Los ‘ ct abe pues, ua - isis ete See fest ae Sree 5 3 zf BG ona iors ie eee: Ze = : eee . Ze S Sai sin sooth eran g 2 aces Sr apa ete Sey sith . i wn] | SBE). |OZO THE NATU | HISTORY MUSEUN 12 } PRESENTED Geology Series HISTORY MUSEUM VOLUME 50 NUMBER 1 23 JUNE 1994 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. 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: (0264) 334748 Fax: (0264) 334058 World List abbreviation: Bull. nat. Hist. Mus. Lond. (Geol.) © The Natural History Museum, 1994 Geology Series ISSN 0968-0462 Vol. 50, No. 1, pp. 1-104 The Natural History Museum Cromwell Road London SW7 5BD Issued 23 June 1994 Typeset by Ann Buchan (Typesetters), Middlesex Printed in Great Britain at The Alden Press, Oxford Bull. nat. Hist. Mus. Lond. (Geol.) 50(1): 1-103 Issued 23 June 1994 Systematics of the melicerititid cyclostome bryozoans; introduction and the genera Elea, Semielea and Reptomultelea PAUL D. TAYLOR Department of Palaeontology, The Natural History Museum, Cromwell Road, London SW7 5BD, UK CONTENTS i err ee Introduction | ust0 NATURAL 2 IRASIODy Oli TESSEMED, cccsnpessedsonopsocsapab adoocobeascSsonCoReAssERbEboCondEoKRccoDed HebBorcbaec : RY. MUSEUM" ABs. toodwosess 2 IMI@7e] NOLO). cos6053 cnanosébackobbaoddapadsusssaqnDbeadpSroniancaseeaeessbsondanceor sods Geababe6 feb! 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GHARHOAE CUOMO) cossocosccobeeendcaosisc c0ccos SABBBEBBDND Go ad965~00n006 ocaccuobedbe oeacoouEdegncoCeeLboocdodseaeooLbuace0c 45 IREDIOTAANAIED, 3a0c-nqneseee7c0!.a20086608000nIee2or 6p ap0r1or Codes EEEEeUECEEOODEEC aobd0dgqcAbcdCcndeedsoceOoduGECe cor sco syoboseesosooo0C 52 46 IR: SEITE AIS (GOTT TSN) caaecqedhocodoonenecesc ceo cose paeBEeE-B=c16000¢ 0 JonneG3905 0 s90909nuEandso255etaresenA93gz4dueGecaaace9C 49 12S CARES 0 SOX scaceoocepecceaaccecdhb0c-000e0800e0 Seg gdo050 00080086 40eo ce Borodin scisqe SecA coséedechnar 9 BeSceCos30 00000000 54 RIRCUAT ISIS PPTIOV a rete teenarese socee soeee/sticise slo onlsaste vin wos edleieetacp en sae eis ses oan eerenis eine eetseen aeromite-ceelefin aneie vases alse 57 TRE INAV OL WO) tis INO ose chorasnsnnecacussasa-celnndoscaBOBeppe: cc cnsoddoorecdace ten aoe Claes Seamer wieiters Mo anae wawemeeesicecks vee 59 Rip DIELLD CF OSGIS Pa TOW ogee ay erste Seieei-d= = oeislelda sais: coe > vse romeb tence de (aid et ed ania dean's ileble\iob oneness seein sis 63 TRS EETIEE (OSM@OIEQN)" Cobc eoseec necdns eect adedos es ee sekececH bec SeBondoC bricdeebeE euctocceckis een code dar pe omace: occcUdr oqagEooeroccq sca 65 IR, CONVEX SP) DOV = ceaeeence eno an ee rnrdehennosssesdacesccoeseasenesesi cme sseiin ohare odsiaissabssin>oasnssceraeascrmavesieriiaaci 66 [RIC ieqoy a (WUETIY)’ “aeedebccdecodeder Soe RRa+Apoaes se eneeedeee seeeean. oo S000 256s Jodaadde6 aAisdaccee o-onsosseapo crane Hoda saastC ONO 1995002 69 RenfilioZati (IMeViNSCU) eens ara esess cocececgsccandedte des - once e eee eae assay oases ose sees ese soe eaasar semen 71 R. gold fussi Sp. DOV. ...............000cccscccecsecececrensnesseccccceecesersnnnrsccecessscesecesesseerecsccsesecesannaneecececesens 73 [Rol BMRBSGIS OS TO\ie tesoccecedocnacoseocecbeeseece506d00800 268606 ce rod doddloroneigadan Hada odosseselee seesoson soouoruccooalse 74 R. MAtUtina SP. NOV. ......-.0222200ceccreeeeeeceeeecsnnencceccccenaneceseccecscaneccesssescaesnesseccessceeeensececesececaneeeeers 77 IRUTMIUSISPEMIOVS sete ertecceesecacrecccecccsc-ccccccncaa41-73 mm ca 1:28 mm Elea hexagona d Orbigny, 1853 Figs 49-56 1853 Elea hexagona d’Orbigny: 633, pl. 738, figs 1-4. 1889 Elea hexagona d’Orbigny; Pergens: 398. non 1897 6 Elea hexagona d’Orbigny; Canu: 753. 1899 Elea hexagona (d’Orbigny); Gregory: 303. 1912 Meliceritites hexagona (d’Orbigny); Levinsen: 43, pl. 5, figs 3-5. MATERIAL. Lectotype (herein designated): MNHN d’Orbigny Collection 8192 (Fig. 49), fragment in tube labelled ‘Type’ by E. Voigt (Voigt photocard 5791), Senonian [Santonian], Vendome, Loir-et-Cher, France. Paralectotype: one of the two other fragments registered with lectotype as MNHN 8192 (the third fragment is ?Foricula). Other mate- rial: ZMC Levinsen Colln M38 and M39, Santonian, Ven- déme, France; ZMC Levinsen Colln, about ten un-numbered topotypes. DESCRIPTION. Colony bifoliate with branches (Figs 49-50) bifurcating, strap-like, narrow, about 1-7-2:3 mm wide. Autozooids usually arranged in approximate quincunx; orga- nization fixed-walled. Colony base not observed. Over- growths produced by eruptive budding onto the branch surface present. Autozooids (Fig. 51) moderately large; frontal surface usually hexagonal with edges parallel to long axis or diamond-shaped, elongate, about twice as long as wide; frontal wall small in area, with subcircular pseudopores; zooecial boundaries formed mainly by apertural rims, raised. Apertures (Fig. 52) medium-sized, longitudinally elongate, about 1:2 x longer than wide, attaining maximum width P.D. TAYLOR Figs 44-47 Elea flabellata sp. nov., VH 10448, holotype, Lower Santonian, Grube Lengede-Broistedt, near Braunschweig, Germany; 44, autozooids and three eleozooids, < 50; 45, autozooids, an eleozooid, and an intramural autozooid (top centre left), x 70; 46, autozooidal operculum, Xx 200; 47, autozooidal aperture and reduced frontal wall, x 160. between hinge line and mid-length, rounded distally; aper- tural shelf variably developed; apertural rim moderately prominent proximally, distally forming a conspicuous, deep tubercle; hinge line bowed. Opercula (Fig. 56) often pre- served in-situ, surface convex, pseudopores not evident in poorly-preserved available material. Terminal diaphragms not observed. Intramurally budded autozooids (Figs 52-53) common, some having apertures in same plane as host aperture but significantly smaller and more rounded distally, others having apertures in an oblique plane facing proximally relative to branch orientation and possessing a very promi- nent tubercle distal to the aperture. Eleozooids (Figs 51, 53-54) abundant, located particularly at branch margins and in bifurcations, about 2-5-3 x longer than wide, frontally slightly wider and significantly longer than autozooids. Aperture elongate, about 2-2-5 x longer than wide, attaining maximum width at the hinge line and tapering distally to a long, narrow rostrum with a rounded end. Opercula not observed in-situ. Gonozooid unknown. MEASUREMENTS (estimated from SEM micrographs). autozooids frontal wall length: ca 0:43-0:50 mm frontal wall width: ca 0:22-0:26 mm apertural length: ca 0-18-0-21 mm apertural width: ca 0:16-0:18 mm eleozooids frontal wall length: frontal wall width: apertural length: apertural width: ca 0:62-0:83 mm ca 0:25-0:27 mm ca 0:36-0:50 mm ca 0:17-0:23 mm REMARKS. The supposed Elea hexagona described by Canu (1897b) from the Cenomanian of Saint-Calais is here included in E. subhexagona sp. nov. This species more closely resembles E. hexagona than any other species of Elea but lacks the very deep and prominent tubercle distal to the SYSTEMATICS OF MELICERITITID CYCLOSTOME BRYOZOANS 25 Fig. 48 lea flabellata sp. nov., VH 10448, holotype, Lower Santonian, Grube Lengede-Broistedt, near Braunschweig, Germany; eleozooid, x 185. autozooidal aperture, which is a characteristic feature of E. hexagona. The larger size of the autozooidal apertures enables distinction between E. hexagona and E. labyrinthica. Filliozat (1908) records E. hexagona from the Calcaire grav- eleux, Assise 4 Crania ignabergensis, Craie de Vend6me. Unfortunately, the species is unrepresented in both the BMNH and VH collections. DISTRIBUTION. Santonian of Venddme, Loir-et-Cher, France. Elea labyrinthica (Michelin, 1843) Figs 57-65 1843 Eschara labyrinthica Michelin: 124, pl. 32, fig. 2. 1843 Eschara neustriaca Michelin: 124, pl. 32, fig. 3. 1853 Elea rhomboidalis d’Orbigny: 631, pl. 737, figs 21-24. 1890 Elea rhomboidalis d’Orbigny; Pergens: 399. 1899 Elea labyrinthica (Michelin); Gregory: 303. 