoC HINA) cess ees ane Sor bene re COREE pepe er EEE See CEE eee Eee RR Pr kOe 51 SYNOPSIS. The well-preserved skeleton of a large theropod dinosaur, Baryonyx walkeri Charig & Milner, 1986, from the Wealden (Barremian, Lower Cretaceous) of Surrey, is described in detail. It is a large theropod with some resemblance to Megalosaurus or Allosaurus, but is sufficiently different to merit its earlier designation as the type of the new family Baryonychidae. Its distinguishing characters include: the prenarial extension of its snout into a spatulate rostrum, a unique increase in the number of the dentary teeth (which are more than twice as numerous per unit length of jaw as the opposing | maxillary teeth), an unusually robust fore-limb, and at least one pair of huge manual talons. Lack of fusion between the components of both skull and vertebrae suggests that this 10 metre long animal was immature. Few other specimens might be referred to Baryonyx: a maxilla fragment of B. walkeri is recorded from the Barremian of Spain; two snout fragments from the Aptian of Niger, which are virtually identical to the conjoined premaxillae of Baryonyx; two isolated tooth crowns, one from the Hauterivian of East Sussex, and one from Surrey; and seven from the Barremian of the Isle of Wight are compared to the genus. On the evidence of jaws and teeth only, the families Baryonychidae and Spinosauridae (Spinosaurus, Angaturama, and perhaps Irritator) are placed in the superfamily Spinosauroidea. An investigation of the wider affinities of Baryonyx, based on a modified version of Holtz’s 1994a data-matrix on the Theropoda, suggests that the Spinosauroidea is a basal member of the Tetanurae and sister-group to the whole of the Neotetanurae (i.e. the Coelurosauria sensu Gauthier plus a variable collection of ‘allosauroids’), with Megalosaurus and Torvosaurus as progressively more distant outgroups. The associated fossil fauna is dominated by Jguanodon and many insects, which lived in the vicinity of a sub-tropical delta. Baryonyx was terrestrial, a fish-eater, probably a scavenger, and possibly an active predator of small to medium-sized land animals. It made greater use of its fore-limbs and talons in attack and defence than its jaws and teeth, which were used mainly for seizing fish and entrails. The taphonomy of the holotype suggests that the animal is unlikely to have been transported from elsewhere and probably died where found. Its skeletal remains lay in sediments that were mostly submerged in shallow water but exposed to the air for brief periods; the bones were trampled and broken before fossilization. | | | | | | | i Natural History Museum, 1997 A.J. CHARIG AND A.C. MILNER | Frontispiece Baryonyx walkeri, holotype, BMNH R9951; restoration of a living animal in the Wealden landscape. Painted by John Sibbick. De _BARYONYX WALKERI INTRODUCTION Discovery The discovery of Baryonyx was a major event in the history of British dinosaur palaeontology (Milner & Croucher 1987). Extant faunas of large terrestrial vertebrates invariably include only a small proportion of carnivores, these being at the apex of the food pyra- mid, and fossil faunas were probably no different. Dinosaur discovery in England goes back more than 300 years (Plot 1677) and includes several finds of a few bones or teeth of carnivores. Until 1983, 10wever, the only significant find of a carnivorous dinosaur had been he partial skeleton of Eustreptospondylus oxoniensis (originally dlaced in the genus Megalosaurus), found in Oxfordshire in 1871 ind now in the Oxford University Museum. Since the discovery of 3aryonyx, the partial skeleton of an allosauroid has been described rom the Wessex Formation (Wealden) of the Isle of Wight (Hutt, Martill & Barker 1996). | Details of the discovery of Baryonyx were published by Milner & -roucher (1987). The enlarged ungual phalanx, the famous ‘Claw’ originally missing the tip), a second smaller, less complete ungual ihalanx and an incomplete haemapophysis, were found on 7 Janu- ry 1983 by Mr William J. Walker at the Ockley Brick Company’s jlaypit near Ockley, Surrey, and a week later he found the missing tip f the claw. After examining this material at the British Museum Natural History), we visited the site on 7 February and found large ark-brown bones from the pelvis and hind-limb, lying just beneath 1e surface of the clay. The physical conditions in the claypit revented us from collecting the skeleton until the period 25 May to 0 June 1983, when a team of eight from the Department of eo ey and a number of volunteers excavated two tonnes of vatrix containing bone. | All the remains were found within an oval excavation measuring oproximately 5 metres by 2 metres (Fig. 49). Most were encased in ia of sideritic siltstone, which had been deposited around the i but some lay unprotected in soft clay. The skeleton was rgely disarticulated and the elements gently scattered, but most of em were still lying approximately in their skeletal position, with € animal’s skull at one end and its tail at the other. Some nodules ere lifted untreated, some were encased in plaster of Paris and hers in expanded polyurethane foam. reparation ‘eparation of the material was especially difficult because of the wdness of the siltstone matrix, made more intractable by the jesence of siderite. A few pieces were subjected experimentally to (emical treatment with thioglycollic acid (which dissolves iron Its but has little or no action on siltstone and other non-ferrous icks), but it had practically no effect, so most of the matrix had to removed by mechanical methods. Some field cocoons, the bulk of : tock and the underlying clay were removed rapidly with an iustrial shot blaster, a Vacu-Blast Nova 150PB, using plastic shot ‘the abrasive; as far as is known (R. Croucher and W. Lindsay, pers. (mms) this is the first recorded instance of the use of such equip- Int for preparing fossils. Rock was also removed with hand tools éd with tools powered by compressed air, including diamond- Gted circular saws and chisels. For more delicate work close to the Ls surface, finely pointed engravers were used under binocular toscopes. Other details of the laboratory work on Baryonyx were i by Milner & Croucher (1987). SYSTEMATIC DESCRIPTION Suborder THEROPODA Marsh, 1881 Superfamily SPINOSAUROIDEA Stromer, 1915 Family BARYONYCHIDAE Charig & Milner, 1986 TYPE-GENUS. Baryonyx Charig & Milner, 1986. DERIVATION OF NAME. From Greek Bapuc, heavy; ovv6, claw. DIAGNOsIs. As for genus Baryonyx. Genus BARYONYX Charig & Milner, 1986 TYPE-GENUS. B. walkeri Charig & Milner, 1986; Early Creta- ceous, Surrey, England. DIAGNOSIS. Monotypic genus, as for species B. walkeri. Baryonyx walkeri Charig & Milner, 1986 DERIVATION OF NAME. the original claw-bone. For Mr William J. Walker, who discovered HOLOTYPE. Natural History Museum, London: Department of Palaeontology, BMNH R9951. MATERIAL. The’holotype alone; consisting of conjoined premax- illae, conjoined vomers, anterior part of left maxilla, conjoined nasals, left lacrimal, left prefrontal, left postorbital, anterior end of braincase (right frontal, right parietal, right orbitosphenoid, right laterosphenoid), posterior end of braincase together with occiput (both prootics, both opisthotics, basisphenoid, supraoccipital, both exoccipitals, basioccipital), left jugal, both quadrates; both dentaries, both splenials, right surangular, both angulars, right coronoid: some upper teeth in situ and many isolated teeth of unknown position in the jaws; axis and 4 further cervical vertebrae, 12 dorsals, 3 or 4 basal caudals, 3 more distal caudals; one axial rib, three other cervical ribs, many dorsal ribs, abdominal ribs, 5 haemapophyses, sternum; both scapulae, both coracoids; both humeri, left radius, left ulna, left pollex with huge ungual, left digit II (complete) or digit Ill, ,, other isolated phalanges of both sides; right ilium, both pubes, left ischium; proximal end of left femur and distal end of right, right fibula, right calcaneum, metatarsal fragments, | pedal ungual. LOCALITY. Smokejack’s Brickworks (Ockley Building Products Limited, formerly Ockley Brick Company Limited), Wallis Wood, Ockley, near Dorking, Surrey, England. A little below the top of the south-east face, nearest to the works buildings. National Grid refer- ence TQ 113373. HORIZON. Lower Cretaceous, Wealden Series, Upper Weald Clay; zone of Cypridea clavata, near the base of the Barremian; about 7 m below the base of BGS Bed Sc (See Ross & Cook 1995). DIAGNOSIS. Prenarial region of snout extended into extremely narrow rostrum with spatulate, horizontal expansion at end (‘term- inal rosette’); snout slightly downturned in lateral view, jaws with sigmoidal margins. Long, low external naris far back on side of Fig. 1 snout. Complex articulation of premaxilla and maxilla unfused above subrostral notch. Small median knob at posterior end of conjoined nasals, cruciform in dorsal view, with anterior limb of cross drawn forwards into low thin median crest. Occiput deep, with paroccipital processes directed horizontally outwards. Basipterygoid processes descend far below basioccipital, diverging laterally only slightly. Anterior end of dentary upturned in lateral view. 6/7 pre- maxillary teeth; only 8 maxillary teeth preserved, but probably about 15; 32 dentary teeth, generally smaller than those in the upper jaw, more than twice as numerous per unit length of jaw. Prominent bony wall on lingual side of all teeth. Tooth crowns flattened only slightly labio-lingually, lightly fluted on lingual side; anterior and posterior carinae finely serrated (about 7 denticles per millimetre). Tooth roots exceptionally long and slender. Axis small, with well devel- oped hyposphene. Cervical vertebrae with flat zygapophyses and well developed epipophyses; ends of centra not offset, no upward curve to neck; neurocentral sutures unfused; neural spines generally short, but those of basal caudals expanded into large flattened plates. Cervical ribs short, crocodiloid, slight overlap. Humerus relatively well developed; both ends broadly expanded but flattened, offset 35° against each other; shaft massive, almost straight. Radius stout, a little less than half as long as humerus. Ulna also stout, somewhat longer than radius, with powerful olecranon. Exceptionally large manual ungual phalanx, not laterally compressed, probably from digit I. Pubic foot not expanded. Ischium with obturator flange proximally continuous with anterior margin. Skull GENERAL DESCRIPTION (Fig. 1). Despite the limited amount of material available, several interesting observations may be made on the skull. It appears to have been long, at least in its anterior portion. Whereas in other theropods the extreme anterior end of the premax- illa gives rise to the steeply ascending nasal process, forming the anterior margin of the external naris, in Baryonyx there is no trace of such a process, and the anterior 170 mm of the conjoined premax- illae form a long low rostrum with a smoothly rounded dorsal surface. Even behind this rostrum there is no ascending process, the Baryonyx walkeri, holotype, BMNH R9951; reconstruction of skull, from left side. x 0.135. A.J. CHARIG AND A.C. MILNER |} confluent external nares appearing merely as an extensive opening lying well back from the tip of the rostrum and passing horizontally from one side of the snout to the other. The first 130 mm of the rostrum are expanded laterally into a spatulate ‘rosette’, not unlike that found in modern gavials; the first 70 mm of its lower margin are turned down in lateral view. Behind the ‘rosette’ the rostrum is remarkably narrow from side to side as seen in dorsal view. The maxilla fits on to the hind part of the ventral margin of th premaxilla by a complex articulation. The resulting effect is that th line of the upper tooth row, passing backwards from the front of the snout, first rises a little and then, at the junction of the two elements, curves strongly downwards and finally flattens out to run horizon- tally along the ventral margin of the maxilla. The net effect of this sigmoid curvature is that the front of the rostrum is elevated aboy the level of the maxillary tooth row, not depressed beneath it. This elevation is reflected in the shape of the upper margin of the dentary, the front 140 mm of which slope strongly upwards towards its anterior end. Nevertheless, in the region of the 5" to 6" premaxillary teeth, the gap between the upper and lower jaws seems to have been much wider than elsewhere and may be termed the subrostral notch. Other noteworthy general features of the skull are: 1. The deep, narrow shape of the occiput, especially deep below the condyle, the basipterygoid processes greatly lengthened. 2. The great length of the dentary, its dentigerous part very shallow. with well-marked Meckelian groove on the medial surface. 3. The presence of a prominent bony wall on the lingual side of eac tooth row. 4. The exceptionally high tooth count (6 premaxillary teeth on th left side and 7 on the right, as contrasted with a more usual count of 5, and 32 dentary, as contrasted with the usual 16 or so). Thi only other theropods known with more teeth per unit length 0 lower jaw than in the corresponding length of upper jaw are Troodon(Osm6lska & Barsbold 1990) and Pelecanimimus (Pérez Moreno eft al. 1994), and even there the disparity is far less|~ extreme. . The much closer packing of the teeth in the dentary than in the) opposing part of the maxilla; as a corollary of this the dentary © teeth must have been generally smaller (it should be noted, Nn BARYONYX WALKERI showing also vomers. x 0.5. however, that our knowledge of the maxillary dentition is restricted to the first 7 teeth). The extremely fragile, laminar nature of the post-dentary bones in the lower jaw. The lack of co-ossification between the elements, except in the midline and in the braincase. The last character suggests that R9951, despite its great size, was juvenile, from which we might deduce that the fully mature animal as probably much larger still. The possibility that a member of the dinosauridae (see p. 55) might attain a truly vast size is confirmed Stromer’s illustrations (1915) of the dentary of Spinosaurus gyptiacus, an element twice the linear size of that of Baryonyx. REMAXILLA (Fig. 2). Both premaxillae are preserved, firmly med.r id.p i g-2 Baryonyx walkeri, holotype, BMNH R9951; snout. A, conjoined premaxillae in dorsal view; B, same, right lateral; C, premaxillae in palatal view, sutured together; the median suture between them is clearly visible on the dorsal surface, though it fades out towards the rear. Both elements are complete anteriorly but broken off behind. The pre- maxillae (as preserved) are not fused to any other elements; each, however, has (or had) a complex articulation with the maxilla. The snout is unlike that of most other known dinosaurs, in which an ascending process rises from the very front of the conjoined premaxillae and forms the rounded anterior border of the external nares. Here, by contrast, the snout is long and low; the equivalent process originates 172 mm behind the tip of the snout and is directed almost horizontally backwards. Thus the heavy, robust snout formed by the conjoined premaxillae is seen in lateral view to bifurcate posteriorly into two rami on each side, a short one below (the main body of the premaxilla) and a longer one above (the 16 ‘ascending process’); the external naris between them terminates anteriorly in an acute angle. In palatal or dorsal view the snout appears extremely narrow from side to side, and there is a ‘waist’ between the anterior, dentigerous portion expanded horizontally into a ‘terminal rosette’ (Charig & Milner 1990) and the two rami, lower and upper, behind. The lower ramus is broken off transversely not far behind the anterior corner of each external naris, but its full extent on each side of the snout is demonstrated by clear indications of its overlap onto the laterodorsal surface of the left maxilla, indications which sug- gest that that too was tapered to a point. The dorsal surface of this ramus, forming the floor of the nasal cavity, slopes steeply down- ward on either side (i.e. it is merely a posterior extension of the side of the snout), but between these slopes is a deep sagittal V-shaped groove. The upper ramus tapers to a thin spike which presumably met the nasals; it has a smooth outer (i.e. dorsal) surface, but internally (i.e. ventrally), where it borders the nasal cavity, it bears on each side two longitudinal ridges, a low lateral ridge and a stronger medial ridge. Together these produce a shallow lateral groove on each side and a deeper median groove, in the bottom of which the latter suture between the paired premaxillae may still be traced intermittently. These ridges would somehow have supported the internasal septum and associated soft tissues, and the lateral grooves doubtless served for articulation with a forwardly projecting prong from the nasal on each side. In lateral view the ventral, dentigerous margin of the premaxilla is distinctly downturned, both anteriorly towards the tip and posteriorly towards the articulation for the maxilla; it is smoothly concave between. There is extensive pitting on the outside of the snout, especially towards the tip, which doubtless served for the exit of blood vessels and nerves; a row of nutritive foramina lies just above the bases of the teeth. In ventral view the terminal rosette has an elongated spatulate shape, around the edge of which lie six dental alveoli on the left side and seven on the right. In these thirteen alveoli only six teeth remain in place, four of them complete to the tip. The first four alveoli on each side are large; nos. 2 and 3 are the largest; nos. 5 to 7 decrease in size progressively. Interdental plates are present, each interrupted by a narrow gap which is rather more posterior than anterior in position. When the premaxilla is seen in palatal view, the area between the tooth rows on either side appears to be fully occupied by a pair of narrow, transversely rounded ‘elements’, sutured together in the midline and broken off short just behind the level of the last premaxillary tooth. Each tapers anteriorly to a combined point which abuts against the rear end of a median groove between the medial walls of the first alveolus on each side. Towards the rear each ‘element’ expands ventrally. From the anterior apex a strongly interdigitating median suture between the two sides pursues a sinuous course backwards for some 30 mm; farther back the interdigitations cease and the two sides are separated by a narrow, elongated, median gap, on either side of which the ‘elements’ curve out laterally away from each other and then, more posteriorly, almost rejoin. In the region of the anterior half of the median gap their ventral surface is somewhat rugose, suggesting the possible presence in life of a horny pad in the roof of the mouth. These, however, are not separate elements (despite their close resemblance to vomers) but seem to be integral parts of the premax- illa; they form a pair of stout ridges that support the adjacent tooth rows, lying lateral to them. Posteriorly from the level of the 4" tooth they expand into two vertical flanges, roughly triangular in shape and extending ventrally to an apex just behind the level of the last A.J. CHARIG AND A.C. MILNER | (7") alveolus and then descending again to terminate just behind the subrostral notch. At the posterior end of the premaxillary tooth row, just below the anterior end of the ventral margin of the lower ramus, is a deep notch | which received a rectangular peg-like process on the anterior end of the maxilla. Just in front of this notch lies the subrostral foramen, | while just above the notch, in the outer surface of the premaxilla, lie three well-defined pits. A vertical bony buttress separates the notch | in front from the anterior end of the deep U-shaped trough behind. | VOMER (Fig. 2C). The ventral (i.e. palatal) surface of the con- joined two lower rami of the premaxilla bears what appears to be a! very deep median septum; this divides the concavity of the premax- | illary trough into a pair of broad, U-shaped, ventrally facing troughs. each of which received the dorsal edge of the front maxilla. We interpret this as the extreme anterior portion of the paired vomers, fused together to form an extremely slender, vertical lamina which’ lies in the midline and is clasped between the vertical flanges of the premaxilla (described above). It extends forwards as far as the level of the alveoli for the 4" premaxillary tooth on each side. The central part of this lamina is attached to the roof of the anterior part of the premaxillary trough (though the suture is partly obliterated); anteriorly and posteriorly there is a gap between the dorsal edge of | the lamina and the roof of the (inverted) trough. The extreme anterior end of the conjoined vomers tapers to a point and appears to project - downwards from the roof of the mouth, in the midline between the) premaxillae, an unlikely position that suggests postmortem dis- placement. | MAXILLA (Fig. 3). All that is preserved is the anterior part of the left maxilla, broken off through the 8" maxillary alveolus. Seen in lateral view, both its upper and lower margins are smooth curves, concave above, and with the lower (dentigerous) margin continuing without interruption around the anterior end; the two margins) approach each other posteriorly so that the whole element, though broad from top to bottom in front, narrows appreciably towards the posterior break. Just below the anterodorsal corner the rectangular’ | peg (referred to above under Premaxilla) projects forwards; the maxillary tooth row begins immediately below it and then curves! }; smoothly ina posteroventral direction; when premaxilla and maxillaj are placed together in natural articulation there is a small gap of] about 20 mm between the last premaxillary tooth and the first)» maxillary. In Dilophosaurus (Welles 1984) this gap in the tooth row is some 40 mm, in an animal which is only about two-thirds the size of the Baryonyx holotype. The maxilla extends anteriorly beneath) | the external naris but is separated from it by the ventral ramus of the! { premaxilla, so that its most anterior extremity lies some 45 mm ) farther forward than does the anterior corner of the external naris (in this it contrasts with the condition in Dilophosaurus, where there 1s very little overlap between the two elements and where the mos! | anterior point of the maxilla lies more or less beneath the middle ol} the naris). Just behind the rectangular peg the medial surface of thd = maxilla bears a shallow channel running in an anteroventral direc tion, forming the lateral margin of the external portion of thd » subrostral canal. Along the dorsal margin of the lateral face is the long, slightly » rugose articular surface for the lower ramus of the premaxilla; i) » tapers to a point 135 mm behind the front tip of the maxilla and little way below its dorsal margin. Behind and medial to this, the dorsal margin is smoothly rounded and forms the posterior part 0 the ventral border of the external naris. The lower margin of the ». medial face of the maxilla is formed by a stout rounded ridge, lyin{| > immediately medial to the tooth row; running above this throughou| its length is the articulating surface for the palatine, the anterio} BARYONYX WALKERI mxX.S Portion bearing four prominent longitudinal ridges. Just below the orsal margin of the medial face lie two large, deep fossae, one Above the septum between the 3” and 4" alveoli and the other above he 6" alveolus. Comparison with other theropod skulls suggests that these may have housed maxillary sinuses. The most anterior part of the medial face is in fact an anteriorly jlirected flange (on to the lower half of which run the longitudinal tidges already mentioned). The anterior tip of this flange articulates with a notch on the lateral surface of the anterior process of the palatine (q.v.). The anterodorsal border of the flange forms the edial wall of the internal portion of the subnarial canal. Seen from below, the anterior/ventral surface of the maxilla is of ore or less uniform width. The lateral half of this is occupied by the ental alveoli; the medial half consists posteriorly of the stout unded ridge referred to above, tapering forwards slightly; anterior the level of the 4" maxillary tooth this ridge tapers further into the in medial flange and a less prominent shelf on the inner wall of the ange immediately bordering the medial walls of the alveoli. Also n the medial wall of the flange, behind the shelf, a bony wall Jurrounds what may well be another maxillary sinus. ‘| On the anterior surface of the rectangular peg is a vertical depres- “ion, behind which is the first maxillary alveolus. Altogether there € seven alveoli in the preserved portion of the maxilla; the first four e almost contiguous, separated from each other by only a thick ny wall, but from no. 4 backwards they are more widely spaced. ‘\lveoli nos. 2, 3 and 4 are the largest; nos. 1, 5 and 6 are smaller, no. 7 smaller still. The alveoli are subcircular; no. 4 contains a com- pletely erupted, slightly worn tooth, no. | contains only the base of a smaller, broken-off tooth only partly erupted, and no. 6 shows the tip of a newly erupting crown. We suggested (Charig & Milner 1986, 1990) that there was a mobile articulation on each side between the premaxilla in front and the maxilla behind. We now believe that that suggestion was prob- ably wrong. The anterior end of the maxilla can be fitted neatly into the inverted trough formed laterally by the premaxilla and, medially, by the thin sagittal lamina which appears to be an anterior prolonga- tion of the paired vomers; the rectangular peg on the maxilla then locks into the notch on the premaxilla (Charig & Milner 1986, fig. 1; 1990, fig. 9.2). This complex articulation between premaxilla and maxilla would not permit any movement between those elements, for which, in any case, there is no obvious functional requirement. NASAL (Fig. 4). The paired nasals are fused together into a single element, although it is possible to detect intermittent traces of the median suture. Three fragments were recovered: the major, poste- rior, fragment (Fig. 4) was found lodged against the left lacrimal. As preserved, the posterior fragment in dorsal view resembles an arrowhead: it is a narrow triangle, with its apex directed forwards, and from the base (hind end) of this triangle a stubby median process — the ‘shaft’ of the ‘arrowhead’ — projects farther posteriorly and must have articulated with the frontals. The dorsal (external) surface of the conjoined nasals is raised in 18 Fig. 4 Baryonyx walkeri, holotype, BMNH R9951; posterior part of conjoined nasals. A, dorsal view; B, left lateral; C, ventral. x 0.5. the midline into a sagittal crest which begins a short way behind the anterior end of the fragment as preserved and continues to its hinder extremity. This is crossed, just anterior to the base of the triangle, by a much lower, rounded transverse ridge; where the two intersect they rise to a tall, prominent peak at the centre of a cruciform excresence. The anterior part of this sagittal crest is narrow and sharp. The ventral (internal) surface of the nasals is relatively flat. Paired shallow parasagittal channels, separated by a low median ridge, run the whole length of the fragment and are delimited laterally by sharp ridges running parallel to the mid-line. The parasagittal channels are fairly smooth anteriorly, but more posteriorly bear a series of parallel longitudinal ridges which indicate a firm sutural connexion. The anterior region must have articulated with the ascending process of the maxilla, and the posterior part with an underlying, forwardly projecting process of the frontal; the most posterior part of this surface curves round dorsally so that it faces posteroventrally rather than directly downwards. The lateral extremities are wing-like triangular flat surfaces, the undersides of the lateral portions of the ‘arrowhead’. At the extreme posterior end of each lateral ‘wing’ a very short parasagittal ridge divides the ventral surface into two; the area lateral to this ridge, and, farther forward, the entire width of the surface together form the articulation for the dorsal ramus of the lacrimal. The posterior termination of the lateral wing of the nasal abuts against the anterior face of the dorsal protuberance of the lacrimal (see below). Between the back of the ‘arrowhead’ and its A.J. CHARIG AND A.C. MILNER | ‘shaft’ there lies, on either side, an embayment which must have been part of the margin of a small but distinct fenestra between the nasal, the lacrimal and the prefrontal and frontal. The only theropod | we know of that has a fenestra in this position is Syntarsus (Raath 1977), for which fenestra Raath proposed the name nasal fenestra! | although we suggest postnasal fenestra as a more appropriate alter- | native. The two more anterior fragments, found in a nearby block (30, | Fig. 49) are parallel-sided and triangular in cross-section, the apices being a continuation of the narrow, sharp sagittal crest, little taller than the height at the anterior end of the large fragment. The parasagittal ridge and paired channels continue along the ventral | surfaces in line with those on the larger posterior fragment. None of | the three fragments join but they indicate that the total length of the © nasal was more than 280 mm. Neither fragment shows any trace of | a premaxillary articulation at the anterior end. LACRIMAL (Fig. 5). Only the left lacrimal is preserved. It consists | of two rami meeting posteriorly at an angle of about 35°, which} encloses the posterodorsal corner of the antorbital fenestra and is considerably more acute than the same angle in other theropods. The nasal (dorsal) ramus is 60 mm long, slender, and tapers to a point; the jugal (ventral) ramus is more than twice as long (155 mm) and stouter. The dorsal edge of the nasal ramus articulated with the more lateral part of the underside of the nasal bone. The ramus bears a short, horizontal, medially directed lip which underlies the posterior’ end of its articulation with the nasal. A second, more pronounced lip lies beneath the first one, so that a long narrow groove, some 3 mm wide, is enclosed between the two. The jugal ramus of the lacrimal projects anteroventrally; its) thickened, rounded anterodorsal margin is complete, but its distal! half appears to be drawn out posteroventrally into a thinner flange’ (maximum width as preserved 45 mm), the central part of which is) broken off. Its medial surface bears a strong ridge running its entire length and parallel to its dorsal margin. A second ridge is present on the posterodorsal half only, running just below the first ridge andj parallel to it, thereby enclosing a deep narrow groove. This groove,) continuous with the groove on the nasal ramus, might be supposed to, have enclosed the two rami of a V-shaped prefrontal like that found in such theropods as Allosaurus (Madsen 1976) and Sinraptor, (Currie & Zhao 1993). However, the prefrontal is quite different in|), Baryonyx, small and compact without rami, and could not have} articulated with these grooves. The distal end of the jugal ramus of) the lacrimal articulated with the main body of the maxilla and, more! | posteriorly, with the jugal itself. In the axil of the two rami, on their lateral side, is the deep lacrimal)" vacuity. Towards the base of this vacuity is the external lacrimal) foramen, the outer opening of the lacrimal duct; the internal opening) sits in a small pit on the posteroventral surface of the jugal ramus some 40 mm below the posterior termination of the bone. Farthei)): down the same surface are two deep pits, probably foramina, on = below the other. These may be the equivalent of the single posterio) _ lateral foramen in Allosaurus (Madsen 1976: 20) and fe 'The literature is somewhat confused with regard to the position of this fenestra. It ij unfortunate that, at present, Raath’s dissertation of 1977 is not available to us. Colbery gave a dorsal view of the skull of Syntarsus rhodesiensis (1989, fig. 42B: ‘adapted fron te Raath 1977’) which shows the nasal fenestra bordered by nasal, lacrimal and prefrontall_ th but he did not mention the fenestra in his text. However, Rowe (1989: 129) wrote ‘At thi” junction between nasal, prefrontal and frontal [no mention of lacrimal] is a diamond)” shaped opening, termed the nasal fenestra by Raath (1977),,, that lies just above ami P rostral [i.e. anterior] to the orbit. This structure is known only in the two species 0} Syntarsus. All this was confirmed by Rowe & Gauthier (1990) and by figs 1C and 1} ty of Rowe’s (1989) paper. = S BARYONYX WALKERI vig. 5 Baryonyx walkeri, holotype, BMNH R9951; left lacrimal. A, lateral view; B, medial view. x 0.5. Torvosaurus (Britt 1991: 19); another opening on the lateral surface »robably connected with one or both of these. At the posterior corner f the lacrimal is a rugose articulation for the prefrontal. Just anterior 0 the articulation and separate from it, is a dorsal rugose excres- ‘fence, about 5 mm high, semicircular in shape, measuring some 20 m anteroposteriorly and 12 mm across. This may represent the ase of a horn core; a similar structure is found in many other . heropods, e.g. Allosaurus. FRONTAL (Fig. 6). The small, compact and apparently com- lete left prefrontal proved difficult to identify. It does not have the jharacteristic V-shape of the typical theropod prefrontal with its two jlongated rami (as in, for exampleAllosaurus or Sinraptor), nor does jither of its rugose articulating surfaces fit convincingly against that f the lacrimal. However, we could not envisage it as any other bone om any other part of the skeleton, and, more persuasively, it was iscovered in close association with the nasals and lacrimal in block 2A (Fig. 49; Appendix B). Those two elements had been displaced nly a little from their proper relative positions, and the prefrontal as lying, as it should, against the posterior end of the lacrimal. 