1899 Elea rhomboidalis d’Orbigny; Gregory: 304. MATERIAL. Type: the syntypes of this species (and of the contemporaneous Eschara neustriaca) are not among the Michelin types in the MNHN (see Walter 1975); Michelin (1843) gives Cap de la Héve, Honfleur, Villers-sur-Mer and Vaches-Noires as localities, and Craie Chloritée [= Craie Glauconieuse, Lower Cenomanian] as the horizon. Other material: MNHN d’Orbigny Collection 6627 [= Voigt photo- Fig. 49 Elea hexagona d Orbigny, 1853, photograph of MNHN d’Orbigny Collection 8192 (Voigt photocard 5791), lectotype, - Senonian [Santonian], Vend6éme, Loir-et-Cher, France; colony is fouled by serpulids and an oyster; x 10. card 7475] (presumed type specimen of E. rhomboidalis), Cenomanian, Le Havre, France; BMNH D31139, VH 10460, Cenomanian, Cap de la Héve, Seine Maritime, France; BMNH D58900-2, Lower Cenomanian, Craie Glauconieuse, Cap de la Heve; BMNH D58763-4, D58890-2, Craie Glauco- nieuse, Port d’Antifer, Seine Maritime, France; BMNH D59156 (sample), Craie Glauconieuse, Villers-sur-Mer, Cal- vados, France; BGS GSM 118097-8, Lower Cenomanian, Warminster Greensand, Warminster, Wiltshire, England, Cunnington Collection; BGS GSM Rh 4582, Cenomanian Limestone, ?Bed Al, Hall Rocks to Beer Head, Devon, England; BMNH D59157, Cenomanian Limestone, Bed Al (mantelli Zone), The Pinnacles, near Beer, Devon; BMNH D59158, Cenomanian (float), White Hart Sandpit, Wilming- ton, Devon. VH unnumbered specimens, Lower Cenoma- nian (carcitanensis Zone), Carriére du Billot, Notre-Dame- de-Fresnaye, Calvados, collected by G. Breton. DESCRIPTION. Colony bifoliate with branches of variable morphology, strap-like, bifurcating and about 6 mm wide in some specimens, broad and folded in others (e.g. Michelin 1843, pl. 32, fig. 2); branches about 0-9 mm deep. Zooids arranged in regular to approximate quincunx (Figs 57, 61); organization fixed-walled. Overgrowths extremely common, originating through eruptive budding onto the surfaces of erect branches; growing edges of overgrowths sometimes closed by terminal diaphragms. Colony base extensive, giving rise to more than one erect branch. Possible conspecific ancestrula adjacent to a gonozooid in VH 10460 has a very short distal tube and large protoecium about 0-22 mm wide. Autozooids (Figs 57, 61-62) medium-sized, with frontal walls elongate, over twice as long as wide, typically hexagonal but occasionally diamond-shaped, pierced by circular to Fig. 50 Elea hexagona d’Orbigny, 1853, ZMC Levinsen Collection M39, Santonian, Vendome, France, x 28. slightly longitudinally elongate pseudopores; zooidal bound- aries marked by low, thread-like ridges. Apertures (Fig. 59) small, longitudinally elongate, a little less than 1-5 x longer than wide, attaining maximum width about mid-length, rounded distally; apertural shelf distinct, narrow; hinge line very slightly bowed, teeth not clearly apparent; apertural rim raised. A large and prominent tubercle (Fig. 61) occurs distally of the autozooidal aperture and is shaped like a compressed barkhan dune facing towards the aperture. Oper- cula (Fig. 58) fairly commonly found in-situ, surface convex, bearing about 10 radial ridges peripheral to a central flatter area; pseudopores not seen. Intramural eleozooids often present; apertures variably D-shaped to inverted T-shaped, located within the distal half of the aperture of the host autozooid. Intramural autozooids possibly present but infre- quent. Eleozooids (Figs 57, 60, 62-64) common, scattered or in small groups, with pseudoporous frontal walls having approximately the same area as, though often narrower than, those of autozooids. Apertures highly variable in length, ranging from about 2 to 5 x longer than wide, tapering to become very narrow at the rounded distal extremity; aper- tural shelf wide, beginning just distally of the hinge-line; distal tubercle absent. Opercula (Fig. 63) occasionally found in-situ. Intramural eleozooids (Fig. 64) often present. P. D. TAYLOR Gonozooids known from only one specimen (VH 10460), which has an intact and an abraded example; in the former (Fig. 65) an initially narrow, tubular distal frontal wall becomes densely pseudoporous before dilating into an elon- gate ovoid shape. Ooeciopore circular or slightly longitudi- nally elongate, about the same length as an autozooidal aperture. Atrial ring not apparent in the abraded gonozooid, in which the vertical walls of the more proximally overgrown zooids protrude from the floor of the gonozooid, whereas those of the more distal zooids do not. MEASUREMENTS. autozooids (10 zooids with in-situ opercula from BGS GSM Rh 4582) frontal length: mean = 0-55 mm; SD = 0-033 mm; CV = 6-1; range = 0-50-0-60 mm mean = 0-22 mm; SD = 0-023 mm; CV = 10-2; range = 0-18-0-26 mm mean = 0:14 mm; SD = 0-014 mm; CV = 10-1; range = 0:12-0:17 mm mean = 0-10 mm; SD = 0-010 mm; CV = 10-6; range = 0-08-0-11 mm (10 zooids from BMNH D31139) frontal length: mean = 0-58 mm; SD = 0-043 mm; CV = 7-4; range = 0-53-0-63 mm mean = 0-25 mm; SD = 0-016 mm; CV = 6:3; range = 0-23-0-27 mm mean = 0-15 mm; SD = 0-007 mm; CV = 4-7; range = 0:14-0:17 mm mean = 0-11 mm; SD = 0-007 mm; CV = 6-7; range = 0:09-0:12 mm eleozooids (5 zooids from BGS GSM Rh 4582; 5 zooids from BMNH D31139) frontal length: frontal width: apertural length: apertural width: gonozooid (VH 10460) frontal width: apertural length: apertural width: frontal width: apertural length: apertural width: range = 0-42-1-02 mm range = 0-17-0-27 mm range = 0-20-0-54 mm range = 0-06—0-09 mm total frontal length: 2:72 mm distal frontal wall length: 2:51 mm frontal wall width: 0-98 mm ooeciopore length: 0-11 mm ooeciopore width: 0-11 mm REMARKS. Although type material is lacking, the identity of this species is not in doubt as Michelin’s enlarged figure (1843: pl. 32, fig. 2b) shows very clearly the prominent tubercles located distally of the autozooidal apertures. In no other melicerititid species are the distal tubercles so well- developed or shaped so much like barkhan dunes. Specimens of E. labyrinthica from the type horizon — the Craie Glauconieuse, where it is very common, exhibit vari- able colony forms, some being adeoniform while others are eschariform. One specimen (BMNH D58763) encrusts a sponge. DISTRIBUTION. Lower Cenomanian of northern France and south-west England; known from sandy facies only. Elea mackinneyi sp. nov. Figs 66-71 MATERIAL. Holotype: VH 10474, Lower Cenomanian, Milheim/Ruhr, Westfalia, Germany. Paratypes: VH 10541 (4 SYSTEMATICS OF MELICERITITID CYCLOSTOME BRYOZOANS Figs 51-56 Elea hexagona d’Orbigny, 1853, Santonian, Vendome, France. 51-54, ZMC Levinsen Collection M38; 51, autozooids and a marginal eleozooid (lower left), x 33; 52, intramural operculate autozooid (left) and autozooidal aperture (right), x 135; 53, marginal eleozooid and intramural autozooids, x 70; 54, non-marginal eleozooid, x 98. 55, 56, ZMC M39; 55, intramural autozo (left) and normal autozooid (right), < 135; 56, autozooidal operculum, x 150. P. D. TAYLOR Figs 57-62 Elea labyrinthica (Michelin, 1843). 57-60, BMNH D31139, Cenomanian, Cap de la Héve, Seine Maritime, France; 5 autozooids, eleozooids and intramural eleozooids; distal tubercles abraded, x 55; 58, autozooidal operculum, x 170; 59, autozooidal aperture, X 185; 60, eleozooid with distally broken rostrum, < 175. 61, 62, BGS GSM Rh 4582, Cenomanian Limestone, ?Bed A1, Hall Rocks to Beer Head, Devon, England; 61, autozooids with prominent distal tubercles, x 48; 62, autozooids and eleozooids with long, narrow rostra, X 70. SYSTEMATICS OF MELICERITITID CYCLOSTOME BRYOZOANS 29 Figs 63-65 Devon, England; 63, large eleozooid with slightly damaged operculum, x 145; 64, two large eleozooids housing small intramural eleozooids, x 110. 65, VH 10460, Cenomanian, Cap de la Héve, Seine Maritime, France, gonozooid with crushed frontal wall, x 35. specimens), same horizon and locality as holotype. NAME. In recognition of the numerous fundamental contri- butions made to bryozoology by Dr F.K. McKinney (Appala- chian State University). DESCRIPTION. Colony bifoliate with branches (Fig. 66) nar- row, strap-like, about 1-8 mm wide between bifurcations, and 0-7 mm deep. Autozooids generally arranged in transverse tows whose regularity is disrupted close to bifurcations; organization fixed-walled. Overgrowths and colony base not observed. Autozooids (Fig. 67) medium-sized, with elongate hexago- nal frontal walls on average slightly less than 2 x longer than wide; pseudopores circular or longitudinally elliptical; zoo- ecial boundaries raised. Apertures small, longitudinally elon- gate, about 1-1 xX longer than wide, attaining maximum width a little proximally to mid-length, appreciably narrower at hinge line level; apertural shelf absent or at most very slightly developed; apertural rim moderately raised; hinge line straight or slightly bowed, typically concave in eroded _ Zooids lacking opercula. Opercula (Fig. 69) rarely preserved in-situ, flat or slightly depressed, bearing about 16 pseudopores arranged in a crescent parallel to the distal/ lateral margin. Terminal diaphragms not observed. Intramu- rally budded autozooids represented by a single example with a slightly oblique aperture (Fig. 70). Eleozooids (Figs 67, 68) common, located along branch _Margins, on average about 3 x longer than wide, usually Elea labyrinthica (Michelin, 1843). 