19 Fig.6 Baryonyx walkeri, holotype, BMNH R9951; left prefrontal. A, dorsal view; B, ventral. x 0.5. The prefrontal is a thick nubbin of bone, flattened dorsoventrally, with a smooth, slightly concave dorsal surface; seen from above it is roughly isodiametric but of irregular outline. The ventral surface, by contrast, is highly sculptured into ridges and valleys. Anterolaterally the thick margin forms a large, highly rugose facet, which presum- ably articulated with the posterior end of the lacrimal (although, as stated above, the fit is far from precise). Behind this is a short smooth section of the margin, facing laterally, which would have formed part of the anterodorsal section of the orbital rim. The posterior part of the margin of the prefrontal forms another articulating facet, only slightly rugose and deeper laterally than medially, which would have abutted against that portion of the frontal that extended laterally between the prefrontal in front and the postorbital behind to reach the orbital rim. The medial surface of the prefrontal is more complex. Dorsally is a deep V-shaped groove, wide anteriorly, tapering posteriorly to an apex, and divided within into two portions by a weak oblique ridge; the anterior portion formed the posterior margin of the postnasal fenestra, the posterior portion articulated with the anterolateral portion of the main body of the frontal. The ventral surface is remarkable only for four parallel ridges running transversely across the finished bone near its post- erior margin and for a steep little peak, directed ventrally, at the ventral termination of the lacrimal facet. POSTORBITAL (Fig. 7). The left postorbital as preserved lacks only its most dorsal portion. As seen in lateral view, it has essentially the shape of a thick lower-case ‘y’, with the stem of the ‘y’ (the jugal process) descending posteroventrally and then curving round so that its ventralmost portion is directed steeply downwards. Except dorsally, the margins of the postorbital in lateral or medial view are very thin, almost blade-like. The anterior half of the lateral surface, curving round medially to form the posterodorsal surface of the orbit, is smooth and convexly swollen throughout its height (i.e., both in its broad dorsal portion and in its narrowing ventral portion). Above the orbit it is weakly rugose, but less so than in Allosaurus or Torvosaurus (Britt 1991). The posterior half of the lateral surface is weakly concave and leads towards a posteriorly directed flange; the dorsal portion of this flange is broken off but would have articulated with the squamosal, the ventral portion forms part of the anterior border of the lower temporal fenestra’ The jugal process is short and square-ended (shorter than the corresponding structures in Allosaurus and Torvosaurus), and of more or less uniform width. The thick dorsal end of the postorbital is broken off but would have led to a massive facet, directed medially; the greater part of this articulated with the frontal, but a relatively small part articulated at its posterior end with the parietal. The medial surface is much flatter and, though smooth, is faintly roughened all over. The jugal process bears a distinct vertical Fig.7 Baryonyx walkeri, holotype, BMNH R9951; left postorbital. A, lateral view; B, medial. x 0.5. groove, running parallel to its posterior margin. From its posterodorsal region a large process projects mediad, like a flat-topped shelf supported strongly from below by a stout, rounded buttress; this presumably underlay the parietal. Anterior to this a weak concavity faces dorsomedially. FRONTAL, PARIETAL, AND ANTERIOR END OF BRAINCASE (LATEROSPHENOID, ORBITOSPHENOID) (Fig. 8). A stout plate of bone, very thick and with a smooth surface, and sheared off medially just to the right of the midline, comprises the greater part of the right frontal. It is continuous posteriorly with part of the parietal; a fairly straight line across the bone surface, more or less at right angles to the sagittal plane, may well be the suture between them. The parietal, behind that somewhat dubious suture, extends posteriorly and curves dorsally to form a steep transverse crest, thin from front to back; this crest, some 70 mm high, becomes thinner dorsally and ends in a flat top. The weakly concave ventral surface of the frontal is the ceiling of the posterior part of the orbit. The parietal and frontal together Fig.8 Baryonyx walkeri, holotype, BMNH R9951; right frontal and anterior end of braincase, in lateral view. x 0.5. A.J. CHARIG AND A.C. MILNER extend laterally to end in a large and extremely rugose facet for articulation with the postorbital; the parietal forms only the most posterior part of this facet, about one-sixth of its entire length. Along the anterior half of this facet the dorsal surface of the frontal (and presumably of the postorbital too) is raised into a conspicuous protuberance. Anteriorly, the frontal may have contacted the pre- frontal, but we cannot be sure of this. Ventrolateral to the frontal (and therefore ventral to the parietal) lies the laterosphenoid. The laterosphenoid forms the wall of the anterior part of the braincase, thick dorsally but tapering ventrally; in posterior view itis seen in section, behind which a gap separates its broken surface from the anterior face of the prootic (with which it once articulated). Its) shape, as preserved, is roughly tetrahedral. In lateral view it appears equilaterally triangular, with one apex directed forwards so that it just reaches the back end of the facet for the postorbital; dorsally it sutures with the parietal, and at its posteroventral corner its surface is) % extended outwards into a short shelf running more or less parallel}/ii with its lower margin. In ventral view it is again triangular, but this) time it forms a narrow isosceles triangle with the acute angle directed) m posteriorly; this surface is part of the internal wall of the orbit. They ii base of the triangle, at the anterior end, sutures with the frontal, and) i the anteromedial corner curves round medially to border a large) i notch which may have served for the emergence of cranial nerves II) (oculomotor) and IV (trochlear). Medial to this notch lies a thin.) eroded, subtriangular plate of bone, which, according to its general) ai shape and topographical relations, could well be the orbitosphenoid)} jp but it is too poorly preserved to be described properly. Ventral to the frontal and medial to the laterosphenoid (though ke: separated from the latter by a wide cavity) lies a confused mass olf & bone and matrix, the identity of which is not clear. As it is, quite) m fortuitously, almost symmetrical with the right-hand wall of the} jin braincase, it gives the impression that it might be the left-hand wall ie) preserved mostly as a natural mould in matrix: but it cannot bé i because it lies entirely to the right of the sagittal plane. POSTERIOR END OF BRAINCASE (WITH OTIC ELEMENTS) ANE Ws OCCIPUT (WITH BASISPHENOID) (Fig. 9). The supraoccipital, botl} j), prootics, opisthotics and exoccipitals (the latter two elements of the lefy 9, side detached as a separate fragment), and basisphenoid and basioc¢} p cipital are preserved. They are described from the posterior forwards} ». BARYONYX WALKERI The foramen magnum is semicircular, with a more or less straight lower border and a curved upper border. The lower border lies immediately above the robust, backwardly projecting occipital con- dyle. Most of the backwardly directed, articulating face of the condyle is formed by the basioccipital; the exoccipitals make only a comparatively minor contribution to this face, namely the dorso- lateral corners, each of which is demarcated from the basioccipital by a straight suture running obliquely. (This suture has separated on the left-hand side.) The dorsal surface of the occipital condyle between the exoccipital sutures on either side is also formed by the basioccipital, which bears a shallow longitudinal depression in the midline; this means that the condyle, seen from behind, has a roughly heart-shaped appearance. A vertical apron of bone, also part of the basioccipital, extends ventrally from beneath the condyle; it is of uniform width, as wide as the condyle itself, and terminates below in a broad median tongue with a shorter, much narrower lappet on either side. The basisphe- noid projects even farther ventrally; this has parallel sides for some distance and then terminates in a pair of processes, the basipterygoid processes of the basisphenoid, which splay out ventrolaterally. The lower margin of the basisphenoid, between the splayed processes, forms a smooth concave curve. The posterior surface of the basi- sphenoid, below the median tongue of the basioccipital, is deeply furrowed in the midline; the furrow becomes broader and shallower ventrally and peters out altogether before reaching the ventral margin. On either side of the foramen magnum/occipital condyle complex lies a horizontally directed lateral wing, the paroccipital process. This is more or less triangular in transverse section, with one face directed posteriorly, another face dorsally and a little anteriorly, and a third anteroventrally. Its medial part is formed by the exoccipital and its more lateral part by the opisthotic; in most dinosaurs those wo elements are fused together indistinguishably, but in this speci- en an ill-defined suture still demarcates the exoccipital as a slender riangle with its apex directed laterally. The opisthotic extends also entrally, lateral to the whole length of the basioccipital. At about id-height of the occipital condyle, on the suture between the xoccipital and the opisthotic and a short distance lateral from the asioccipital, lies a fairly large foramen which is divided in its depth nto two separate canals. Recent Crocodylia possess several oramina in the same general region, one of which, with exactly the ame topographical relationships as the foramen in Baryonyx, is ikewise divided into two and is unequivocally the vagus foramen; e therefore give that same identity to the ‘double’ foramen of aryonyx. lordansky (1973: 226) observed that in the crocodilian oramen vagi ‘The medial canal extends to the cerebral cavity and is aversed by the IXth and Xth nerves; the lateral canal extends to the iddle ear cavity and contains the Ramus communicans (N. ympathicus) connecting the VIIth and [Xth nerves.’ [actually Xth nd XIth; see Romer 1956: 66]. In Baryonyx the external aperture of e smaller canal is dorsomedial in relation to its larger, ventrolateral eighbour. There are two internal apertures within the cavity of the rain stem, just anterior to the foramen magnum; the lower one, ying at the junction of the floor and wall of the cavity, connects with ‘he larger, ventrolateral external foramen, while the upper one, lying ehind and a little above its partner, probably connects with the maller, dorsomedial external foramen. Whether these two are pre- isely homologous to their crocodilian counterparts is open to uestion. Meanwhile the dorsomedial part of the anteroventral face f the paroccipital process is overlapped by the prootic (q.v.) below nd the supraoccipital (q.v.) above. The left paroccipital process, i.e. exoccipital plus opisthotic, is resent but has been cleanly detached from the rest of the skull. It 21 lacks the ventral extension and has been distorted to some degree. However, laterally it is more complete than the paroccipital process of the right side, and therefore gives a better indication of the true shape and extent of this structure, of which about 25 mm is missing on the right side (see also Charig & Milner 1986, fig. 2; 1990, fig. 9.4). The dorsal part of the occiput is contributed by the supraoccipital, which forms (a) the central region of the dorsal margin of the foramen magnum, between the medially directed arms of the opisthotics; (b) the central part of the back end of the skull roof; and (c) arising from the latter, a prominent stout process which, though essentially projecting dorsally, is also inclined a little forwards at its upper end. This process, somewhat compressed antero-posteriorly and is rectangular in shape when viewed from front or rear; its dorsal surface is broadly crescentic. On its posterior surface, on either side of its base, is the external aperture of a canal that has its other end in the upper part of the internal wall of the braincase and presumably served for the passage of a large blood vessel. The basal part of the anterior side of the supraoccipital process, together with the dorsal part of the underlying prootics, forms on each side a strongly furrowed surface which must have sutured with the parietal. The prootic on either side is a solid, chunky element, forming the wall and floor of this posterior part of the braincase; each meets its fellow along the midline of the floor. A row of foramina for the passage of the cranial nerves runs along the angle between the wall and the floor, effectively dividing the internal and external surfaces of the prootic into an upper part and a lower part (see below, next paragraph). Anteriorly each prootic bears furrowed surfaces for articulation with the laterosphenoid and, more dorsally, with the parietal. Between those articulating surfaces, below the floor of the braincase, is the posterior part of the pituitary fossa; it appears here as two deep median concavities (a smaller above and a larger below, with a saddle between) which penetrate horizontally backwards into the substance of the bone. The posterior wall of the pituitary fossa is the dorsum sellae. On either side of the top of the upper concavity is a large foramen, through which the VIth (abducent) cranial nerve is presumed to have passed forward from the floor of the braincase into the pituitary fossa (see Romer 1956: 67) and thence continued in the same direction to emerge laterally into the orbit (see Osborn 1912: 17). Externally the upper part of the prootic is applied to the anterior surface of the opisthotic, extending about half-way along the length of the paroccipital process. Its lower margin is a smooth, slightly concave curve, separated from the opisthotic behind by a deep furrow which terminates medially in the fenestra ovalis. Just anteromedial to the latter is a foramen for the VIIth cranial nerve and, farther forward still, a large open notch for the emergence of the Vth. Ventral to these nerve exits the lower part of the prootic forms a large plate-like posterolateral process which is applied to the anterolateral face of the basisphenoid. On the left side, the opisthotic has become detached from the rest of the braincase; this exposes the cavity of the otic capsule, con- tained within the body of the prootic. Inside this capsule are two small foramina, both of which lead through into the endocranial cavity to emerge posterior to the large foramen for the passage of the Vth nerve. One of these could well be the duct for the passage of the VIlIth cranial (auditory or acoustic) nerve. These ducts and foramina serve for the passage, not only of blood vessels and lymph ducts, but also of pneumatic openings. The whole of the ventral portion of this occipital fragment, seen in anterior view, is formed from the basisphenoid with its splayed, downwardly projecting basipterygoid processes. As stated above, the more dorsal part of the basisphenoid is hidden by the postero- tO tN Fig.9 Baryonyx walkeri, holotype, BMNH R9951; occiput and posterior part of braincase, lacking detached left opisthotic. A, posterior view; B, right lateral. x 0.5. lateral processes of the prootics. In the midline of the anterior surface is an extremely prominent, ridge-like septum which divides ventrally (like an inverted Y ) into a pair of ridges, each leading down towards the basipterygoid process of its side; between the two ridges is a deep cavity. On either side the flat surface of the basisphenoid slopes posterolaterally so that the whole resembles a pitched roof. At the dorsomedial corner of each flat surface, just beneath the postero- lateral process of the prootic, is a moderately deep hollow. JUGAL (Fig. 10). The left jugal is represented by an elongate portion from the centre of the element, lacking what was probably a significant length at either end. In lateral view the lower margin appears almost straight, while the upper margin forms a smooth, shallow, concave curve that is the wide ventral border of the orbit; thus the element is beginning to widen towards the break at either end, the posterior widening being much the larger of the two. The lower margin is generally rounded, the upper margin sharper. The central part of the bone, directly beneath the centre of the orbit, is fairly robust; the ends of the fragment, however, become rapidly thinner towards the breaks (forwards, backwards and upwards). The lateral surface is more or less smooth except for a weak ridge running diagonally across the suborbital bar, from the upper margin of the bar beneath the front of the orbit to the lower margin of the bar at about the level of the back of the orbit. The medial surface, by contrast, is distinguished by a series of features. The lower margin is wrapped around to form a strong, A.J. CHARIG AND A.C. MILNER | dorsomedially directed flange; this flange is especially well devel- oped beneath the ascending process, and it passes anteriorly and a} little ventrally, becoming weaker as it does so, to merge into the) lower margin itself beneath the orbit. Beneath the orbit, too, it is) flattened, ridged and grooved parallel to its own length to form an articulation that presumably met the ectopterygoid. Posterior to this, where the flange is developed much mote strongly, it encloses a deep} trough between the main body of the jugal and itself. border of the deep trough enclosed by the ventral flange, the troug 1 being at its narrowest at this point. The flange, and therefore the) trough which it helps to enclose, taper away to disappear entirely just) )_ directed process of the quadratojugal. The missing anterodorsal corner of the jugal presumably articu- lated with the maxilla and the posterodorsal corner with the). postorbital. QUADRATE (Fig. 11). Both quadrates are preserved complete, each with a broad transverse articulation for the lower jaw at its ventral) | BARYONYX WALKERI y | lateral. x 0.5. jend. Viewed posteriorly, the quadrate tapers rapidly upwards from ‘the lower jaw articulation into a narrow shaft that swings a little /anterolaterally and then dorsally. It terminates in an expanded knob- like head that fitted into the underside of the squamosal; on the lateral side of the head is a small, distinct, slightly concave facet. The jlateral surface of the broad basal part of the quadrate forms a large, \rugose, kidney-shaped concavity which provided an immovable )sutural attachment for the quadratojugal. Higher up the shaft is a laterally directed ridge, its outer end forming a narrow sutural surface, also for the quadratojugal. Between these two quadratojugal attachments lay a large, elongated quadrate foramen delimited ‘medially by the quadrate and laterally by the quadratojugal. This foramen is much larger than the equivalent foramen of Allosaurus, which is almost entirely enclosed by the quadrate. The entire anterior side of the quadrate is extended forwards into ja wide pterygoid flange; the medial side of the lower half of this ‘/flange was presumably applied to the lateral face of the pterygoid. In llosaurus this flange is directed obliquely towards the midline. The road quadrate condyle, extending transversely mediad from the uadratojugal facet, is characterized by a screw-like sigmoid swell- ing; this is anterior in position on the medial side but curves round elow the condyle to a posterior position on the lateral side. This surface, which articulated with the lower jaw, is much wider medi- ally than laterally. 23 j Fig. 9 cont Baryonyx walkeri, holotype, BMNH R9951; occiput and posterior part of braincase, lacking detached left opisthotic. C, anterior view; D, left Lower jaw (Fig. 12) Parts of both rami are preserved, including: Left side: dentary (virtually complete); splenial (fragmentary); angular (partial). Right side: dentary (a section of the dentigerous bar containing eight alveoli, probably nos. 18-25); splenial (complete); surangular (fragmentary); angular (partial); coronoid (complete). DENTARY (Figs 13, 14). Of all the elements in the mandible, the left dentary is the best preserved. It was found broken into two: an anterior portion containing the first 26 tooth alveoli and a posterior portion containing the last 6. The broken ends had remained in contact with each other, but the two parts had undergone a relative dislocation to produce a divergence of about 45° from the straight line. Nevertheless, their two ends can be fitted together without the interpolation of any significant missing portion. There were there- fore 32 teeth in each complete dentary, 64 in the entire lower jaw. The anterior two-thirds of the dentary (apart from the terminal expansion) is essentially elongated from front to back and flattened from side to side. Its labial and lingual sides are flat and parallel; the dorsal and ventral margins are nearly parallel, but they do converge a little anteriorly. The dentigerous dorsal surface is also flat and demarcated from the labial and lingual surfaces on either side by a Fig. 10 Baryonyx walkeri, holotype, BMNH R9951; left jugal. A, lateral view; B, medial. x 0.5. right angled corner. The ventral margin is rounded so that it would appear U-shaped in transverse section. Viewed from either side, the dorsal margin is bent significantly upwards as it approaches the anterior end: at about the level of the 9" tooth alveolus it is angled upwards through some 25° and then runs in a more or less straight line to the anterodorsal corner of the bone, 1.e. to the region of the 1* tooth. Forwards of the level of the 9" alveolus the dentary starts to expand both vertically and laterally (thereby creating the enlarged ‘terminal rosette’ ). The ventral margin, at the extreme anterior end, passes smoothly into the anterior margin. This region is where the two rami, left and right, joined at the mandibular symphysis. The symphysial surface, sq Fig. 11 Baryonyx walkeri, holotype, BMNH R9951; left quadrate. A, lateral view; B, posterior; C, ventral. A.J. CHARIG AND A.C. MILNER however, is marked by only a very few short parallel striations on the » | lingual side of the dentary, suggesting that the symphysis must have been effected through connective tissue only, retaining some mobil- ity between the two jaw rami. The posterior third of the dentary is altogether much thinner than} the anterior part; it has a broader, blade-like appearance, expanded vertically, with the dorsal and ventral margins diverging widely posteriorly. Indeed, the Meckelian groove on its lingual surface © widens backwards to the extent that it disappears into a broad, flat)” surface occupying the entire height of the bone between the dorsal and ventral rims. The labial surface is flat and featureless. The dorsal and ventral margins of this posterior dentary ‘blade’ xiOi5: | BARYONYX WALKERI Fig. 12 Baryonyx walkeri, holotype, BMNH R9951; partial reconstruction of lower jaw, in medial (lingual) aspect. x 0.135. are much narrower than are the margins of the stout anterior part of the dentary, but they are still somewhat thickened anteriorly, being backward continuations of the lips of the ever-widening Meckelian groove and continuing to curl round towards each other as distinct overhangs. The dorsal overhang forms a slot for the surangular. As ‘the blade widens posteriorly its ventral lip, which forms a slot for the angular, diminishes and eventually disappears altogether; the form of this fragile posterior end of the dentary is indicated only by an outline that has, in places, been badly damaged and eroded. The ventral margin is nevertheless preserved intact and forms a smooth, slightly convex curve. Its extreme tip (as preserved) is probably not far short of what was once the posterior end of the complete dentary. From that end the dorsal margin of the dentary ascended anterodorsally to the widest point and then descended again for- ‘wards, but this part is extremely thin and, as preserved, has an irregularly broken edge. However, at two places on this broken edge there are small lengths of smooth-edged bone of a curved outline, recessed from the general level of the surface, which are doubtless arts of the margins of dentary fenestrae. In dorsal view the lingual margin of the dentary forms a smooth, ently concave curve, somewhat more marked in front of the 9" alveolus than behind it. The labial margin, by contrast, curves outwards markedly to accommodate the roots of the greatly enlarged ‘rst 5 teeth; this swelling produces the mandibular portion of the inal rosette’. _ The Meckelian groove runs lengthwise along the lingual surface of the fragment, at a little below mid-height. It has a strongly overhanging dorsal lip and a fairly well-marked ventral lip, the gap detween them being 16 mm; the base of the groove is a smooth as Anteriorly the groove becomes shallower and narrower, its ips become less pronounced, and it peters out anteriorly some 60 nm behind the symphysis and beneath the wall separating the 4" and t alveoli. Posteriorly, from the 19" alveolus backwards, the lips of he Meckelian groove are themselves grooved and facetted for irticulation with the splenial. Indeed, the major part of the lingual urface of the posterior end of the dentary, tapering forwards into the eckelian groove, served as a slot for the splenial. The labial surface of the dentary bears a number of foramina, _vhich doubtless served for the passage of blood vessels and/or erves. There are about 25 scattered over the expanded area at the nterior end, with a much smaller number farther back. There is also linear series of mental foramina running parallel to the dorsal dargin of the bone, lying in a shallow groove at approximately the evel of the bases of the alveoli. These do not, however, bear a fixed ‘umerical relationship to the alveoli, being significantly fewer than pe latter. They served for the passage of branches of the inferior Iveolar nerve and also, presumably, blood vessels. The last foramen f this linear series happens to be on the anterior end of the posterior nen behind this the shallow groove continues all the way to the osterior end of the dentary as preserved. The dorsal border of the lingual surface of the dentigerous region of the dentary is, for most of its length, of approximately the same height as the dorsal border of the labial wall and as the interdental plates. The interdental plates are, therefore, barely exposed in lingual view, unlike the usual theropod condition (e.g. Allosaurus). The alveoli themselves vary in form, from squarish/circular in occlusal view (2") to egg-shaped (3") to a rectangle twice as broad as long (9"). They vary also in size (see ‘Dentition’). They are immediately adjacent to each other so that they are separated by no more than a thin interalveolar wall. The lingual wall of the row of alveoli, i.e. the row of interdental plates, is separated from the dorsal rim of the lingual surface of the dentary by a deep paradental groove (the ‘nutrient groove’ of Osborn 1912). However, each alveolus communicates with the nutrient groove by means of a slot in its interdental plate, with a foramen at its base. For most of the tooth row these slots are narrow and situated posterolingually (see Fig. 14), but, in the case of the larger anterior teeth nos. 1-5, they are much wider and more anterior in position. SPLENIAL (Fig. 15). The right splenial is preserved complete save for the fragile dorsal margin and posterior end. The left splenial, by contrast, is very incomplete, the only part of its true margin remain- ing being a small piece of its posterodorsal corner. The splenial is a thin, narrow, subtriangular sheet of bone, its apex directed anteriorly, which was slotted into the Meckelian groove on the lingual surface of the dentary. The medial (or lingual) surface of the splenial is distinctly convex dorsoventrally, the convexity being more marked in the posterior half of the bone. The lateral (or labial) surface, facing the dentary, is correspondingly concave. Midway along the length of the element, just above its ventral border, is the moderately large splenial fenestra (= the anterior mylohyoid foramen of Currie & Zhao 1993 and the splenial foramen of Madsen 1976, the latter labelled Meckelian canal in his Plate 9), elongated in an anteroposterior direction. This fenestra is completely surrounded by bone in Baryonyx and Sinraptor but is in a marginal position in Allosaurus; in Ceratosaurus it is only small, and in Coelophysis and Dilophosaurus it is absent altogether. The medial surface bears a sharp ridge just above its ventral margin, which, by virtue of its overhanging nature, produces a ventral-facing groove that received the dorsally directed, wrapped- around lower margin of the Meckelian groove on the dentary. Anterior to the splenial fenestra this splenial groove is well devel- oped; beneath the splenial fenestra it is very shallow, forming nothing more than a faint lip; behind the fenestra the groove is again distinctly present, but it is narrower and shallower than in the anterior region and posteriorly it fades out altogether into a flat, narrow, ventral margin to the bone. The dorsal margin of the splenial rises to a ‘dorsal process’. Anterior to that the edge of the bone, although a little crushed, is almost straight. Posterior to the dorsal process, the dorsal margin of A.J. CHARIG AND A.C. MILNER Sa “CEO “({ensur]) perpaut “| *(ferqey) Tere] “gq >Me TA (TBSN[I9O) [esiop “y ‘Krequap Ye] :1S66u HNNA ‘ad ojoy “UayIDM xkuoking €l ‘sq | BARYONYX WALKERI ‘4 | 18 Aus pd.g nu.for Fig. 14 Baryonyx walkeri, holotype, BMNH R9951; part of right dentary, in dorsal (occlusal) view. x 0.5. the splenial is preserved complete and forms a smooth concave “curve, extending posteroventrally and then posteriorly; this, pre- sumably, formed the anteroventral border of the internal mandibular fenestra. The posterior end of the splenial which articulated with the _angular is missing. The anterior tip of the splenial is a flattened tongue that bears ‘strong longitudinal striations on its lateral face. The ventral margin ‘of the latter, just posterior to the splenial fenestra, forms a shelf, ‘which would have received the anterior end of the angular. “SURANGULAR (Fig. 16). surangular is preserved. This is the central part of the stout upper border of the bone, which itself constitutes the upper border of the posterior half of the mandible as a whole. It consists essentially of a vertical plate, thin and irregularly broken below, which at its upper | Only a large fragment of the right pe is greatly thickened and folded over medially as a brow-like | | if B S - a ee pi ca Fig. 32 Baryonyx walkeri, holotype, BMNH R9951; left humerus. A, lateral view; B, posterior; C, medial; D, anterior; E, proximal; F, distal. x 0.25. v _ BARYONYX WALKERI | On the lateral side of the proximal expansion, where it begins to ‘narrow into the shaft, are two obvious areas of muscle attachment. Immediately posterior to the apex of the deltopectoral crest is a low, lightly rugose hump; this is presumed to have been the point of insertion of the deltoides clavicularis muscle (cf. Norman 1986: 338). A little lower down and more posterior in position, indeed ‘almost in the centre of the lateral surface, is a large shallow depres- sion with a distinct raised border below and on either side; this probably served as the point of origin of the brachialis muscle (cf. ! orman 1986, loc. cit.). At the distal end of the humerus the ectepicondyle is well ex- Be ided, The lower margins of the entepicondyle and the ectepicondyle form an angle of approximately 45° with each other, : nd their facets for articulation with the radius and ulna respectively are subequal in length (the ectepicondylar facet being just a little i onger). In Dilophosaurus the humeral shaft is less straight than in Baryonyx, and inAllosaurus it is even more curved. The two ends are ore offset in Dilophosaurus than in Baryonyx. ADIUS (Fig. 33). Both radii are preserved, the left complete but the right lacking a section of the shaft so that the two ends are no longer connected. The radius is a short, straight, stout bone, not distinguished by any remarkable features. It is 225 mm long and erefore only 49% of the length of the humerus. The proximal end is flattened and expanded, its profile being convex in side view; it apers smoothly into the roughly cylindrical shaft, which continues to taper slightly until it reaches the less expanded distal end. The jatter differs from the proximal end in that it is narrower and not | attened; in end view it appears triangular with one corner extended, the longest side (in the plane of expansion) being almost perpendicu- ar to the plane of flattening and expansion of the proximal end. The distal end-surface is perpendicular to the axis of the bone and has a Shallow convexity at its centre. 45 ig. 33 Baryonyx walkeri, holotype, BMNH R9951; left radius. A, lateral view; B, posterior; C, medial; D, anterior: E, proximal; F, distal. x 0.25. ULNA (Fig. 34). Only the left ulna is preserved, almost complete and 283 mm long; thus it is considerably longer (61% of the length of the humerus) than is the radius. The ulna, unlike the radius, is distinctly bowed. The proximal end bears an unusually powerful olecranon, projecting posteromedially beyond the head of the bone. The articulating surface for the humerus projects anteriorly (making an angle of about 130° with the olecranon projection when viewed from the proximal end) and has a concavely curved anterior profile distal to its apex. The lateral surface of the proximal end actually consists of two surfaces, one facing obliquely forwards and the other obliquely backwards so that they together make a right angle. This angle forms a powerful projection which, unfortunately, has been severely damaged. The most unusual feature of the ulna is the broad expansion of the distal end. This is essentially in the transverse plane, but the greatest part of the expansion is on the medial side where the bone has been extended anteromedially and, at the same time, somewhat dorsally. Although this medial expansion is distinctly more slender than the central part of the ulna, the bone thickens again towards the medial margin so that the articulating surface is of almost constant width. There is also a rather less prominent anterolateral expansion of the lateral side, but here the bone becomes appreciably thicker, so that this part of the articulating surface is virtually semilunate. The distal profile of the ulna, in anterior or posterior view, is markedly convex. CARPUS. served. No recognisable remains of the carpal bones are pre- MANUS (Figs 35, 36). The metacarpals too have all been lost, but a few phalanges are preserved (including three unguals), and some are in excellent condition. They include the original and famous ‘Claw’, i.e. a huge ungual, together with what is probably the phalanx with which it articulated; three phalanges found in articulation 46 Fig. 34 Baryonyx walkeri, holotype, BMNH R9951; left ulna. A, lateral view; B, posterior; C, medial; D, anterior; E, proximal; F, distal. x 0.25. with each other, of which the most distal is a much smaller ungual; and another four, less complete phalanges. Also preserved are a few more small phalanges which are too incomplete and indeterminate to merit description. We assume that Baryonyx possessed the typical theropod phalan- geal formula of 2: 3: 4: (1 vestigial or 0) : 0. Before describing these phalanges it was desirable to determine: 1. Whether they belonged to the manus or the pes. Non-ungual phalanges from the theropod manus are generally smaller than those of the pes; this comparison, however, requires manual and pedal phalanges from the same individual. Unguals from the manus are more strongly curved than those on the pes. When non- unguals and unguals are found together in articulation, these two indicators should reinforce each other. The precise location of the phalanges within the excavation is another source of helpful information; for example, the only articulated digit found was in Block 27b, which, lying at the head end of the ‘dig’ (see Fig. 49) and containing also part of the left scapula, was more likely to contain parts of the manus than the pes. 2. Which side they belonged to, left or right. Recognition of which side of the phalanx is medial and which is lateral enables us to determine whether the element in question is from the left side or the right. A study of theropod phalanges illustrated in the litera- ture (e.g. Allosaurus figured by Madsen 1976, pls 43 and 44) shows that the lateral profile of each non-ungual phalanx is almost parallel to the sagittal plane but the medial profile is A.J. CHARIG AND A.C. MILNER inclined obliquely, in such a way that the distal breadth is |) significantly broader than the proximal. Further, a longitudinal |) groove divides the distal end into two condyles in such a way that |)! the lateral condyle is slightly wider than the medial. As for the | qu i rated by a ridge which begins in a dorsal swelling and then runs ventrally; the swelling and the ridge are a little closer to the lateral | towards the lateral side. 3. Which digit they belonged to and their position within the digit. | Evidence of this nature suggests that all the well-preserved phalanges | are of the left manus. considerably longer.) The slightly eccentric position of the dorsal swelling between the twin facets of the phalangeal articulation suggests that this ungual may be attributed to the left side. The arc of | curvature is larger than in an Allosaurus of comparable size, i.e. itis less strongly curved. Otherwise this element is a fairly typical} * theropod ungual of average proportions, almost perfectly bilaterally | ' _ BARYONYX WALKERI | A 47 ‘Fig 35 Baryonyx walkeri, holotype, BMNH R9951; ungual attributed to left digit I (pollex). A, lateral view; B, proximal. x 0.5. “symmetrical, very little compressed, smoothly rounded along its j length, and sharply pointed. Grooves for the horny sheath run right to the tip, and the flexor tubercle is pinched off more sharply on the “medial side than on the lateral. A distal end of another large phalanx, broken on one side, is Probably part of the first phalanx of the same digit. If so, it is with this element that the large ungual articulated. It appears to be a left ohalanx, and it compares well with the articulated phalanx described ‘mmediately below as the first phalanx of left digit II or the second of left digit III. It has a deep groove between the two condyles, the nedial condyle projects obliquely and has a very distinct ligament sit, and the lateral condyle is broken off almost entirely. | _EFT DIGIT II OR III (Fig. 36). The three phalanges that make up his digit were found in articulation; they might be either an entire igit II, or a digit III lacking its basal phalanx. We can try to resolve his dilemma by comparing the bones with the corresponding ele- ments of Allosaurus and other theropods. The greater length of the phalanges suggests digit II. On the other hand, the proximal end- surface of the most proximal phalanx of the three (II, or Il =) does 10t have the somewhat irregular, single-facetted form characteristic >f a basal phalanx (which would articulate with the distal end of etacarpal II) but has instead the double-facetted form typical of a Ton. basal phalanx (which would articulate with the pulley-like listal end of phalanx III,). These elements, in most respects, are : tandard for a theropod; the ungual, unlike its neighbour on digit I, not enlarged, its length being estimated (it lacks its tip) at 165 . The middle phalanx of the three is appreciably shorter (91 m) than the most proximal (132 mm). This contrasts with the Ondition in digit II of Allosaurus, where the second phalanx is yery slightly longer than the first (Madsen 1976, pl. 45), and in leinonychus, where it is considerably longer (Ostrom 1969: 104). gain, the elements are almost perfectly symmetrical, but a few inor asymmetries indicate that we are here dealing with a finger f the left hand. The most pronounced asymmetry is seen in distal fiew, where the lateral profile is almost parallel to the sagittal plane ut the medial profile is inclined obliquely. GHT UNGUAL ITORIII. This element is poorly preserved, it lacks {s distal half and its proximal end is badly damaged. Nevertheless, } is apparent that it is a damaged mirror-image of the ungual attributed to left digit II or III, and is therefore identified as the corresponding right ungual. RIGHT DIGIT III ORIV. One of the smaller phalanges (length 65 mm) is presumably from a digit III or IV, and the obliquity of its medial profile in dorsal aspect indicates that it comes from the right side. Pelvic girdle ILIUM (Figs 37, 38). Only the right ilium is preserved. The remains consist of four substantial unconnected fragments of bone and one natural mould: 1. The pubic peduncle, with the anterior third of the acetabulum. 2 The ischiadic peduncle, with the posterior two-thirds of the acetabulum (there can be no more than a narrow gap between this and fragment no. 1). 3.A large part of the anterior portion of the iliac blade, albeit with no natural edges. 4. Part of the posterior process, with the brevis shelf and brevis fossa, and a large part of the iliac blade dorsal to it. 5. A mould of the medial surface of a large part of the iliac blade, lacking the anterior end; the original natural mould, formed on the rock which underlay the blade in the field, has been repro- duced in plaster of Paris via a silicone pull. The broken end of fragment no. 4 certainly joins on to fragment no. 2, but the loss of what appears to be a few millimetres of material prevents a firm connexion between them. The information afforded by all of the above permits us to describe the ilium as a whole. The estimated length of the ilium, as preserved, is 835 mm. The material available suggests that the dorsal outline of the blade was a smooth convex curve. The length and precise form of both the anterior and posterior processes are unclear, but the posterior proc- ess seems to have tapered posteriad to a slightly pointed, spatulate termination. This process bears three diverging ridges separated by three concavities, i.e. it is triradiate in posterior view. Thus, we have a concave lateral surface, a concave medial surface, and a deeply 48 Fig. 36 Baryonyx walkeri, holotype, BMNH R9951; three articulated phalanges from digit II or III of left manus. A, proximal and middle phalanges in dorsal view; B, digit with ungual phalanx, left lateral; C, proximal and middle phalanges, ventral; D, proximal end of ungual; E, distal end of middle phalanx; F, proximal end of middle phalanx; G, distal end of proximal phalanx; H, proximal end of proximal phalanx. x 0.5. excavated ventral surface. This deep excavation is the brevis fossa, and the ventromedially directed ridge which partly encloses it is the brevis shelf. The ventral part of the otherwise concave medial surface is broadly convex, with faint subparallel ridges, and it appears to terminate posteriorly in a pronounced thickening; the rest of the ventromedial blade is very slender. The brevis shelf and the iliac blade are of uniform and approximately equal thickness along their length; the brevis shelf thins towards its ventral edges. The ventro- lateral ridge, by contrast, is much stouter and smoothly rounded, and it thickens anteriorly in the direction of the acetabulum. The iliac blade as a whole is of almost uniform thickness, and its lateral surface is strongly dished anteroventrally where it thickens towards the (missing) anteroventral termination. Further, its medial surface bears two scars for the direct attachment of the truncated diapophyses of the sacral vertebrae: one lies more or less dorsal to the pubic peduncle and is broadly U-shaped, the other lies dorsal to the ischiadic peduncle and is narrow and curved. (The distance between these two scars suggests that there must have been another scar midway between them, where the bone of the blade is missing A.J. CHARIG AND A.C. MILNER and the mould of its medial surface is not well enough preserved toy indicate its presence.) These scars have no distinct border, but are), characterized by their finely rugose surface, easily distinguishable from the smooth surface of the rest of the blade. It would be expected that the most ventral portion of the blade would bear another row of} . scars, marking the distal attachment of what are usually calledy ‘sacral ribs’ but which generally consist of no more than the capitu-/ blade is missing, but the mould of its medial surface shows two)». prominent and well-defined scars which presumably mark the) attachment of the last two sacral ribs. | The pubic peduncle is very robust; its articular surface forms 4} ., narrow triangle, the short side being the acetabular edge. The surface e! | »BARYONYX WALKERI | i } ] } | lateral; D, pubic peduncle, lateral. x 0.25. The ischiadic peduncle is shorter and much more slender. Its icular surface is much smaller, semicircular with its medial corner extended to a point (marking the sharp medial edge of the acetabu- um) and extremely rugose. The supra-acetabular crest is well veloped and has a sharp lateral edge. PUBIS (Fig. 39A). One large elongated fragment, 277 mm long, is proken at one end and another fragment, 248 mm long, at both ends; one broken end on each resembles the other so much in size and shape that they are almost certainly parts of the same element, not mmediately adjacent to each other but separated by a gap of 1 determinate width. Both are stout shafts produced on one side into broad flange that could well represent part of one half of the ‘pubic pron’, concave on one side and flat on the other. The fragment with complete end is probably the distal part of the left pubis, the other \ third fragment, 251 mm long, has the shape of one end (presumed listal) very like that of the proximal end of the central portion and is ikely to be a more proximal piece of the same element. The lengths of the three pieces added together total 776 mm, and we estimate the ength of the entire pubis to have been about 1000 mm; this is not \nreasonable when compared with the overall length of the entire i ium (about 820 mm). The position of all three pieces on the block jlan (Fig. 48) is fully in accord with this identification. The acetabular portion of the left pubis is missing entirely. The jroximal fragment of the shaft has a thick, rounded lateral edge; the edial border of this proximal partis a true edge, slightly downturned. he medial half of the dorsal surface is distinctly convex, bulging ‘orsally, and the lateral half is distinctly concave. The bone itself arrows considerably from its proximal end (from 105 mm at the | teak to a minimum of 71 mm), with its medial and lateral profiles ‘ymmetrical and slightly concave. The distal 100 mm or so of this agment shows, on its medial side, the proximal beginnings of a hedial flange, also with a true medial edge; lower down, on the jroximal part of the central fragment, this could well have united 49 Fig. 37 Baryonyx walkeri, holotype, BMNH R9951; right ilium. A, posterior process in posterior view; B, same, ventrolateral; C, ischiadic peduncle, with its right counterpart to form a pubic apron (see next paragraph). The central part of the left pubis is fairly straight and compressed in an oblique transverse plane. Its anterodorsal surface is more or less flat and has a thick rounded lateral edge which widens regularly towards the proximal end of the bone. Its posteroventral surface, by contrast, is strongly concave in transverse section, for it tapers medially into a thin flange. In the more proximal part of the preserved fragment this flange is thicker and is broken off on the medial side, suggesting that it might well have extended farther in that direction to join its fellow in a median symphysis as part of a pubic apron. The more distal part of the flange, however, is extremely slender and has a blade-like medial edge of finished bone, making it less likely that it extended to the midline. This means that a complete apron would have been formed only in the middle section of the pubis. Fig. 38 Baryonyx walkeri, holotype, BMNH R9951; reconstruction of right ilium in lateral and ventral views. x 0.125. 50 nape: TTT] cs Lg 7] Sl ‘ey Fig. 39 Baryonyx walkeri, holotype, BMNH R9951; ventral elements of pelvis. A, left pubis lacking proximal end, posterior view; B, left ischium lacking) distal end, lateral. x 0.25. The distal part of the left pubis is also flattened, with fairly thick, rounded edges; but it seems likely that the plane of flattening was somewhat different from that of the central fragment, parasagittal rather than transverse, from which we infer that there must have been some degree of torsion between the two pieces. This means that A.J. CHARIG AND A.C. MILNER the two surfaces would have faced medially and laterally respec- tively, and the positions of the two edges would have been anterodorsal and posteroventral; meanwhile the blade shows a distinct curvature towards the midline as it approaches its distal end. The anterodorsal edge thickens towards the distal end, its medial and lateral margins | BARYONYX WALKERI | 51 | Fig. 40 Baryonyx walkeri, holotype, BMNH R9951; proximal part of left femur. A, lateral view: B, proximal; C, medial. x 0.25. Be erging as gentle concave curves; the lateral margin develops a “distinct lip, protruding laterally. The profile of the posteroventral edge is weakly concave, so that the pubis expands a little Jposteroventally towards the distal end. However, the end itself “does not expand extensively either anteriorly or posteriorly to form a ‘pubic boot’, as it does in many other theropods; indeed the distal asic of the medial or lateral surface forms a smooth convex ‘curve. The medial surface is perceptibly concave towards its distal nd. _ISCHIUM (Fig. 39B). A large part of the left ischium is preserved, including most of the shaft (total length of the element as preserved 470 mm). It lacks only the distal end of the shaft and a small part of the anterior edge of the pubic peduncle. Of the right ischium there remain only four fragments of the proximal end, none of which can de fitted together directly; one of them includes the pubic articula- ion with a small part of the acetabulum, another includes the obturator flange (see below), a third small piece is a further length of he acetabular margin, and a fourth piece is a length of the upper part x the shaft. _ The element as preserved is a fairly typical ischium, shaped omewhat like a Y. Between the two peduncles is the most ventral ortion of the acetabular ring, narrower in the centre and widening in either direction towards the articulating surfaces for the pubis and _ lium respectively; the iliac surface is the larger of the two. Both i deduncles (and of course the margins of their articulating surfaces) “are convex on their lateral sides and concave medially. Ventral to this _egion the ischium forms a broad plate which has a stout posterior “Margin; just below the iliac peduncle this plate is weakly concave _detween two longitudinal ridges. Towards its anterior margin, below he pubic peduncle, the plate tapers to a slender flange. The ventral dart of this flange with its distinctive fluted margin might be _/nterpreted as the equivalent of an obturator process, but the evident “absence of any notch or embayment proximal thereto suggests that there was no process projecting anteroventrally, such as is found in Allosaurus. There is, however, a slight notch in the anterior edge of jhe ischium distal to this region; we therefore propose to call this part - of the bone the obturator flange. This condition is very similar to that found in ceratosaurs, in particular Dilophosaurus (Welles 1984) and Carnotaurus (Bonaparte, Novas & Coria 1990). The posterior surface of the shaft is broad and flat, but anteriorly it is produced into a thin flange that widens towards the broken distal end. The medial surface of this flange is more or less planar, but the lateral surface is weakly concave. Hind limb FEMUR (Figs 40,41). Both femora suffered extreme damage from clay-winning operations before the presence of the dinosaur skel- eton was recognized. The only fragments rescued were (from the left femur) the proximal end with the uppermost part of the shaft, together with the tibial condyle, and (from the right femur) the distal 40% or so of the bone. There was no overlap between the two sides, and it is therefore impossible to make an accurate estimate of the length of the entire element; 1,200 mm is probably a reasonable approximation. Unfortunately, in this limited material many of the external features had either been destroyed without trace or would have been on regions of the bone that were not preserved; these include the internal and fourth trochanters. The proximal end of the femur appears not to have been crushed but is nevertheless somewhat flattened anteroposteriorly. The inturned head is oval in outline and its articulating surface is directed, not medially, but rather posteromedially; it is separated from the much larger greater trochanter by a broad U-shaped vertical channel on the posterior side. The greater trochanter is elongated in a mediolateral direction and tapers towards the lateral side, with its anterior surface lightly convex and its posterior sur- face concave. The lateral edge, narrow proximally, swells out more distally into a distinct protuberance with a convex anterior margin and a straight posterior margin. The shaft of the femur is more or less circular in cross-section. The distal end shows the usual two condyles, both well preserved; A.J. CHARIG AND A.C. MILNER uos'qy n u09'qiy 90399 u0d'qly u09"qi} ny"00}99 uos q y » BARYONYX WALKERI the lateral (fibular) extends about 15 mm below the medial (tibial) ' They are separated by deep intercondylar grooves on both anterio1 | and posterior surfaces; the anterior groove is V-shaped, while the | posterior is broader and U-shaped. The tibial condyle, seen in / anterior or posterior view, appears to be displaced medially from the line of the shaft; it is greatly expanded anteroposteriorly (though ) somewhat obliquely) but is relatively narrow from side to side. The / fibular condyle, by contrast, lies more or less on the line of the shaft | and is less expanded anteroposteriorly. On its posteromedial surface arises a prominent longitudinal flange; this is equivalent to the ‘blocky protuberance’ described in Allosaurus by Madsen (1976: 43) and the “ectocondylar tuber’ described in Dilophosaurus by ' Welles (1984: 137). The deep concavity between this flange and the fibular condyle itself is the trochlea fibularis of Currie & Zhao / (1993). A longitudinal groove, deep and V-shaped, runs up the “medial surface of the distal part of the shaft; apart from this, the preserved portion of the shaft is in too poor a condition to merit description. TIBIA. One fragment is recognisable as a distal part of a tibia, but “it is too badly crushed to be described. |FIBULA (Fig. 42). The right fibula is in good condition and lacks jonly its extreme distal end; it is 510 mm long as preserved. The ' proximal end is broad and flat and expanded anteroposteriorly; the expansion is mostly towards the rear, so that the posterior profile is )weakly concave. The lateral surface is weakly convex and the medial ‘surface weakly concave (the latter would have faced the tibia). The elongated, weakly crescentic end-surface is just a little thicker anteriorly than posteriorly. The anteromedial edge of the shaft, just below the head, is slightly thickened for the attachment of the | tibiofibular ligaments, but this thickening is less well developed in Baryonyx than in Allosaurus, Dilophosaurus and certain other ‘theropods. Seen in lateral or medial view, the proximal end tapers ‘rapidly distad (with a very weakly concave profile both anteriorly and posteriorly) into a straight, slender, rod-like shaft; the shaft continues to taper more gently, and its anterior and posterior margins ‘eventually run parallel to each other, after which the fragment is \terminated by the distal break. _ The weak concavity of the medial surface of the proximal end, i.e. \the medial sulcus, is continued distad into the medial surface of the ‘shaft, eventually becoming a relatively deep V-shaped trough that \terminates suddenly about half-way down the bone (as preserved). /Below this the shaft is almost circular in cross-section for a short ‘way but then becomes concave again on the medial side, so that the ‘broken end-surface is clearly crescentic. This medial sulcus is better developed inAllosaurus, Torvosaurus, troodonts and dromaeosaurs; in Allosaurus it is continuous all the way down the shaft. /ASTRAGALUS. A flattened piece of bone compares well with the ‘ascending process of the left astragalus of Allosaurus, but is too broken and irregular in outline to merit description. |CALCANEUM (Fig. 43). The right calcaneum is virtually complete. Un lateral or medial view it is semilunate, with a weakly concave ‘proximal profile and a strongly convex distal profile. In proximal ‘view, the proximal surface is thick and rounded at the anterior end but tapers to a sharp point posteriorly; the surface itself is weakly concave to receive the distal end of the fibula. The lateral margin of this proximal surface is not higher than the medial margin, in which respect Baryonyx is like Torvosaurus, but unlike Allosaurus and \Sinraptor. The lateral surface is almost flat and very slightly dished. The jmedial surface is convex and bears, at its anterior end, two deep jrounded hollows, with a slight protuberance between them. One of Fig. 42 Baryonyx walkeri, holotype, BMNH R9951; right fibula. A, medial view; B, lateral. x 0.25. these is above and slightly anterior to the other; they are the facets for the two lateral tuberosities of the astragalus, the proximal facet being the larger of the two. The remaining part of the medial surface forms a Shallow facet for the reception of the tibia; this facet is much deeper and more cup-shaped in Allosaurus. PEs. The collection includes two damaged distal ends of metatar- sals, with typical broad, rounded articular surfaces and with fairly well-developed ligament pits. They could be metatarsals II, III or 1V of either foot. Another fragment is the basal part of an ungual from the foot, as indicated by its large radius of curvature and flattened ventral (internal) surface. A.J. CHARIG AND A.C. MILNER Fig. 43. Baryonyx walkeri, holotype, BMNH R9951; right calcaneum. A, proximal view; B, lateral; C, medial; D, distal. x 0.5. OTHER MATERIAL REFERRED TO THE SAME FAMILY Taquet (1984) described two fragments from the Elrhaz Formation (Aptian) of Gadoufaoua, Niger, as mandibular symphyses of a spinosaurid (Musée National d’Histoire Naturelle, Paris, MNHN GDF 365 and 366); they were refigured more clearly in Kellner & Campos 1996 (fig. 7). Each is virtually identical to the conjoined premaxillae of the holotype of Baryonyx (Charig & Milner 1986; 1990: 139) except in that they possess seven alveoli on each side, not six on the left and seven on the right as in R9951. We consider that these snouts, despite their much younger age, are referable to Baryonyx sp. indet. More recently, Viera & Torres (1995: figs 1-2) referred a left maxilla fragment (Museo de San Telmo, San Sebastian, GA-2065) from the Enciso Group, Barremian of Igea, La Rioja, Spain, to Baryonyx walkeri. It is less complete and about 75% of the size of the holotype maxilla but otherwise indistinguishable. The tip, ante- rior to the 3rd alveolus is missing but complete 8th, 9th and partial 10th alveoli are present confirming that the pattern of evenly spaced subcircular alveoli, gradually decreasing in size, continues further posteriorly than is preserved on the holotype maxilla. The following isolated tooth crowns are referred to cf. Baryonyx; their fragmentary nature and the consequent lack of further informa- tion precludes a more precise identification. Wessex Formation (=Wealden Shales, Barremian), Isle of Wight (Isle ofWightCounty Museum, SandownIWCMS 3642 and5120from Hanover Point; IWCMS 5122, IWCMS 1995 207-209, University of Portsmouth, UOP.97, all unlocalised) described as possible baryonychid by Martill & Hutt (1996). All these crowns are virtually identical with the teeth of the holotype R995 | intheirshape, ornamental pattern, enamel texture, and possession of finely serrated anterior and posterior carinae with seven or eight denticles per millimetre. } Upper Weald Clay (Barremian), former Ewhurst Brickworks, | Surrey (National Grid ReferenceTQ 108379), a single crown (Maid- stone Museum MNEMG 1996.133) collected by Dr E A Jarzembowski in the mid 1980’s and reported recently from just below BGS Bed Sc, equivalent to the top of the Smokejacks beds. Ashdown Sand, (Hauterivian) at Redlands Bricks, Ashdown Works, Scallets Wood, Turkey Road, Bexhill-on-Sea, East Sussex, National Grid reference TQ60/70 721097, a single crown (Bexhill | Museum BEXHM:1993.485), collected by Mr D. Brockhurst in | 1993. This crown differs from the Barremian material only in that i the carinae do not extend the full distance to the base of the crown. | \ PHYLOGENETIC RELATIONSHIPS AND SYSTEMATIC POSITION Our first publication on Baryonyx (Charig & Milner 1986) made no / suggestions as to the relationships of the genus, beyond the claim thatit | - ‘was a typical large theropod in certain respects, resembling, for i example, Allosaurus’. We nevertheless considered it to be sufficiently | distinctive to merit designation as the type-genus of a new family, } Baryonychidae. Our second, more detailed publication (1990) con- | firmed our view of its separate family status; our only other conclusions were that it was not a spinosaurid and that it could not be fitted into Gauthier’s (1984, 1986) cladistic classification of the theropods. Other workers have referred to Baryonyx in various contexts. | Their assessments of its phylogenetic position were made in the light | of one or both of our preliminary descriptions, which were incom- plete and contained some errors. Some of those assessments, because ' BARYONYX WALKERI / of this inadequate basis and because of their brevity, scarcely merit discussion. Nevertheless the authors concerned (including ourselves) ‘have generally considered Baryonyx to be a close or not-so-close Telative of Spinosaurus, so that the two genera have often been placed together as a single family (Spinosauridae Stromer, 1915). This has led certain authors (notably Sereno ef al. 1994) to employ that family as a single unit for the purpose of character distribution analysis. Bonaparte (1991: 18) suggested a relationship between the Spinosauridae, later fragmentary theropods from Africa (Carcharodontosaurus and Bahariasaurus), and the South Ameri- can Abelisauridae; he claimed that the femur of all these forms possessed an anteromedially directed head and a plesiomorphically low lesser trochanter and, if this were true, then they would together represent a major diverse clade (named Neoceratosauria by Novas in 1991), distributed mainly in Gondwana. Holtz (1994a: 1105) cited Bonaparte’s work, but misleadingly stated that the Spinosauridae comprised Spinosaurus and Baryonyx; the latter genus, in fact, was never mentioned by Bonaparte. Holtz also questioned Bonaparte’s suggestion, quoting our claim (1990) that Baryonyx possessed the tetanuran synapomorphy of an obturator process on the ischium; we now know that to be incorrect — the structure is only a flange. Nevertheless, we agree with Holtz because Baryonyx has the femo- ral head directed medially, not anteromedially, and its other characters are tetanuran rather than neoceratosaurian (see below). Meanwhile Sereno ef al. (1996), on the basis of new material from the Cenomanian of Morocco, have interpreted Carcharodontosaurus as a member of the Allosauroidea. Elzanowski & Wellnhofer (1992, 1993) proposed a monophyletic group of theropods consisting of Baryonyx, Spinosaurus, the Troodontidae, Lisboasaurus (tentatively identified as a manuraptoran by Milner & Evans in 1991 but reidentified more recently as a crocodilomorph by Buscalioni et al. 1996), and Archaeornithoides (a tiny fragmentary skull from the Late Cretaceous of Mongolia, described by Elzanowski & Wellnhofer in 1992). This suggestion was based on the following shared characters: 1. Interdental plates absent. 2. Paradental groove present (separating interdental septa from lingual wall of dentary). 3. Lingual wall of dentary lower than labial wall (Baryonyx, Spinosaurus and Archaeornithoides only). 55 Fig. 44 Baryonyx walkeri, holotype, BMNH R9951; reconstruction of entire skeleton, x 0.020. 4. Details of articulation between premaxilla and maxilla (Baryonyx and Archaeornithoides only). 5. Ridge in midline of anterior part of palate (Baryonyx and Archaeornithoides only). Elzanowski & Wellnhofer suggested that Archaeornithoides was the closest known theropod relative of birds, and that its consistent similarities to Baryonyx, Spinosaurus and the troodontids narrowed down the ancestry of birds to those theropods that lacked interdental plates and possessed paradental grooves. However, interdental plates are unequivocally present in the dentary of Baryonyx (Figs 13, 14), forming a barrier between the alveoli and the paradental groove. Further, since Baryonyx does not possess any apomorphous charac- ters of the Manuraptora' (in which taxon is placed the Troodontidae), both it and Spinosaurus must be far removed from the origin of birds. Another argument against the hypothesis of Elzanowski & Wellnhofer is that Archaeopteryx too had interdental plates (as shown by the dentary of the seventh specimen in lingual view, Elzanowski & Wellnhofer 1995). The same is true (Currie 1995) of Dromaeosaurus, type-genus of the family Dromaeosauridae, which many authors (most recently Holtz 1994a) regard as the sister-group of Archaeopteryx. Claims of relationship at family level A minor misunderstanding has arisen over the systematic relation- ship between (a) Baryonyx, (b) Spinosaurus, and (c) a partial maxilla from the Upper Cretaceous of Morocco, described and figured by Buffetaut (1989) as Spinosaurus. The maxilla of the Spinosaurus holotype is known only from a poorly preserved fragment now lost (see below p. 56). Buffetaut was therefore obliged to refer the Moroccan maxilla to Spinosaurus on the basis of the only structures common to both specimens, namely the teeth and their alveoli (using, for Spinosaurus, dentary teeth and alveoli and isolated teeth). Buffetaut claimed also that Baryonyx must be closely related to Spinosaurus (basing his evidence both on the original Spinosaurus 'The name ‘Maniraptora’, proposed by Gauthier (1986: 30) and used by several workers since then, is etymologically wrong. Manus (which is Latin, not Greek as stated by Gauthier), meaning hand, is a fourth-declension feminine noun with the root manu- (see International Code of Zoological Nomenclature, 3rd edition, 1985: 215), not man- as Gauthier obviously supposed. His replacement of the English adjective manual with ‘manal’, op. cit., is presumably based on the same incorrect supposition. 56 and on the referred Moroccan fossil). He believed that both genera might be in the same family, but concluded that, for the time being, the Spinosauridae and the Baryonychidae should be regarded as separate but closely related. We subsequently argued (Charig & Milner 1990: 131-133, 139) that the Moroccan maxilla resembled Baryonyx more closely than it did Spinosaurus, and we stated unequivocally that it should be referred to the Baryonychidae as ‘baryonychid gen. et sp. indet.’. Buffetaut (1992), however, listed certain objections to our argu- ments: 1. We had claimed that his 1989 illustrations of the Moroccan maxilla showed a ‘peg-like anterior extremity’ and noted also that the posterodorsal rim of the fragment was the ‘anteroventral portion of the rim of the external naris’, despite the fact that neither of those structures was mentioned in the legend or in his description. Buffetaut subsequently stated (1992) that no evi- dence of either structure could be seen on the specimen; it seems that we had misinterpreted as true edges the broken edges of the fragment (as seen in the illustrations). 2. We had wrongly accused Buffetaut of misinterpreting the broken posterodorsal rim of the fragment as evidence of an antorbital fenestra, failing to note his comment (1989: 82) that “no evidence of an antorbital fenestra is visible’. . We stated (1990: 32) that the characters common to the Moroc- can specimen and to Spinosaurus ‘can hardly be considered on their own as diagnostic of a particular genus or even a particular family’, but we nevertheless included most of them in our list of the important characters which, in combination, typify Baryonyx. These statements, however, do not really contradict each other; characters that are insufficiently diagnostic on their own might yet be helpful in combination with others. Ww Moreover, further preparation and research have shown that our interim accounts of the partly developed Baryonyx contained some errors, e.g. we had stated wrongly (1990) that the animal possessed at least one elongated neural spine. The matter was broken down by Buffetaut into two separate problems. Should Spinosaurus and Baryonyx be included in the same family? (If the answer is yes, then the family would have to be called by the older family name, viz. Spinosauridae.) The resolution of this problem depends only on material that may be included with near-certainty in one genus or the other, i.e. only on the two holotypes; the Moroccan maxilla is wholly irrelevant. Secondly, is it possible to tell whether the Moroccan maxilla is more similar to Baryonyx or to Spinosaurus, and if so, which? Buffetaut (1992) was right in pointing out the incorrectness of the similarities that had caused us to ally the Moroccan maxilla with Baryonyx rather than with Spinosaurus (Charig & Milner 1990); without those alleged similarities the only structures on which the Moroccan maxilla may be compared with either Baryonyx or Spinosaurus are the teeth. Table 3 summarizes all relevant compari- sons between the teeth of those three forms (as far as is possible with the extremely limited material at our disposal). On teeth alone it can be seen that the Moroccan maxilla resembles Spinosaurus more closely than it does Baryonyx, which is what Buffetaut (1989) had originally claimed. Sereno et al. (1994: 270) also proposed a unique family relation- ship between Baryonyx and Spinosaurus, based on a cladogram containing 44 characters in 9 terminal taxa, including the Spinosauridae (specifying Baryonyx and Spinosaurus) and Afrovenator, a new theropod of Early Cretaceous age. This relation- ship was based on the following proposed synapomorphies: A.J. CHARIG AND A.C. MILNER Table 3. Comparisons of teeth of Spinosaurus with those of related forms. Baryonyx Moroccan maxilla Spinosaurus All teeth: curvature recurved, slightly ? hardly recurved but consistently at all serrations 7 to the mm none none fluting lingual sides of not mentioned sometimes very faint, fine vertical striping on enamel towards base crowns fluted Upper teeth only: spacing widely spaced widely spaced dh interdental plates on both none ‘ pmx and mx labial edge of Wavy in wavy in maxilla ventral view ventral view ? Lower teeth only: number 32 ? not more than 15 spacing close together ? well separated interdental plates present ? absent (fide Stromer and Buffetaut) 1. Elongate prenarial snout. In Baryonyx the whole of the snout is elongate; in Spinosaurus it is unknown. 2. Specialized anterior dentition, manifested by increase in number of premaxillary teeth. Baryonyx has 6/7 premaxillary teeth, | which is 1/2 more than the 5 usual in theropods; but, in any case, | their number is unknown in Spinosaurus. 3. Specialized anterior dentition manifested also by terminal rosette. We coined that term for the horizontally expanded tip to the upper | jaw in Baryonyx, almost completely surrounded by dental al- | veoli. The upper jaw of Spinosaurus is unknown except for a | small fragment of maxilla. 4. Increase in height of dorsal neural spines. This was mistakenly reported in our 1990 publication and is incorrect. In contrast, the | dorsal neural spines of Spinosaurus are greatly elongated (hence | the generic name). Thus, of the four allegedly synapomorphic character-states, three | are unknown in Spinosaurus and may therefore be discounted. The | last is present in Spinosaurus but absent in Baryonyx. This means | that the synapomorphies of Sereno er al. (1994), considered on their | own, provide no justification for placing the two genera concerned in the same family. Consequently, we are now in agreement with Buffetaut in so far as we accept a particular relationship between Spinosauridae and) Baryonychidae. However, in the absence of decisive evidence in) either direction, the following factors predispose us against synonymising the two families: 1. The material of Baryonyx is far more complete than that of Spinosaurus, and Baryonyx would therefore make a much more informative type-genus for the family in which it is included. More} importantly, all of Stromer’s (1915) Spinosaurus material, housed in Munich Museum, was destroyed in an Allied bombing raid in| 1944, and is therefore no longer available for comparisons; nor is there any other material that could be designated as a neotype. 2. The most important distinguishing feature of the Spinosauridae, | as suggested by the name, is the elongation of the neural spines |~ on the vertebrae. Baryonyx has no such elongated spines. The presence or absence of such spines may have little or no BARYONYX WALKERI phylogenetic significance, and it would not be against the Inter- national Code of Zoological Nomenclature to call Baryonyx a spinosaurid; nevertheless it could be very misleading. 3. The similarities between the two genera, as known at present, are not (in our opinion) sufficiently close to justify their treatment, for cladistic analysis, as asingle O.T.U. (operational taxonomic unit). : Also of importance in this connexion are two recently discovered theropods from the Romualdo Member of the Santana Formation : (Albian) of the Araripe Basin of north-eastern Brazil. Kellner & _ Campos (1996) described a fragmentary snout as a new member of _the Spinosauridae and named it Angaturama. Angaturama is espe- cially useful in that it shares certain apomorphous characters with _ Spinosaurus, yet, at the same time, possesses other apomorphous characters that it shares with Baryonyx but which could not be demonstrated in Spinosaurus itself because of the latter’s incom- _pleteness. We therefore believe that, using Angaturama as a link, we can justify a closer relationship between the Spinosauridae | (Spinosaurus, Angaturama and the Moroccan maxilla) and the | Baryonychidae (Baryonyx) than between either of those two fami- | lies and any other. We further believe that this relationship might be reflected in the classification by placing those two families together in the same superfamily Spinosauroidea' Stromer, 1915, excluding the other genera previously placed there by Sereno ef al. in 1994 (Torvosaurus, Eustreptospondylus). The Spinosauroidea as defined here is characterized by the fol- _ lowing apomorphous characters, unique among the Theropoda: } 1. The elongation of the jaws, especially in the prenarial region. 2. A greater or lesser tendency to develop a terminal rosette (greater in the upper jaw than in the lower). }3. The shape of the front of the lower jaw, turned upwards at the | extreme anterior end and constricted transversely just behind that. | 4. An increase in the number of premaxillary teeth to seven. 