63, 64, BGS GSM Rh 4582, Cenomanian Limestone, ?Bed A1, Hall Rocks to Beer Head, occupying two transverse rows of autozooids. Aperture long, widest at the hinge line but broad throughout, sometimes very slightly spatulate, well-rounded distally, on average twice as long as wide. In-situ opercula and intramural buds not observed. Kenozooids (Fig. 68) occasionally present at branch mar- gins, especially within bifurcations. Gonozooids (Fig. 71) of unknown abundance (one fully developed and one partially developed example present in holotype VH 10474); distal frontal wall dilating soon after emerging from maternal aperture, longitudinally ovoidal in shape, about 2 x longer than wide. Ooeciopore not observed. Atrial ring present. MEASUREMENTS. autozooids (10 zooids from holotype VH 10474) frontal length: mean = 0-49 mm; SD = 0-029 mm; CV = 6-0; range = 0-45—-0-54 mm mean = 0:26 mm; SD = 0-018 mm; CV = 6-7; range = 0:24-0:29 mm mean = 0-18 mm; SD = 0-006 mm; CV = 3-6; range = 0-17-0-18 mm mean = 0-16 mm; SD = 0-005 mm; CV = 2-9; range = 0-15-0-17 mm frontal width: apertural length: apertural width: eleozooids (10 zooids from holotype VH 10474) frontal length: mean = 1:00 mm; SD = 0-184 mm; CV = 18-5; range = 0-75-1-26 mm 30 Fig. 66 Elea mackinneyi sp. nov., VH 10474, holotype, Lower Cenomanian, Milheim/Ruhr, Westfalia, Germany; branch with broken gonozooid and fouling foraminifer (top left), x 20. mean = 0-36 mm; SD = 0-050 mm; CV = 13-9; range = 0-30-0-44 mm mean = 0-49 mm; SD = 0-102 mm; CV = 20-7; range = 0-33-0-65 mm mean = 0-24 mm; SD = 0-034 mm; CV = 13-8; range = 0-18-0-29 mm frontal width: apertural length: apertural width: gonozooid (from holotype VH 10474) total frontal length: 2-34 mm dilated frontal wall length: 2-07 mm frontal wall width: 1-04 mm REMARKS. Although known from only a few specimens, this is a very distinctive species. The broad shape of the eleozooid rostra serve to distinguish Elea mackinneyi from sympatric Elea elegantula sp. nov., and it differs from E. viskovae sp. nov. from the Turonian of the Kazakhstan in having longer rostra and in lacking an appreciable autozooidal apertural shelf. DISTRIBUTION. Lower Cenomanian of Germany. P.D. TAYLOR Figs 72-78 MATERIAL. Holotype: VH 10462, Upper Campanian, St Severin d’Uzel, Charente Maritime, France. Paratypes: VH 10461, same horizon and locality as holotype; VH 10471, Senonian, La gare de Soullans, Vendée, France. BMNH BZ 441-4 , Upper Campanian, Cailleau, near Talmont, Charente Maritime, France. Other material: VH unnumbered topo- types. NAME. With reference to its similarity to Elea lamellosa (d’Orbigny). Elea pseudolamellosa sp. nov. DESCRIPTION. Colony bifoliate, consisting of broad, folded fronds about 0-9 mm deep. Growing edge may be occluded by kenozooids and eleozooids. Colony base not observed. Overgrowths may be present. Zooids arranged in approxi- mate quincunx; organization fixed-walled. Autozooids (Figs 72, 76) medium-sized, frontally hexago- nal in outline, on average 1-6 X as long as wide, distal edge prolonged by aperture; frontal wall slightly convex, with circular pseudopores; zooecial boundary wall inconspicuous, thin, little raised. Apertures (Fig. 74) small, longitudinally elongate, about 1-2 x longer than wide, reaching maximum width a little distally of the hinge line, well-rounded distally; apertural shelf narrow; apertural rim narrow but prominent, continuous with zooecial boundary wall; hinge line bowed. Opercula (Fig. 73) infrequently preserved in-situ, prominent, surface convex, seemingly with radially elongate pseudopores widely distributed. Terminal diaphragms not observed. Intra- mural eleozooids present, their D-shaped apertures occupy- ing the distal part of the host aperture, which is proximally covered by a slightly concave pseudoporous exterior wall. Eleozooids (Figs 72, 76-77) common, often clustered, especially near occluded growing edges; frontally variable in size, typically only slightly smaller than autozooids; frontal wall with circular pseudopores. Apertures (Fig. 77) small, D-shaped, equidimensional or transversely elongate. In-situ opercula and intramural buds not observed. Kenozooids observed in association with eleozooids and gonozooids (Fig. 75). Gonozooids (Fig. 78) large, longitudinally elongate; frontal wall with a short parallel-sided portion emerging from the maternal zooidal aperture (Fig. 75), dilating into an ovoidal distal portion about twice as long as wide. Ooeciopore not observed (destroyed during abrasion of frontal wall in avail- able specimens). Atrial ring not observed. MEASUREMENTS. autozooids (10 zooids from holotype VH 10462) frontal length: mean = 0:53 mm; SD = 0-029 mm; CV = 5-6; range = 0-48-0-57 mm mean = 0-33 mm; SD = 0-024 mm; CV = 7-2; range = 0:30-0:38 mm mean = 0-18 mm; SD = 0-014 mm; CV = 7-7; range = 0:17-0:21 mm mean = 0-15 mm; SD = 0-007 mm; CV = 4-7; range = 0-14-0-17 mm from SEM micrographs of VH frontal width: apertural length: apertural width: eleozooids (estimated 10461-2) apertural length: apertural width: ca 0-05—0-07 mm ca 0-06—0-07 mm gonozooid (one zooid from VH 10462) frontal length: 2:52 mm _ SYSTEMATICS OF MELICERITITID CYCLOSTOME BRYOZOANS 31 69 oblique intramural autozooidal aperture, x 265. 2:31 mm 1:20 mm distal frontal length: frontal width: REMARKS. This species is extremely similar to Elea lamellosa d Orbigny (see p. 17). The principal point of distinction is the D-shaped form of the eleozooid aperture, which contrasts with the inverted T-shaped aperture characteristic of the eleozooids in E. lamellosa. (i.e. the eleozooids of E. pseudolamellosa are demizooids, whereas those of E. lamel- losa are trifoliozooids; compare Figs 77 and 19) This differ- ence appears to reflect the presence of a rostral shelf in E. lamellosa which is absent in E. pseudolamellosa. Autozooidal frontal walls may be a little more convex and zooecial boundary walls rather less well-developed in E. pseudolamel- losa than in E. lamellosa. The stratigraphical distribution of the two species differs, E. lamellosa occurring in the Coniacian-Santonian (and possibly the Turonian), whereas E. pseudolamellosa is known with certainty only from the Upper Campanian; a paratype (VH 10471) from the Vendée Figs 67-70 Elea mackinneyi sp. nov., VH 10474, holotype, Lower Cenomanian, Milheim/Ruhr, Westfalia, Germany; 67, autozooids and marginal eleozooid, x 55; 68, two marginal eleozooids and kenozooids at a bifurcation, x 44; 69, autozooidal operculum, x 265; 70, assigned to E. pseudolamellosa is of unknown age within the Senonian. DISTRIBUTION. Upper Campanian of Charente Maritime, and undifferentiated Senonian of the Vendée, France. Figs 79-86 non 1846 Escharites dichotoma Reuss: 66, pl. 15, fig. 31. 1874 Echarites dichotoma (Reuss); Reuss: 135, pl. 25, fig. 8. 1897b Elea hexagona d’Orbigny; Canu: 753. Elea subhexagona sp. nov. MATERIAL. Holotype: DM un-numbered, Voigt photocard 2060 (specimen figured by Reuss 1874, pl. 25, fig. 8), Upper Turonian, ‘Oberen Planers’, Strehlen, Dresden, Germany. Paratypes: ZMC M53-5, ‘Turonien’ [probably Cenomanian], St Calais, Sarthe, France. Other material: MNHN Canu Collection unnumbered. Fig. 71 Elea mackinneyi sp. nov., VH 10474, holotype, Lower Cenomanian, Miilheim/Ruhr, Westfalia, Germany; gonozooid with broken frontal wall exposing atrial ring, x 48. NAME. With reference to the resemblance to Elea hexagona. DESCRIPTION. Colony bifoliate with branches (Fig. 79) bifur- cating, strap-like, narrow, about 1-3—2-3 mm wide. Autozoo- ids generally arranged in approximate quincunx; organization fixed-walled. Colony base not observed. Overgrowth present in ZMC M54, consisting of kenozooids spreading between apertures of underlying autozooids, possibly free-walled (Fig. 79). Autozooids (Figs 80, 81) medium-sized; frontal surface usually hexagonal, elongate, about twice as long as wide; frontal wall occupying about half of frontal surface, flat with circular pseudopores; zocecial boundaries raised. Apertures (Fig. 83) small, longitudinally elongate, on average 1-1-1-2 x longer than wide, attaining maximum width between hinge line and mid-length, rounded distally; aper- tural shelf narrow; apertural rim well-developed; hinge line with a subdued median bar. Opercula (Fig. 82) often pre- served in-situ, surface convex, sometimes with a flattened median area close to the hinge line, pseudopores not visible in studied material. Terminal diaphragms not observed. Intramurally budded autozooids may be present, their aper- tures typically shorter and more rounded than normal auto- zooids. Eleozooids (Figs 84-85) common, especially at branch margins and in bifurcations, about 3-4 x longer than wide, frontally slightly wider but appreciably longer than autozoo- P! Di TAYLOR ids. Aperture elongate, about 2-3 x longer than wide, widest at the hinge line and tapering to a long narrow rostrum with a rounded end. Opercula (Fig. 84) observed in-situ, surface convex. Intramurally budded eleozooids (Fig. 81) present. Kenozooids sometimes present in branch bifurcations. Gonozooids (Fig. 86) with a short parallel-sided proximal frontal wall emerging from the maternal zooid aperture, becoming longitudinally ovoidal, distal dilated frontal wall approximately 1-5 x longer than wide. Ooeciopore removed by abrasion; atrial ring present. MEASUREMENTS. autozooids (10 zooids with in-situ opercula from the holo- type) frontal length: mean = 0:53 mm; SD = 0-040 mm; CV = 7-6; range = 0-48-0-60 mm mean = 0:27 mm; SD = 0-019 mm; CV = 7-1; range = 0-26-0-30 mm mean = 0-19 mm; SD = 0-007 mm; CV = 4-1; range = 0-18-0-20 mm mean = 0-17 mm; SD = 0-007 mm; CV = 4:3; range = 0-15—0-18 mm frontal width: apertural length: apertural width: eleozooids (5 zooids from the holotype) frontal length: range = 0-74-0-81 mm frontal width: range = 0:23-0:29 mm apertural length: range = 0-36-0-42 mm apertural width: range = 0-15—0-20 mm gonozooids (2 zooids estimated from SEM micrographs of ZMC MS3 