5. Teeth that show a reduction in: , (a) the compression of the whole tooth in a labio-lingual direc- | tion; | (b) the recurvature of the crown (only slight in Baryonyx, practi- : cally non-existent in Spinosaurus); (c) the size of the denticles on the anterior and posterior carinae of the crown (very fine in Baryonyx, absent altogether in Spinosaurus and Angaturama). | The otherAlbian theropod from Brazil, /rritator Martill, Cruickshank, \Frey, Small & Clarke 1996, is a partial skull lacking the end of the ‘snout; this makes it difficult to compare with Angaturama, except in ')so far as both genera obviously had elongated jaws, external nares set far back, and teeth that share all the unique tooth characters of )Spinosauridae (5. above). It seems very likely that /rritator too is a | spinosaurid, and even possible that it is a senior synonym (by one -month!) of Angaturama. Its authors, however, assigned it to the /Bullatosauria Holtz, 1994a, a new taxon of manuraptorans that . includes the Troodontidae and the Ornithomimosauria. | These characters could, of course, represent independent adapta- tions to similar diets. On the other hand, it is more parsimonious to : ‘Some confusion surrounds the superfamilial name Spinosauroidea (which, according 0 the Principle of Coordination laid down in Article 36(a) of the International Code of ological Nomenclature, must retain the same author and date as existing family- roup names based upon the same type-genus). Sereno er al. (1994) proposed a taxon including Baryonyx, Spinosaurus, Torvosaurus and Eustreptospondylus, naming it orvosauroidea. However, Sereno et al. (1996) changed the name of that nominal taxon (© Spinosauroidea, presumably because the family name Spinosauridae Stromer, 1915 is Senior to Torvosauridae. Si) interpret the unique distribution of all seven in the Spinosauridae and the Baryonychidae as indicating a sister-group relationship between those two families. This latter interpretation is consistent with the fact that the degree of reduction shown in the tooth characters (b) and (c) above is significantly less in Baryonychidae than in the much later Spinosauridae, i.e. the characters reflect a trend which seems to have increased greatly during the time interval between the Barremian and the Cenomanian. The systematic position of Baryonyx and its allies within the Theropoda By far the most detailed analysis of theropod relationships published to date is Holtz’s work of 1994a. His computer analysis of the data, based ona matrix of 19 O.T.U.s and 126 characters, produced a single most-parsimonious cladogram (his fig. 4), and a strict consensus cladogram of the six equally parsimonious ‘runners-up’ (his fig. 5). Holtz’s cladogram agrees in many respects with the cladograms of earlier workers, notably Gauthier’s. In some details, however, it differs greatly. The most important difference lies in the positions assigned to the Tyrannosauridae and the Troodontidae, among the most highly derived Coelurosauria (within the Manuraptora and close to the Ornithomimosauria). We assessed the phylogenetic relationships of Baryonyx by incor- porating it as an additional, twentieth taxonomic unit into Holtz’s data-matrix. It seemed to us, however, that a few of the characters that Holtz had used in his analysis were unsatisfactory and that others were scored wrongly. This opinion has been confirmed independently through the recent publication of criticisms by Clark, Perle & Norell (1994) of several of Holtz’s characters (see Appen- dix C) leading to scepticism of his main conclusion, that the Tyrannosauridae should be placed within the Manuraptora. Our relatively minor modifications of Holtz’s analysis have not affected that conclusion. We deleted 8 characters from Holtz’s data-matrix and re-scored a few others, thereby producing a modified data-matrix (Appendix C). All available information on Baryonyx (on 57 characters, informa- tion on the 61 others being unavailable or too uncertain to be of any value) was added and the analysis run with both the same computer programme (PAUP Version 3.0, Swofford 1990) and with Hennig86 (Farris 1988) and the results compared. PAUP was run under Heuris- tic and Branch & Bound algorithms and produced six equally parsimonious trees of 228 steps, with a consistency index (C.I.) of 51% and a retention index (R.I.) of 70%. Hennig86, using mh* and Branch & Bound options, gave similar results: six equally parsimo- nious trees of 234 steps, witha C.I. of 50% and an R.I. of 70%. These figures resemble Holtz’s (51% and 71% respectively). Holtz’s best cladogram had 244 steps, a few more, but it should be remembered that he used 126 characters as against our 118. All trees were rooted to a hypothetical ancestor on unordered characters. All these trees (and the corresponding consensus trees) produced by both programmes display topologies that are generally similar to each other and which, in their broad outlines, resemble Holtz’s single most-parsimonious cladogram. In all of them there is a monophyletic Ceratosauria arising from the basal node, and, much farther away from the root, a monophyletic Neotetanurae. Between them are three other nodes that give rise to a polyphyletic assem- blage that might be referred to collectively and informally as ‘basal Tetanurae’ (among which Baryonyx is placed). The more detailed arrangement of the O.T.U.s was constant within the Ceratosauria but varied from tree to tree within the Neotetanurae; those clades, however, do not concern us here, and discussion of such problems is beyond the scope of the present work. Zs) Ceratosauria Fig. 45 Cladogram of the Theropoda, based on our modified version of Holtz’s data-matrix and including also Baryonyx. In all our cladograms (summarized in Fig. 45) Baryonyx is placed as the sister-group of the Neotetanurae Sereno ef al., 1994 (=Avetheropoda Paul, 1988), arising from node 5 (numbering of nodes and characters are as used in this work, not Holtz 1994a; for details seeAppendix C).The nearest outgroup of the combined clade is Megalosaurus, arising from node 4, and the second nearest Torvosaurus, arising from node 3. All these taxa (1.e., Zorvosaurus, Megalosaurus, Baryonyx and Neotetanurae) together comprise the Tetanurae Gauthier, 1986. If Baryonyx were inserted into Holtz’s cladogram (his fig. 4) it would split off from the main stem of his Hennigian comb, leading to the Coelurosauria sensu Gauthier, between his nodes 6 and 7. The specific evidence for the position of Baryonyx on our cladogram (Fig. 45) is assessed critically below. At node 3 (Jorvosaurus dichotomy, Holtz’s node 5, Tetanurae) there are fourteen synapomorphies. Baryonyx provides positive evidence of only three of those, namely: 50. Dorsal vertebrae pleurocoelous (“unambiguous synapomorphy’ of Tetanurae, fide Holtz loc. cit.). 4. Dorsal vertebrae with transverse processes that are not strongly backturned or triangular in dorsal view (supposedly reversed character). 110. All three pelvic elements not fused together in adults (suppos- edly reversed character). At node 4 (Megalosaurus dichotomy, Holtz’s node 6, unnamed) « a) . w = y Ses et ne) ees Bin wots ey Cees: ow « $ $ s | S f ¢ F Seat Se S & S S S cy 9 S 5 & $ § S g ss RS 2 S S S 9 y G gy 9 gS 8s o ~~ x 4 y 9 Oe ee Re ee ee ee Sine exQhol C initio; aaeQagean Bie ett mite, Sa A.J. CHARIG AND A.C. MILNER there are eight synapomorphies, but unfortunately Baryonyx affords © evidence of none of them. At node 5 (Baryonyx dichotomy, new node, unnamed) there are | five synapomorphies, on only one of which does Baryonyx provide data, namely: 96. Coracoid tapers posteriorly (‘unambiguous synapomorphy’ of | Holtz). We thought it unwise to propose a new name for the taxon based on this node; the phylogeny is not yet sufficiently well established. This evidence is not very convincing. More helpful is the list of | synapomorphies for node 6 (dichotomy between Allosaurus/ Acrocanthosaurus and Coelurosauria sensu Gauthier; Holtz’s node 7; Neotetanurae orAvetheropoda); there are four of them upon which ! diagnosis of the Neotetanurae is based. In this case, fortunately, Baryonyx provides clear evidence to show that it has none of them: 105. Axis with spine table. 117. Manual digit I reduced in length. 88. Obturator process present. 92. Pubic boot pronounced (‘unambiguous synapomorphy’ of | Neotetanurae, fide Holtz loc. cit.). Certain characters of Baryonyx, though used in Holtz’s and our analyses and seeming to be apomorphous for that O.T.U., also occur; elsewhere in the cladogram. If our suggested phylogeny is correct, | BARYONYX WALKERI this means that they must have evolved independently in Baryonyx and in the other forms concerned. Examples are: 5. Pronounced subnarial gap, indicating a possibly mobile premax- illary-maxillary joint. This gap (renamed the subrostral notch in Baryonyx, see p. 14) occurs also in certain ceratosaurs, namely Coelophysidae and Dilophosaurus, and was therefore consid- ered by Holtz (1994a: 1104) tobe anunambiguous synapomorphy | for the Coelophysoidea. Rowe & Gauthier (1990: 154) claimed that articulated material shows the premaxillary-maxillary joint | of ceratosaurs to bea firm junction, as is also the case inBaryonyx. | 77. Nasals narrow. This occurs also in Dromaeosaurus, Archaeopteryx, Tyrannosaurus, Troodon and Ornithomimus. 13. Parietals projected dorsally. This occurs also in Ceratosaurus and Abelisaurus. __ We considered also the partial analysis of Sereno er al. 1994 (in _ which the synapomorphies are listed only under “References and | Notes’). Those authors did consider Baryonyx, though not as a | separate O.T.U. but as part of the taxon Spinosauridae (see above, p. | 56). Their cladogram (Fig. 46) shows the Spinosauridae as the sister- group of Torvosauridae, arising from node 4 (unnamed, our / numbering); those two O.T.U.s together as the sister-group of Afrovenator, arising from node 3 (Torvosauroidea); and the _ Torvosauroidea as the sister-group of the Neotetanurae. The | Torvosauroidea and the Neotetanurae together constitute the _ Tetanurae. We replaced Spinosauridae as an O.T.U. by the genus | Baryonyx on its own, in an otherwise unmodified data-matrix, then _ tan the programme as described above (p. 57). This produced a single most-parsimonious tree of 58 steps, C.I. 93%, R.I. 90%, | identical to the tree figured by Sereno et al. except in that Baryonyx . occupies the position that was previously occupied by Spinosauridae. _ This cladogram requires critical scrutiny. Sereno et al’s (1994) piinking of the Spinosauridae (which included Baryonyx; _ Spinosauroidea sensu this work) with the Torvosauridae | (Torvosaurus + Eustreptospondylus) to form an unnamed taxon is | supported by only two synapomorphies: "1. Radius less than 50% of humeral length. This is true of Baryonyx (49%), but in Spinosaurus neither humerus nor radius is known. (1) Neotheropoda Fig. 46 Cladogram of the Theropoda, according to Sereno et al. (1994). 59 N Manual digit I ungual elongate (three times height of proximal articular end). This is essentially true of Baryonyx, where the length of the ungual is more than four times the height of the proximal end: length of ungual measured around the outside of the curve 310 mm, height of end 73 mm. Again, the manus is unknown in Spinosaurus. Thus, while there is no evidence either way for the condition in Spinosaurus, it seems that Baryonyx and Torvosaurus do share two characters. However, such similarities in relative proportions could easily have developed homoplastically and therefore lack phylogenetic significance. Further, while the reduction or absence of a quadrate foramen links Torvosaurus with Afrovenator (see below), that same foramen is prominent in Baryonyx; indeed, it is more of a fenestra than a foramen. Thus there does not seem to be an adequate demonstration of a close phylogenetic link between Baryonyx and the Torvosauridae. Sereno etal. (1994) also claimed that this unnamed Torvosauridae + Spinosauridae taxon was the sister-group of their new Saharan theropod Afrovenator, ranking the combined taxon as a superfamily and naming it Torvosauroidea (though those same authors, in their 1996 publication, called it Spinosauroidea). This was based on five synapomorphies, all characters of the skull: 1. Anterior ramus of maxilla as long anteroposteriorly as tall. In Baryonyx the whole of the maxilla as preserved is anterior to the antorbital fenestra and may therefore be presumed to be part of the ‘anterior ramus’ of Sereno ef al.; in which case it is several times longer than tall. In Spinosaurus practically nothing of the maxilla is preserved and this character-state is therefore un- known. Anterior ramus of lacrimal dorsoventrally narrow. In Baryonyx this structure, more usually referred to as the nasal or dorsal ramus, shows a condition intermediate between those of Torvosaurus and Allosaurus. Nothing is known of the condition in Spinosaurus. 3. Lacrimal foramen small and positioned at mid-length along the jugal ramus. This is confusing in so far as there are three lacrimal foramina in Jorvosaurus (Britt 1991: 17-19); Britt’s photograph of the lacrimal in medial view clearly shows that the large one, which he called the posterior lacrimal foramen, lies one-third of the way down the orbital side of the jugal ramus. In Baryonyx also there are three foramina, one large and, close together, two very small ones; the large one lies beneath the junction of the two rami, i.e. at the dorsal extremity of the jugal ramus. The corre- sponding character-state is unknown in Spinosaurus. This proposed synapomorphy is altogether unclear. 4. Ventral process of postorbital broader transversely than anteroposteriorly. This character-state is not found in Baryonyx and is unknown in Spinosaurus. 5. Quadrate foramen reduced or absent. Baryonyx has a large, prominent quadrate foramen between the quadrate and the quadratojugal. The condition is again unknown in Spinosaurus. N In summary, two of the relevant character-states (nos | and 2) may be present in Baryonyx but it is difficult to have much confidence in them. For the other three (nos 3, 4 and 5) the character is either unclear or in the opposite state. All five characters are unknown in Spinosaurus; and none appears in the abbreviated description and diagnosis of Afrovenator abakensis published by Sereno et al. (1994: 270). We can find no justification for the placing together of these alleged sister-groups. The net result of our re-working of the analysis of Sereno et al. 60 (1994), with Baryonyx replacing Spinosauridae as an O.T.U., is to confirm the systematic position of Baryonyx as a ‘basal tetanuran’, and, at the same time, to cast serious doubts upon any particularly close connexion with Torvosaurus and/or Afrovenator. Another cladistic analysis of the Theropoda that deserves brief consideration is that of Russell & Dong (1993: 2122-2125). Many of the O.T.U.s (mostly at family level, but including also Baryonyx) were novel, as were many of the synapomorphies proposed, and this resulted in a cladogram that was unusual in several respects. One group, including Baryonyx, Yangchuanosaurus, Allosaurus, dromaeosaurids and tyrannosaurids appeared to be defined ‘by the acquisition of horn-like tuberosities along the dorsolateral edge of the skull and relatively large teeth which are few in number (Russell & Dong, 1993: 2121). In fact, all that appears to pertain to Baryonyx in the former connexion is one small tuberosity on the lacrimal. Its teeth (in the lower jaw at any rate) are more numerous than in any other theropod except Troodon (Osmolska & Barsbold 1990) and Pelecanimimus (Pérez-Moreno et al. 1994), both of which are systematically remote from Baryonyx. Russell & Dong’s analysis does not seem to shed any additional light upon the systematic position of the latter. Finally, Holtz (1995) published a new phylogeny of the Theropoda in which he proposed a weakly supported monophyletic Megalosauroidea comprising Afrovenator + (Megalosaurus + Baryonyx + Torvosaurus). Conclusions Our final conclusions on the systematic position of Baryonyx are that it should remain as the type-genus of the family Baryonychidae; that its nearest relatives (to be placed in the same superfamily Spinosauroidea) are the poorly known Spinosaurus, Angaturama and perhaps /rritator; that this group lies within the Tetanurae, close to the base of that taxon, with the Neotetanurae as its sister-group; and that Megalosaurus and Torvosaurus respectively are progres- sively more distant outgroups to the combined Spinosauroidea+Neotetanurae clade. This is the best that we can do in our present state of knowledge. Our conclusions are subject to the usual caveats: 1. Lists of characters for the same group, drawn up by different authors, are often very different’ (Charig 1993) and are bound to be far from complete (Panchen 1982). 2. The itemisation of the characters and the scoring of the character- states for the data-matrix require a great number of difficult subjective decisions. 3. Small changes in selection of characters, itemisation and scoring can produce drastic changes in the results of the analysis. 4. Our knowledge of those same character-states in Baryonyx 1s very incomplete, and in certain cases the scoring was likewise difficult. 5. Other authors, generally using very different lists of characters and different O.T.U.s, have produced different cladograms for the Theropoda. 6. The most parsimonious cladogram does not necessarily repre- sent the actual course of evolution (Charig 1982: 414). At the same time we must point out that, when two or more analyses based on very different lists of characters produce remarkably ‘Holtz (1994a) used 126 characters in his analysis of the Theropoda; Sereno et al. (1994) used 80 (excluding the Ceratosauria). Even the most generous interpretation could find no more than 17 characters common to both, of which only 4 pertained to the skull and vertebral column. A.J. CHARIG AND A.C. MILNER similar cladograms, the fact that the shared topology is supported by different rafts of evidence greatly enhances our confidence in its correctness. PALAEOECOLOGY In Early Cretaceous times the Weald of Surrey, Sussex and Kent was partly occupied by the Wealden Lake, a large lake of fresh to’ brackish water that extended westwards into Hampshire. To the north lay dry land in the region of what is now the London conur- bation; to the south lay the Anglo-Paris Basin and the open sea. Two large rivers flowed down from the north and north-east to) discharge their waters into the lake through a common delta, with 7 shallow creeks and oxbows. The climate was, in modern terms, sub-tropical. Ross & Cook (1995) have recently reported on the stratigraphy and sedimentology of the Smokejack’s locality in particular. The exposure at that site consists of 23 m of Upper Weald Clay, all of it of early Barremian age (estimate of absolute age approximately 130 million years). The Baryonyx remains were in fine silty clays of non- marine origin, light olive-grey in colour and containing large irregular nodules of bioturbated sideritic siltstone. The sediments give indica- tions of shallow-water facies (e.g ripple-marks at the base of the dinosaur level) but show no signs of complete drying out (mud- cracks, erosion etc.), nor is there evidence of braided river deposition: they had obviously settled out of still water (a “low energy” environ- ment). This lithology, both at the Baryonyx horizon and in the bed: immediately below, has been interpreted by Ross & Cook as indicat- ing a fluvial and/or mudplain environment with areas of shallo water, lagoons and marsh. The fossil flora and fauna of the Smokejack’s locality as a whole give some idea of the environment in which Baryonyx live (Frontispiece), although it should be remembered that the flora, fauna and total environment all varied significantly at differen levels in the sequence. Common plant remains include the fer Weichselia reticulata (Stokes & Webb), concentrated locally in layers within the clay and in sideritic lenses, and an aquatic o1” marsh-dwelling herbaceous plant that grew in monotypic stands Bevhalstia pebja (Hill 1996). Also present were filicopsid ferns horsetails, club mosses and conifers. Of great importance to th environmental interpretation are the abundance and variety oj fossil insects at Smokejack’s, 10 orders of which were recorded by) Jarzembowski (1984). The exact stratigraphical positions of mam of the older finds are not known, but some insects have now bee: recovered in situ from sideritic siltstone lenses at a number 0 horizons below Baryonyx (Ross & Cook 1995). Other elements 0 the invertebrate fauna were ostracods, isopods, conchostracan! and bivalves. The vertebrate fauna includes sharks, bony fishes (notably Lepidotes mantelli Agassiz), crocodiles, pterosaurs, and dinosaurs Apart from Baryonyx walkeri, the only other dinosaur remaini yielded by this locality consist of a considerable quantity of materi referred to the relatively common ornithopod /guanodoi atherfieldensis Hooley, 1925, and a very few isolated bones of smal sauropods. In 1982 we excavated the hind portion of an articulat Iguanodon atherfieldensis skeleton at a distance of only about 1007 from the Baryonyx site and at approximately the same stratigraphic level. Fragmentary remains of a large Iguanodon bernissartensi Boulenger, 1881, were collected in 1988 from a higher level in th sequence. —E BARYONYX WALKERI FUNCTIONAL MORPHOLOGY AND MODE OF LIFE We (Charig & Milner 1986) have suggested for Baryonyx: (a) ichthyophagy (a diet consisting mainly of fish) but with the further possibility of a scavenging habit; (b) a terrestrial existence; and (c) quadrupedality, facultative at least. DIET AND METHODS OF FEEDING. Kitchener (1987) noted that ‘The alternative lifestyle of a scavenger [had] received very little attention’ in our paper; he followed this with the full details of our case for a scavenging habit (as presented by us at meetings in Drumheller and Belfast in 1986 but omitted from our publication because of lack of space). Kitchener then went on to state his preference for the idea of a scavenging habit over that of ichthyophagy, a preference which we do not share. The characters suggesting a scavenging habit, as published by Kitchener, may be summed up as follows: . Stance facultatively quadrupedal. . Neck long (both 1 & 2 suitable for feeding at ground level). 3. Fore-limbs massively developed, with huge claws (ideal for breaking into a carcase). 4. Snout narrow (well suited to investigating the body cavity of the carcase). 5. External nares placed far back from tip of snout (permitting simultaneous feeding and breathing). 6. Premaxilla and maxilla connected by flexible hinge (allowing freer movement in the restricted space in the body cavity). 7. Teeth slender and finely serrated (for processing the soft viscera). Noe However, we now know that there is no evidence for characters nos 1, 2 and 6. Kitchener also put forward one character as evidence against a typical macropredacious habit: 8. Mandible and teeth weakly developed (particularly unsuitable for killing and feeding on large herbivorous dinosaurs). Even crocodilians, after the birds the nearest living relatives of Baryonyx, have great difficulty in breaking through the skin of large prey. e€ also made two arguments against ichthyophagy: It is unlikely that such a heavy creature ‘could have been sufficiently manouverable [sic] to catch fast-moving fish’. 10. Baryonyx has too many adaptations for fish feeding, viz. fore- limbs for hooking fish and teeth and jaws similar to the fish-eating gavial’s; ‘one adaptation would suffice’. N a critical reply to Kitchener, Reid (1987) cited three counter- ‘guments: 0 3 above: Most available carcases would probably already have been broken into by the primary predator. © 8 above: The teeth of modern crocodilians are more or less conical, not adapted to slicing, and are in no way comparable to the bilaterally compressed “steak-knife” teeth of typical theropod dinosaurs. (In fact, the condition in Baryonyx is intermediate between the two.) © 9 above: Large animals, e.g. grizzly bears, are capable of 61 catching fast-moving fish, at least in shallow water. Techniques analogous to ‘gaffing’ or trout-tickling could also have been employed. We add two further comments: To 7 above: The crowns of the teeth were not slender; in fact, they were less compressed laterally than those of other, more ‘typical’ theropods. To 10 above: There is no logic in the argument that, because an animal has two different adaptations that appear to have served the same purpose, neither of them can in fact have served that purpose. The information that may help us assess the diet and feeding methods of Baryonyx is as follows: 1. There is direct evidence of what the animal had been eating. Within its smashed-up rib-cage, in its stomach region, were found: a. Acid-etched scales and teeth of the common Mesozoic fish Lepidotes (Fig. 47). These are of especially great significance. b. The disarticulated skeletal remains of a young Iguanodon [See Appendix B] showing some evidence of abrasion (and/or etching). 2 There is also circumstantial evidence of the ichthyophagous habits of Baryonyx. Modern crocodilians have certain adapta- tions which are clearly effective in catching and swallowing fishes; analogous characters are found in Baryonyx: a. The jaws are long and very narrow from side to side; they are expanded horizontally at the anterior end, with enlarged teeth around the margins of the expansions (Fig. 2). In the upper jaw this spatulate expansion forms a ‘terminal rosette’, not unlike the corresponding region of the skull of a modern gavial. b. Seen from the side, both upper and lower jaws have sigmoid dentigerous margins (Fig. 1); the upper jaw has a downturned tip and a ‘subrostral notch’. Altogether this particular aspect of the animal has an appearance that is distinctly crocodile-like, albeit only superficially so. These similarities support the idea that Baryonyx caught small and moderate-sized fishes in a crocodilian manner, i.e., it seized them with the end of its pincer-like jaws and gripped them transversely in its subrostral notch and lateral teeth; the shape of the jaws and the nature of the teeth accord well with the suggestion that they some- how helped the grasping and manoeuvring of slippery prey. The animal might then have tilted its head back and manoeuvred the fish around so that the fish slid head-first down the gullet into its stomach, as do modern crocodilians. We believe that Baryonyx ate fish habitually, though not necessarily exclusively. Whatever the nature of Baryonyx’s preferred diet, the animal seems ill-equipped to have been a typical theropod macropredator of the type that relied largely on short, powerful jaws and blade-like teeth with serrated edges to capture, kill and dismember its prey. This is because: 1. The middle part of each bony ramus of the mandible is wafer- thin. 2. The teeth are only slightly compressed from side to side, thus differing from those of typical macropredacious dinosaurs like Allosaurus. 3. The denticles on the carinae of the teeth are remarkably fine (about 7 per millimetre). A A.J. CHARIG AND A.C. MILNER Fig. 47 Lepidotes scales, scanning electron micrographs. A, from rib-cage region (Block 44) of the Baryonyx skeleton, showing acid-etching of the enamel, x 8; B, from the same horizon and locality but not associated with the Baryonyx, showing smooth unetched enamel, x 11. Conversely, Baryonyx possesses characters that suggest different methods of acquiring its food: 1. The jaws are long and narrow, not unlike a pair of forceps. They could have been dipped, either into water to seize relatively small fish, or into the body cavity of a large dead animal to seize the entrails. 2. The external nostrils were lateral and far posterior in position. This would have enabled the animal to continue breathing com- fortably even while its snout was still deep in the water or in the body cavity of a carcase. 3. The large and presumably heavy head would have limited the mobility of the neck. 4. The ends of the cervical centra are not “offset”; it is therefore unlikely that the neck would have been held in a sigmoid curve. 5. The cervical vertebrae have well developed epipophyses, but their neural spines are low and lack spine tables. This suggests that the intervertebral muscles were strong and the neck highly mobile, which again would have helped its feeding activities. 6. The humerus is very robust, with both ends broadly expanded. The proximal end in particular is expanded anteriorly into an unusually long and high deltopectoral crest and posteriorly into a well-developed internal tuberosity (compare with other theropods, including Jorvosaurus). The pectoral girdle too is well developed and there was also an ossified sternum. All this would have facilitated the powerful adduction, abduction (and perhaps rota- tion?) of the fore-limb as a whole. 7. The radius is relatively short and that too is robust and powerful. The same applies to the ulna without the olecranon, but the olecranon itself is remarkably long; thus the ratio ‘length of olecranon/length of ulnar shaft’ is exceptionally high. This pro- duced a mechanical advantage (1.e., leverage) when the fore-arm was extended. 8. One manual ungual is enormous; it is of typical theropod form, though somewhat more slender than in Al/osaurus, and it is not modified into a “sickle” like the enlarged pedal ungual of Deinonychus (Ostrom 1969). This was clearly an extremely powerful offensive weapon. The characters of the fore-limb and manus suggest that the fore- | ~ limbs of Baryonyx were exceptionally powerful, the fore-arm being | capable of exerting great force at the wrist when extended. By | ~ activating the enormous claw on the thumb, this would have ena- | bled the animal not only to catch and kill its prey (if necessary), but |” also to rip and tear it to pieces. In short, the active predation of larger animals and the breaking up of its food were probably \ performed by the fore-limbs and claws rather than by the jaws and | teeth. The enlarged claws could also have been used for ‘gaffing’, | i.e. hooking or flipping fishes out of the water as is done today by | ~ grizzly bears. The fine tooth denticles of Baryonyx do not seem to be suitable for the ‘rip and grip’ cutting action demonstrated by Abler (1992) for more coarsely serrate theropod teeth. Farlow, Brinkman, Abler & |) Currie (1991) suggested that theropod lateral teeth with very fine denticles might function in a manner indistinguishable from that of | smooth-bladed teeth; this might be true of Baryonyx. Farlow et al. | also showed that the lateral blade-like teeth of most theropods | displayed a consistent relationship between tooth size and denticle , size. They reasoned that a serrated blade might inflict as much | damage as a smooth-edged thinner blade but would be less likely to break. Since the blade-like teeth of Baryonyx are less compressed than those of most other theropods, the reduction in the size of their denticles might be correlated with their greater robustness. If we consider the function of the dentition as a whole, all the teeth | seem suitable for piercing prey and cutting it smoothly. The long, 4 comparatively straight teeth in the terminal rosette and expanded tip | of the dentary also possessed a stabbing function. Those teeth on the lower jaw that lie below the preserved portion |_, of the maxilla were (on the evidence of the alveoli) smaller, more} \ crowded together, and more than twice as numerous per unit length | of jaw as the maxillary teeth that opposed them. A somewhat similar discrepancy is known not only in Troodon, which has 15—20 maxil- | . lary teeth opposed by 35 in the dentary (Currie et al. 1990), but has | also been observed in the strange new ornithomimid Pelecanimimus (Pérez-Moreno et al. 1994). This discordance between the number f of teeth in the upper jaw and the corresponding number in the lower is very unusual and, as far as we know, has no analogue among living | | BARYONYX WALKERI reptiles. One might speculate that its functional significance in ' Baryonyx might be somehow connected with the gripping and | manipulation of food, the lower teeth forming a closely spaced series _ of piercing and holding points that opposed the more widely spaced | upper teeth. _ In connexion with the animal’s feeding habits, it should also be mentioned that an apparent gastrolith was found within the rib-cage | of the Baryonyx. _ On balance, we still envisage Baryonyx as mainly a fish-eater. It probably crouched on the banks of lakes, creeks and rivers or waded in the shallows (Frontispiece), and it secured its prey by direct ‘seizure with the jaws and perhaps also by ‘gaffing’. Small fishes would have been swallowed whole, larger ones broken up by the e fore-limbs with their huge claws. Fishing, however, was | not the only source of food; there is also: 1. Circumstantial evidence that it may well have been both an active predator (using its powerful fore-limbs and claws rather than its jaws and teeth) and/or an opportunistic scavenger. 2. Positive evidence (i.e. recognisable bony remains within its rib- cage) that it had eaten a small /guanodon, though whether this was the result of active predation or scavenging cannot be determined. \TERRESTRIALITY. If we accept that fish formed a significant part of the diet of Baryonyx, then we must consider the possibility that the Janimal led an aquatic or semi-aquatic existence. Nevertheless, its anatomy gives no indication of any modifications towards that mode of life. For example, it certainly had no flipper-like modifications of the limbs and it lacked the dorso-ventrally flattened skull with dorsally situated external nares typical of crocodiles. Indeed, the UF sition of the nostrils on the side of the skull would be disadvanta- ous to an animal spending much of its time in the water. However, a could probably swim, as can most land vertebrates, de- pte their lack of any special adaptations. STANCE AND GAIT. Despite our previous suggestions of ‘quadrupedality (Charig & Milner 1986), the anatomy of Baryonyx 7 ; affords no evidence of a gait any different from that of any other poe The length of the humerus of Stegosaurus, which is most ‘certainly a quadruped, varies between 43% and 55% of the length of ‘its femur; the corresponding ratio for Baryonyx is only 39% (based . n an unprejudiced estimate of the femoral length). On the other i] and, if Baryonyx really was a fish-eater, then it would have been pbliged to capture its aquatic prey from a crouching or quadrupedal dosition, either on the edge of the water or actually in it; and its "massive fore-limbs certainly possessed sufficient mechanical strength nd adequate musculature for the quadrupedal posture. he details of the fossilisation of the Baryonyx walkeri holotype _jsome of which have already been published in Charig & Milner, 986) are as follows: ANNER OF OCCURRENCE. Most of the bones were encased in jideritic siltstone nodules of irregular shape and were directly jurrounded by uneven accumulations of extremely fine sand and ilts; such accumulations are not found anywhere else. The rest of : : bones lay unprotected in clay. IEGREE OF ARTICULATION. d somewhat scattered. The bones were largely disarticulated 63 METRES - Skull - lower jaw = teeth - dorsal ribs - abdominal ribs pectoral girdle - pelvic girdle - cervical vertebrae - dorsal vertebrae - caudal vertebrae - hind limb Fig. 48 Plan of the excavation at Smokejack’s Brickworks, Ockley, May- June 1983, indicating the in situ positions of the numbered blocks and the distribution of skeletal elements. O —Q fe) = 4 & Z pr e 2 ) Q [o Oo 88 known; sagittal length approximately 60% of maximum width; axis with width about 70% of sagittal length; first three pleural furrows deeply incised; interpleural furrows nearly obscure; articulating half-ring set off from axis by prominent ring furrow; posterior axial rings and ring furrows difficult to discern; subdued but definite pygidial border developed around margin; posterior margin evenly arcuate; lateral and posterior aspects of cranidium defined as subvertical wall (PI. 4, fig. 7b), which is turned out again in a second, ventral, marginal rim; doublure relatively broad. DISCUSSION. We have not formally named this species because the two best cranidia (Pl. 4, figs 8, 11) were lost or broken following photography, and there are no replacement specimens suitable to serve as types. . The Tourmakeady species is obviously related to previously described species of Jsocolus, indicated particularly by the slot-like glabellar furrows that are greatly shallowed adaxial to the contact with the axial furrow. However, the laterally concave glabella of the Irish form is unique in the entire group. Other autapomorphic features include the apparently lacking dorsal sculpture, versus prominent raised lines in other species, the significantly broader posterior fixigena, and the posterolaterally directed, versus subtransverse, glabellar furrows. The ventral morphology of the /socolus pygidium has not previ- ously been described. The Irish material reveals the unexpected presence of a vertically oriented wall beneath what appears dorsally to be the pygidial margin. This ‘wall’ is flared ventrally into a second, ventral rim (PI. 4, fig. 7b). Reference to articulated material (Whittington 1956, 1963) indicates the larger, dorsally placed rim is almost certainly the true pygidial margin, and not analogous to the fulcral processes, rim, or spines often seen in groups like Entomaspidae Ulrich in Bridge, 1931 (e.g., Ludvigsen & Westrop in Ludvigsen ef al., 1989). The ventral wall and rim may therefore be doublural in origin. Family CATILLICEPHALIDAE Raymond, 1937 DISCUSSION. See remarks under discussion of Isocolidae above. Catillicephalid gen. et sp. nov. PI. 3, fig. 4; Pl. 4, figs 1-6 MATERIAL. Assigned specimens It. 25966, 25977-25982. DISCUSSION. The forward-expanding glabella, very short preglabellar field, tiny palpebral lobe set near to the axial furrow, anterior eye position, broad posterior fixigena, and distally elon- gated (exsag.) posterior cranidial border all indicate relationship of this unusual Irish species to Sunwaptan-Ibexian forms presently assigned to Catillicephalidae. The size and attitude of the posterior border as well as the position and inclination of S1, position and size of the palpebral lobe, and size of the cephalic border are further similarities to the stratigraphically nearest species, ie. Distazeris adoceta Ingham (in Ingham et al., 1986), from the Highland Border Complex of Scotland (Pratt 1992: 73 has criticized the generic assignment of this species). PLATE 5 J.M. ADRAIN AND R.A. FORTEY The Irish species possesses a feature that distinguishes it from all related taxa, and apparently from all other trilobites: the develop- ment at the junction of the axial, posterior border, and occipital furrows of an exoskeletal hood that encloses a cone-shaped space opening laterally. The homology of this structure is very difficult to determine. The posterior border furrow runs directly into it, and appears to terminate beneath the hood (PI. 4, fig. 2d). The occipital furrow meets the axial furrow on the adaxial side of the hood, and the junction of these furrows continues to circumscribe the hood posteriorly (PI. 4, fig. 2b). Family ILLAENIDAE Hawle & Corda, 1847 Genus ILLAENUS Dalman, 1827 TYPE SPECIES. Entomostracites crassicauda Wahlenberg, 1818, p. 27, from the Llandeilo of Fyacka, Dalarna, Sweden; by subsequent designation of Pictet (1854: 515). Illaenus weaveri Reed in Gardiner & Reynolds, 1909 Pl. 2, figs 7, 10-12; Pl. 3, figs 6-8, 10, 11; Pl. 5, figs 1-11; Pl. 6, figs 1-12 1909 Illaenus weaveri Reed in Gardiner & Reynolds: 142, pl. 6, figs 1-3. 