and M55) total frontal length: distal frontal wall length*: ca 1-24-1-44 mm frontal width: ca 0:85-0:96 mm (* i.e. that part of the frontal wall distal to the narrow origin) ca 1-61-1-88 mm REMARKS. This new species was identified by Reuss (1874) as Escharites dichotoma Reuss, 1846. Unfortunately, the type specimen of E. dichotoma, from the Exogyrensandstein of Bohemia, was destroyed in 1956 (E. Voigt pers. comm. December 1986). However, it is clear from the figure of Reuss (1846), which depicts a vinculariiform colony resem- bling Filicea, that it is not the same species as E. dichotoma — sensu Reuss, 1874. Canu (1897b) identified specimens here regarded as‘Elea | subhexagona as E. hexagona d’Orbigny, a Santonian species which is very similar to E. subhexagona but which can be distinguished by the deep and prominent tubercle situated | distally to the autozooidal aperture. Canu apparently realized | later that this identification was mistaken because material of his in the MNHN and in the ZMC (donated to G.M.R. Levinsen) is labelled with the MS name ‘Elea fissurata’. The | exact stratigraphical horizon of these specimens from St | Calais (Sarthe) is questionable; Canu originally ascribed his | St Calais material to the Cenomanian but subsequently | referred it to the Turonian (Canu and Bassler, 1922). It seems | possible that the specimens are from the late Cenomanian | plenus Zone, although E. subhexagona is not represented | among material of this age from St Calais in the VH and BMNH collections. DISTRIBUTION. Upper Cenomanian/Lower Turonian of St } Calais, Sarthe, France, and Upper Turonian of Dresden, Germany. | SYSTEMATICS OF MELICERITITID CYCLOSTOME BRYOZOANS 33 Figs 72-77 Elea pseudolamellosa sp. nov. 72-74, VH 10461, Upper Campanian, St Severin d’Uzel, Charente Maritime, France; 72, autozooids and eleozooids (top left), x 47; 73, autozooidal operculum, x 250; 74, autozooidal aperture, x 250. 75, VH 10462, holotype, Upper Campanian, St Severin d’Uzel, gonozooid origin with kenozooid above and to the left of the maternal zooid, x 65; 76, 77, VH 10471, Senonian, Le gare de Soullans, Vendée, France; 76, eleozooids and autozooids, x 65; 77, apertures of two eleozooids, x 250. 34 Fig. 78 Elea pseudolamellosa sp. nov., VH 10462, holotype, Upper Campanian, St Severin d’Uzel, Charente Maritime, France; gonozooid with broken roof, x 42. Elea triangularis (Michelin, 1841) Figs 87-94 1841 Eschara triangularis Michelin: 5, pl. 1, fig. 6. 1850 Eschara triangularis Michelin; d’Orbigny: 140. 1851 Eschara triangularis Michelin; d’Orbigny, pl. 602, figs 4 and S. 1853 Elea triangularis (Michelin); d’Orbigny: 630, pl. 737, figs 17-20. 1890 Elea triangularis (Michelin); Pergens: 397. 1899 Elea triangularis (Michelin); Gregory: 305. 1975 Elea triangularis (Michelin); Walter: 314, pl. 29, figs 1, >: MATERIAL. Holotype: _MNHN _ Michelin Collection un-numbered (Fig. 87), Albian, Grandpré, Ardennes, France. Other material: MNHN d’Orbigny Colln 6020 (5 pieces mounted on a tablet with the mark of a sixth, lost piece), same horizon and locality as holotype; BMNH 60541 (large colony and several fragments), ‘Gault Chloritic’ (prob- ably Lower Albian mammillatum Zone according to H.G. Owen pers. comm.), Folkestone, Kent, UK; BMNH D52080 (SEM stub), Lower Albian (tardefurcata or mammillatum Zone, see Owen, 1972; Rawson et al., 1978), Shenley Lime- stone, Leighton Buzzard, Bedfordshire, UK, R.J. Hogg Collection. DESCRIPTION. Colony bifoliate (Fig. 87) with narrow, folded branches which bifurcate occasionally; branches generally 6 P. D. TAYLOR Fig. 79 Elea subhexagona sp. nov., ZMC Levinsen Collection M54, ‘Turonien’ [probably Cenomanian], St Calais, Sarthe, France; branch with autozooids, eleozooids and distally, an overgrowth of kenozooids occupying the areas between zooidal apertures; x 39. mm wide and 1 mm deep. Zooids arranged in approximate quincunx; organization fixed-walled. Colony base hollow (in holotype). Growing edge often arcuate. Overgrowths some- times developed on erect branches. Zooidal buds may be occluded by terminal diaphragms located well proximal to the interzooidal wall ends. Autozooids (Fig. 89) medium-sized, generally rhombic with relatively short, convex frontal walls pierced by large, circular, slightly countersunk pseudopores; zooidal bound- aries raised. Apertures (Fig. 90) very large, elongate, almost 1-5 x longer than wide, attaining maximum width a little distal to the hinge line, well-rounded distally; apertural shelf present only in the mid-part of the aperture, indenting the margins of the aperture and producing the bell-shape typical of the species; hinge line with a pair of prominent hinge teeth. Apertural rim thick, continuous with variably extensive but low tubercles at the lateral corners of the frontal wall. Opercula (Fig. 88) occasionally preserved in-situ, proximal edge a little concave, surface slightly convex, fissured and with about 25 elongate pseudopores arranged in an irregular crescent; internally (Fig. 92), prominent, inward-sloping sclerites form a continuous arch immediately outward of the pseudopores. Intramurally budded autozooids may be present. A tubular structure apparently composed of SYSTEMATICS OF MELICERITITID CYCLOSTOME BRYOZOANS WwW n Figs 80-83 Elea subhexagona sp. nov. 80, DM un-numbered, Voigt photocard 2060, holotype, Upper Turonian, “Oberen Planers’, Strehlen, Dresden, Germany, autozooids, mostly with opercula, and eleozooids, x 45. 81-83, ZMC Levinsen Collection M54, ‘Turonien’ [probably Cenomanian], St Calais, Sarthe, France; 81, autozooids and a primary eleozooid containing an intramural eleozooid, x 70; 82, autozooidal operculum, X 210; 83, autozooidal aperture, x 210. pseudoporous exterior wall, which can be seen within the aperture of one zooid (Fig. 91), is interpreted as a bioclaus- tration (i.e. an embedment structure formed by the bryo- zoan; cf. similar structures in Cretaceous onychocellids attributed to the presence of folliculinid protists by Ernst, 1985). Interzooidal walls with conspicuous pores. Mural spines present in great abundance in the proximal parts of zooids, long, and either simple or branched (Fig. 93). Eleozooids unknown, presumed absent. Kenozooids sometimes present. Gonozooid known from only one partly developed example (Fig. 94) showing the tubular, densely pseudoporous proximal part of the frontal wall emerging from the aperture of the maternal zooid. MEASUREMENTS. autozooids (10 zooids from BMNH 60531) frontal length: mean = 0-55 mm; SD = 0-036 mm; CV = 6-6; range = 0-50-0-60 mm frontal width: - mean = 0-35 mm; SD = 0-027 mm; CV = 7:8; range = 0-32-0-39 mm mean = 0:29 mm; SD = 0-015 mm; CV = 5-2; range = 0:27-0:32 mm mean = 0-20 mm; SD = 0-011 mm; CV = 5:2; range = 0-20-0-23 mm apertural length: apertural width: REMARKS. The large size and characteristically bell-shaped autozooidal apertures immediately distinguish this uncom- mon species from others in the genus. E. triangularis is the only bifoliate melicerititid described from the Lower Creta- ceous. It is also currently unique in being the only melicer- ititid known to possess mural spines (= zooecial or intrazooecial spines) (Fig. 93). Similar structures are, how- ever, common and widely distributed taxonomically among non-melicerititid cyclostomes (see Farmer, 1979) and further investigations of the internal morphology of melicerititids will probably reveal more examples. DISTRIBUTION. Albian (?Lower Albian only) of France and England. 36 P. D. TAYLOR Figs 84-86 Elea subhexagona sp. nov. 84, DM un-numbered, Voigt photocard 2060, holotype, Upper Turonian, “Oberen Planers’, Strehlen, Dresden, Germany, eleozooid with operculum, x 145. 85, 86, ZMC Levinsen Collection M53, ‘Turonien’ [probably Cenomanian], St Calais, Sarthe, France; 85, eleozooid, x 120; 86, damaged gonozooid, x 53. Fig. 88 Elea triangularis (Michelin, 1841), BMNH D52080, Lower 87 Albian (tardefurcata or mammillatum Zone), Shenley Limestone, Fig. 87 Elea triangularis (Michelin, 1841), photograph of the Leighton Buzzard, Bedfordshire, UK; secondary electron image holotype colony, MNHN Michelin Collection un-numbered, of coated specimen showing autozooidal operculum in slightly Albian, Grandpré, Ardennes, France; x 2-5. oblique view, x 117. SYSTEMATICS OF MELICERITITID CYCLOSTOME BRYOZOANS | Figs 89-94 Elea triangularis (Michelin, 1841), BMNH 60531, ‘Gault Chloritic’ [probably Lower Albian mammillatum Zone], Folkestone, | Kent, UK; 89, group of autozooids, including one with an intramural autozooid (upper centre), < 62; 90, autozooidal aperture, x 125; 91, autozooidal aperture containing enigmatic tubular structure, x 120; 92, inner side of an autozooidal frontal wall and operculum showing sclerites and pseudopores, x 118; 93, fractured specimen exposing mural spines within the proximal chamber of an autozooid, x 355; 94, proximal part of a broken gonozooid, x 37. Si 38 Figs 95-99 MATERIAL. Holotype: VH 10450, Turonian (‘C22’ division), Kyzylsaj, ca 180 km east of Fort Shevchenko, Kazakhstan, Najdin Collection. Paratypes VH 10542 (6 specimens), details as for holotype. Elea viskovae sp. nov. NAME. In honour of the Russian bryozoologist L.A. Viskova. DESCRIPTION. Colony bifoliate with branches (Fig. 95) nar- row, strap-like, bifurcating, about 1-3-2-0 mm wide and 0-5—0-6 mm deep. Autozooidal arrangement variable, both autozooids and eleozooids divergent close to branch margins; organization fixed-walled. Overgrowths common. Colony base unknown. One of the paratypes grows against and partially envelops a ?serpulid tube. Autozooids (Fig. 96) large; frontal walls irregularly hex- agonal, distally rounded, elongate, 2-5-3 = longer than wide, with circular pseudopores; zooecial boundaries raised. Aper- tures (Fig. 99) of moderate size, longitudinally elongate, on average 1-1 x longer than wide, attaining maximum width just proximal to mid-length, appreciably narrower at hinge line level, well-rounded distally; apertural shelf broad, 0:02-0:03 mm wide, causing the aperture to have the shape of a high isosceles triangle with rounded corners; apertural rim Fig.95 Elea viskovae sp. nov., VH 10450, holotype, Turonian ((C22’ division), Kyzylsaj, ca 180 km east of Fort Shevchenko, Kazakhstan, Najdin Collection; branch with autozooids and marginal eleozooids, x 37. P: DI TAYEOR moderately prominent; hinge line bowed. Opercula (Fig. 97) often preserved in-situ, slightly convex, with about 16 pseudopores arranged in a cresent parallel to the distal/lateral margin. Terminal diaphragms (Fig. 97) positioned proximally to the apertural shelf, ?non-pseudoporous. Intramural buds not observed with certainty. Eleozooids (Figs 95, 96) common, located close to branch margins, about 2—2-5 longer than wide, their frontal walls a little longer and significantly wider than those of the autozoo- ids. Aperture elongate, attaining maximum width slightly distally to the hinge-line, well-rounded distally; apertural shelf not clearly developed but rostral shelf present. Opercula known from only one partial example in-situ, surface strongly convex at preserved edges. Intramural buds not observed. Kenozooids commonly present at branch margins. Gonozooids not observed. MEASUREMENTS. autozooids (10 zooids with in-situ opercula from holotype VH 10450) frontal length: mean = 0-64 mm; SD = 0-070 mm; CV = 10-9; range = 0:54-0:77 mm mean = 0-24 mm; SD = 0-024 mm; CV = 9-6; range = 0-21-0-29 mm mean = 0-19 mm; SD = 0-009 mm; CV = 4-4; range = 0-18-0-21 mm mean = 0-17 mm; SD = 0-010 mm; CV = 6-0; range = 0-17-0-20 mm eleozooids (5 zooids from holotype VH 10450) frontal length: range = 0-63—0-80 mm frontal width: range = 0-32-0-35 mm apertural length: range = 0-27-0-30 mm apertural width: range = 0-21-0-24 mm frontal width: apertural length: apertural width: REMARKS. The marginal eleozooids of Elea viskovae, which resemble slightly-enlarged autozooids, distinguish this new species from others within the genus Elea. They have less well-developed rostra than those of E. mackinneyi, a species with smaller autozooids. Eleozooid morphology recalls that present in such species as Meliceritites semiluna and Biforicula filicosa. Many of the autozooidal opercula (and some terminal diaphragms) in the holotype are pierced by small circular borings (Figs 96-97) like those described previously in other melicerititids (Taylor, 1982). DISTRIBUTION. Turonian of Kazakhstan. Figs 100-109 MATERIAL. Holotype: VH 10538, top Lower Cenomanian (orbignyi Zone) — basal Middle Cenomanian (costatus Zone), Saint-Germain-la-Campagne, descente sur Orbiquet, Eure, France, collected by G. Breton 1981. Paratypes: VH 10439, 10539-40, same horizon and locality as holotype. BMNH D55027-8, Cenomanian [?Lower Cenomanian, Craie Glau- conieuse], Bruneval, Seine-Maritime, France, S. Whiteley Collection. Other material: VH, numerous un-numbered topotypes. Elea whiteleyi sp. nov. NAME. In honour of the late S. Whiteley, an enthusiastic amateur geologist who collected paratypes of this species. DESCRIPTION. Colony bifoliate with bifurcating, narrow strap-like branches about 1-1—2-3 mm wide and 0-6 mm deep, elliptical in cross-section. Autozooids arranged more or less SYSTEMATICS OF MELICERITITID CYCLOSTOME BRYOZOANS 39 in quincunx or in ill-defined transverse rows (Fig. 100), separated from adjacent autozooids by demizooids; organiza- tion fixed-walled. Overgrowths moderately common, at least some formed by eruptive budding onto surfaces of erect branches, sometimes consisting predominantly of demizooids and kenozooids . Colony base (Figs 105-107) extensive, giving rise to several erect branches whose median budding laminae are parallel to local orientation of basal zooids; demizooids of base have higher, more pointed apertures than those of erect branches, and many possess non-inclined apertures orientated parallel to adjacent autozooids. Autozooids (Figs 102, 104) small, with longitudinally ellip- tical pseudoporous frontal walls, about twice as long as wide, depressed with respect to the surrounding demizooids. Aper- tures small, longitudinally elongate, about 1-1 x longer than wide, attaining maximum width at a level between the hinge line and mid-length; apertural shelf slight; apertural rim prominent, sometimes prolonged at the mid-point of the aperture; hinge line with a curved median bar. Opercula (Fig. 101) very often preserved in-situ, surface moderately convex, a flattened median proximal area often evident, bearing an estimated 14~18 slit-shaped pseudopores in a crescent close to — Re Ce gs rae Figs 96-99 Elea viskovae sp. nov., VH 10450, holotype, Turonian (“C22’ division), Kyzylsaj, ca 180 km east of Fort Shevchenko, Kazakhstan, Najdin Collection; 96, autozooids with opercula, some bored, and three marginal eleozooids, x 62; 97, bored autozooidal operculum (upper left) and terminal diaphragm (lower right), x 160; 98, autozooidal operculum, x 250; 99, autozooidal aperture, 250. the lateral/distal edge. Intramural buds and terminal dia- phragms not observed. Rostrozooids (Figs 105, 108) moderately common, usu- ally occurring singly close to the branch mid-line just before a bifurcation, but sometimes located close to branch margins, elongate, about 3-5 x longer than wide. Aper- ture long, attaining maximum width at the hinge line, very slightly spatulate, rounded distally, about 3-4 x longer than wide. In-situ opercula not seen. Rostrozooids and/or demizooids may be budded intramurally within rostrozoo- ids (Fig. 108). Demizooids (Fig. 103) numerous, normally about 2 per autozooid, often paired disto-laterally on either side of an autozooidal aperture; frontal walls usually long and narrow, forming raised areas between frontal walls of laterally adja- cent autozooids, their boundaries, however, poorly-defined; apertures variously orientated, many in oblique or reverse orientation with respect to colony growth direction, typically in a plane inclined about 60° to the colony surface, small, semicircular to longitudinally elongate in shape, rounded or slightly arched distally. Opercula often preserved in-situ; terminal diaphragms observed. PDF TAWIEOR Figs 100-103 Elea whiteleyi sp. nov., top Lower Cenomanian (orbignyi Zone) — basal Middle Cenomanian (costatus Zone), Saint-Germain-la-Campagne, descente sur Orbiquet, Eure, France. 100-101, VH 10539; 100, branch surface, x 53; 101, autozooidal operculum, x 330. 102, 103, VH 10538, holotype; 102, autozooids, kenozooids, and demizooids with obliquely-directed, inclined apertures, X 95; 103, operculate autozooid and two demizooids, x 220. Kenozooids (Figs 102, 105) observed commonly in colony base, overgrowths, and at the margins of erect branches. Gonozooids (Fig. 109) fairly common. Distal frontal wall emerging from maternal aperture initially parallel-sided before becoming bulbous and longitudinally ovoidal in shape. Ooeciopore located beyond inflated part of frontal wall, transversely elliptical, about 1-5 x wider than long. Atrial ring not observed. MEASUREMENTS. autozooids (10 zooids with in-situ opercula from holotype VH 10538) frontal length: mean = 0-33 mm; SD = 0-015 mm; CV = 4-5; range = 0:30-0:35 mm mean = 0:17 mm; SD = 0-008 mm; CV = 4-5; range = 0-17-0-18 mm mean = 0-15 mm; SD = 0-005 mm; CV = 3-1; range = 0:15-0:17 mm mean = 0:13 mm; SD = 0-006 mm; CV = 4-8; range = 0-12-0-:14 mm frontal width: apertural length: apertural width: rostrozooids (5 zooids from 5 colonies) frontal wall length: range = 0-63-0-92 mm frontal wall width: range = 0-20-0-21 mm apertural length: range = 0-33-0-57 mm apertural width: range = 0-11-0-15 mm demizooids apertural width: ca 0-06-0-09 mm gonozooids (one zooid from holotype VH 10538) total frontal wall length: 2-36 mm dilated frontal wall length*: 1-13 mm ooeciopore length: 0-08 mm ooeciopore width: 0-12 mm (* i.e. that part of the frontal wall distal to the narrow origin) REMARKS. The presence of numerous small eleozooids enables this new species to be readily distinguished from all other species of Elea, including E. elegantula which it most resembles in the morphology of the other polymorphs. These ' eleozooids and their distribution surrounding the autozooids invite comparison between E. whiteleyi and the Upper Cen- /omanian bifoliate melicerititid Biforicula multicincta, the ‘oldest known species of Biforicula (see Voigt, 1989) The principal qualitative difference between E. whiteleyi and B. _multicincta is the existence of cancelli in the latter. It there- ' fore seems possible that E. whiteleyi belongs in the stem- | group of Biforicula. DISTRIBUTION. Lower-Middle Cenomanian of Normandy, France. Genus SEMIELEA d’Orbigny, 1853 TYPE SPECIES. Entalophora vieilbanci d’Orbigny, 1850, des- ignated by Bassler (1935: p. 198) as ‘S. vielbanci D’Orbigny, 1853’ [sic]. | SYSTEMATICS OF MELICERITITID CYCLOSTOME BRYOZOANS 4] Figs 104-107 Elea whiteleyi sp. nov. 104, BMNH D55027 Cenomanian [?Lower Cenomanian, Craie Glauconieuse], Bruneval, Seine-Maritime, France, operculate autozooids (demizooids obscured by diagenetic cement), x 98. 