1910 Illaenus weaveri Reed in Gardiner & Reynolds: 272. 1945 Illaenus weaveri Reed: 63. 1968 Illaenus weaveri Reed; Whittington: 56. 1988 Illaenus weaveri Reed; Morris: 115. DIAGNOSIS. Terrace lines on rear of cranidium, restricted to ante- rior part of librigenal field; librigenal flange only moderately developed; vincular furrow only impressed posteriorly on librigenal doublure; pygidium with sagittal length 55-60% of maximum width. MATERIAL. Lectotype, selected here, SM A10387, pygidium (PI. 2, fig. 7), original of Reed in Gardiner & Reynolds (1909, pl. 6, fig. 2); paralectotype SM A10316, cranidium and right librigena (PI. 2, fig. 10); topotypes It. 25959, 25960, 25967-25971, 25992-26014. DESCRIPTION. Due to the varying amounts and vectors of distor- tion of much of the available material, measured ratios are. approximate at best. The large pygidium of Pl. 6, fig. 7, is considered to be nearly undistorted, and pygidial ratios are based upon this specimen. Cranidium with length (sag., measured in sagittal profile) 75— 88% of maximum width across midlength (exsag.) of palpebral lobes; maximum anterior width about 90% of width across palpebral lobes; posterior sections of facial sutures declined nearly vertically when palpebral lobe is oriented in horizontal plane, diverging sharply posteriorly, nearly transverse; anterior sections of facial suture declined at about 30 degrees from horizontal when palpebral lobe is oriented in horizontal plane, slightly anteriorly divergent immedi- ately in front of palpebral lobe, then forming even, laterally convex arc to converge near anterior margin; palpebral lobes relatively large and elongate; axial furrow strongly effaced, impressed only posteriorly, behind midlength of palpebral lobe; entire cranidium Figs 1-11 //laenus weaveri Reed in Gardiner & Reynolds, 1909 1a-b, It. 25992, cranidium and left librigena, anterior and left lateral views, x10. 2a-b, It. 25993, cranidium and right librigena oblique and anterior views, x4.5. 3a-e, It. 25994, cranidium, dorsal, anterior, and left lateral views, x7.5. 4a-e, It. 25995, cranidium, dorsal, anterior, and right lateral views, x10. 5a-c, It. 25996, cranidium, dorsal, anterior, and left lateral views, x15. 6, It. 25998, rostral plate, ventral view, x10. 7a-b, It. 25997, rostral plate, ventral and dorsal views, x7.5. 8a-b, It. 25999, rostral plate, posterior and ventral views, x10. 9a-b, It. 26000, cranidium, dorsal and anterior views, x15. 10a-c, It. 26001, cranidium, dorsal, anterior, and left lateral views, x15. 11, It. 26002, cranidium, ventral view, x10. ORDOVICIAN TRILOBITES FROM THE TOURMAKEADY LIMESTONE 89 90 vaulted, maximum sagittal curvature achieved slightly posterior to slightly anterior of palpebral lobes; anterior border and border furrow not evident; prominent terrace lines running subparallel to anterior margin, finer and more closely spaced near margin, coarser and more widely spaced dorsally; fine subparallel, transverse terrace lines developed across rear of glabella, occipital region, and rear of palpebral lobes, originating with close spacing laterally, spacing greater sagittally, forming ellipsoid pattern (Pl. 5, fig. 3a); occipital and posterior border doublure very short (sag., exsag.). Librigena with prominent, closely spaced terrace lines on antero- lateral aspect; coarser lines matching those on cranidium only expressed on anterior part of field; posterolateral librigenal corner broadly lobate, ventrolateral margin bowed in, single prominent terrace line forming sharp posterolateral rim; field with moderate dorsal convexity, sculpture excluding anterior terraced lines smooth; posterior margin with strong posterior convexity; eye relatively large, exsagittal length slightly more than twice the width (tr.), doublure broad and robust; vincular furrow strong posteriorly be- neath posterolateral corner, becoming effaced anteriorly. Rostral plate subtrapezoidal; length (sag.) 40% of width; anterior margin with gentle, even anterior convexity; posterior margin nearly transverse, with slight sagittal posterior bulge; connective sutures obliquely inclined at about 45 degrees, laterally concave; ventral aspect with prominent, coarse, terraced lines, larger and more widely spaced (sag., exsag.) anteriorly; reentrant dorsal flange incompletely known, but robust. Hypostome with sagittal length about 80% of maximum width across anterior wings; anterior margin (hypostomal suture) with lobate ‘M’ shape; anterior wings broad and nearly spatulate; wings grading into middle body posteriorly, but separated anteriorly by trough-like furrow delineating anterior part of body; lateral border narrow, sharply defined and ridge-like; lateral border furrow narrow and deeply incised; middle furrow deep, deepest laterally, fully impressed medially, with strong posterior curvature; middle body moderately inflated, lacking sculpture; maculae small but promi- nent, set just behind middle furrow; lateral border furrow grading without interruption into posterior border furrow; lateral border grading without interruption into posterior border; posterior border with posterior convexity nearly identical to that of middle furrow. Thoracic segments poorly known; articulating half ring and ring furrows not discernible; axial furrow defined only as break in slope from pleura; axis with broad transverse convexity; prominent ful- crum on pleural lobe, 70-80% distance abaxially. Pygidium with length (sag.) 55-60% of maximum width; axis with anterior width just under 40% of pygidial width; anterior margin transversely straight between fulcra; fulcrum set at 75% of distance between sagittal plane and lateral margin; prominent, subtriangular articulating facet forming obliquely inclined, anterolaterally directed plane distal to fulcrum; posterior margin nearly semicircular in plan view, subelliptical in posterodorsal view; pygidium with sagittal profile nearly flat anteriorly, prominently vaulted posteriorly, set at nearly 90 degrees to anterior part posteriorly near margin; plane of margin declined about 10 degrees from that of PLATE 6 J.M. ADRAIN AND R.A. FORTEY anterior flat part of sagittal profile; axis only slightly raised from pleura, becoming increasingly less differentiated posteriorly; doublure broad, extended forward to rear of axis (about 60% of sagittal length from front of pygidium); doublure notched medially around termination of axis, protruding forward on either side of axis, then evenly arcuate distally. DIscuUSSION. //laenus weaveri belongs to Jaanusson’s (1957: 110) I. sarsi group, which includes the taxa/. consimilis Billings, 1865, I. fraternus Billings, 1865 (see Whittington 1965 for both), J. auriculatus Ross, 1967, and I. oscitatus Fortey, 1980 (see also Nielsen 1995). The group is characterized particularly by the form of the pygidial doublure, with its median notch flanked by anterior projections (e.g., Pl. 6, fig. 12), and all species show to greater or lesser extent the development of posterolateral ‘flanges’ on the librigenae. Illaenus weaveri is perhaps most similar to J. auriculatus, from the basal Whiterockian (Zone L) of the Antelope Valley Limestone, Pyramid Peak, California. However, the species differ in that /. weaveri has a relatively longer pygidium, much less pronounced medial pygidial doublural notching, a rostral plate that is much longer medially, and librigena with a less pronounced lateral flange. The vincular furrow of J. weaveri is restricted to the posterior part of the librigenal doublure (Pl. 6, figs 3b, 4b), whereas that of J. auriculatus is continued anteriorly (Ross 1967: pl. 5, fig. 29). Illaenus weaveri differs from I. oscitatus, from the Whiterockian of Spitsbergen, in the lack of that species’ prominently pitted sculpture and fully defined cranidial anterior border. Additionally, the cranidial sagittal profile of J. oscitatus is much more evenly convex than that of /. weaveri (compare Fortey 1980: pl. 10, figs 2, 6, with Pl. 5, figs 3c, 4c, 5c). Both J. consimilis and I. fraternus, from the Whiterockian of Newfoundland, are distinguished from /. weaveri in the possession of prominent terrace lines over the entirety of their dorsal surface, including medially on the cranidium, on the librigenal field, and on all of the pygidial axial and pleural region. The size and shape of the librigenal flange of J. fraternus, however, is similar to that of /. weaveri (e.g., Whittington 1965: pl. 45, fig. 17). Family CHETRURIDAE Hawle & Corda, 1847 Subfamily uncertain DISCUSSION. Cheirurid subfamilial classification is in a state of flux. Several subfamilies (including Cheirurinae, Acanthoparyphinae, Heliomerinae, and Deiphoninae) are undoubtedly monophyletic. Others (e.g., Eccoptochilinae, Sphaerexochinae, Cyrtometopinae, Areiinae) are more problematic, and their status as natural groups has yet to be convincingly established. Two of the genera dealt with herein (Kawina Barton, and Mayopyge gen. nov.) would be as- signed, by current convention, to the subfamily Sphaerexochinae. This taxon, however, is particularly problematic because Sphaerexochus itself is a highly autapomorphic genus, the sister Figs 1-12 I/laenus weaveri Reed in Gardiner & Reynolds, 1909 1, It. 26003, right librigena, external view, x7.5. 2, It. 26004, left librigena, external view, x5. 3a-b, It. 26005, right librigena, external and internal views, x10. 4a-b, It. 26006, left librigena, external and internal views, x7.5. 5, It. 26007, thoracic segment, dorsal view, xS. 6, It. 26008, thoracic segment, dorsal view, x5. 7a-c, It. 26009, pygidium, dorsal, posterior, and left lateral views, x3.5. 8, It. 26010, pygidium, dorsal view, x5. 9a-b, It. 26011, right librigena, ventrolateral and external views, x10. 10a-b, It. 26012, pygidium, dorsal and posterior views, x7.5. 1la-b, It. 26013, pygidium, dorsal and posterior views, x5. 12, It. 26014, pygidium, ventral view, x10. Figs 13-16 Kawina divergens (Reed, 1945). 13, It. 26015, left librigena, external view, x6.5. 14, It. 26016, hypostome, ventral view, x10. 15, It. 26017, cranidium, oblique view, x10. 16, It. 26018, cranidium, dorsal view, x10. ORDOVICIAN TRILOBITES FROM THE TOURMA 09 taxon of which is unknown. Until such time as we are able to base the subfamilial classification of such genera on explicit and well-sup- ported hypotheses of cladistic relationship, we prefer to regard their subfamilial affinities as uncertain. Genus KAWINA Barton, 1916 Cydonocephalus Whittington, 1963: 97. TYPE SPECIES. Cheirurus vulcanus Billings, 1865, from the Cow Head Group (lower Whiterockian), western Newfoundland; by origi- nal designation. DISCUSSION. Whittington (1963: 97) distinguished his new Cydonocephalus (type species C. griphus Whittington, 1963, lower Whiterockian, western Newfoundland) from Kawina Barton, 1916, on the assertion that the ‘glabella is most convex anteriorly (not posteromedially) and juts forward, lobe 1p, and in some species 2p and 3p, are gently inflated, and occipital furrow curves forward (not back) medially.’ Of these features, only the glabellar convexity holds for all six of the western Newfoundland species assigned to Cydonocephalus. The glabellar lobes of large specimens of C. torulus Whittington, 1963, are not markedly more inflated than those of Kawina vulcanus (Billings, 1865) (compare Whittington 1963: pl. 28, figs 5, 16), and the occipital furrows of both C. torulus and C. griphus both clearly curve backward (Whittington 1963: pl. 27, figs 3, 10, 13, 16; pl. 28, figs 1, 5; pl. 29, figs 1, 4). All of the pygidia (Whittington 1963: pl. 31) which likely belong to species assigned by Whittington to Cydonocephalus have their pleural ribs fused along almost their entire length. Kawina arnoldi Whittington, 1963, however, has ribs with distally free tips (Whittington 1963: pl. 26, fig. 14). This is similar to the Irish species (Pl. 7, fig. 2), the pygidium of which differs from that of K. arnoldi only in proportions. However, the sagittal profile of the Irish cranidia is obviously like that of Cydonocephalus, with the point of max1- mum convexity anterior, not posterior. Kawina arnoldi, however, lacks the strong posterior convexity of the type species, and has a nearly evenly arcuate sagittal cranidial profile (Whittington 1963: pl. 26, fig. 8). Considering all the species it is not possible to specify synapomorphic characters that would distinguish the two genera as separate monophyletic groups. Rather, species presently assigned to one or the other show overlapping variation in characters considered by Whittington to be diagnostic of Cydonocephalus, as well as in pygidial morphology. For these reasons, Cydonocephalus is placed in subjective junior synonymy of Kawina herein. Kawina divergens (Reed, 1945) Pl. 6, figs 13-16; Pl. 7, figs 1-5, 7 1909 Pliomera aff. fischeri (Eichwald); Reed in Gardiner & Reynolds: 144; pl. 6, fig. 4. 1925 Kawina sp., Raymond: 144. 1945 Kawina divergens Reed: 59. 1971 Kawina? divergens Reed; Lane: 56; text-fig. 9a. 1988 Kawina? divergens Reed; Morris: 119. DIAGNOSIS. Dorsal sculpture of very fine, densely spaced gran- ules; short, thorn-like genal spine retained in large holaspides; glabella with nearly even sagittal convexity, point of maximum convexity anterior; pygidium wide, with splayed, subquadrate ribs bearing free tips. J.M. ADRAIN AND R.A. FORTEY HOLOTYPE. Pygidium, SM A10396 (PI. 7, fig. 2); topotypes It. 26015-26022, 26200. DESCRIPTION. Cranidium with sagittal length 70-75% of maxi- mum width across posterior border; glabella with width across midlength (exsag.) of L2 subequal to or slightly narrower than width across midlength of L1; maximum glabellar width subequal to length (measured in sagittal profile) excluding LO; anterior border short but complete medially; preglabellar furrow deeply incised, grading abaxially into axial furrow of similar depth; anterior fixigena — a narrow, laterally convex strip, widest opposite midlength (exsag.) of L3, narrowing posteriorly in front of palpebral lobe; palpebral lobe narrow, entirely set off from interocular fixigena by very sharply incised palpebral furrow; interocular fixigena broadening posteriorly, smooth, with moderate dorsal convexity; interocular fixigena grading smoothly into broad posterior fixigena, held in plane declined about 60 degrees from horizontal; anterior sections of facial sutures short (exsag.), parentheses-shaped, with strong ante- rior convergence from midlength of L3 to front of glabella; glabella strongly inflated, sagittal profile with nearly even dorsal convexity, slightly more pronounced anteriorly; S1 nearly transverse distally, curved in nearly subcircular arc proximally, similar in depth to axial furrow but shallowing abruptly in front of SO so that L1 is not quite fully isolated; L1 with only slight independent inflation, length (exsag.) subequal to width (tr.); S2 similar distally to S1, proximal part shorter and less posteriorly inclined; L2 with no independent inflation, length (exsag.) 80-85% of that of L1; S3 as deeply incised as S| and S2, but not reaching as far adaxially, with less posterior curvature, and shallowed slightly near contact with axial furrow; L3 with slight anterolateral inflation, length about 80% of that of L2; frontal glabellar lobe with slight lateral inflation immediately ante- rior to S3, anterior margin with blunt anterior convexity; SO composed of three distinct posteriorly bowed regions, two lateral ones behind LI, and a medial one between the L1 lobes, depth similar to axial furrow, medial region slightly longer (sag., exsag.) than lateral parts (exsag.); LO similar in length to distal parts of posterior border, with nearly flat top in sagittal profile; posterior border sharply incised, very short (exsag.), and running nearly transversely to genal angle, where it is bowed anteriorly; posterior border very short proximally, but lengthening greatly distal to fulcrum to form lobate genal angle; short, thornlike, subtriangular genal spine retained on even largest specimens; entire dorsal cranidial surface with very fine, unimodal granular sculpture. Librigena poorly known; field and lateral border both with fine granular sculpture similar to that of cranidium; lateral border well defined posteriorly, but grading into field anteriorly; lateral border furrow with concomitant shallowing anteriorly; field apparently quite narrow (tr.); eye unknown. Rostral plate unknown. Hypostome with sagittal length 75% of maximum width across anterior wings; width across shoulders 80% of width across anterior wings; width across posterolateral corners two thirds that across anterior wings; anterior margin (hypostomal suture) with strong anterior convexity; sharp marginal rim and furrow developed laterally, middle body extended to margin medi- ally; strong antennular notch between shoulder and anterior wing; lateral border broad and inflated, with fine granular sculpture; lateral border furrow shallow and broad (tr.); posterior margin with only slight posterior convexity; posterior border and posterior border furrow similar in dimensions to lateral border and furrow; middle body with sagittal length slightly greater than maximum width, moderate ventral inflation, strongest anteriorly; sculpture of fine granules somewhat more subdued than that of lateral border; middle furrow impressed only laterally, strongly declined posteriorly. ORDOVICIAN TRILOBITES FROM THE TOURMAKEADY LIMESTONE # is PLATE 7 Figs 1-5,7 Kawina divergens (Reed, 1945). la-e, It. 26019, cranidium, dorsal, anterior, and right lateral views, x5, x4, x4. 2, SM A10396, pygidium, holotype, dorsal view, x4. 3a-b, It. 26020, cranidium, dorsal and right lateral views, x4. 4a-b, It. 26021, cranidium, oblique and dorsal views, x4. 5a-b, It. 26022, cranidium, dorsal and anterior views, x4. 7a-c, It. 26200, cranidium, dorsal, right lateral, and oblique views, x7.5. Figs 6a-b Mayopyge zapata gen. et sp. nov. It. 26023, cranidium, dorsal and left dorsolateral views, x15. J.M. ADRAIN AND R.A. FORTEY o's aw ae’ ,epe ‘ igs ee es n “i ae *? —- ay > i a ORDOVICIAN TRILOBITES FROM THE TOURMAKEADY LIMESTONE Thorax unknown. Pygidium (see also Lane 1971: 56) with sagittal distance from articulating half ring furrow to rear of medial spines 44% of maximum width; pygidium composed of three distinct rings, pleural segments, and a small subtriangular terminal piece; interpleural furrows deeply incised, but first and second ribs fused to just over half distance abaxially, second and third ribs fused to about two thirds distance abaxially; pleural furrow shallow but incised on proximal part of first rib, very faint furrow visible on corresponding part of second rib; axial furrow moderately deep opposite first ring, progressively shallower posterior, but deepened around small termi- nal piece; first axial ring with significant sagittal convexity and bowed posteriorly in plan view; second ring less convex and more transverse; third ring not inflated and nearly transversely oriented; ring furrows shallowed medially; ribs with spatulate, subquadrate free tips. DISCUSSION. Kawina divergens has previously been known only from its holotype pygidium (PI. 7, fig. 2). The species is distin- guished from K. scrobiculus (Whittington, 1963) and K. prolificus (Billings, 1865) in its less inflated lateral glabellar lobes and posteriorly versus anteriorly bowed occipital furrow. It differs from K. mercurius (Billings, 1865) in the lack of the autapomorphic tranversely impressed S1 of that species. Kawina divergens is quite similar to both K. griphus and K. torulus (see generic discussion above), but differs from both in the presence of a much finer cephalic sculpture and an S1 that is much better impressed proximally to more fully isolate L1. The closest comparison among described species is with K. arnoldi. Pygidia of the two species are discussed above. Shared cephalic features include a similar sagittal convexity, similarly subdued dorsal sculpture (although that of K. arnoldi is slightly more robust), and similar posterior curvature of SO. The species differ particularly in the presence in K. arnoldi of an S1 that is strongly sigmoidal in lateral view. Genus MAYOPYGE gen. nov. TYPE SPECIES. Mayopyge zapata sp. nov., from the Tourmakeady Limestone, Co. Mayo, western Ireland. OTHER SPECIES. ?Pseudosphaerexochus tuberculatus Warburg, 1925, Leptaena Limestones, Ashgill, Dalarne, Sweden. and the Chair of Kildare Limestone, Ashgill, eastern Ireland (Dean 1971). ETYMOLOGY. After Co. Mayo, in which the type locality is situ- ated, and the Greek noun pyge, tail. DIAGNOSIS. Prominent eye ridge and relatively wide interocular fixigena; strong sutural ridge along anterior section of facial suture of librigena; swollen knob-like structure proximally on thoracic pleura; pygidium with three segments, anterior of which is similar to posterior thoracic segment; shallow, ‘v’-shaped anterior pygidial doublural margin, with arcuate posterior embayment; densely and coarsely tuberculate dorsal sculpture. DISCUSSION. The subfamilial affinities of Mayopyge gen. nov. are exceptionally difficult to judge. The densely tuberculate dorsal 95 exoskeleton is most similar to that developed in many acanthoparyphine clades. The prominent peak in convexity at the rear of the glabella, seen in many specimens, has obvious compari- sons with the hypertrophied structure present in the same topological position in species of the primitive acanthoparyphine Nieskowskia Schmidt, 1881. Most species of Nieskowskia also have prominent tuberculate sculpture. However, Mayopyge lacks any of the acanthoparyphine apomorphies. Most importantly, M. zapata dis- plays the primitive three-segmented pygidial condition, in contrast to the reduction to two segments characteristic of Acanthoparyphinae. In addition, the maximum glabellar width in the Irish species is achieved across L2, rather than across L1 as in acanthoparyphines. Mayopyge does show some similarities to early species of Sphaerexochus. The hypostome of M. zapata, for example (Pl. 9, figs 17-20), is nearly identical to that of the upper Whiterockian S. arenosus (Chatterton & Ludvigsen, 1976, pl. 13, figs 32, 33, 37, 41, 42). Mayopyge also shares with Sphaerexochus fully isolated L1 with strong independent inflation. In addition to several plesiomorphic features, including its rela- tively elongate anterior border and very prominent eye ridge, Mayopyge zapata displays several seemingly autapomorphic morphologies. The strong inflation of L2 and its near isolation from the median glabellar lobe in many specimens (PI. 8, figs 1b, 2c, 9b) is not seen in any other species with a strongly inflated glabella. The thoracic pleural structure is also apparently unique. In contrast to the transverse furrow or row of pits common to most non-cheirurine cheirurids, Mayopyge shows only a faint, obliquely inclined furrow (Pl. 10, figs la, 2a, 4a), with the anterior pleural band swollen into a hemispherical knob and the posterior band greatly reduced. The structures seem analogous to those present in Cheirurinae, but in that taxon both the anterior and posterior pleural bands are swollen and the pleural furrow, though obliquely inclined, runs in a direction opposite to that seen in Mayopyge. In cheiruines, the pleural furrow contacts the axial furrow anteriorly, and runs posterolaterally. In Mayopyge, the contact is posterior and the furrow runs anterolaterally. In summary, it does not seem possible at present to relate Mayopyge with confidence to other cheirurids. Potentially synapomorphic comparisons can be made with acanthoparyphines and with Sphaerexochus, but additional relevant diversity will probably be necessary to resolve the systematic position of the genus. Two cranidia from the Ashgill Chair of Kildare limestone, eastern Ireland, figured by Dean (1971: pl. 11, figs 1-3, 9, 10) as Pseudosphaerexochus? tuberculatus Warburg, 1925, have coarse, dense, bimodal tuberculate sculpture. The species also agrees with Mayopyge zapata in its unusually well-impressed S2 and S3. Swed- ish type material of P. tuberculatus has never been photographically illustrated, but Warburg’s (1925: pl. 10) cranidia are densely tuber- culate. Pseudosphaerexochus tuberculatus possibly represents a species of Mayopyge, to which it is assigned with reservation herein, but it could equally prove to be an acanthoparyphine and confirma- tion will require more complete material. Mayopyge zapata sp. nov. Pl. 7, fig. 6; Pl. 8, figs 1-10; Pl. 9, figs 1-21; Pl. 10, figs 1-17 PLATE 8 Figs 1-10 Mayopyge zapata gen. et sp. nov. la-d, It. 26024, cranidium, dorsal, right lateral, oblique, and anterior views, x7.5. 2a-e, It. 26025, cranidium, dorsal, ventral, right lateral, anterior, and posterodorsal views, x10. 3a-b, It. 26026, cranidium, left lateral and dorsal views, x10. 4a-b, It. 26027, cranidium, left lateral and dorsal views, x10. 5, It. 26028, cranidium, oblique view, x7.5. 6a-b, It. 26029, cranidium, dorsal and right lateral views, x10. 7Ta-b, It. 26030, cranidium, dorsal and right lateral views, x10. 8a-c, It. 26031, cranidium, dorsal, left lateral, and anterior views, x10. 9a-b, It. 26032, cranidium, dorsal and oblique views, x7.5. 10a-c, It. 26033, cranidium, dorsal, left lateral, and anterior views, x10. J.M. ADRAIN AND R.A. FORTEY A an ORDOVICIAN TRILOBITES FROM THE TOURMAKEADY LIMESTONE ETYMOLOGY. The pygidial spines droop in the style of a mous- tache. DIAGNOSIS. Occipital spine absent; L1 swollen and fully circum- scribed by deep S1; S2 deep and L2 with prominent inflation; librigenal field with large, coarse pitting; first pair of pygidial spines longest. HOLOTYPE. Pygidium It. 26059 (Pl. 10, fig. 6); paratypes It. 26023-26058, 26060-26071. DESCRIPTION. Specimens of this taxon are so subject to varying amounts and vectors of distortion that relative cranidial dimensions are not meaningful. Anterior border relatively long (for subfamily), sculpture of numerous densely distributed fine tubercles; anterior margin with median part shaped like shallow inverted ‘v’ in plan view, with second distinct change in course in front of junction of axial and preglabellar furrow; anterior sections of facial sutures nearly straight, with considerable anterior convergence; axial fur- rows subparallel from SO to opposite S2, anteriorly convergent in front of L2 to grade smoothly into preglabellar furrow; axial and preglabellar furrows deep; small trapezoidal frontal area with sculp- ture similar to anterior border; eye ridge prominent, running obliquely from opposite L3 to front of palpebral lobe, subparallel with lateral margin, defined by furrows, anterior of which is deepest, seen best ventrally (Pl. 8, fig. 2b); palpebral lobe very narrow, posteriorly contiguous with small sutural ridge; posterior furrow of eye ridge grading into incised palpebral furrow, continued posteriorly along- side posterior sutural ridge; interocular fixigena broad (tr.) for subfamily, with tuberculate sculpture somewhat coarser than that of frontal area; posterior fixigena with significant area and lateral development, subtriangular, sculpture of moderate sized tubercles; posterior border furrow similar in depth to axial furrow, of same length (exsag.) adaxially and abaxially; posterior border with strong dorsal convexity, adaxial length similar to adjoining LO, becoming longer abaxially, sculpture of fine, evenly spaced tubercles similar to that of anterior border; prominent tubular spine running posterolaterally from fulcrum of posterior border; glabella strongly inflated, prominent sculpture of fine to coarse tubercles, coarse, _ closely spaced tubercles becoming predominant posteriorly; sagittal convexity of most specimens showing strong peak posteriorly in cone-shaped dorsal projection in front of LO; L1 fully isolated by deep S1; L2 slightly smaller than L1; S2 well impressed, but shallower than $1; smooth band present on many specimens around anterior edge of S1 and S2; L3 strongly defined; S3 short (exsag.) and with reduced transverse course, but still quite strongly incised; SO deep, of even length sagittally and exsagittally; LO quite short (sag., exsag.), dorsally convex (sag., tr.), and with dense tuberculate sculpture similar to rear of pre-occipital glabella; tiny fossula present in axial furrow opposite midlength of eye ridge (PI. 9, fig. 5b). Librigenal lateral margin with strong, even lateral convexity; lateral border about 40% of width (tr.) of librigena, broadly inflated, with sculpture of dense tubercles, finer anteriorly and laterally, becoming coarse posteriorly along lateral border furrow; lateral 97 border furrow broad and deep, shallowing abruptly both posteriorly and anteriorly, terminated anteriorly by very pronounced sutural ridge along anterior section of facial suture; field with large, irregu- larly distributed pits and mixture of large, coarse tubercles and greater number of fine tubercles; single exsagittal row of very fine tubercles beneath eye; eye small (Pl. 9, figs 14, 16); doublure slightly narrower than lateral border, essentially flat and smooth, narrowing slightly posteriorly. Rostral plate unknown. Hypostome with sagittal length about 55% of maximum width (excluding anterior wings) across shoul- ders; moderately strong sutural ridge along hypostomal suture; anterior margin anteriorly convex with more abrupt change in slope sagittally, bowed anteriorly around anterior wings; middle body separated from sutural ridge by very short (sag., exsag.) furrow, sagittal length about 60% of maximum width, sculpture of very fine tubercles, slightly coarser anteromedially; middle furrow deep later- ally, running posteromedially, in some specimens shallowing but meeting sagittally to fully circumscribe anterior and posterior lobes; lateral border broad, with tuberculate sculpture slightly coarser than that of middle body; anterior wing tab-shaped, set at slightly oblique (30-45 degrees) angle; shoulder small but sharply protruded; lateral and posterior border furrows broad and very shallow; posterior border long (sag., exsag.), sculpture smooth, posterolateral corners lobate, embayed sagittally; doublure forming sharp dorsal fold and ridge around shoulder, lacking sculpture. Thoracic segments with large articulating half-ring, set off posteriorly by sharp break in slope; axial ring longer (exsag.) laterally, shortened sagittally, sculpture of dense, fine to moderate sized tubercles; axial ring separated from articulating half-ring by broad, long preannular lobe, always developed as a depressed area, never with independent inflation; axial furrow very shallow, ring nearly contiguous with pleura; pleura proximal to fulcrum com- posed of subrectangular base topped by semicylindrical rib; rib with prominent knob-like swelling on anteroproximal part, with shallow, oblique furrow set posterior to swelling (Pl. 10, fig. 4a); pleura distal to fulcrum composed of free, tubular spine, lengthening and more posteriorly inclined on more posterior segments, with dorsal sculp- ture of moderately sized tubercles. Pygidium composed of three segments, anterior segment with morphology essentially identical to that of posterior thoracic seg- ment; articulating half-ring separated from first axial ring by narrow furrow, preannular lobe absent; axial furrows subparallel and deep on first segment, very shallow on second, entirely effaced on third; first axial ring with robust tuberculate sculpture; second ring very short (sag., exsag.), with four or five tubercles of same size as those on first ring; ring furrow between first and second rings broad and deep, forming pit laterally; second ring furrow slot-like (Pl. 10, figs 11a, 14a), in many specimens reduced to lateral pits (Pl. 10, figs 7, 9); third segment expressed as pair of median spines united by small, tuberculate terminal piece; first spine pair elongate, posteriorly recurved; second and third pairs progressively shorter; all spines with dorsal and dorsolateral tuberculate sculpture similar in size and | PLATE 9 | Figs 1-21 Mayopyge zapata gen. et sp. nov. 1, It. 26034, cranidium, dorsal view, x6. 2, It. 26035, cranidial fragment, dorsal view, x10. 3a-b, It. 26036, cranidium, dorsal and left lateral views, x7.5. 4, It. 26037, cranidium, dorsal view, x7.5. 5a-e, It. 26038, cranidium, dorsal, ventral, and anterior views, x10. 6a-c, It. 26039, cranidium, left lateral, dorsal, and anterior views, x10. 7a-b, It. 26040, cranidium, dorsal and left lateral views, x7.5. 8, It. 26041, cranidium, dorsal view, x7.5. 9a-b, It. 26042, right librigena, external and internal views, x10. 10, It. 26043, left librigena, external view, x7.5. 11, It. 26044, right librigena, external view, x7.5. 12, It. 26045, right librigena, external view, x10. 13, It. 26046, left librigena, external view, x7.5. 14, It. 26047, left librigena, external view, x10. 15, It. 26048, right librigena, external view, x7.5. 16, It. 26052, right librigena, external view, x7.5. 17a-b, It. 26049, hypostome, ventral and dorsal views, x7.5. 18, It. 26050, hypostome, ventral view, x10. 19, It. 26051, hypostome, ventral view, x7.5. 20, It. 26053, hypostome, ventral view, x7.5. 21a-b, It. 26054, cranidium, oblique and posterior views, 7.5. J.M. ADRAIN AND R.A. FORTEY ORDOVICIAN TRILOBITES FROM THE TOURMAKEADY LIMESTONE density to that of first axial ring; doublure forming narrow, convex ventral rim, with fine tuberculate sculpture; anterior doublural mar- gin ‘v’-shaped, with median posterior embayment. DISCUSSION. This species is the second most common in our collections and, like other common taxa, displays several morphotypes due to tectonic distortion. In Mayopyge zapata the distortion seems to be nearly bimodal. Overwhelmingly common among the cranidia is the type illustrated on Pl. 8, which displays a conical dorsal swelling of the glabella in front of the occipital ring. Whether this is a biological structure or a the result of distortion and enhancement of an original convexity peak is not known. The structure is so pervasive that it seems likely that at least some original inflation was present. The second cranidial type is that illustrated on Pl. 9, figs 1, 3a, 4, 5a, in which this projection is entirely absent and the glabella roundly and evenly inflated. An undistorted calcareous crackout cranidium (PI. 7, fig. 6) shows an inflated convexity peak at the rear of the glabella. It is conceivable that different vectors and amounts of distortion could produce either of the silicified morphotypes from such an original morphology. Pygidial types do not show as strong a disjunct occurrence as do the cranidia, but are even more morphologically discrete. The slightly more common form (PI. 10, figs 6-10, 12, 13, 17) includes the holotype. It is relatively long versus wide (sagittal length 45— 50% of anterior width), has long, posteriorly directed spines, a second ring furrow that is typically reduced to two lateral pits, anda rather sharp doublural embayment (PI. 10, figs 6b, 8b). The second, slightly less common, form (PI. 10, figs 11, 14-16) is much shorter (sagittal length 27—33% of anterior width), has more laterally splayed spines, with the median pairs apparently shorter, a second ring furrow that is a medially continuous slot (Pl. 10, figs 11a, 14a, 15a), and a relatively shallow doublural embayment (PI. 10, figs 11b, 15b). Given the apparent presence of two cranidial and pygidial morphotypes, there are three possibilities. First, the variation may be genuine and reflect sexual dimorphism. Second, the variation may be genuine and reflect the presence of two closely related species. And third, the dimorphism may be artefactual, and a result of the tectonic distortion affecting the entire fauna. Sexual dimorphism is improbable; it remains unproven in the trilobites as a whole (Adrain & Kloc 1997; Hughes & Fortey 1996; Ramsk6ld & Chatterton 1991; Ramsk6old & Werdelin 1991). If sexual dimorphism were the case it might be expected that other acathoparyphines would also exhibit it, which species known from abundant silicified material manifestly do not. It cannot be entirely disproved that there are two, closely related species: since cheirurid pygidia are distinctive the name is attached to the best specimen of the commonest morph. Overall, we consider that bimodal tectonic distortion is responsible for the Variation, since ‘long’ and ‘short’ forms have also been recognised in | Illaenus weaveri and Celmus michaelmus, where the cause is de- | monstrably tectonic. However, the pygidial differences remain a cause for concern, as the different ring furrows are not readily accounted for by distortion alone. PLATE 10 99 Subfamily CHETRURINAE Hawle & Corda, 1847 Genus CERAURINELLA Cooper, 1953 TYPE SPECIES. Ceraurinella typa Cooper, 1953, from the Edinburg Formation (Mohawkian), Virginia, U.S.A.; by original designation. Ceraurinella sp. Pl. 11, figs 1-9 MATERIAL. Assigned specimens It. 26072—26080. DISCUSSION. The fragmentary nature of the available material does not allow a determination of the species, and several important morphological differentia (e.g., eye position) are not fully pre- served. Nevertheless, the Tourmakeady species is clearly a primitive member of the Ceraurinella group (including Sycophantia Fortey, 1980). The Irish species differs from the Spitsbergen plesiomorph S. seminosa Fortey, 1980, in its much shorter anterior cranidial border (in which it resembles later and presumably more advanced species) and in the complete loss of the small median pygidial spine retained in S. seminosa. Both the available librigenae and fixigenal frag- ments, however, indicate that the Irish species retained very wide posterior fixigenae, similar to S. seminosa. Of other early species, the Tourmakeady taxon resembles C. polydorus (Billings, 1865) (see Whittington 1965: pl. 60) in the length of the anterior cranidial border, but differs in possessing less inflated lateral glabellar lobes (compare Pl. 11, fig. 1 with Whittington 1965: pl. 60, fig. 4), a broader posterior fixigena, and a narrower pygidium lacking an independently defined median spine. Family ENCRINURIDAE Angelin, 1854 Subfamily CYBELINAE Holliday, 1942 Cybeline indet. (not figured) MATERIAL. Assigned specimen It. 26201. DISCUSSION. A single very poorly preserved cybeline cranidium has been recovered from the silicified residues. It is obscured by silicified debris, and identifiable only to subfamily level. It is noted here for completeness. Family LECANOPYGIDAE Lochman, 1953 Genus BENTHAMASPIS Poulsen, 1946 TYPE SPECIES. Benthamaspis problematica Poulsen, 1946, Ibexian, Ellesmere Island, Canadian Arctic; by monotypy. Benthamaspis aff. B. diminutiva Hintze, 1953 Pl. 11, figs 17-19, 21-23 MATERIAL. Assigned specimens It. 26089-26094. Figs 1-17 Mayopyge zapata gen. et sp. nov. 1a-b, It. 26055, thoracic segment, dorsal and anterior views, x7.5. 2a-b, It. 26056, thoracic segment, dorsal and anterior views, x7.5. 3a-b, It. 26057, thoracic segment, dorsal and anterior views, x7.5. 4a-b, It. 26058, thoracic segment, dorsal and anterior views, x7.5. 5a-b, It. 26060, thoracic segment, oblique and dorsal views, x7.5. 6a-c, It. 26059, holotype, pygidium, right lateral, dorsal, and ventral views, x7.5. 7, It. 26061, pygidium, dorsal view, x10. 8a-b, It. 26064, pygidium, dorsal and ventral views, x7.5. 9, It. 26062, pygidium, dorsal view, x7.5. 10, It. 26063, pygidium, dorsal view, x10. 11a-b, It. 26066, pygidium, dorsal and ventral views, x7.5. 12, It. 26065, pygidium, dorsal view, x10. 13, It. 26067, pygidium, dorsal view, x10. 14a-b, It. 26068, pygidium, dorsal and posterior views, x7.5. 15a-b, It. 26069, pygidium, dorsal and ventral views, x7.5. 16, It. 26070, pygidium, dorsal view, x7.5. 17, It. 26071, pygidium, dorsal view, x10. 4 [o4 ie) x < jaa ( ZZ < 4 < low QA piodynayuopy Z€0'°0-10'0 10-F0'0 660-610 670-900 6I-%I 10-00 I10--00 610-10 £€0-€10 $60-860 b-£ XMIDNUIS = = (y) Z0'0 = = (Loo (r)seo (hr) Ero (SI (bh) 900 () 60 Elo )8I0 ()r0 () E¢ pdt yopiuq 06 91-01 900-100 10-200 8L0-E1'0 [10-200 ZI-Ol 10-10 €10-80'0 EZ0-11'0 670-€1'0 Pll Sb aapjod ‘J9 (1) 0€ = (1) (1) 810°0 = = () soo (1) 8€0 (1)900 (Lor @ito @Wdto WMos9t0 WMt0 Writ 1) sr Dwojsojvg LL-OS 91-01 90'0-F10°0 €1'0-0:0 L9'0-1'0 9+'0-Z0'0 OZ-O01 €10-20'0 LI0-+00 €EZ0-110 +E 0-1'0 €b'I-8h'0 ord asuauvsf{3oj9 ()EL9 (97 (MD EOO 3 = (cD LOO0 (ED87ZO (STO 6) 9EI (D800 (ID60 (ESTO (EDEIo (2z)80 (EZ) GE DUOjsoIDg €10-900 I10-r00 LS0-10 170-900 ZI-6 00-200 €10-F00 LZ0-LI'0 +€0-£70 18'T Z1-01 pupisinquasam = - - ~ @soo0 (1400 @WMo9z0 (110 @MZ0l WMe0 Wio Mito Muto (ist Ot ‘ye piodoyvy 11'0-80'0 80°0-S0'0 61'0-L0'0 10-600 £0970 9S siupynzad = - - - (10 (jdL00 - (9) LV'0 - (r) 10 - - (S) 6v70 - (1) 9°¢ piodoyvy LI-01 6S 710-S0'0 170-800 S1'0-90°0 SZ 10400 T1000 60-0 Ss0-760 877 ZI (€—) $7 (€) LL (€) 60°0 - (%) €10 =) 110 = = (ive (€)900 () blo (2) LEO (€) sro (1) 8z7z (1) ZI ‘ds { pwajo01aT7 ££-97 Z-1 ZE0-EZ0'0 90'0-Z0'0 SS-6E 10--00 900-200 I1'0-800 €1'0-80:0 L9'0 cI (1) $6z@ =) ZI (1) Lz0°0 = = (1) 600 = = (D) Lp (1) 800 (1) 00 §=(1) €60°0 Ivo @ L490 (1) ST q ‘ds pwaj01a7 be-SZ 9% —_¥0'0-20'0 610-1110 10-900 69 LIO-10 I10-900 €70-E10 ¥€0-170 7-81 8 (ORG (CO MIES (2) £00 = @ sto @WtT10 = - Mrs Mero WMeoo Wsio W970 Wel @ 8 V ‘ds pwajs01aT 650°0-St0'0 610-110 11'0-90'0 6 800-00 SI0-1'0 €Z'0-€1'0 Ls0 v - - (1) S00 = tl0 = (1) 600 $ = (1) 6 (1) sso0 Z10 = (1) 610 (1) LS0 (I) > ‘ds pd(yosajayy L8'0-91':0 6¥'0-67'0 cr Z0°0 9F'0-ZE'0 LS0-8E0 Se - - ~ - - - (1) 70 (1)9E0 (Ebr (1 700 - (1) 6€0 (1) Lr0 = @ sz ‘ds pwspsydiuay EZ S-p LLO0-Sr0'0 I10-r0'0 LZ0-11'0 £7'0-80'0 SI-OI 10-200 €1'0-900 SZO-SIO 6Z70-LI'0 EE I-FI'l s+ wnapusdd (1) €Z (@ L9b (2) 1s0'°0 - - 800 (Lzt0 @Msto MLA WML0 Wito W600 WEZO MrzI Wsr puspiydiwapzy €Z-b1 L-1 ¥S0'0-€20'0 LV'0 00 670-900 8-9 €1'0-200 I70-F0'0 620-E1'0 ZE0-LI'0 7Z9'I-LS°0 OI SISUALI]IUDS (st (€)tr Wtr00 = = (6) €80°0 = (6) elo (4) sz78 (8)seo0 )3=6©9) 110 02) 70 SCO 970 SC TI SCO 8's psodoniq WNV ZV dv WAND NAHWNdC XN Nad xXxdd WAZ LMZ GWXW CUZNW dCZXN MXd MOZ sarsadg ORDOVICIAN BRYOZOA FROM THE LLANDEILO LIMESTONE Maesyrwyn vs 500 metres rT Pantygwenin exploratory * trench Ir. 16h SLY Clog-y-fran Fig. 1 Locality map showing position of exploratory trench where the limestone blocks containing the fauna were collected. SYSTEMATIC PALAEONTOLOGY The terminology in all descriptions is that of Boardman et al. (1983). All genera are placed in families based on the following sources: Trepostomata —Astrova (1978); Cystoporata—Utgaard (in Boardman et al., 1983). Family level classification is generally unsatisfactory in Palaeozoic trepostomes and is currently being revised for the Treatise on Invertebrate Paleontology by R.S. Boardman. Biometric details for all trepostome species are tabulated (Table 1). Each measurement was made up to seven times per specimen, and the means and ranges calculated for each parameter. Raw data can be found in an unpublished Ph.D. thesis (Buttler 1988). All specimens are represented by thin sections or acetate peels. Meas- urements given in the systematic descriptions are all mean values unless otherwise stated. All material is deposited in The Natural History Museum, London. Phylum BRYOZOA Ehrenberg, 1831 Class STENOLOAEMATSA Borg, 1926 Order TREPOSTOMATA Ulrich, 1882 Suborder HALLOPOROIDEA Astrova, 1965 Family HETEROTRYPIDAE Ulrich, 1890 Genus DITTOPORA Dybowski, 1877 Dittopora sanclerensis sp. nov. Figs 2-3 HOLOTYPE. NHM PD 8338. PARATYPES. NHM PD 8333-8337, 8339-8341. NAME. The species is named after St. Clears (Sancler in Welsh), the nearest town to the type locality. DIAGNOSIS. Colony large, ramose. Autozooecia, with very thin, slightly wavy walls in endozone, which curve out from branch axis to intersect zoarial surface; polygonal in zoarial transverse section, rounded-circular in shallow zoarial tangential sections. Circular 119 mesozooecia present, originating in outer endozone/inner exozone. Partial and complete diaphragms in exozonal autozooecia; numer- ous diaphragms in mesozooecia. Acanthostyles large and abundant in exozone. DESCRIPTION. Zoaria erect with thick cylindrical branches, on average 5.8 mm in diameter. Autozooecia curve out gently from the branch axis in the endozone to meet the zoarial surface at 90°. Autozooecia within the endozone have very thin, slightly wavy walls. The exozone has an average diameter of 1.1 mm (although the range is large: 0.57—1.62 mm) and is recognised by a thickening of the zooecial walls. Autozooecia all originate in the endozone where they are polygonal in transverse section, becoming rounded-circu- lar in the exozone, as seen in tangential sections of branches. Autozooecial in the exozone contain abundant partial diaphragms and complete diaphragms (spaced on average 0.13 mm apart) which increase in thickness distally along the autozooecia. All diaphragms are basal and are deflected orally at their junctions with zooecial walls. Their laminae are continuous with the autozooecial linings. Mesozooecia are common and originate in the outer parts of the endozone and inner parts of the exozone. They are circular in shallow tangential sections and contain numerous thick, orally deflected basal diaphragms, spaced on average 0.83 mm apart and generally increasing in thickness distally along the mesozooecia. Acanthostyles are abundant and large, with an average diameter of 0.04 mm and density of four per mm?*. They originate deep within the exozone and can on rare occasions indent the zooecial apertures. A hyaline core is surrounded by steeply dipping conical laminae. Autozooecial wall thickness averages 0.08 mm in the exozone. Wall microstructure is composed of steeply inclined, U-shaped laminae. Zooecial boundaries are indistinct due to the presence of large acanthostyles which disrupt the wall structure. Frequently the autozooecia, and virtually all of the mesozooecia, are infilled with laminar calcite close to the zoarial surface. In longitudinal section this infilling consists of broad U-shaped laminae. Often large areas of adjacent zooecia are infilled. An intrazoarial overgrowth is recognised in one specimen (PD 8334). It is continuous with the exozone and has endozonal and exozonal components. REMARKS. Dittopora sanclerensis sp. noy. is characterised by extremely thin endozonal walls which thicken markedly in the exozone. Partial and complete diaphragms are present in the exozonal autozooecia. Circular mesozooecia with numerous diaphragms are present. Acanthostyles are large and abundant in the exozone. Dittopora annulata (Eichwald, 1860), from the Orthoceras Lime- stone (Llanvirn) in Estonia and the Glauconite Limestone in Russia, is similar internally to Dittopora sanclerensis. However, D. annulata has alternating bands of autozooecia and mesozooecia, whereas in D. sanclerensis they are not arranged in bands. Modzalevskaya (1953) described two new species of Dittopora, D. sokolon and D. ramosa, from the western Russian Platform: D. sokolon has similar autozooecia, diaphragms and acanthostyles to D. sanclerensis but thicker endozonal walls; D. ramosa has similar diaphragms but smaller and more abundant acanthostyles. A common feature of D. sanclerensis is that zooecia close to the zoarial surface are filled with U-shaped laminar calcite (Fig. 2). This may be because the studied material consists mainly of the basal parts of colonies with ontogenetically older zooids. 120 C. BUTTLER ORDOVICIAN BRYOZOA FROM THE LLANDEILO LIMESTONE Genus HEMIPHRAGMA Ulrich 1893 Hemiphragma pygmaeum Bassler, 1911 Figs 4-5 1911 Hemiphragma pygmaeum Bassler: 289, fig. 176. non 1970 Hemiphragma pygmaeum Bassler; Nekhorosheva: 84; pl. vii, figs 5-6. SYNTYPES. NHM D 22829, D 22536; Chasmops Limestone (up- per Caradoc), Oland, Sweden. MATERIAL. NHM PD 833 1a, b. DESCRIPTION. Zoaria erect with quite thick cylindrical branches, on average 4.5 mm in diameter. Autozooecia curve out gradually from the branch axis to meet the zoarial surface at 90°. The autozooecia within the endozone have thick straight walls. The exozone has an average diameter of 1.24 mm. It is recognised by a thickening of the zooecial walls. Autozooecia all originate in the endozone, where they are rounded and occasionally petaloid in transverse section, becoming rounded to circular as seen in tangen- tial sections of branches. Autozooecial diameters average 0.19 mm by 0.23 mm within the exozone. Diaphragms are found throughout the colony but are not common. Partial diaphragms are, however, very abundant everywhere. They are spaced on average 0.17 mm apart in the endozone and 0.15 mm apart in the exozone, and tend to occur on alternating sides of the autozooecia. The direction of deflection of the laminae of the diaphragms at their junctions with zooecial walls cannot be distinguished. Mesozooecia are present, although not common, and originate in the outer parts of the endozone. They are rounded in tangential section and have an average maximum diameter of-0.1 mm. Mesozooecia contain orally deflected basal diaphragms, spaced on average 0.08 mm apart. Acanthostyles are large and abundant with an average diameter of 0.05 mm and density of 23 per mm’. They originate throughout the colony and can indent the autozooecial apertures to produce a petaloid shape. In the outer exozone they are normally found in the centre of the thick walls. A large hyaline core is surrounded by steeply dipping conical laminae. Autozooecial wall thickness averages 0.07 mm in the exozone. Wall microstructure is composed of steeply inclined, U-shaped laminae. Adjacent zooecial wall boundaries are occupied by wide granular areas. The wall structure is hard to distinguish because it is greatly disrupted by the large acanthostyles. Frequently zooecia are infilled with laminar calcite close to the zoarial surface. In longitu- dinal section this infilling consists of broad U-shaped laminae. REMARKS. Hemiphragma pygmaeum was originally described by Bassler (1911) from the Chasmops Limestone (upper Caradoc) of Oland, Sweden but has not hitherto been recognised elsewhere. Bassler characterised the species by its ‘mushroom’-shaped colo- nies, thick zooecial walls, large acanthostyles throughout the colony and the abundant partial diaphragms. The specimens from Clog-y-fran are known only in section. They differ from the Swedish H. pygmaeum in having a straight-sided, 121 more erect colony form and slightly smaller acanthostyles. Also the mesozooecia do not appear to originate as deeply in the colony. At present the Welsh material is placed within H. pygmaeum and the differences with the Swedish material are considered to represent intraspecific variability until more material can be examined. The genus Hemiphragma has not be previously described from Great Britain. A second species from Clog-y-fran, Hemiphragma sp., 1s also described here. The two species are very different, H. pygaeum being ramose in form, with thick walls and large acanthostyles, and Hemiphragma sp. being hemispherical, with thin walls and ring-diaphragms. H. pygmaeum was described from the middle Ordovician of Pai- Khoi and Vaigach Island, Russia (Nekhorosheva 1970). The illustrations, however, show thin-walled specimens with abundant diaphragms and small acanthostyles; these are considered not to be H. pygmaeum. Hemiphragma sp. Fig. 6 MATERIAL. NHM PD 8327. DESCRIPTION. Zoaria large and hemispherical, on average 2.5 mm in diameter. Autozooecia all originate from the basal lamina and curve outwards towards the zoarial surface. Autozooecial walls are straight throughout the colony; there is no differentiation between endozone and exozone. Autozooecia are large with an average diameter of 0.39 mm x 0.47 mm and are polygonal/rounded to rounded in transverse section throughout the colony. Thin dia- phragms and ring diaphragms are present in all zooecia, spaced 0.62 mm apart in the endozone and 0.36 mm in the exozone. These are basal diaphragms which are orally-deflected at their junctions with the zooecial walls and have laminae continuous with the autozooecial linings. Possible acanthostyles occur in the outer parts of the colony, but are rare. Autozooecial wall thickness averages 0.02 mm at the periphery of the colony. It is not possible to identify the microstruc- ture from available peels and thin sections. REMARKS. ‘The specimen described herein is characterised by a hemispherical colony shape. Autozooecia have straight, thin walls throughout the zoarium, and the autozooecial apertures are polygo- nal-rounded to rounded in transverse section. Diaphragms and ring diaphragms are present in all autozooecia. Bassler (1911: 282, fig. 170) described and illustrated specimens of H. tenuimurale Ulrich, 1893 from the type locality, the Clitambonites and Nematopora Beds, Lower Trenton, Minnesota and Iowa; and from the Wassalem Beds (Caradoc), Uxnorm, Esto- nia. The Welsh specimen is similar to the Estonian material: it has thin walls and mesozooecia are lacking, but differs in having a hemispherical rather than ramose colony, diaphragms within the endozone, and by the presence of ring diaphragms rather than partial diaphragms. H. tenuimurale described by Ulrich (1893) from Min- nesota (see Bassler, 1911: fig. 171) has fewer diaphragms in the endozone compared to the Estonian material. Brown (1965) des- cribed specimens of H. tenuimurale from the Logana and Jessamine Figs 2-3 Dittopora sanclerensis sp. noy. 2, NHM PD 8338 (holotype); 2a, longitudinal section, x22; 2b, transverse section, x22; 2e, tangential section, x53; 2d, tangential section, showing infilled zooecia, x53. 3, NHM PD 8333 (paratype); longitudinal section, showing infilled autozooecia, x37. Figs 4-5 Hemiphragma pygmaeum Bassler, 1911. 4, NHM PD 833 1a; longitudinal section, showing partial diaphragms, x35. 5, NHM PD 8331b; 5a, tangential section, x35; 5b, tangential section, x94. Fig. 6 Hemiphragma sp., NHM PD 8327; 6a, longitudinal section, x23; 6b, longitudinal section, showing ring diaphragms, x37; 6c, tangential section, x30. Fig. 7 Heterotrypa sp., NHM PD 8314; 7a, longitudinal section, x26; 7b, longitudinal section, x47; 7c, transverse section, showing infilled zooecia in the exozone, x37; 7d, tangential section, x70. Fig. 8 Leioclema sp. A. NHM PD 8307; longitudinal section, x22. Figs 9-10 Leioclema sp. A. 9, NHM PD 8308; transverse section, x22. 10, NHM PD 8307; tangential section, showing petaloid autozooecia, x101. Fig. 11 Leioclema sp. B., NHM PD 8306; 11a, longitudinal section, x86; 11b, transverse section, x55; 11c, tangential section, x86. Figs 12, 13 Leioclema? sp. 12, NHM PD 8138; longitudinal section, x22. 13, NHM PD 8137; 13a, tangential section, x30; 13b, tangential section, showing large acanthostyles indenting autozooecial walls, x112. Fig. 14 Hallopora peculiaris Pushkin (in Ropot & Pushkin, 1987). NHM PD 8396; longitudinal section, x22. Fig. 15 Hallopora atf. wesenberginia (Dybowski, 1877), NHM PD 8312; 15a, longitudinal section, x22; 15b, tangential section, x37. Limestones (middle Ordovician) of Kentucky which are very similar to those from Minnesota and Iowa. Restudy is needed to assess the variability within the species, and it is possible that the Estonian and American forms are different species. Until further material can be examined, this one incomplete specimen is left in open nomenclature. C. BUTTLER Genus HETEROTRYPA Nicholson, 1879 Heterotrypa sp. Fig. 7 MATERIAL. NHM PD 8314. DESCRIPTION. Zoarium erect with thin cylindrical branches, on average 4 mm in diameter. ORDOVICIAN BRYOZOA FROM THE LLANDEILO LIMESTONE Autozooecia curve outwards gradually from the branch axis to meet the zoarial surface at 90°. Autozooecia within the endozone have thin, slightly wavy walls. The exozone is narrow with an average width of 0.57 mm. It is recognised by a thickening of the zooecial walls. Autozooecia all originate in the endozone where they are rounded-polygonal in transverse section, becoming rounded to slightly petaloid in the exozone as seen in tangential sections of branches. Autozooecial diameters average 0.19 mm in the exozone. Diaphragms are rare in the autozooecia, and when present only one or two are found in the outer endozone and exozone. These basal diaphragms are all orally- deflected at their junctions with the zooecial walls. Mesozooecia are common and originate throughout the endozone; their shape in shallow tangential section has not been observed. Orally deflected basal diaphragms are found along the entire length of the mesozooecia. Vertical walls are extensively constricted at the positions of the diaphragms, producing a pronounced beaded (in some cases vesicular) appearance. Acanthostyles are large and abundant, with an average diameter of 0.05 mm. They originate throughout the colony and commonly indent the autozooecia. Acanthostyle microstructure is difficult to distinguish, but seems to consist of a hyaline core surrounded by steeply dipping conical laminae. Autozooecial wall thickness averages 0.06 mm. Wall microstruc- ture is composed of inclined laminae, but is hard to distinguish because the walls are greatly disrupted by acanthostyles. Frequently, the zooecia are infilled with laminar calcite close to the zoarial surface. The zooecia in large parts of the colony have been found to be all infilled. In shallow tangential sections of these areas, all that is seen is the laminar calcite wall pierced by acanthostyles. REMARKS. Only one poorly preserved specimen of this species has been found. It is characterised by the ramose colony form and meandering endozonal walls. Autozooecia are rounded to slightly petaloid in shallow tangential sections, and beaded mesozooecia are common throughout the colony. Diaphragms are rare in autozooecia but abundant in mesozooecia throughout the colony. The irregularly shaped, beaded zooecia make this a particularly characteristic spe- cies and distinguish it from H. sladei described by Buttler (1991b). Identification is, however, left in open nomenclature until better preserved material can be examined. Heterotrypa sp. is similar to the Russian species H. ovata Astrova, 1957, but has more weakly beaded zooecia and less prominent acanthostyles. Genus LEIOCLEMA Ulrich, 1882 Leioclema sp. A Figs 8-10 MATERIAL. NHM PD 8307-8308. DESCRIPTION. Zoaria erect with cylindrical branches on average 8 mm in diameter. Autozooecia curve outwards gradually from the branch axis to meet the zoarial surface at 80°-90° and have moderately thick, slightly wavy walls within the endozone. The exozone is quite large with an average diameter of 1.9 mm. It is recognised by an extensive thickening of the zooecial walls. Autozooecia all originate in the endozone, where they are rounded- petaloid in transverse section becoming extensively petaloid in the exozone as seen in tangential sections of branches. Autozooecial diameter averages 0.18 mm by 0.26 mm within the exozone. Dia- phragms are rare and frequently wholly absent in autozooecia. 123 Mesozooecia are very common and originate in the outer parts of the endozone and inner parts of the exozone. They are rounded and have a maximum diameter averaging 0.09 mm. The mesozooecia contain abundant orally-deflected basal diaphragms, spaced on average 0.15 mm apart in the endozone and 0.1! mm apart in the exozone, with successive diaphragms often increasing in thickness distally along the mesozooecia. Acanthostyles are large and very abundant with an average diam- eter of 0.03 mm and density of 29.5 per mm?. They originate throughout the colony and are very abundant in the exozone, where they indent the autozooecial apertures, producing a petaloid shape. Acanthostyles are larger in size in the outer exozone than in the rest of the colony. In longitudinal section acanthostyles can be identified protruding into the zooecial chambers. Acanthostyles are composed of a broad hyaline core surrounded by steeply dipping conical laminae. Autozooecial wall thickness averages 0.13 mm in the exozone. Wall microstructure is composed of steeply inclined, U-shaped laminae. In tangential sections a thick granular layer can be identi- fied between adjacent zooecia. Virtually all mesozooecia, and some autozooecia, are infilled with laminar calcite close to the zoarial surface. In longitudinal section this infilling consists of broad U- shaped laminae. The infilling of the mesozooecia commences in the middle exozone, whereas the autozooecia are infilled in the outer exozone. REMARKS. Only two poorly preserved specimens of this species have been found at Clog-y-fran. The colonies are primarily recog- nised by the erect form, and by autozooecial walls that are thick throughout the colony. Autozooecial apertures are rounded-petaloid in transverse section and markedly petaloid in shallow tangential section. Rounded mesozooecia are common with abundant dia- phragms; diaphragms are rare in the autozooecia. Acanthostyles are present throughout the colony; they are large and abundant in the exozone. The identification of this species is difficult because of the poor quality of the two known specimens. Leioclema sp. A is placed in open nomenclature until better preserved material can be obtained and a complete description of this possibly new species can be made. Leioclema sp. B Fig. 11 MATERIAL. NHM PD 8306. DESCRIPTION. Zoaria erect with very thin cylindrical branches, on average 1.5 mm in diameter. Autozooecia curve away gradually from the branch axis in the endozone and then more abruptly in the exozone to meet the zoarial surface at 90°. Autozooecia within the endozone all have straight thin walls. The exozone is extremely broad with an average width of 0.67 mm. It is recognised by a simultaneous thickening of zooecial walls and a change in zooecial orientation. Autozooecia all originate in the endozone where they are rounded in transverse section. They retain this shape in the exozone, as seen in tangential sections of branches. Autozooecial diameters average 0.09 mm by 0.11 mm within the exozone. Diaphragms are absent in the autozooecia. Occasional cystiphragms can, however, be found in the outer exozone. Mesozooecia are common and originate in the inner parts of the exozone. They are rounded in shape in tangential sections and have a maximum diameter averaging 0.04 mm. The mesozooecia contain abundant orally deflected, thick, basal diaphragms, spaced on aver- age 0.04 mm apart, and generally increasing in thickness distally along the mesozooecia. 124 Acanthostyles are common and have an average diameter of 0.03 mm and density of 29.5 per mm?*. They originate deep in the exozone, extend the length of the exozone, and are composed of a hyaline calcite core surrounded by steeply dipping conical laminae. Autozooecial wall thickness averages 0.08 mm in the exozone. Wall microstructure is composed of steeply inclined, V-shaped laminae. The zooecial boundaries are, however, indistinct. Zooecia are frequently infilled with laminar calcite close to the zoarial surface. In longitudinal section this infilling consists of broad U- shaped laminae. REMARKS. Leioclema sp. B is primarily characterised by: the small size of the erect branches; autozooecial apertures rounded in transverse section throughout the colony; an exozone which is very wide in comparison with the endozone; common rounded mesozooecia containing abundant diaphragms; and small but long acanthostyles. Only one specimen is known from Clog-y-fran. Leioclema sp. B can be distinguished from Leioclema sp. A by the smaller colony size. Owen (1962, 1965, 1969) described numerous species of Leioclema and the related Asperopora from the Silurian of the Welsh Borderland and Shropshire. L. halloporoides Owen, 1962 was de- scribed from the Ludlow Aymestry Limestone of Shropshire. This also has very small colony branches (2 mm diameter) and mesozooecia containing diaphragms. Leioclema sp. B, however, has more abundant diaphragms within the mesozooecia and larger acanthostyles Leioclema sp. B is possibly a new species but, as it is represented only by one poorly preserved specimen, no specific name will be assigned until further material is obtained. Leioclema? sp. Figs 12-13 MATERIAL. NHM PD 8316-8318. DESCRIPTION. Zoaria erect with thick cylindrical branches, on average 12 mm in diameter. Autozooecia curve outwards gradually from the branch axis to meet the zoarial surface at 90°. The autozooecia within the endozone have very thick walls. The exozone has an average diameter of 2.28 mm. It is recognised by a slight thickening of the zooecial walls. Autozooecia all origi- nate in the endozone where they are rounded-polygonal in transverse section; they become irregular-rounded in the exozone, as seen in tangential sections of branches. Autozooecial diameters average 0.37 mm x 0.45 mm within the exozone. Thin basal diaphragms, which are orally-deflected at their junctions with the zooecial walls, are rare. Mesozooecia occur in the exozone, on shallow tangential sec- tions. They are rounded with a maximum diameter averaging 0.14 mm, and contain abundant, thick, orally-deflected, basal diaphragms, which are spaced on average 0.13 mm apart in the endozone and 0.1 mm in the exozone. Acanthostyles are large and abundant with an average diameter of 0.09 mm and density of 12.5 per mm’. They vary greatly in size, ranging in diameter from 0.05 mm to 0.12 mm, and originate throughout the colony. They are often wider than the zooecial walls. The acanthostyles are composed of a broad hyaline core; no sheath- ing laminae can be identified. Autozooecial wall thickness averages 0.06 mm in the exozone. Wall microstructure is composed of inclined, V-shaped laminae, but is exceedingly poorly preserved. REMARKS. Only three fragmentary specimens of this species have C. BUTTLER been found. The colonies are erect with thick autozooecial walls. The autozooecia are rounded-polygonal to slightly petaloid in shal- low tangential section. Rounded mesozooecia are present and have abundant diaphragms. Diaphragms are rare in autozooecia. Acanthostyles are large and abundant throughout the colony, occa- sionally inflecting autozooecial walls. Identification of the species is difficult because of the poor preservation of the material. The abundance of diaphragms in the mesozooecia and the lack of them in the autozooecia fit within the generic concept of Leioclema followed here. However, the acanthostyles are large (often wider than the zooecial walls), abun- dant and originate throughout the colony. A thick hyaline core is identifiable but no surrounding laminae are present, as found in the acanthostyles of other species of Leioclema. The acanthostyles are similar to those observed in the early Ordovician genus Orbipora Eichwald, 1856, illustrated in Astrova (1978: pl. 11, fig.i) and Taylor & Cope (1987: fig.1). Orbipora is, however, characterised by its massive form and absence of mesozooecia. The precise taxonomic placing of the species is uncertain because of the poor quality of the material available. The specimens are therefore tentatively designated Leioclema? sp. Family HALLOPOROIDAE Bassler, 1911 Genus HALLOPORA Bassler, 1911 Hallopora peculiaris Pushkin, 1987 Fig. 14 1987 Hallopora wesenbergiana peculiaris Pushkin in Ropot & Pushkin: 153; pl. 8, fig. 5, pl. 9, fig. 1. 1991b Hallopora peculiaris Pushkin; Buttler: 86; pl. 3, figs 3-8. MATERIAL. NHM PD 8396. OTHER OCCURRENCES. Piriguskii Stage (Lower Ashgill, upper Ordovician), Shikipi, Latvia (see Pushkin in Ropot & Pushkin, 1987), Slade and Redhill Beds (upper Rawtheyan, Ashgill), A40 Pengawse Hill diversion, W. of Whitland, Dyfed, Wales (SN 164170) (see Buttler 1991b). DESCRIPTION. Zoarium erect with cylindrical branches on average 5.7 mm in diameter. Autozooecia curve gradually away from the branch axis in the endozone and meet the zoarial surface at approximately 80—90°. In the endozone the zooecial walls are very thin. The exozone, recognised by a thickening of the zooecial walls, has an average width of 1.52 mm. Autozooecia are circular in transverse section throughout the colony and average 0.29 mm in diameter in the exozone. Diaphragms are rare within the autozooecia and when present, usually occur closely spaced in the distal exozone. These basal diaphragms are orally-deflected at their junctions with the zooecial walls and their laminae are generally continuous with the zooecial linings. The average spacing between diaphragms is 0.17 mm in the exozone. Mesozooecia are common throughout the whole zoarium, often originating in the inner parts of the endozone. Mesozooecial walls are thin in the endozone and thicken in the exozone. They are polygonal to polygonal-rounded in shallow tangential sections. Basal diaphragms are present throughout their length, spaced on average 0.1 mm apart in the endozone and 0.07 mm in the exozone. Diaphragms tend to increase in thickness distally along the mesozooecia. In some colonies mesozooecial walls are constricted at the position of the diaphragms, producing a slightly beaded appearance. ORDOVICIAN BRYOZOA FROM THE LLANDEILO LIMESTONE Autozooecial wall thickness averages 0.1mm in the exozone. Wall microstructure is composed of steeply inclined, V-shaped laminae. The precise contact between the zooecia is indistinct. The thickened exozonal diaphragms in the mesozooecia are also laminar and are continuous with the wall laminae. Maculae composed of a concentration of mesozooecia can be recognised in thin sections. REMARKS. Hallopora peculiaris is primarily characterised by the extensive beaded mesozooecia which originate in the inner endozone. The autozooecia are circular throughout the colony, and diaphragms are rare in the endozone, becoming more abundant in the outermost regions. H. peculiaris has also been described from the Slade and Redhill Beds (Ashgill) of South Wales (Buttler, 1991b). Hallopora aff. wesenbergiana (Dybowski, 1877). Fig. 15 MATERIAL. NHM PD 8310-8313. DESCRIPTION. Zoaria erect with thick cylindrical branches, on average 11 mm in diameter. Autozooecia are parallel to the branch axis in the inner endozone and then curve outwards gradually to meet the zoarial surface at 80°-90°. Within the endozone autozooecial walls are thin and straight. The exozone is difficult to distinguish; it can be recognised by a slight thickening of the zooecial walls. Autozooecia all originate in the endozone where they are polygonal-rounded in transverse sec- tion, becoming circular in the exozone as seen in tangential sections of branches. Autozooecial diameters average 0.21 mm by 0.27 mm within the exozone. Diaphragms are very abundant throughout the autozooecia. In the endozone there are periodic concentrations of diaphragms which occur throughout the colony at the same level. Within the concentrations, diaphragms are spaced on average 0.1 mm apart; elsewhere they are spaced on average 0.6 mm apart. In the exozone, diaphragms are very abundant and on average spaced 0.11 mm apart. All the diaphragms are basal and orally-deflected at their junctions with the zooecial walls. Mesozooecia are present, originating in the outer parts of the endozone and inner parts of the exozone. They are rounded in shallow tangential sections and have a maximum diameter averag- ing 0.1 mm. They contain abundant orally deflected basal diaphragms spaced on average 0.07 mm apart in the exozone. Autozooecial wall thickness averages 0.03 mm in the exozone. The wall microstructure is poorly preserved but vague laminations can be recognised in one tangential section. REMARKS. Hallopora aff. wesenbergiana (Dybowski, 1877) is characterised by the ramose colony form with thick branches. Thin- walled zooecia have rounded apertures in shallow tangential sections. Diaphragms are abundant throughout the whole colony and are periodically concentrated in bands, which possibly indicate periods of slow growth. Bassler (1911) illustrated H. wesenbergiana from the Wesenberg Limestone and Wassalen Beds (Caradoc) in Estonia. This material is similar to the Welsh; it has thin-walled autozooecia with abundant diaphragms, although these do not occur in bands. The banding, or lack of it, may not be a specific feature but may relate to an environmental influence acting on the colonies. The poor quality of the Welsh material does not allow a more precise specific identification. 