105-107, VH 10439, top Lower Cenomanian (orbignyi Zone) — basal Middle Cenomanian (costatus Zone), Saint-Germain-la-Campagne, descente sur Orbiquet, Eure, France; encrusting colony base; 105, irregular arrangement of autozooids, kenozooids, small eleozooids and a large eleozooid, x 40; 106, large eleozooid (rostrozooid), x 100; 107, operculate autozooid and small eleozooid, x 135. OTHER SPECIES. Semielea dichotoma d@’Orbigny, 1853. REVISED DIAGNOSIS. Eleid with tubular colony-form; axial lumen of subequal diameter lined by an inferred exterior wall; zooids in primary layer orientated parallel to tube length; autozooids fixed-walled; cancelli lacking. REMARKS. D’Orbigny (1853) referred three species to Semie- lea. Only the first two are here retained in the genus; the third, Semielea plana d’Orbigny, 1853 is regarded as the base of Elea lamellosa (d’ Orbigny, 1850) (see p. 19). Diagnostic of Semielea is the possession of branches with an axial lumen, the bounding wall of which forms a lamina for zooidal budding. The axial lumen is intermediate in size between the narrow axial canals present in some species of Meliceritites (e.g. M. semiclausa (Michelin)) and the wide tubular interiors of certain specimens of Reptomultelea with cavariiform colonies. The growth of the axial lumen in Semielea also seems distinct. Although no specimens have P.D. TAYLOR Fig. 108 Elea whiteleyi sp. nov., VH 10539, top Lower Cenomanian (orbignyi Zone) — basal Middle Cenomanian (costatus Zone), Saint-Germain-la-Campagne, descente sur Orbiquet, Eure, France; primary rostrozooid containing intramural rostrozooid in turn containing intramural demizooid, x 120. been studied with their microstructure sufficiently well- preserved to prove that the wall which lines the lumen is an exterior wall rather than an interior wall, the transverse folding and wrinkling of this wall (Fig. 119) strongly suggests that it is an exterior wall. Rare examples of colony bases viewed from the underside (Fig. 122) show that the lumen of each branch opens at the colony base and that the lumen wall is smoothly continuous with the basal lamina. As the basal lamina is an undoubted exterior wall, this observation is further evidence that the lumen walls are exterior. In S. vieilbanci occasional transverse platforms partition the axial lumen and seem to have been formed by progressive constric- tion of the lumen. The presence of these platforms, which would have periodically sealed the entrance to the lumen at branch growth tips, may explain why fouling of the apparent exterior walls lining the lumen has not been observed (cf. exterior walls lining the tubes of cavariiform Reptomultelea colonies). The platforms also imply that branches could not have grown around another erect organism (cf. some species of Reptomultelea), but instead grew freely into space. The phylogenetic relationship is unclear between Semielea and species of Meliceritites on the one hand and of Reptomul- telea on the other. Assuming Semielea to be a relatively advanced genus, it could have derived from either of the two latter genera. Derivation from Meliceritites would necessitate Fig. 109 Elea whiteleyi sp. nov., VH 10538, holotype, top Lower Cenomanian (orbignyi Zone) — basal Middle Cenomanian (costatus Zone), Saint-Germain-la-Campagne, descente sur Orbiquet, Eure, France; gonozooid, x 50. the incorporation of tubular prolongations of the basal lamina into the centre of the erect cylindrical branches, together with loss of the normal endozonal budding pattern. Derivation from Reptomultelea would require formation of tubular pro- longations of the basal lamina to give erect cylindrical branches. On the grounds of parsimony, the latter alternative seems the more likely, but future phylogenetic analysis, including zooid-level characters, might provide better evi- dence of affinity. DISTRIBUTION. Turonian (?Upper) — Lower Santonian, northern France. Semielea vieilbanci (d’Orbigny, 1850) 1850 Entalophora vieilbanci d’Orbigny: 200. 1851 Diastopora vieilbanci (d’ Orbigny); d’Orbigny, pl. 637, figs 7-8. 1851 Diastopora arborescens d’Orbigny: pl. 638, figs 1-5. 1853 Semielea vieilbanci (d’Orbigny); d’Orbigny: 636, pl. 738, figs 5—9. 1853 Semimultelea arborescens (d’Orbigny); d°Orbigny: 652, pl. 741, fig. 5. 1890 Semielea vieilbanci (d’Orbigny); Pergens: 392. 1893 Semielea vieilbanci (d’Orbigny); Pergens: 211. 21897a Semielea vieilbanci (d’Orbigny); Canu: 155. 21897b Semielea vieilbanci (d’Orbigny); Canu: 750. 1899 Elea vieilbanci (d’Orbigny); Gregory: 300, fig. 33. Figs 110-122 SYSTEMATICS OF MELICERITITID CYCLOSTOME BRYOZOANS 43 Fig. 110 Semielea vieilbanci (d’Orbigny, 1850), photograph of MNHN d’Orbigny Collection 6965 (part), Turonian, France; x 8-S. 1912 Meliceritites vieilbanci (d’Orbigny); Levinsen: 47, pl. 1, figs 15 and 16. 1953 Semielea vielbanci [sic] (d’Orbigny); Bassler: G77, fig. 40, 3a, b. 1981 Semielea vieilbanci (d’Orbigny); Voigt: 453, fig. 5a. MATERIAL. Type: there is no material of this species from the type locality (Turonian of Tourtenay; see d’Orbigny 1850, p. 200) in the d’Orbigny Collection, MNHN, although over a hundred specimens from the Turonian of several other locali- ties are registered under number 6965 (e.g. Fig. 110). In view of the absence of topotype specimens, together with the unequivocal identity of the species, no neotype designation is made. Other material: MNHN d’Orbigny Colln 6965, over 100 pieces from the Turonian of various localities in France, including Troot, St Maure, Tours, Angouleme, Villavard. MNHN d’Orbigny Colln 8200 (Figs 121, 122), Senonian, ?Villedieu, ?Vendéme, syntypes of Diastopora arborescens d’Orbigny, 1851, several specimens including Voigt Photo- card 8312 and an un-numbered specimen corresponding to that illustrated by d’Orbigny (1851: pl. 638, fig. 2). ZMC Levinsen Colln unnumbered, Bruillé Poncé and Villedieu, France. VH unnumbered: U. Turonian, Souge; Turonian, Les Ulmes, Eure; U. Turonian, Bois de Gareau, nr Ecom- moi, Sarthe; U. Turonian, Angoumien, La Charte sur la Loire; U. Turonian, Sougé, Loir-et-Cher; Turonian, Luynes, Indre et Loire; Turonian, Les Ulmes, Maine; U. Turonian, _Troot, Loir-et-Cher; U. Turonian, Les Roches, Loir-et-Cher; -Turonian, Les Ulmes, Maine; U. Turonian/Coniacian, La Barre, nr St Maure, Indre et Loire; Coniacian, St Paterne, Indre et Loire; Coniacian, Tours, Indre et Loire; Coniacian, Fécamp, Seine Maritime. BMNH D3747 (4), ‘Senonian’, Lavardin, Gamble Colln; D8959-60, ‘Senonian’, Lavardin; D4807, Craie de Villedieu [Coniacian or Santonian] , S. of les Roches; D4672, Craie de Villedieu, Luynes; D4679, D4800, D4806, D4860 (4), [Turonian], Montloire; D4675, D4676 (5), D4677, D4678 (4), D4683 (6), D4684 (3), D4715, D4799, D4811 (2), D4839, D4840 (3), D4861, D4862 (5), D4864 (2), D4865, D4868, D13573, Craie Marneuse, Villardin; D4937, [?Turonian], “Nodula Chalk’, Villardin Castle, purchd F.H. Butler; D4674 (figd Gregory, 1899: fig. 33), D4685, Craie Marneuse, Chinon; D33939-40, Turonian, St Calais, Sarthe, E. Darteville Colln; D36151—5, ‘Senonian’, Lavardin, Loir- et-Cher, Pergens Colln; D36156, Turonian, Lavardin, Loir- et-Cher, Pergens Colln; D53444, Turonian, Les Ulmes, Maine, Voigt Colln; D59368, Santonian, Crate de Villedieu, Bouchardiére Mbr, Bed 20 of Jarvis et al. (1982), La Boucha- rdiére, Villedieu, MLoir-et-Cher; D59369 (3), Craie de Villedieu [Coniacian or Santonian], Luynes; D59370-2, D59373 (3), D59374 (6), D59375 (sample), Turonian, Lavar- din, Loir-et-Cher, F. Canu Colln; D59376 (6), Turonian, Chatellerault, Vienne, France; BZ 729 (sample), Turonian, St Maure, Indre-et-Loire, France, presd E. Voigt, November 1991. DESCRIPTION. Colony tubular (cavariiform) with branches (Fig. 110) bifurcating, normally subcircular in cross-section but becoming elliptical immediately prior to bifurcation, varying from about 1-0 to 4-4 mm in diameter, typically 3-5-4-5 mm. Axial lumen lined by transversely wrinkled, inferred exterior wall (Fig. 119) and generally 0-6—-1-4 mm in minimum diameter, becoming elliptical in cross section prior to dividing at branch bifurcations. Single layer thickness about 0-3-0-5 mm. Budding apparently occurs only on the lumen-lining wall. Overgrowths occasionally present, origi- nating through intrazooecial fission, either disc-shaped and multidirectional or fan-shaped and unidirectional (Fig. 118); pseudoancestrula an autozooid, aperture about 0-09 mm long by 0-08 mm wide, initiating a secondary zone of astogenetic change of increasing zooid size. Zooid apertures arranged in rough quincunx or in rows transverse to growth direction. Organization fixed-walled. Autozooids (Figs 111, 115) of moderate size, frontally elongate, subhexagonal or subrhomboidal in outline, on average 1-7—1-8 < longer than wide, well-rounded distally; boundary wall salient, forming a widened wall with contigu- ous apertural rim at distal end of zooid; frontal wall flat proximally, sometimes slightly convex distally, pseudopores subcircular. Apertures (Fig. 113) small, occupying less than a third of the frontal area, approximately equidimensional or a little wider than long, attaining maximum width about mid- length, well-rounded distally; apertural rim salient; apertural shelf of moderate width, tapering proximally; hinge line poorly preserved in all scanned specimens, bowed. Opercula (Fig. 112) often preserved in-situ, convex, prominent; pseudopores radially elongate, numbering about 16, arranged in a crescent. Terminal diaphragms (Fig. 117) common, located just proximally to the apertural shelf, with a central depression and pore. Intramurally budded autozooids not observed. Intramurally budded eleozooids (Fig. 111) very common; aperture (Fig. 114) trifoliate, prominent. Kenozooids (Fig. 116) uncommon, developed close to branch bifurcations. P. D. TAYLOR Figs 111-114 Semielea vieilbanci (d’Orbigny, 1850), BMNH D59370, Turonian, Lavardin, Loir-et-Cher, France; 111, group of autozooids, some containing intramural eleozooids, X 75; 112, autozooidal operculum, x 330; 113, autozooidal aperture, a) 184 Figs 181-184 Reptomultelea spp., photographs of colonies. 181-183, R. canui (Voigt, 1924), VH 10305, Lower Santonian, Vallstedt, Westfalia, Germany, < 2-9; 181, upper side of the colony showing chimney-like growth around an unpreserved object; 182, profile; 183, underside showing original substrate (centre), two secondary substrates (lower right), and free colony growth. 184, R. goldfussi sp. nov., EM RE 551.763.31.A745/1, holotype, Cenomanian, Essen, Westfalia, Germany, worn upper surface of colony, X 3-3. mean = 0:30 mm; SD = 0-023 mm; CV = 7-8; range = 0:27-0:32 mm mean = 0-17 mm; SD = 0-010 mm; CV = 6-0; range = 0-15-0-18 mm mean = ():15 mm; SD = 0-007 mm; CV = 4-7; range = 0-15-0-17 mm eleozooids (5 zooids from VH 10305) frontal length: range = 0-56-0-60 mm frontal width: range = 0-30-0-33 mm apertural length: range = 0-12-0-15 mm apertural width: range = 0-06—0-08 mm frontal width: apertural length: apertural width: REMARKS. Reptomultelea canui co-occurs with the com- moner R. polytaxis (Voigt, 1924) in the Santonian of Westfa- lia and can be distinguished from the latter by its larger autozooids and less numerous kenozooids. The combination of well-rounded autozooidal apertures and narrow arch- shaped eleozooidal apertures can be used to recognize R. canui among other species of Reptomutltelea. DISTRIBUTION. Santonian of Westfalia, Germany. Reptomultelea convexa sp. nov. Figs 191-196 MATERIAL. Holotype: VH 10467, Turonian (?), Chenu, between Le Mans and Tours, Sarthe, France, collected by G. Breton. Paratypes: VH 10443, 10444, 10550 (8 fragments), Turo- nian, Bois de Gareau, near Ecommoi, Sarthe, France. TEMATICS OF MELICERITITID CYCLOSTOME BRYOZOANS Figs 185-190 Reptomultelea canui (Voigt, 1924), VH 10305, Lower Santonian, Vallstedt, Westfalia, Germany; 185, autozooids and eleozooids, x 40; 186, autozooidal operculum, x 300; 187, autozooidal aperture closed by terminal diaphragm, x 250; 188, autozooids and eleozooids, X 85; 189, primary eleozooid aperture, X 250; 190, intramural eleozooid aperture, x 250. Figs 191-196 Reptomultelea convexa sp. nov. 191-193, VH 10467, holotype, Turonian (?), Chenu, between Le Mans and Tours, Sarthe, France; 191, autozooids and eleozooids, x 42; 192, autozooidal operculum, x 200; 193, autozooidal aperture with broken terminal diaphragm, x 200. 194-196, Turonian, Bois de Gareau, near Ecommoi, Sarthe, France. 194, VH 10444, partly-formed, broken gonozooid (upper left) and overgrowth (upper right), x 33. 195-196, VH 10443; 195, intramural eleozooids and autozooids, X 95; 196, aperture of intramural eleozooid, Xx 200. NAME. With reference to the marked convexity of the distal part of the frontal wall. DESCRIPTION. Colony unilamellar or multilamellar, each layer between 0-23 and 0-30 mm thick, often cavariiform and generally with a free-growing, undulose basal lamina. Over- ‘growths (Fig. 194) originate through intrazooecial fission. Organization fixed-walled. Apertures arranged roughly in -quincunx. Autozooids (Figs 191, 195) medium-sized, frontally elon- igate, less than twice as long as wide, usually subhexagonal or subrhomboidal in outline, subrounded distally; frontal wall iconvex, especially close to the hinge line; boundary wall ‘salient. Aperture (Fig. 193) of moderate size, occupying about a third of the frontal surface, longitudinally elongate, sabout 1-2-1-3 x longer than wide, arched distally, attaining ‘maximum width about mid-length (or a little proximally of ‘mid-length); apertural rim slightly raised; hinge line poorly preserved in all available specimens; apertural shelf conspicu- ‘ous, wide to moderately wide, tapering proximally. Opercu- lum (Fig. 192) convex; pseudopores not visible in poorly ‘preserved specimens. Intramurally budded eleozooids com- ‘mon; aperture often cowl-like, directed proximally (Fig. 190). Kenozooids infrequent. Eleozooids (Figs 185, 188) common, often clustered in groups of 2 or 3, frontally elongate, less than twice as long as wide and smaller than the autozooids, pointed distally. Aper- jture (Fig. 189) of small size, trifoliate, about twice as long as wide. Opercula not observed. Intramurally budded eleozoo- lids apparently present, distinguished from ‘non-regenerated’ ‘eleozooids by proximal inclination of apertural plane. Gonozooids known from one complete and one broken example (Fig. 194). Frontally of small size, elongate, over twice as long as wide, a parallel-sided tube of variable length emerging from the maternal aperture and dilating into a subcircular distal portion. Ooeciopore transversely elongate, |twice as wide as long. Atrial ring not observed. q MEASUREMENTS. autozooids (10 zooids from holotype VH 10467) frontal length: mean = 0:55 mm; SD = 0-053 mm; CV = 9-6; range = 0-48-0-62 mm mean = 0-33 mm; SD = 0-017 mm; CV = 5-0; range = 0:30-0:35 mm mean = 0-20 mm; SD = 0-009 mm; CV = 4:3; range = 0-18-0-21 mm mean = 0-16 mm; SD = 0-007 mm; CV = 4-5; range = 0:15-0:17 mm { apertural length: apertural width: mean = 0:49 mm; SD = 0-052 mm; CV = 10-6; range = 0-44-0-57 mm mean = 0-28 mm; SD = 0-035 mm; CV = 12-4; range = 0:21-0:32 mm mean = 0-12 mm; SD = 0-019 mm; CV = 16-6; range = 0-09-0-14 mm mean = 0:06 mm; SD = 0-005 mm; CV = 8-6; range = 0-06-0-08 mm gonozooids (1 zooid from VH 10550) frontal length 1-62 mm distal frontal wall 1-26 mm length: frontal width: 0-71 mm oeciopore length: 0-08 mm ooeciopore width: 0-15 mm SYSTEMATICS OF MELICERITITID CYCLOSTOME BRYOZOANS 69 REMARKS. This species is represented only by fragmentary specimens which are either cavariiform or free-growing lamellar. The high convexity of the distal frontal wall close to the aperture is particularly characteristic. Eleozooids have trifoliate apertures like those of Reptomultelea betusora nom. nov., R. polytaxis (Voigt) and R. auris sp. noy., but R. convexa has much larger autozooids than R. polytaxis and is distinguished by its convex frontal walls from all three species. The intramurally budded eleozooids in the paratypes (Figs 194-196) are more strongly cowled, autozooidal apertures larger and their shelves broader than in the holotype (Figs 191-193). These differences are not considered to warrant species level separation, however, until sufficient material is available to assess species variability. DISTRIBUTION. Turonian of Sarthe, France. Reptomultelea dixoni (Lang, 1906) Figs 197-201 1899 Reptomultisparsa rowei Gregory: 121 (partim) [non pl. 7, fig. 1]. 