125 Genus BATOSTOMA Ulrich, 1882 Batostoma clogyfranense sp. nov. Figs 16-18 HOLOTYPE. NHM PD 8362. PARATYPES. NHM PD 8353-8361, 8363-8375. NAME. After the type locality. DIAGNOSIS. Colony small, ramose. Autozooecia curve out gradu- ally from branch axis to zoarial surface. Autozooecial walls thin in endozone. Autozooecia polygonal-rounded in transverse section, circular in shallow tangential sections. Small polygonal-rounded mesozooecia present, originating in outer endozone. Diaphragms present in all zooecia. Acanthostyles small, abundant in exozone. DESCRIPTION. Zoaria erect with narrow cylindrical branches, on average 3.9 mm in diameter. Autozooecia curve out gradually from the branch axis to meet the zoarial surface at an angle of 70°-80°. The autozooecia within the endozone have thin walls. The exozone is narrow with an average diameter of 0.8 mm; it is characterised by a thickening of the zooecial walls. Autozooecia originate in the endozone where they are polygonal-rounded in transverse section, becoming circular in the exozone as seen in tangential sections of branches. Autozooecial diameters average 0.15 mm by 0.18 mm in the exozone. Diaphragms are found throughout the autozooecia, although they are less common in the inner exozone. They are spaced on average 0.28 mm apart in the endozone, and 0.15 mm apart in the exozone. These basal dia- phragms are all orally-deflected at their junctions with the zooecial walls and their laminae are continuous with the zooecial linings. Mesozooecia are present and originate in the outer parts of the endozone. They are polygonal-rounded in shallow tangential sec- tions with a maximum diameter which averages 0.09 mm. Orally-deflected basal diaphragms are found along the length of the mesozooecia, spaced on average 0.07 mm apart. Acanthostyles are small, often irregularly shaped and highly abundant, with an average diameter of 0.03 mm and density of 67 per mm7?. They originate in the exozone and usually form a ring around the autozooecia, consisting of approximately ten acanthostyles. The acanthostyles are composed of a circular, or sometimes irregular hyaline core surrounded by indistinct dipping conical laminae. Autozooecial wall thickness averages 0.08 mm in the exozone. Wall microstructure is composed of inclined, U-shaped laminae; however, it is poorly preserved. Frequently, zooecia are infilled with laminar calcite close to the colony surface; in longitudinal section this infilling consists of broad U-shaped laminae; large sections of zoaria often have all the zooecia infilled in this way. REMARKS. Batostoma clogyfranense sp. nov. is primarily charac- terised by thin, straight zooecial walls, a narrow exozone, and diaphragms regularly spaced throughout the colony. Autozooecial apertures are rounded in shallow tangential section, and polygonal- rounded mesozooecia occur. Acanthostyles are abundant and occasionally irregular in shape. Another species of Batostoma, B. cf. polare Astrova, 1965, that is described here can be distinguished from B. clogyfranense by the thicker exozonal walls and the less abundant diaphragms in the exozone. The middle Ordovician species B. subtile Astrova (1965: pl. 50, fig. 2, pl. 51, fig. 1), from Vaigach Island, Novaya Zemlya, Russia, has a similar pattern of diaphragms within the endozone to B. clogyfranense. Diaphragms are, however, more abundant in the C. BUTTLER Figs 16-18 Batostoma clogyfranense sp. nov. 16, NHM PD 8362 (holotype), longitudinal section, x22. 17, NHM PD 8374 (paratype), longitudinal section, x30. 18, NHM PD 8362 (holotype); 18a, transverse section, x30; 18b, tangential section, showing infilled zooecia, x37; 18c, tangential section, x86. Fig. 19 Batostoma cf. polare Astrova 1965, NHM PD 8324; 19a, longitudinal section, x22; 19b, longitudinal section, x70; 19¢, tangential section, x41; 19d, tangential section, x96. exozone, and mesozooecia are less common in the Russian species. Two species of Batostoma have been previously described from the Lower Palaeozoic of the Welsh Basin. B. murchisoni was de- scribed by Spjeldnaes (1957) from “Horderley’ in Shropshire. This species has few diaphragms and mesozooecia, and acanthostyles are absent, suggesting that the species may not belong to Batostoma. A re-examination of the type material is required. Owen (1962) described Batostoma sp, from the Aymestry Limestone (Ludlow Series, Silurian), Wenlock. This species has a very small exozone and mesozooecia are absent. Batostoma cf. polare Astrova, 1965 Fig. 19 MATERIAL. NHM PD 8324. DESCRIPTION. Zoarium erect with narrow cylindrical branches, on average 4.5 mm in diameter. Autozooecia are parallel to the branch axis within the endozone and curve abruptly outwards in the exozone to meet the zoarial surface at 90°. The autozooecia within the endozone have thin, slightly wavy walls. The exozone is thick with an average diameter of 1.1 mm. It is recognised by an extensive thickening of the zooecial walls and a simultaneous change in zooecial orientation. Autozooecia all origi- nate in the endozone, where they are polygonal-rounded in transverse section. They become circular in the exozone, as seen in tangential sections of branches. Autozooecial diameters average 0.16 mm by 0.2 mm within the exozone. Diaphragms are present throughout the autozooecia, spaced on average 0.06 mm apart in the endozone and increasing greatly to 0.38 mm apart in the exozone. The majority of these are basal diaphragms, which are deflected orally at their junctions with the zooecial walls. Successive diaphragms increase in thickness distally along the autozooecia. Several subterminal, aborally deflected diaphragms have been recognised at the distal end of the colony. Mesozooecia are present, although not abundant, and originate in the inner parts of the exozone. They have a maximum diameter averaging 0.1 mm. In shallow tangential sections they are rounded. The mesozooecia contain abundant, thick, orally deflected, basal diaphragms, which are spaced on average 0.05 mm apart. Acanthostyles are small and abundant with an average diameter of 0.02 mm and density of 30 per mm”. They originate in the exozone ORDOVICIAN BRYOZOA FROM THE LLANDEILO LIMESTONE and usually form a ring around the autozooecia, approximately 14 acanthostyles surrounding one autozooecium. The acanthostyles have a hyaline core surrounded by steeply dipping conical laminae. Autozooecial wall thickness averages 0.11 mm in the exozone. Wall microstructure is composed of steeply inclined, V-shaped laminae. Zooecial boundaries are dark, crenulated and granular. Some zooecia are infilled with laminar calcite close to the colony surface. In longitudinal section this infilling consists of broad U- shaped laminae. An intrazoarial overgrowth has been recognised which 1s continu- ous with the underlying branch and is composed of outer endozonal/ inner exozonal components. REMARKS. Only one specimen of Batostoma cf. polare Astrova, 1965, has been found during this study. It is characterised by the ramose colony form and particularly thin autozooecial walls in the endozone, which thicken extensively in the exozone. Autozooecial apertures are circular in shallow tangential sections, and rounded mesozooecia, which originate in the outer endozone, are present. Thick diaphragms are abundant in the exozone and thin diaphragms occur in the endozone. Acanthostyles are small and numerous in the exozone. B. polare Astrova, 1965, described from the Varnek Stage, Vaigach and Novaya Zemlya, Russia, is very similar to the specimen from Clog-y-fran. They both have thin zooecial walls in the endozone which thicken in the exozone, abundant basal exozonal diaphragms, small mesozooecia and acanthostyles. Measurements for the Soviet and Welsh specimens are similar. The major difference between the Welsh specimen and type B. polare is the presence of the dia- phragms within the endozone of the former. Genus ERIDOTRYPA Ulrich, 1893 Eridotrypa simulatrix (Ulrich, 1890) 1890 Batostoma simulatrix Ulrich; 432, pl. 35, fig. 1. 1893 Monticulopora simulatrix (Ulrich); James: 194. 1908 Eridotrypa simulatrix (Ulrich); Cummings: 828, pl. 16, fig. 4. 1928 Eridotrypa simulatrix (Ulrich); Bassler: 152 1987 Eridotrypa simulatrix (Ulrich); Ropot & Pushkin: 171, pl. 15, fig. 2. MATERIAL. NHM PD 8342-8352. Fig. 20 OTHER OCCURRENCES. Cincinnati Group, Savanna, Illinois, USA; English Head and Vaureal Formations, Anticosti Island, Quebec, Canada; Waynesville Formation, Harmons Station, Indiana, USA; Pirguskii Stage (Caradoc), Yuzhnoi, Pribaltiki, Russia. DESCRIPTION. Zoaria erect with narrow cylindrical branches, on average 3.3 mm in diameter. Autozooecia meander roughly parallel to the branch axis within the endozone and then curve slightly in the exozone to meet the zoarial surface at 50°. Within the endozone they have thin walls. The exozone is narrow with an average diameter of 0.64 mm. It is recognised by an extensive thickening of the zooecial walls. Autozooecia all originate in the endozone and are rounded-polygo- nal in transverse section, becoming oval-rounded in the exozone as seen in tangential sections of branches. In branch transverse section the autozooecia are larger in diameter in the inner endozone than in the outer endozone. Autozooecial diameters average 0.13 mm by 0.18 mm within the exozone. Diaphragms are occasionally present in the endozone and exozone, spaced on average 0.35 mm apart in 127 the endozone and 0.13 mm in the exozone. In some specimens they are very abundant (PD 8348). These basal diaphragms are all deflected orally at their junctions with zooecial walls and their laminae are continuous with the zooecial linings. Mesozooecia are present and originate in the endozone. They are rounded in shallow tangential section, with a maximum diameter averaging 0.09 mm. Abundant orally deflected diaphragms are found along the length of the mesozooecia, spaced on average 0.07 mm apart. Acanthostyles are small, occasionally irregular, abundant, with an average diameter of 0.02 mm. They are composed of a hyaline calcite core surrounded by indistinct dipping conical laminae. Autozooecial wall thickness averages 0.06 mm in the exozone. Wall microstructure is rather indistinct (only peels of these speci- mens are available) and is composed of steeply inclined, V-shaped laminae. Zooecial boundaries have not been distinguished. Autozooecia, and more especially mesozooecia, are frequently infilled with laminar calcite close to the zoarial surface. In longitu- dinal section this infilling consists of broad V-shaped laminae; large areas of the colony can be infilled. REMARKS. Eridotrypa simulatrix is characterised by the ramose colony with narrow branches. Autozooecial walls are thin and meandering in the endozone and thicken in the exozone. Autozooecial apertures are large and rounded/polygonal in transverse section, and small and oval in shallow tangential section. Diaphragms are present and abundant small acanthostyles occur in the exozone. It is not easy to compare the Russian and American specimens of E. simulatrix because those from North America are only illustrated by line drawings. The Welsh and Russian specimens appear to be identical but together may prove to represent a distinct species from the American specimens when direct comparisons have been made using the actual material. Genus MONTICULIPORA @ Orbigny, 1850 Monticulipora aff. compacta Coryell, 1921. Fig. 21 MATERIAL. NHM PD 8328, 8331c. DESCRIPTION. Zoaria erect with narrow cylindrical branches, on average 4 mm in diameter. Autozooecia curve out gradually from the branch axis in the endozone and meet the zoarial surface at 90°. The autozooecia within the endozone have thin, slightly wavy walls. The exozone is moderately broad with an average width of 1.05 mm. It is recognised by a thickening of the zooecial walls. Autozooecia all originate in the endozone, where they are circular in transverse section. They become rounded in the exozone as seen in tangential sections of branches. Autozooecial diameters average 0.16 mm by 0.13 mm within the exozone. Diaphragms are present throughout the autozooecia. They are spaced on average 0.15 mm apart in the endozone and increase to 0.07 mm apart in the exozone. The majority of the diaphragms are basal, deflected orally at their junctions with the autozooecial walls. Some diaphragms are possi- bly subterminal, but the poor preservation of the specimen does not allow this to be confirmed. Cystiphragms are numerous along the whole length of the autozooecia, especially in the exozone. The cystiphragms are normally restricted to one side of an autozooecium. Mesozooecia are uncommon, and originate in the outer parts of the endozone and inner parts of the exozone. They have an average maximum diameter of 0.08 mm, are rounded in shallow tangential section and contain abundant, orally deflected basal diaphragms. C. BUTTLER Shes RE Ace BN eed Ps if ‘ : al a * saa Be RANA Fig. 20 Eridotrypa simulatrix (Ulrich, 1890), NHM PD 8343; 20a, longitudinal section, x25; 20b, transverse section, x25; 20c, tangential section, x53; 20d, tangential section, x103. Fig. 21 Monticulipora aff. compacta Coryell, 1921, NHM PD 8328; 21a, longitudinal section, x30; 21b, transverse section, x22; 21c, tangential section, x86. Fig. 22 Homotrypa cf. similis Foord 1883, NHM PD 8323; 22a, longitudinal section, x30; 22b, transverse section, x30. Acanthostyles are abundant with an average diameter of 0.03 mm and density of 47 per mm’. They originate throughout the whole colony and occasionally indent autozooecial apertures. The acanthostyles are composed of a hyaline calcite core surrounded by steeply dipping conical laminae. Autozooecial wall thickness averages 0.05 mm in the exozone. Wall microstructure is composed of steeply inclined, V-shaped laminae. It is, however, indistinct due to the presence of the acanthostyles. Occasional zooecia are infilled with laminar calcite close to the zoarial surface. In longitudinal section this infilling consists of broad U-shaped laminae. REMARKS. This species is characterised by the ramose colony form and autozooecial apertures which are rounded in shallow tangential section. Zooecial walls are thin in the endozone, thicken- ing in the exozone. Diaphragms and/or cystiphragms occur in the autozooecia, while the mesozooecia contain only diaphragms. Acanthostyles are abundant throughout the colony. This is the only species of Monticulipora encountered in the present study and the material is poorly preserved and visible only in section. Monticulipora compacta Coryell, 1921, described from the Pierce Limestone (Caradoc), Tennessee, USA (Coryell 1921: 283), Novaya Zemlya, Vaigach, and Stodolbskgo (middle Ordovician), Zapadno- Arkticheska Province, Russia (Astrova 1965: 197), has a pattern of diaphragms and cystiphragms, and rare mesozooecia similar to the Welsh specimens. In tangential section (Coryell, 1921: pl. tv, fig. 6) the autozooecial apertures are polygonal-rounded. This figured section is, however, quite deep and so cannot be accurately com- pared with the shallow Welsh sections. In M. compacta acanthostyles are reported from within the axial region, but they are not identified from the exozone, in contrast to the Welsh material. The specimens described herein are therefore assigned to M. aff. compacta until better preserved material can be examined. Genus HOMOTRYPA Ulrich, 1882 Homotrypa cf. similis Foord, 1883 Fig. 22 MATERIAL. NHM PD 8323. DESCRIPTION. Zoarium erect with cylindrical branches, on aver- age 8 mm in diameter. ORDOVICIAN BRYOZOA FROM THE LLANDEILO LIMESTONE Autozooecia are roughly parallel to the branch axis within the endozone, and gradually curve outwards to meet the zoarial surface at approximately 70°. Walls are thin and slightly wavy within the endozone. The exozone has an average diameter of 1.6 mm, and is recog- nised by a slight thickening of the zooecial walls. The autozooecia originate within the endozone, where they are polygonal in trans- verse section. No tangential sections are available. Diaphragms are present throughout the autozooecia. They are spaced on average 0.26 mm apart in the endozone and 0.1 mm apart in the exozone. The diaphragms are basal and are orally deflected at their junctions with the zooecial walls. Many diaphragms in the endozone are inclined and some are sigmoidal. Cystiphragms are numerous in the exozone where there are, on average, ten present per mm. The cystiphragms are normally restricted to one side of the autozooecia. Mesozooecia are rare and originate in the exozone when present, with an average maximum diameter of 0.08 mm. They contain abundant orally deflected basal diaphragms, spaced on average 0.04 mm apart. Small inconspicuous acanthostyle-like structures have been ob- served in the exozone; tangential sections are needed for their precise identification. Autozooecial wall thickness averages 0.04 mm in the exozone. Wall microstructure is composed of inclined V-shaped laminae. The zooecial boundaries are dark and crenulated. Some zooecia are infilled close to the zoarial surface. In longitudinal section this infilling consists of broad U-shaped laminae. REMARKS. Only one incomplete specimen of Homotrypa has been found in this present study; this has made identification difficult, especially as no tangential section could be obtained. The specimen is characterised by thin autozooecial walls in the exozone and rare mesozooecia. The autozooecia contain abundant diaphragms, espe- cially in the exozone, often sigmoidal in shape in the endozone. Cystiphragms are extremely numerous in the exozone. Ross (1963, 1965) described three Ordovician species; Homo- trypa sp. A, Homotrypa sp. B and H. oweni from the Hoar Edge Group (Caradoc Series), Shropshire. Homotrypa sp. A of Ross (1963) has widely spaced diaphragms and cystiphragms; meso- zooecia and dense acanthostyles are present but not always easy to observe in tangential section. The autozooecial apertures are po- lygonal to subpolygonal in shallow tangential sections. Homotrypa sp. B of Ross (1963) has thin autozooecial walls within the endozone and subpolygonal zooecial apertures. Diaphragms are present throughout the colony and acanthostyles are long and thin. The specimen from Clog-y-fran differs from Homotrypa sp. A in having more abundant diaphragms and cystiphragms; and from Homotyrpa sp. B by the absence of long thin acanthostyles. Homotrypa oweni Ross, 1965 differs from the Clog-y-fran specimen in having a cone- shaped or encrusting colony form, mesozooecia and rare diaphragms. The arrangement of the cystiphragms and diaphragms in the Welsh specimen is similar to that found in Homotrypa similis Foord, 1883 (e.g. Karklins 1984: pl. 5, figs 2, 3). This species is well known in North America and Eastern Europe. H. similis has acanthostyles in the exozone but in the specimen from Clog-y-fran their presence is questionable. Bork & Perry (1968: 1053) recognised H. similis (from the Guttenberg and Ion Formations, middle Ordovician, Iowa, USA) in longitudinal section ‘by extremely gradual curvature of the zooecia towards the zoarial surface, diaphragms throughout most of the axial region and well-developed cystiphragms and diaphragms in the mature zone’; the Welsh specimen fits this description. Karklins (1984: 29) noted a difference between specimens of H. similis from the Trenton Beds (middle Ordovician), Ottawa, Canada, 129 and the Lexington Limestone (middle/upper Ordovician), Ken- tucky, USA, and those from the Wassalen Beds (Caradoc) of Estonia (described by Bassler 1911). Specimens from Estonia have rela- tively broadly serrated autozooecial boundaries and well-defined acanthostyles which commonly indent the autozooecial apertures. North American specimens have narrower serrated boundaries and poorly-defined acanthostyles. The cystiphragms in the Estonian specimens are more closely spaced in the exozone that those of North America. Middle Ordovician specimens from Vaigach Island in Russia, illustrated by Astrova (1965: pl. 35), do not, however, have large acanthostyles, and their cystiphragms first occur in the outer endozone and become closely-spaced in the exozone. Thus, there appears to be a wide range of variation within the species H. similis. The Clog-y-fran specimen is compared herein with H. similis rather than positively identified as this species because the incom- plete specimen does not provide sufficient information. Genus MONOTRYFPA Nicholson, 1879 Monotrypa sp. Figs 23-24 MATERIAL. NHM PD 8329, 8330. DESCRIPTION. Zoaria hemispherical, on average 13.5 mm in diam- eter. The majority of autozooecia originate from the basal lamina and curve gently outwards to meet the zoarial surface. Autozooecial walls are straight throughout the colony. No differentiation between endozone and exozone can be recognised. The autozooecia are polygonal-rounded in transverse section, with an average diameter of 0.29 mm by 0.32 mm. Diaphragms are rare, usually only one per autozooecium. In one specimen (PD 8329), however, there is a small area of the colony with relatively numerous diaphragms which are thin, basal and orally deflected at their junctions with the zooecial walls. Autozooecial wall thickness averages 0.02 mm at the periphery of the colony. Wall microstructure is composed of inclined U-shaped laminae; the zooecial boundaries are indistinct. Occasionally zooecia are infilled with laminar calcite close to the zoarial surface. In longitudinal section this infilling consists of broad U-shaped laminae. REMARKS. The specimens described herein are characterised by the hemispherical colony form and polygonal-rounded autozooecia. Autozooecial walls are thin and straight, with no differentiation between endozone and exozone. Diaphragms are generally uncom- mon. Only two specimens are known, both in peels. Several species of Monotrypa with thin straight walls and sparse diaphragms have previously been described, e.g. M. testudiformis and M. cantarelloidea described by Dreyfuss (1948: pl. 2, figs 4, 5, 8-10) from the upper Ordovician of the Montagne Noire. The poor quality of the Welsh specimens prevents detailed com- parisons with other species, so their identification is left in open nomenclature. Genus AMPLEXOPORA Ulrich, 1882 Amplexopora sp. Figs 25-26 MATERIAL. NHM PD 8325, 8326. DESCRIPTION. Zoaria erect with thick cylindrical branches, on average 8 mm in diameter. C. BUTTLER ORDOVICIAN BRYOZOA FROM THE LLANDEILO LIMESTONE Autozooecia curve out gradually from the branch axis in the endozone and change direction abruptly in the exozone to meet the colony surface at 90°. Autozooecia within the endozone all have very thin walls. The exozone is wide, with an average diameter of 1.9 mm. It is recognised by a thickening of the zooecial walls and a simultaneous change in zooecial orientation. Autozooecia all originate in the endozone where they are rounded in transverse section, becoming irregularly rounded in the exozone as seen in tangential sections of the branches. Autozooecial diameters average 0.24 mm by 0.3 mm within the exozone. Diaphragms are very abundant and closely spaced along the whole length of the autozooecia. They are spaced on average 0.23 mm apart in the endozone, decreasing to 0.09 mm apart within the exozone. All diaphragms are basal and are orally deflected at their junctions with the zooecial walls. In the mid exozone of specimen PD 8325 there is a large interval (0.34 mm) between two adjacent diaphragms, which is found in the same position throughout the colony. The first diaphragms on the distal side of this interval are greatly deflected orally. In the majority of the colony, growth resumes as normal after the interval; however, in some small sec- tions the thickened exozonal wall terminates and is replaced by one much thinner. Mesozooecia are present, although not abundant, and have a maximum diameter averaging 0.12 mm. They originate in the exozone, are oval in shape in shallow tangential sections, and contain abundant orally deflected basal diaphragms, spaced on average 0.05 mm apart. Acanthostyles are large and abundant, with an average diameter of 0.05 mm and density of 14 per mm?. They originate throughout the exozone, commonly extending the entire length of the exozone, and can slightly indent the zooecial apertures. The acanthostyles are composed of a hyaline core surrounded by steeply dipping conical laminae. Autozooecial wall thickness averages 0.08 mm in the exozone. Wall microstructure is composed of inclined U-shaped laminae. Zooecial boundaries are indistinct. Some zooecia (especially mesozooecia) are infilled with laminar calcite close to the zoarial surface; in longitudinal section this infilling consists of broad U- shaped laminae. REMARKS. This species is characterised by the ramose colony form, thin autozooecial walls and rounded apertures in shallow tangential section. Oval mesozooecia are present and originate in the outer endozone/inner exozone. Basal diaphragms are abundant throughout the colony, and large acanthostyles are abundant in the exozone. Three species of Amplexopora have been previously described from the Welsh Basin. All were described by Ross (1963, 1965) from the Hoar Edge Limestone, Hoar Edge Group (Caradoc), Evenwood Quarry, Shropshire, and all vary markedly from the species de- scribed herein. A. thomasi Ross, 1963 is a bifoliate species with 131 small acanthostyles and lacking diaphragms in the endozone. A? evenensis Ross, 1965 and Amplexopora? sp. A of Ross, 1965 both have crenulate walls, diaphragms confined to the exozone and small acanthostyles. The specimens of Amplexopora from Clog-y-fran are similar to A. septosa (Ulrich, 1879) (redescribed by Boardman 1960) from the Fairview Formation (Ashgill), Covington, Kentucky. The major differences in A. septosa are the absence of diaphragms in the endozone and the presence of numerous short, off-set acanthostyles, as well as the long acanthostyles which occur throughout the exozone. Off-set acanthostyles can be identified in specimen PD 8326, but have not been recognised in PD 8325. Other examples of Amplexopora containing abundant diaphragms in the endozone have been described by Brown & Daly (1985) from the Dillsboro Formation (Cincinnati Series) of SE Indiana; they were identified as A. cf. septosa. Numerous specimens (over 150) of A. septosa were collected from this formation, including a few atypical specimens with abundantly spaced diaphragms in the endozone. Brown & Daly (1985: 24) suggested that because the specimens were similar in all other respects to A. septosa the differences may be due to environmental factors. The specimens from Clog-y-fran are very similar to those from the Dillsboro Formation, except that the short acanthostyles are less common and the diaphragms more abundant. Genus HALLOPORINA Bassler, 1911 Halloporina cf. crenulata (Ulrich, 1893) Fig. 27 MATERIAL. NHM PD 8315, 8394. DESCRIPTION. Zoaria erect with cylindrical branches, on average 4.5 mm in diameter. Autozooecia are parallel to the branch axis within the endozone and then curve outwards gradually in the exozone to meet the zoarial surface at 70°. The autozooecia within the endozone have very thin wavy walls. The exozone is narrow with an average diameter of 0.76 mm. It is recognisable by a thickening of the zooecial walls and a simultane- ous change in zooecial orientation. Autozooecia all originate in the endozone where they are polygonal-rounded in transverse section, becoming oval-rounded in the exozone, as seen in tangential sec- tions of branches. Autozooecial diameter averages 0.18 mm by 0.22 mm within the exozone. Basal diaphragms are rare or even wholly absent in the autozooecia and, if present, only one or two are found in the exozone. They are all deflected orally at their junctions with the zooecial walls and their laminae are continuous with the autozooecial linings. Exilazooecia are present and originate in the exozone. They are rounded in shape in shallow tangential sections, with a maximum diameter averaging 0.08 mm. They occasionally contain orally Figs 23, 24 Monotrypa sp. 23, NHM PD 8330; longitudinal section, x30. 24, NHM PD 8329; tangential section, x30. Fig. 25 Amplexopora sp., NHM PD 8325; longitudinal section, x12. Fig. 26 Amplexopora sp., NHM PD 8325; 26a, longitudinal section, showing large interval between adjacent diaphragms, x35; 26b, transverse section, x12; 26c, tangential section, x30. Fig. 27 Halloporina cf. crenulata (Ulrich 1893), NHM PD 8315; 27a, longitudinal section, x22; 27b, tangential section, x61. Figs 28, 29 Graptodictya bonnemai Bassler 1911. 28, NHM PD 8392b, longitudinal section, x30. 29, NHM PD 8389; 29a, transverse section, x55; 29b, tangential section, x55. Fig. 30 Pushkinella sp., NHM PD 8376; 30a, tangential section, x22; 30b, tangential section, x53. Fig. 31 Phylloporina sp., NHM PD 8384; 31a, tangential section, x12; 31b, tangential section, x22. 132 deflected basal diaphragms, and are therefore mesozooecia (sensu stricto), but the term exilazooecia is retained for consistency with the genus description. Autozooecial wall thickness averages 0.05 mm in the exozone. Wall microstructure is composed of steeply inclined, V-shaped laminae; the zooecial wall boundaries are dark and granular. Some exilazooecia are infilled with laminar calcite close to the zoarial surface. In longitudinal section this infilling consists of broad U- shaped laminae. REMARKS. Halloporina crenulata, the type species of the genus, has been described from the Black River and Trenton Formations (middle Ordovician) of the U.S. mid-west but has not been recog- nised elsewhere. Only one other species of Halloporina has hitherto been identified: H. parva (Ulrich & Bassler, 1904), also from the Black River and Trenton Formations of the U.S. mid-west. H. crenulata is distinguished from H. parva in having a larger zoarium, larger rounded zooecial apertures (H. parva has polygonal zooecial apertures) and more abundant exilazooecia. Nothing is known about the range of variation within either species of Halloporina. Ulrich & Bassler (1904: pl. xiv) illustrated the two species. Colony size appears to be the only major difference between them as shown in the illustrations. The drawings indicate little difference in zooecial aperture shape; the tangential section of H. crenulata is, however, deeper than in H. parva. A re-examination of the species and further material, is needed for a greater understanding of these species. The specimens from Clog-y-fran are identified only as H. cf. crenulata. The identification is tentative due to the presence of occasional diaphragms within the outer exozone. The rounded zooecia and relatively abundant exilazooecia suggest similarity to H. crenulata rather than H. parva. Order CRYPTOSTOMATA Vine, 1884 Suborder PTILODICTICTYINA Astrova & Morozova, 1956 Family ESHAROPORIDAE Karklins, 1983 Genus GRAPTODICTYA Ulrich, 1882 Graptodictya bonnemai Bassler, 1911 Figs 28-29 1911 Graptodictya bonnemai Bassler: 122, pl. 8, fig. 3, text-fig. 48. 1921 Graptodictya bonnemai jaervensis Bekker: 58, pl. 8, figs 1. 1952 Graptodictya bonnemai Bassler; Toots: 126, pl. 7, figs 5, 8, pl. 8, fig. 3, pl. 9, figs 1, 2, pl. 10, fig. 1. 1965 Graptodictya bonnemai Bassler; Astrova: 2252, pl. Ix, fig. 2, pl. xi, fig. 1. 1970 Graptodictya bonnemai Bassler; Nekhorosheva: 86, pl. vit, fig. 1. MATERIAL. NHM PD 8389-8391, 8392b, 9393. OTHER OCCURRENCES. kKuckers Shale (Kukruse Stage, Llandeilo), Baron Toll’s Estate, Estonia; Jarve, Kukersite Quarry, Wesenberg Limestone (Kukruse Stage, Llandeilo), Wesenberg, Estonia; Vaigach Island and Pai-khoi (Yugorskiy Stage, Llandeilo/Caradoc), Urals, Russia. DESCRIPTION. Zoaria erect with thin branches, on average 1.85 mm wide by 0.69 mm deep. The margins of the branches are striated. Mesothecae are thin and sinuous. In the exozone the autozooecia form 90° angles with the mesothecae. Autozooecial apertures are oval in shallow tangential sections and average 0.48 mm by 0.3 mm C. BUTTLER in the exozone. Short superior hemisepta are commonly present. Autozooecial boundaries are slightly serrated. Zooecial wall micro- structure is composed of broadly U-shaped laminae. Exilazooecia and diaphragms are absent. REMARKS. Graptodictya bonnemai was first described by Bassler (1911: 122) from Estonia, as being very similar to the type species of Graptodictya (G. perelegans). The two species were distinguished by G. bonnemai branching less frequently and having more elongate autozooecial apertures. Order FENESTRATA Elias & Condra, 1957 Suborder PHYLLOPORININA Lavrentjeva, 1979 Family ENALLOPOROIDAE Miller, 1889 Genus PUSHKINELLA Lavrentjeva, 1979 Pushkinella sp. Fig. 30 MATERIAL. NHM PD 8376-8378, 8380-8383. DESCRIPTION. Zoaria are reticulate and anastomosing; only frag- mentary specimens have been found at Clog-y-fran. No exterior frontal views of the colonies are available because frontal surfaces are all embedded in sediment. Fenestrules are oval-rounded, with diameters averaging 0.58 mm by 0.42 mm. Branches are rounded and average 0.38 mm diameter. The exozone 1s distinguished by a change in the orientation of the autozooecia and considerable thickening of the zooecial walls. Autozooecia are rounded in the endozone, becoming rounded- slightly petaloid in the exozone, where they average 0.16 mm in diameter. Across one branch one to three autozooecia are present. Wall microstructure is hard to distinguish. In one specimen (PD 8378) longitudinal laminar microstructure can be identified. Short, narrow acanthostyles are abundant throughout the colony; in the exozone they occasionally indent the autozooecial apertures. The acanthostyles are composed of a hyaline core surrounded by conical laminae, and are on average 0.1 mm in diameter. REMARKS. The genus Pushkinella was previously known only from the Baltic region of the former Soviet Union. Two Ordovician species have been recognised: P. mirabilis Lavrentjeva 1979, and P. robusta Lavrentjeva 1979, from Estonia, and one Silurian, P. acanthroporoides Pushkin 1976, from Byelorussia. The Welsh specimens of Pushkinella are characterised by the anastomosing colony form, the small oval-rounded fenestrules and rounded branches. Autozooecial walls are extensively thickened in the exozone and autozooecial apertures are rounded to slightly petaloid. Short and narrow acanthostyles are abundant. The Welsh specimens differ from P. robusta in having more zooecial apertures per branch, and from P. acanthroporoides in having a greater number of acanthostyles. P. mirabilis has similar- sized apertures and acanthostyles to the Welsh material, but differs in having occasional basal diaphragms, which are absent in the Welsh specimens. Family PHYLLOPORINIDAE Ulrich, 1890 Genus PHYLLOPORINA Foerste, 1887 Phylloporina sp. Fig. 31 MATERIAL. NHM PD 8384. ORDOVICIAN BRYOZOA FROM THE LLANDEILO LIMESTONE DESCRIPTION. Zoaria are reticulate and anastomosing, only frag- mentary specimens have been found at Clog-y-fran. No exterior frontal views of the colonies are available because frontal surfaces are all embedded in sediment. Fenestrules are oval, with diameters averaging 0.78 mm by 1.67 mm. Branches are rounded and average 0.48 mm diameter. The reverse side of the colony is striated. The exozone is distinguished by a change in the orientation of the autozooecia and a thickening of the zooecial walls. Rounded zooecial apertures are present on the frontal side of the colony. The apertures average 0.09 mm by 0.12 mm in diameter in the exozone. Three to four zooecial rows occur on each branch. Zooecial walls are thin and straight. Occasional basal diaphragms have been observed which are deflected orally at their junction with the zooecial walls and in some cases have laminae continuous with the zooecial linings. The microstructure is laminar, although hard to distinguish. Pos- sible acanthostyle-like structures are present. REMARKS. The specimen from Clog-y-fran is fragmentary and embedded in sediment, which makes identification difficult. It is characterised by rounded branches and oval fenestrules. Zooecial walls are straight in the endozone, becoming thickened in the exozone. Zooecial apertures are rounded and three to four rows occur on each branch. Occasional diaphragms are present and acanthostyle-like structures have been recognised. Phylloporina hillistensis, described from Estonia by Lavrentjeva (1985: pl. ii, fig. 2), has similar thick straight endozonal walls and occasional diaphragms but differs from the Clog-y-fran specimen in having more abundant zooecia per branch and lacking striae on the reverse of the colony. ACKNOWLEDGEMENTS. I would like to thank Dr. P.D. Taylor and Dr. J.C.W. Cope for supervising this project, which was carried out under the tenure of a Natural Environment Research Council Studentship. I am grateful to Dr. D.H. Evans and Mr. F. Cross for assistance in the field. I would also like to thank Mrs D.G. Evans for drafting Table | and Fig. 1, and Mrs L. Weaver for re-typing the manuscript. REFERENCES Astrova, G.G. 1965. Morfologiiya, istoria razvitiya i sistema ordovikskikh1 siluriiskikh mshanok. Trudy Paleontologicheskogo Instituta Akademiya Nauk SSSR, Moscow, 106: 432 pp, 84 pls (In Russian). 1978. Istoriya razvitiya, sistema i filogeniya mshanok, Otryad Trepostomata. Trudy Paleontologicheskogo Instituta Akademiya Nauk SSSR, Moscow, 169; 240 pp.. 46 pls (In Russian: Engl. transl. by D.A. Brown, 307 pp, no pls.). — & Morozova, IP. 1956. K sistematike mshanok otryada Cryptostomata. Doklady Akademii Nauk Soyuza Sovetskikh Sotsialisticheskikh Republik. St Petersburg, 110: 661-664 (In Russian). Bassler, R.S. 1911.The Early Paleozoic Bryozoa of the Baltic Provinces, Bulletin of the United States National Museum, Washington, 77: 11-137. 1928. Bryozoa. Jn, Twenhofel, W.H., Geology of Anticosti Island, Memoirs of the Geological Survey of Canada, Ottawa, 154: 143-168. Bekker, H. 1921. The kukers stage of the Ordovician rocks of N.E. Estonia. Acta et Commentations Universitatis Tartuensis, Dortpat, A2: 92 pp. Bengston, P. 1988. Open nomenclature. Palaeontology, London, 31: 223-227. Boardman, R.S. 1960. A revision of the Ordovician bryozoan genera Batostoma, Anaphragma and Amplexopora. Smithsonian Miscellaneous Collections, Washing- ton, 140: 28 pp. , Cheetham, A.H., Blake, D.B., Utgaard, J., Karklins, O.L., Cook, P.L., Sandberg, P.A., Lutaud, G. & Wood, T.S. 1983. Bryozoa (revised). Jn, Robison, R.A. (ed), Treatise on Invertebrate Paleontology, G (1): 625 pp, Boulder, Colorado and Lawrence, Kansas. Borg, F. 1926. Studies on recent Cyclostomous Bryozoa. Zoologiska Bidrag fran Uppsala, 10: 181-507. 133 Bork, K.B. & Perry, T.G. 1968. Bryozoa (Ectoprocta) of Champlainian age (Middle Ordovician) from northwestern Illinois and adjacent parts of Iowa and Wisconsin. Part II, Homotrypa, Orbignyella, Prasopora, Monticulipora and Cyphotrypa. Jour- nal of Paleontology, Menasha, 42: 1042-1065, pls 133-138. Brown, G.D. 1965. Trepostomatous Bryozoa from the Logana and Jessamine Lime- stone (Middle Ordovician) of the Kentucky Bluegrass region. Journal of Palaeontology, Tulsa, 39: 974-1006. & Daly, E.J. 1985. Trepostome Bryozoa from the Dillsboro Formation (Cincinnatian Series) of south-eastern Indiana. Special report of the Geological Survey of Indiana. Bloomington, 33: 1-95. Buttler, C.J. 1988. Studies on Ordovician bryozoans from Wales and the Welsh Borderland. Ph.D. thesis. University of Wales (unpublished). 1991a. Bryozoans from the Llanbedrog Mudstones (Caradoc), north Wales. Bulletin of British Museum of Natural History (Geology Series), London, 47: 153- 168. 1991b. Studies on Ordovician bryozoan fauna from the Slade and Redhill Beds, South Wales. Palaeontology, London, 34: 77-108. Coryell, H.N. 1921. Bryozoan faunas of the Stones River Group of central Tennessee. Proceedings of the Indiana Academy of Sciences, Brookville, 1919: 261—340, 14 pls. Cummings, E.R. 1908. The Stratigraphy and Palaeontology of the Cincinnati Series of Indiana. Annual Report. Indiana Department of Geology and Natural Resources, Indianapolis, 32 (for 1907): 605-1188, 55 pls. Dreyfuss, M. 1948. Contribution a l’etude geologique et paleontologique de !’Ordovicien superieur de la Montagne Noir. Memoires de le Societe Geologique de France. Paleontologie, Paris, 58: 63 pp. Dybowski, W. 1877. Die Chaetetiden der ostbaltischen Silurformation. Verhandlungen der Russisch-Kaiserlichen Mineralogischen Gesellschaft zu St. Petersburg, St. Petersburg, 14: 1-134. Ehrenberg. C.G. 1831. /n, Hemprich, FG. & Ehrenberg, C.G., Symbolae physicae, seu icones et descriptiones animalium evertebratorum. 4 (Evertebrate 1). 126 pp., 10 pls. (1828), Stockholm. Eichwald, E. 1856. Beitrag zur geographischen Verbeitung der fossilenThiere Russland. Bulletin de la Societe Imperiale de Naturalistes de Moscou. Moscow, 29: 91-96. — 1860. Lethaea rossica ou Paleontologie de la Russie. Ancienne Periode. Stuttgart, 1: 355-419, 434-435, 450-452, 475495. Elias, R.J. & Condra, G.E. 1957. Fenestella from the Permian of West Texas, Memoirs of the Geological Society of America, Washington, 70: 158 pp. Foerste, A.F. 1887. The Clinton Group of Ohio, Pt.UI, Bulletin of the Scientific Laboratories of Denison University, Granville, Ohio, 2: 140-176. Foord, A.N. 1883. Contribution to the micropalaeontology of the Cambro-Silurian rocks of Canada. Geological and Natural History Survey, Canada, Ottawa, 26 pp. Fortey, R.A., Harper, D.A.T., Ingham, J.K., Owen, A.W. & Rushton, A.W.A. 1995. A revision of the Ordovician series and stages from the historical type area. Geological Magazine, 132: 15-30. James, J.F. 1893. Manual of the palaeontology of the Cincinnati Group. Journal of the Cincinnati Society of Natural History, Cincinnati, 15: 144-159. Karklins, O. 1983. Ptilodichtyoid Cryptostomata Bryozoa from the Middle and Upper Ordovician rocks of Central Kentucky. Journal of Paleontology, Tulsa, 57: (Memoir 14); 1-31. 1984. Trepostomate and cystoporate bryozoans from the Lexington Limestone and Clays Ferry Formation (Middle and Upper Ordovician) of Kentucky. Profes- sional Paper, United States Geological survey, Washington, 1066-1: 105 pp. Lavrentjeva, V.D. 1979. Novii portirad paleozoishich mshanok. Paleontologicheskii Zhurnal, Moscow, 9-68. (In Russian). — 1985. Mshanki Podotryada Phylloporinina. Trudy Paleontologicheskogo Insti- tute. Akademiya Nauk SSSR. Moscow, 214: 102 pp. Miller, S.A. 1889. North American Geology and Palaeontology, Western Methodist Book Concern, Cincinnati, 664 pp. Modzalevskaya, E.A. 1953. Trepostomaty Ordovika Pribaltiki i ikh stratigragicheskoe znachenie. Trudy Veseouznogo Neftyanogo Nauchno-Issledovatel ‘skogo Geologo- Razvedochnogo Instituta, (VNIGRI), Leningrad & Moscow, Y 78: 91-167, 14pls (In Russian). Nekhorsoheva, L.V. 1970. (Ordovician bryozoans from the north of Pai-khoi, Vaigach, and the south of Novaya Zemlya). In, Bondarev, V.I. (ed.), Opornyi razrez Ordovika Pai-khoya, Vaigacha i yugo Novoi Zemli: 63-95, 9 pls. Leningrad, Izd-vo NIIGA, (In Russian). Nicholson, H.A. 1879. On the Structure and Affinities of the ‘Tabulate Corals’ of the Palaeozoic Period with Critical Descriptions of Illustrative Species. 342 pp, Edin- burgh. Orbigny, A, d’ 1850. Prodrome de paleontologie stratigraphique universelle. 1: 394 pp-, Paris. Owen, DE. 1962. Ludlovian Bryozoa from the Ludlow District. Palaeontology, London, 5: 195-212. 1965. Silurian Polyzoa from Benthall Edge, Shropshire. Bulletin of the British Museum of Natural History (Geology), London, 10: 95-117. 1969. Wenlockian Bryozoa from Dudley, Niagara and Gotland and their palaeogeographic implications. Palaeontology, London, 12: 621-636. Pushkin, V.I. 1976. Novye vidy mshank iz Ordovika i Silura brestskoi vpadiny. V Kn.: 134 Novye vidy iskopaemykh zhivotnykh i rastenii Belorussii. Nauka i tekhika, Minsk, 40 pp. (In Russian). Ropot, I.V. & Pushkin, V.I. 1987. Ordivik Belorussky. 234 pp., 23 pls. Minsk, Institut Geochmimii i GeofizikiAcademii Nauk Belorusskoy S.S.R. & Belorusskiy Nauchno- Issledovatel’skiy Geologorazvedochniy Institut. (In Russian). Ross, J.R.P. 1963. Trepostome Bryozoa from the Caradoc Series, Shropshire. Palaeon- tology, London, 6: 1-11. 1965. Homotrypa and Amplexopora? from the Caradoc series, Shropshire. Palaeontology, London, 8: 5-10. Scotese, C.R. & McKerrow, W.S. 1990. Revised world maps and introduction. Jn, McKerrow, W.S. & Scotese, C.R. (eds), Palaeozoic palaeogeography and biogeogra- phy. Memoir of the Geological Society of London, 12: 1-12. Spjeldnaes, N. 1957. A redescription of some type specimens of British Ordovician Bryozoa, Geological Magazine, Hertford, 94: 364-376, pls. 12-13. 1963. Some silicified Ordovician fossils from south Wales. Palaeontology, London, 6: 254-263, pls.36—37. Strahan, A., Cantrill, T.C., Dixon, E.E.L. & Thomas, H.H. 1909. The geology of the South Wales coal-field, Part X, The county around Carmarthen. Memoirs of the Geological Survey, London, 229: 177 pp. Taylor, P.D. & Cope, J.C.W. 1987. A trepostome bryozoan from the Lower Arenig of South Wales: implications of the oldest described bryozoan. Geological Magazine, Cambridge, 124: 367-371. Toots, H. 1952. Bryozoan des estruschen Kukersits. Mitteilungen aus dem geologischen Staatsinstitut in Hanburg, 21: 113-137. Ulrich, E.O. 1879. Descriptions of anew genus and some new species of Bryozoa from the Cincinnati Group. Cincinnati Society of Natural History Journal, 2: 119-131, pl. xii. 1882. American Palaeozoic Bryozoa. Cincinnati Society of Natural History Journal, 5: 121-175, 232-257. 1890. Palaeontology of Illinois. Section VI. Palaeozoic Bryozoa, Geological Survey of Illinois, Geology and Palaeontology, Chicago, 8: 283-688, pls. 29-78. 1893. On Lower Silurian Bryozoa of Minnesota. Minnesota Geological and Natural History Survey, Final Report, 3: 96-332. & Bassler, R.S. 1904. A revision of the Palaeozoic Bryozoa, Pt. Il — on genera and species of Trepostomata, Smithsonian Miscellaneous Collections, Washington, 47: 15-55. Vine, G.R. 1884. Fourth report of the Committee appointed for the purpose of reporting on fossil Polyzoa. Report of the British Association for the Advancement of Science 1883, London, 161-209. TREPOSTOME IDENTIFICATION KEY In order to identify trepostome bryozoans from Clog-y-fran thin sections are needed. Ideally at least two oriented sections (longitudi- nal and tangential) are required. However, specimens can sometimes be identified from randomly oriented sections. Identification can be hindered when specimens are fragmented and abraded. The key aims to aid identification, but results should be carefully checked against the complete descriptions and the illustrations of the species. 1. Zoaria massive and hemispherical i2 ZR ATIANCTO CE sca cease cea cectineSces cat adie Soe teers eae ates nantes een ad ear CHU 3 10. Hil. 14. C. BUTTLER Ring diaphragms present . Hemiphragma sp Ring diaphragms and mesozooecia absent ................4+- Monotrypa sp Hemiphragms present Ia (S049) AVEYSTROTS GIDSTESAL | se ccacoecacoeroasonecandocctucenosceedne caso sporeccScossoaceecsadeououc Mesozooecia abundant with numerous diaphragms along their length af caves sayccqtemeeseaneataze rvarautnas dune dn dese catines meateuceeaneeees Dittopora sanclerensis Mesozooecia present, but not common. ...... Hemiphragma pygmaeum Cystiphrasms|presemtrese stot seeeee ee era cee eee eee ere ae 6 Gystiphra smStaDSen tye snc: cvecsects ss dees asp usec seuaes wesc eee ere eee ee one 7 Abundant acanthostyles present ............ Monticulipora aff. compacta Small indistinct acanthostyles present .............. Homotrypa cf. similis INGEVNINOS HPS [OSS cccesonecccececo scenes cooseo sno Lee so SaoeReeee LALoScEboDRO ASN SOSEOS 8 AcanthostylestabSemtieeee: seeere seen este ee eee eee 15 DiaphrapmsirareinlautaZOOcclabereseeeee eter eee een ene 9 Diaphragms abundant in autoZO0ecia ..............eeceeseeeeeeeeeeeeeseesenees 12 Mesozooecial walls constricted at the position of the diaphragms pro- ducingia beaded appearance see ee eee eee Hetrotrypa sp Mesozooecial walls straight in appearance ..............:eseeseeseeeeeeeeeees 10 BYPAVINS ASS 333) AEN THM CHANTS cpopceccacosco ecocc cceccecercoseoscoscascoosece/nsoscets 11 Branches <3 mm in diameter ...........:..:c:ceseeseeceeeereee Leioclema sp. B Acanthostyles composed of broad hyaline core with no sheathing TAMMAIT AC iy set cdaek pace veereseestea see hes vb eatesseuesvecateeecesmeenseeeee Leioclema? sp. Acanthostyles composed of broad hyaline core surrounded by steeply dippinpiconicalilaminde lesen eee ene Leioclema sp. A Autozooecial basal diaphragms very abundant and regularly spaced {BaTCOYUYSA NOBUO STINT cecaeconaencocosecscneenceuccnencoacoseccadeab bc30 Amplexopora sp. Autozooecial basal diaphragms present throughout colony but more abundantymngthe/exOZOme peeceeaeeeee eentee teeter see eet 13 Meandering autozooecia roughly parallel to branch axis in endozone then curving very slightly to meet zoarial surface ..............::.:ssceeeeee aukestauccead kieee totes as teetoeeeack cua secsuagh eect omer ne ay Eridotrypa simulatrix Straight autozooecia roughly parallel to branch axis in endozone curve to;mectizoariall SUGLACE: ..2-<.cccewcscuss «hevses-au-crseeeun eee eee eee 14 Thick exozonal walls with abundant diaphragms sa baeee vices acvnts doe inst eden essa aan socd eee Batostoma cf. polare QUST WASE as. eee. ceece nce coccesc hens c te sesegee sc sateccnee Batostoma clogyfranense MesoZO0€ClaNpTeSe mb ivsicseness ssessceeesbslencanesecsbessvtsesseuntataavusteeesseenteataaes 16 Mesozooecia absent, exilazooecia present . Halloporina cf. crenulata Mesozooecia beaded in appearence ................... Hallopora peculiaris Mesozooecia straight-walled .............. Hallopora aff. wesenbergiania Bull. nat. Hist. Mus. Lond. (Geol.) 53(2): 135-138 Issued 27 November 1997 New information on Cretaceous crabs C.W. WRIGHT The Old Rectory, Seaborough, Beaminster, Dorset DT8 3QY SYNOPSIS. Re-examination of the supposedly Jurassic, Tithonian crab fauna from Klement in Austria shows that it is Cretaceous, Cenomanian, thus removing the puzzling record of Diaulax from the Jurassic. A new species of Paranecrocarcinus is described from the Lower Cretaceous, Barremian of Zululand, South Africa. New material from the English Lower Cretaceous is described, including a new species of Rathbunopon from the Lower Aptian and important new information about Withersella. THE KLEMENT ‘TITHONIAN’ CRAB FAUNA In 1931 Glaessner listed a small fauna of Crustacea from a block of presumed Tithonian limestone in a conglomerate at Klement in Lower Austria. It is of importance because it included a species of Diaulax, a relatively advanced genus otherwise known only from the Cretaceous, Lower Albian to Cenomanian. Shortly before his death Glaessner entrusted me with his Klement specimens with a view to joint description, since he had doubts about their Jurassic date. These doubts were fully justified, since revised identifications indi- cate that the fauna is almost certainly of Cenomanian date. The original (Glaessner, 1931) and the new identifications are: Original: Revised: Prosopon verrucosum Reuss Rathbunopon obesum (Van Straelen) Pithonoton marginatum Meyer Pithonoton cenomanense Wright & Collins Cyphonotus oxythyreiformis (Gemmellaro) Diaulax sp. Palaeodromites incertus (Bell) Diaulax oweni (Bell) The *Prosopon verrucosun’ (BMNH IC 6, Fig. 1) resembles very closely the fragmentary English Cenomanian specimen identified by Wright & Collins (1972: 23, pl. 1, fig. 8) as Rathbunopon obesum (Van Straelen), a species originally described from the Cenomanian of Navarre, Spain. A second, minute, specimen (BMNH IC 14, Fig. 2) probably belongs to the same species but is too juvenile for certain attribution. The ‘Pithonoton marginatun’ (BMNH IC 17, Fig. 3) conforms well with P. cenomanense Wright & Collins in the outline of the cephalothorax, the course of the cervical and branchiocardiac grooves, and the disposition of the granules. Differences between species of Pithonoton are generally fine, but identity with P cenomanense seems highly probable. The ‘Cyphonotus oxythyreiformis’ (BMNH IC 8, Fig. 4), though incomplete, is beautifully preserved and is undoubtedly identical with Palaeodromites incertus, of which an English specimen of the same size is figured for comparison (Fig. 5). Species of Palaeodromites were shown by Wright & Collins to have a short range in the Cretaceous and this Klement specimen alone is suffi- cient to demonstrate the Cenomanian age of the fauna. The ‘Diaulax sp.’ (BMNH IC 7, Fig. 6) is certainly a Diaulax and differs in no way from the abundant English material of D. oweni from the Lower Albian to the Cenomanian. The immediate ancestor of D. oweni has not been identified but Wright & Collins (1972: 55) referred to the origin of Diaulax in “broad flat species of Pithonoton’ ; © The Natural History Museum, 1997 they commented (p. 56) on the supposed Upper Jurassic occurrence from Klement as representing ‘a very early development of a relatively advanced carapace form’, an anomaly now removed by the revised dating of the Klement fauna. A NEW SPECIES OF PARANECROCARCINUS FROM THE BARREMIAN OF ZULULAND Genus Paranecrocarcinus Van Straelen, 1936 TYPE SPECIES. Paranecrocarcinus hexagonalis Van Straelen, 1936 (p. 36, pl. 4, figs. 6, 7) from the Hauterivian of Auxerre, France, by monotypy. DISCUSSION. Wright & Collins (1972) differentiated Paranecrocarcinus from Necrocarcinus by the bifid rostrum of the former and the trifid rostrum of the latter. They then united Forster’s (1968) Protocarcinus, as a synonym, and Pseudonecrocarcinus as a subgenus of Paranecrocarcinus, separating the two subgenera partly on the basis that P (Paranecrocarcinus) did not have and P. (Pseudonecrocarcinus) did have post-rostral slits in the carapace. This distinction was false, since the type species P. hexagonalis does have post-rostral slits. The remaining diagnostic character of Pseudonecrocarcinus, the many small rounded tubercles on the surface of the carapace, as seen both in the Maastrichtian type species P. (Pseudonecrocarcinus) quadriscissus (Noetling) and in the Cenomanian P. (P) biscissus Wright & Collins, might be thought sufficient to justify the two subgenera. However, some doubt is cast on this idea by the juvenile specimen of P. biscissus discussed in the last section of this paper below. Provisionally I am inclined to abandon the distinction of two subgenera. Paranecrocarcinus kennedyi sp. nov. Figs 7, 13 NAME. For Professor W J Kennedy who found the specimen. HOLOTYPE. BMNH IC 16, from the Barremian Makatini Forma- tion, Mlambongwenya Spruit, Zululand, South Africa. DIAGNOsIS. A Paranecrocarcinus with a transverse row of nine tubercles across the gastric regions and a single one on each metabranchial lobe and with two prominent spines on the anterola- teral border; apparently without post-rostral slits. DESCRIPTION. The holotype consists of an internal mould with the rostrum and margins only partially preserved, together with the counterpart showing the central area of the cephalothorax in hard 136 C.W. WRIGHT : 6 a “i a Figs 1,2 Rathbunopon obesum (Van Straelen). Limestone boulder in conglomerate, Klement, Lower Austria, Cenomanian. 1, BMNH IC6; 1a, upper, 1b, right side, x2; 2, BMNH IC 14, 2a, upper, 2b, right side, x4. Fig. 3 Pithonoton cenomanense Wright & Collins. As for Figs. 1, 2. BMNH IC 17. x2. Figs 4,5 Palaeodromites incertus (Bell). 4, as for Figs. 1, 2, BMNH IC 8, x2. 5, Cenomanian Sands, Lower Cenomanian, Mantelliceras dixoni Zone, White Hart Pit, Wilmington, Devon, BMNH IC 9, x2. Fig.6 Diaulax oweni (Bell). As for Figs. 1, 2. BMNH IC 7; 6a, upper, 6b, right side, x2. Fig. 7 Paranecrocarcinus kennedyi sp.nov. Holotype. Makatini Formation, Barremian, Mlambongwenya Spruit, Zululand, South Africa. BMNH IC 16; 7a, upper, 7b, front, x2. Fig. 8 Galathea sp. Lower Greensand, Crackers Bed, Deshayesites forbesi Zone, Atherfield, Isle of Wight. BMNH IC 13, x4. Fig.9 Rathbunopon ? atherfieldense sp.nov. Holotype. As for Fig. 8. BMNH IC 11; 9a, upper, 9b, front, x4. Fig. 10 Paranecrocarcinus biscissus Wright & Collins. Cenomanian Limestone, Bed A or B, Whitecliff, Seaton, Devon. BMNH IC 10, x2. Fig. 11 Paranecrocarcinus digitatus Wright & Collins. As for Fig. 5. BMNH IC 5; 11a, upper, 11b, front, x2. Fig. 12 Withersella crepitans Wright & Collins. As for Fig. 8. BMNH IC 15, upper, x2. NEW INFORMATION ON CRETACEOUS CRABS 14 16 Fig. 13. Paranecrocarcinus kennedyi sp.nov. Makatini Formation, Barremian, Zululand, South Africa. Reconstruction, based on the holotype, Fig. 7. x ca. Bi Fig. 14 Rathbunopon ? atherfieldense sp.noy. Lower Greensand, Crackers Bed, Lower Aptian, Atherfield, Isle of Wight. Reconstruction, omitting the granulation, based on the holotype, Fig. 9, and paratype. The orbito-frontal margins are diagrammatic, since details of fissures and teeth are not preserved. x ca. 8. Fig. 15 Paranecrocarcinus biscissus Wright & Collins. Cenomanian Limestone, Whitecliff, Seaton, Devon. Diagrammatic reconstruction, based on specimen in Fig. 10. x ca. 4. Fig. 16 Withersella crepitans Wright & Collins. Lower Greensand, Crackers Bed, Lower Aptian, Atherfield, Isle of Wight. Diagram of left frontal margin, based on specimen in Fig. 12. x ca. 5. matrix and visible only from underneath. The cephalothorax is roughly pentagonal in outline with slightly convex anterolateral, straight posterolateral and slightly concave posterior margins. It is weakly arched in transverse and longitudinal sections, with appar- ently deeply undercut sides. The front is produced into a broad sulcate rostrum, incompletely preserved but showing upwardly directed rostral spines. The orbital margins are not well-preserved but the orbits appear to have been moderately wide with a fissured upper rim and an outer orbital spine; the orbito-frontal width was about half that of the carapace. The anterolateral margin ends in a spine at the lateral angle, and there is one between this and the outer orbital spine. The long posterolateral margins are almost straight and converge towards the slightly concave posterior margin. The cervical sulcus is bent strongly round the rear of the mesogas- tric lobe and then takes a sinuous oblique course to the margins. Distinct epibranchial sulci branch obliquely to the rear and define small triangular epibranchial lobes. The branchiocardiac sulci are weaker than the cervical and are more or less parallel to it in their outer part; they run back between the small triangular cardiac lobe and a small parallel ridge on either side. DISCUSSION. The Hauterivian P. hexagonalis has two large tuber- cles on the mesogastric lobe but no others forward of the cervical sulcus and has a pair of post-rostral slits. The Cenomanian P. mozambiquensis Forster, 1970, has a single large tubercle on each protagastric lobe. P. libanoticus Forster, 1968, from the Cenomanian of Lebanon has a single small tubercle on the mesogastric lobe, two large ones on each protogastric and two smaller ones on each anterior branchial lobe; it also has two post-rostral slits. The Turonian Povalis Stenzel from Texas has an aligned row of large tubercles across the hepatic and protogastric lobes as in P. kennedyi, but the cephalothorax is much broader than long and has a less pentagonal outline, as does the Upper Albian P. graysonensis Rathbun, 1935 with weaker tuberculation. P digitatus Wright & Collins, 1972, from the English Cenomanian has a pair of post-rostral slits (Collins et al., 1995: 198), and is characterised by its elongated radiating ridges on the protogastric lobes. P. foesteri Wright & Collins, 1972, differs in having strongly granulated posterolateral margins and posterior edges of the branchiocardiac furrows. However, none of these species is known by more than a very few specimens and the extent of intraspecific variation is unknown. SOME NEW ENGLISH CRETACEOUS CRABS Fig. 8 A poorly preserved Galathea has been found in Lower Aptian Crackers material from Atherfield, supplied by Prof. W.J. Kennedy. It is inadequate for proper description, but it is worth recording since, with a specimen from the Aptian of Spain (Via Boada, oral communication), it is probably the oldest known species of the genus. Galathea sp. nov.? Rathbunopon? atherfieldense sp. nov. Figs 9, 14 TYPES. The holotype is BMNH IC 11 and paratype IC 12, both from Lower Greensand, Crackers Bed, Lower Aptian, D. callidiscus Zone, Atherfield Point, Isle of Wight. DIAGNOSIS. A presumed primitive Rathbunopon, longer than wide, with the paired bosses at the rear of the mesogastric lobe small and close together; the urogastric lobe feebly developed, divided by a shallow longitudinal groove. DESCRIPTION. Small, 7 mm long, about 25% longer than wide, narrowed in front and with slightly convex margins; strongly arched in transverse section, less so in longitudinal; front turned down and deeply furrowed; orbitofrontal margins oblique at about 45°. The furrows delimiting the mesogastric lobe are shallow in front but deepen as they approach the cervical furrow, which is wide and deep laterally. The branchiocardiac furrows are shallower than the cervi- cal. There is a strong circular epigastric boss on either side of the medial furrow, a feeble longitudinal oval one on the anterior process 138 of the mesogastric lobe, a large round one on the middle of the lobe and a transverse pair of small ones to the rear; there is a large round boss on the middle of each hepatic lobe and an outer small one, all forming a transverse line with the anterior mesogastric boss. All the raised areas of the cephalothorax have well-separated small granules between the bosses. DISCUSSION. The holotype is incomplete, lacking nearly all of the frontal and lateral margins. The arrangement of the lobes, except for the urogastric which is weakly bilobed longitudinally rather than divided transversely into two bars, is close to that of R. polyakron Stenzel and R. woodsi Withers. The epigastric, mesogastric and hepatic bosses also are similar in disposition. It is highly probable that the present species is a primitive Rathbunopon but in the absence of evidence of the characteristic orbits attribution must remain uncertain. There is some resemblance to the fragmentary holotype of the Hauterivian Homolopsis tuberculata Van Straelen, 1936, which may also be a Rathbunopon. Paranecrocarcinus biscissus? Wright & Collins, 1972 Figs 10, 15 An incompletely preserved internal mould from the Cenomanian of Whitcliff, Seaton, Devon (BMNH IC 10) has the same arrangement of outer orbital spines and fissures as P. biscissus Wright & Collins (1972: text-fig. 10b) and a multiplicity of small tubercles, including three on the urogastric lobe. However the number and arrangement of the other tubercles is not exactly as in the holotype of P. biscissus. The present specimen has an estimated breadth of 9 mm, against 12 mm of the holotype, and is probably an earlier moult of the same species. Paranecrocaricinus digitatus Wright & Collins, 1972 Fig. 11 A further specimen from Wilmington (BMNH IC 5) confirms the restoration given by Wright & Collins (1972: text-fig. 10a). Hemioon elongatum (Milne-Edwards, 1862) A poorly preserved specimen has been found in Bed C of the Devon Cenomanian Limestone, thus extending the range of this species to the Calycoceras guerangeri Zone of the Upper Cenomanian. C.W. WRIGHT Withersella crepitans Wright & Collins, 1972 Figs 12, 16 Wright & Collins (1972: 91) established W.crepitans on the basis of 14 specimens of a delicate crab from the Crackers Bed at Atherfield. They gave a restored diagrammatic view of the cephalothorax showing the frontal margin with broad rectangular indentations and teeth. Subsequently a specimen was found (BMNH IC 15) with the left frontal margin almost perfectly preserved indicating that the diagram in Wright & Collins was based on a broken edge of the thin carapace. The actual frontal margin (Figs 12, 16), is bounded by large outer orbital spines and is rather concave, interrupted only by paired oblique supraorbital fissures and a marked inner orbital spine on either side of a bifid rostrum. In effect the front of Withersella is extremely close to that of Carcineretes walcotti Withers, except for the greater projection of the rostrum in Withersella, thus confirming the attribution to Carcineretidae by Wright & Collins, which Glaessner (1980: 180) had regarded as unconvincing. Also, the front of Withersella more closely resem- bles that of Binkhorstia than was apparent in 1972, although there are significant differences in the latter’s peculiar spatulate rostrum, third supraorbital fissure and less oblique fissures (Collins, Fraaye & Jagt, 1995: figs 12a-c). REFERENCES Bell, T. 1863. A monograph of the fossil malacostracous Crustacea of Great Britain. Part II, Crustacea of the Gault and Greensand. Monograph of the Palaeontographical Society of London. viii + 40 pp., 11 pls. Collins, J.S.H., Fraaye, R.H.B. & Jagt, J.W.M. 1995. Late Cretaceous anomurans and brachyurans from the Maastrichtian type area. Acta Palaeontologica Polonica, 40: 165-210, 12 figs. Glaessner, M.F. 1931. Geologisches Studien in des aiisseren Klippenzone. Jahrbuch der geologischen Bundesanstalt, Wein, 81: 1-23. Forster, R. 1968. Paranecrocarcinus libanoticus n. sp. (Decapoda) und die Entwicklung der Calappidae in der Kreide. Mitteilungen der Bayerischen Staatssammlung fiir Paldontologie und historische Geologie, 8: 167-195, pl. 13. 1970. Neue Decapoden Reste aus der Oberkreide von Mocgambique, Norddeutschland und den bayerischen Alpen. Paldontologische Zeitschrift, 44: 134— 144, pl. 17. Stenzel, H.B. 1945. Decapod crustaceans from the Cretaceous of Texas. Bulletin of the University of Texas Bureau of economic Geology and Technology, 4401: 401-476, pls. 34-45. Withers, T.H. 1928. New Cretaceous crabs from England and Syria) Annals and Magazine of Natural History, (10) 2: 456-462, pl. 13. Wright C.W. & J.S.H. Collins, 1972. British Cretaceous Crabs. Monograph of the Palaeontographical Society of London. 114pp., 22 pls. CORRIGENDUM Correction to: Barrett, P.M. 1996. The first known femur of Hy/aeosaurus armatus and re-identification of ornithopod material in the Natural History Museum, London. Bull. nat. Hist. Mus. Lond. (Geol.) 52 (2): 115-118. In the Synopsis (p. 115), line 1, the words ‘ornithopod dinosaur’ should be replaced by ‘nodosaurid ankylosaur’. Bulletin of The Natural History Museum Geology Series Earlier Geology Bulletins are still in print. The following can be ordered from Intercept (address on inside front cover). Where the complete backlist is not shown, this may also be obtained from the same address. Volume 34 No. 1 No. 2 No. 3 No. 4 Volume 35 No. 1 No. 2 No. 3 Relative dating of the fossil hominids of Europe. K.P. Oakley. 1980. Pp. 1-63, 6 figs, 17 tables. £8.00 Origin, evolution and systematics of the dwarf Acanthoceratid Protacanthoceras Spath, 1923 (Cretaceous Ammonoidea). C.W. Wright & W.J. Kennedy. 1980. Pp. 65-107, 61 figs. £6.25 Ashgill Brachiopoda from the Glyn Ceiriog District, north Wales. N. Hiller. 1980. Pp. 109-216, 408 figs. £14.75 Miscellanea Type specimens of some Upper Palaeozoic Athyridide brachiopods. C.H.C. Brunton. 31 figs. 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Pp. 1-30. 11 figs. 0 565 07015 0. £5.50 Lower Turonian (Cretaceous) ammonites from south-east Nigeria. P.M.P. Zaborski. 1987. Pp. 31-66. 46 figs. 0 565 07016 9. £6.50 The Arenig Series in South Wales: Stratigraphy and Palaeontol- ogy. I. The Arenig Series in South Wales. R.A. Fortey & R.M. Owens. II. Appendix. Acritarchs and Chitinozoa from the Arenig Series of South-west Wales. S.G. Molyneux. 1987. Pp. 67-364. 289 figs. 0 565 07017 7. £59.00 Miocene geology and palaeontology of Ad Dabtiyah, Saudi Arabia. Compiled by P.J. Whybrow. 1987. Pp. 365-457. 54 figs. 0 565 07019 3. £18.00 Cenomanian and Lower Turonian Echinoderms from Wilmington, south-east Devon. A.B. SMith, C.R.C. Paul, A.S. Gale & S.K. Donovan. 1988. 244 pp. 80 figs. 50 pls. 0 565 07018 5. £46.50 Volume 43 No. 1 A Global Analysis of the Ordovician—Silurian boundary. Edited by L.R.M. Cocks & R.B. 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Pp. 1-163. 0 565 07025 8. £40.00 No. 2 A review of the Tertiary non-marine molluscan faunas of the Pebasian and other inland basins of north-western South America. C.P. Nuttall. 1990. Pp. 165-371. 456 figs. 0 565 07026 6. £52.00 Volume 46 No. | Mid-Cretaceous Ammonites of Nigeria—new amphisbaenians from Kenya—English Wealden Equisetales—Faringdon Sponge Gravel Bryozoa. 1990. Pp. 1-152. 0 565 070274. £45.00 No. 2 Carboniferous pteridosperm frond Neuropteris heterophylla— Tertiary Ostracoda from Tanzania. 1991. Pp. 153-270. 0565 07028 2. £30.00 Volume 47 No. 1 Neogene crabs from Brunei, Sabah & Sarawak—New pseudosciurids from the English Late Eocene—Upper Palaeozoic Anomalodesmatan Bivalvia. 1991. Pp. 1-100. 0 565 07029 0. £37.50 No. 2 Mesozoic Chrysalidinidae of the Middle East—Bryozoans from north Wales—Alveolinella praequoyi sp. nov. from Papua New Guinea. 1991. Pp. 101-175. 0 565 070304. £37.50 Volume 48 No. | ‘Placopsilina’ cenomana d@ Orbigny from France and England—Revision of Middle Devonian uncinulid brachiopod—Cheilostome bryozoans from Upper Cretaceous, Alberta. 1992. Pp. 1-24. £37.50 No. 2 Lower Devonian fishes from Saudi Arabia—W.K. Parker’s collection of foraminifera in the British Museum (Natural History). 1992. Pp. 25-43. £37.50 Volume 49 No. 1 Barremian—Aptian Praehedbergellidae of the North Sea area: a reconnaissance—Late Llandovery and early Wenlock Stratigraphy and ecology in the Oslo Region, Norway— Catalogue of the type and figured specimens of fossil Asteroidea and Ophiuroidea in The Natural History Museum. 1993. Pp. 1-80. £37.50 No. i) Volume 50 No. 1 No. 2 Volume 51 No. 1 No. 2 Volume 52 No. 1 No. 2 Volume 53 No. 1 Mobility and fixation of a variety of elements, in particular, during the metasomatic development of adinoles at Dinas Head, Cornwall—Productellid and Plicatiferid (Productoid) Brachiopods from the Lower Carboniferous of the Craven Reef Belt, North Yorkshire—The spores of Leclercqia and the dispersed spore morphon Acinosporites lindlarensis Riegel: a case of gradualistic evolution. 1993. Pp. 81-155. £37.50 Systematics of the melicerititid cyclostome bryozoans; introduction and the genera Elea, Semielea and Reptomultelea. 1994. Pp. 1-104. £37.50 The brachiopods of the Duncannon Group (Middle-Upper Ordovician) of southeast Ireland. 1994. Pp. 105-175. £37.50 A synopsis of neuropteroid foliage from the Carboniferous and Lower Permian of Europe—The Upper Cretaceous ammonite Pseudaspidoceras Hyatt, 1903, in north-eastern Nigeria—The pterodactyloids from the Purbeck Limestone Formation of Dorset. 1995. Pp. 1-88. £37.50 Palaeontology on the Qahlah and Simsima Formations (Cretaceous, Late Campanian-Maastrichtian) of the United Arab Emirates-Oman Border Region—Preface—Late Cretaceous carbonate platform faunas of the United Arab Emirates-Oman border region—Late Campanian-Maastrichtian echinoids from the United Arab Emirates-Oman border region—Maastrichtian ammonites from the United Arab Emirates-Oman border region—Maastrichtian nautiloids from the United Arab Emirates-Oman border region—Maastrichtian Inoceramidae from the United Arab Emirates-Oman border region—Late Campanian-Maastrichtian Bryozoa from the United Arab Emirates-Oman border region—Maastrichtian brachiopods from the United Arab Emirates-Oman border region—Late Campanian-Maastrichtian rudists from the United Arab Emirates-Oman border region. 1995. Pp. 89-305. £37.50 Zirconlite: a review of localities worldwide, and a compilation of its chemical compositions—A review of the stratigraphy of Eastern Paratethys (Oligocene—Holocene)—A new protorichthofenioid brachiopod (Productida) from the Upper Carboniferous of the Urals, Russia—The Upper Cretaceous ammonite Vascoceras Choffat, 1898 in north-eastern Nigeria. 1996. Pp. 1-89. £43.40 Jurassic bryozoans from Balt6w, Holy Cross Mountains, Poland—A new deep-water spatangoid echinoid from the Cretaceous of British Columbia, Canada—The cranial anatomy of Rhomaleosaurus thorntoni Andrews (Reptilia, Plesiosauria)—The first known femur of Hylaeosaurus armatus and re-identification of ornithopod material in The Natural History Museum, London—Bryozoa from the Lower Carboniferous (Viséan) of County Fermanagh, Ireland. 1996. Pp. 91-171. £43.40 The status of ‘Plesictis’ croizeti, ‘Plesictis’ gracilis and ‘Lutra’ minor: synonyms of the early Miocene viverrid Herpestides antiquus (Mammalia, Carnivora)—Baryonyx walkeri, a fish- eating dinosaur from the Wealden of Surrey—The Cretaceous- Miocene genus Lichenopora (Bryozoa), with a description of a new species from New Zealand. 1997. Pp. 1-78. £43.40 CONTENTS 79 Ordovician trilobites from the Tourmakeady Limestone, western Ireland J.M. Adrain and R.A. Fortey 117 Ordovician Bryozoa from the Llandeilo Limestone, Clog-y-fran, near Whitland, South Wales C. Buttler 135 New Information on Cretaceous crabs C.W. Wright Bulletin of The Natural History Museum GEOLOGY SERIES Vol. 53, No. 2, November 1997 sos Riley Dunn & Wilson Ltd EXPERT CONSEFVATORS & BOOKBINDER'S sapeea shar eae sires z oo " siete = cries oe eens Z z cea = = 5 . pists oe Sears wees’ ee : E Se one eraraesenoa Sth ‘¢ = <5 i. mae ee = a oo ; = etre - lieteseet : ie ae Faris