1906 Semimultelea dixoni Lang: 64, figs 4 and 12. MATERIAL. Holotype: BMNH D7845, [Coniacian], cortestu- dinarium Zone, Pit No. 32 of Dibley (1900), opposite the Rose and Crown Inn, Kenley, S. of Croydon, Surrey, England, Withers and Chatwin Collection. Other material: BMNH D3031 (2 fragments), Upper Chalk, Offham Pit, Lewes, Sussex, Capron Colln (originally described as Reptomultisparsa rowei Gregory, 1899). BMNH D31844—50 (fragments of one colony), [Turonian], Dunton Green, Sevenoaks, Kent, England, Dibley Colln. BMNH D43660, [Coniacian], cortestudinarium Zone, Seaford Head, Sussex, England, Rowe Colln. BMNH D46061, D46066, Turonian, /ata Zone, White Cliff to the Hooken, Devon, Rowe Colln. BMNH D57505-9, Coniacian, Craie 4 M. normanniae, above Tilleul No. 3 Hardground (see Kennedy & Juignet, 1974), Etretat, Seine Maritime, France, Taylor & Hammond Colln. DESCRIPTION. Colony unilamellar or multilamellar, each layer about 0-27-0-41 mm thick, occasionally growing freely with an exposed basal lamina. Overgrowths (Fig. 197) origi- nate through intrazooecial fission from one or several closely- spaced parental zooids; pseudoancestrula either an autozooid (aperture 0-15 x 0-12 mm) or a kenozooid, initiating a secondary zone of astogenetic change of increasing zooid size. Organization fixed-walled. Zooidal apertures variably arranged, often irregular. Autozooids (Figs 198, 199) large, frontally elongate, about 1-6-2 x longer than wide, variable in shape; frontal wall slightly convex, pseudopores apparently small and subcircu- lar; boundary wall salient. Aperture of moderately large size, a little longitudinally elongate, on average 1-1 x longer than wide, attaining maximum width about mid-length, slightly pointed distally; apertural rim raised, drawn to a projecting point distally; apertural shelf narrow, tapering proximally; hinge line short, teeth and bar not clearly visible in the inadequately preserved material available. Operculum (Fig. 200) often preserved in-situ, convex; pseudopores not observed due to poor surface preservation. Terminal dia- phragms (Fig. 193) may be present just beneath apertural shelf. Intramural buds not observed. Kenozooids (Figs 198, 199) common, variable in number, intercalated among other zooids in the normal budding 10 P. D. TAYLOR ee Rd oo 199 200 Figs 197-200 Reptomultelea dixoni (Lang, 1906). 197, BMNH D7845, holotype, [Coniacian], cortestudinarium Zone, Kenley, Surrey, England, overgrowth origin, x 23. 198, BMNH D31844, [Turonian], Dunton Green, Sevenoaks, Kent, England, autozooids, kenozooids and an eleozooid, x 55. 199-200, BMNH D57506, Coniacian, Craie 4 M. normanniae, Etretat, Seine Maritime, France; 199, autozooids, kenozooids and eleozooids, x 37; 200, autozooidal operculum, x 215. sequence and also developed at anastomoses between over- frontal length: mean = 0-73 mm; SD = 0-073 mm; growths; frontally polygonal in outline and invariably shorter CV = 10-0; range = 0-62-0-84 mm and narrower than the autozooids. frontal width: mean = 0-41 mm; SD = 0-050 mm; Eleozooids (Figs 198, 199, 201) moderately common, fron- CV = 12-3; range = 0:36-0:50 mm tally elongate, 2-3 x longer than wide, longer but about the apertural length: mean = 0-24 mm; SD = 0-015 mm; same width as the autozooids. Aperture elongate, spatulate, CV = 6:3; range = 0-21-0-26 mm parallel-sided or slightly tapering, widest at the hinge line, apertural width: mean = 0-22 mm; SD = 0-014 mm; rounded distally; rostral shelf extensive, depressed distally. CV = 6:2; range = 0-20-0-24 mm pena (Fig. 201) often preserved in-situ, pseudopores not (5 zooids from BMNH D57506) observed due to poor surface preservation. Terminal dia- ; ins frontal length: range = 0-62-0-72 mm phragms observed. Intramural buds not seen. cares na Gass waknBee frontal width: range = 0-36-0-41 mm apertural length: range = 0-23-0-26 mm MEASUREMENTS. apertural width: range = 0-20-0-24 mm autozooids eleozooids (5 zooids from BMNH D57506) (10 zooids from holotype BMNH D7845) SYSTEMATICS OF MELICERITITID CYCLOSTOME BRYOZOANS 71 range = 0-77-1-05 mm range = 0-32-0-42 mm apertural length: range = 0:38-0:59 mm apertural width: range = 0-17-0-29 mm kenozooids (6 zooids from BMNH D57506) frontal length: range = 0-48-0-60 mm frontal width: range = 0-32-0-38 mm REMARKS. The holotype specimen lacks unequivocal eleo- zooids, as noted by Lang (1906) in his original description of Reptomultelea dixoni which is based entirely on this speci- men. However, eleozooids are present in most other colonies (e.g. Figs 198, 199, 201). Among congeneric species, R. dixoni closely resembles R. sarissata Gregory, 1899 in the morphology of the eleozooid which has an elongate aperture and is often spatulate. However, although similarly pointed, the autozooidal apertures of R. dixoni lack the distal prolon- gations of R. sarissata apertures which give them a distinctive ogee arch-shape (Figs 282, 283). The proportion of kenozooids to other zooids varies greatly between colonies of this species. In some colonies (e.g. BMNH D31844), kenozooids occupy about the same surface area and are more numerous than autozooids. The high proportion of kenozooids in colonies like this one are unre- lated to disturbances in growth caused by anastomoses of overgrowths. Colonies vary from lamellar to nodular in shape. Specimen ,BMNH D57505 bioimmures a perished substratum of unknown identity which was roughly cylindrical in outline and ornamented by tubercles. _ frontal length: frontal width: DISTRIBUTION. Turonian (/ata Zone)-Coniacian (cortestudi- | narium Zone) of southern England and northern France. Reptomultelea filiozati (Levinsen, 1912) Figs 202-208 1912 Meliceritites filiozati Levinsen: 34, pl. 6, figs 7-10. 1990 Reptelea filiozati (Levinsen); Taylor: fig. 11.2. MATERIAL. Lectotype (herein designated): ZMC Levinsen Collection M13, ‘Middle Senonian’, Fécamp, Seine- Maritime, France; the specimen shown in Levinsen’s plate 6, figure 7. Paralectotypes: ZMC Levinsen Colln M12, M14—20, | same details as lectotype. Other material: BMNH D54286-7, Coniacian, Fécamp, Voigt Colln. VH un-numbered specimens from same horizon and locality. DESCRIPTION. Colony unilamellar or multilamellar, each layer about 0-3 mm thick, often becoming free with a transversely wrinkled basal lamina. Overgrowths apparently originate through intrazooecial fission, often from several closely-spaced pseudoancestrulae (Fig. 206); pseudoancestru- lae are autozooids with apertures about 0-11 mm long and wide, sometimes operculate (Fig. 207); peri-pseudo- ancestrular zooids number 5-6 and commence a zone of secondary astogenetic change marked by increasing zooid size. Organization fixed-walled. Zooidal apertures arranged roughly in quincunx. Autozooids (Figs 202, 205) of large size, elongate, on average a little less than twice as long as wide, diamond- shaped, pentagonal, rhomboidal or hexagonal in outline, well-rounded distally; frontal wall gently convex, occupying about two-thirds of the frontal surface; pseudopores small, circular; boundary wall salient, clearly visible. Apertures (Fig. 204) of moderate size, elongate, about 1-25 x longer Fig. 201 Reptomultelea dixoni (Lang, 1906), BMNH D57505, Coniacian, Craie 4 M. normanniae, Etretat, Seine Maritime, France; eleozooid with in-situ operculum, x 105. than wide, attaining maximum width about mid-length, well- rounded distally; apertural rim slightly raised; apertural shelf narrow, tapering proximally; hinge line with a median bar. Operculum (Fig. 203) often preserved in-situ, slightly convex; pseudopores not clearly visible in studied material. Dia- phragms and intramural buds not observed. Kenozooids (Figs 202, 205) common, especially clustered at anastomoses between overgrowths. Eleozooids (Figs 202, 206, 208) moderately common, large, frontally elongate, about 2-5-3 x longer than wide, longer but slightly narrower than the autozooids. Aperture elongate, about twice as long as wide and twice the length of an autozooidal aperture, more or less parallel-sided, well- rounded distally with an extensive rostral platform depressed beneath the level of the apertural rim; hinge line with a long median bar. Opercula not observed in-situ. Intramural buds not seen. Gonozooids unknown. MEASUREMENTS. autozooids (10 zooids from lectotype ZMC M13) frontal length: mean = 0-62 mm; SD = 0-049 mm; CV = 8-0; range = 0:56-0:71 mm Figs 202-207 Reptomultelea filiozati (Levinsen, 1912), ‘Middle Senonian’, Fécamp, Seine-Maritime, France. 202-204, ZMC Levinsen Colln M12; 202, autozooids, eleozooids and kenozooids, x 55; 203, autozooidal operculum, x 245; 204, autozooidal aperture, x 250. 205-207, ZMC Levinsen Colln M13, lectotype; 205, area of coalescent growth with kenozooids and autozooids, x 55; 206, four closely-spaced pseudoancestrulae surrounded by radiating autozooids, eleozooids and kenozooids, x 55; 207, operculate pseudoancestrula surrounded by five autozooidal buds, x 78. SYSTEMATICS OF MELICERITITID CYCLOSTOME BRYOZOANS 73 Fig. 208 Reptomultelea filiozati (Levinsen, 1912), ZMC Levinsen Colln M12, Middle Senonian’, Fécamp, Seine-Maritime, France; eleozooid, x 155. > 4 mean = 0-34 mm; SD = 0-036 mm; CV = 10-7; range = 0-30-0-42 mm mean = 0-20 mm; SD = 0-009 mm; CV = 4-8; range = 0-18-0-21 mm mean = 0-16 mm; SD = 0-006 mm; | CV = 3-9 range = 0-15-0-17 mm | eleozooids (5 zooids from lectotype ZMC M13) frontal length: range = 0-68-0-80 mm _ frontal width: range = 0-27-0-30 mm | apertural length: range = 0-38-0-41 mm apertural width: range = 0-17-0-20 mm | frontal width: | apertural length: / apertural width: REMARKS. The shape of the eleozooidal aperture is very characteristic in Reptomultelea filiozati and enables immedi- jate distinction from most other species of the genus. Only R. pegma sp. nov. (see p. 85) and R. scanica sp. nov. (see p. 96) have similarly broad eleozooidal apertures, but both species ‘are readily distinguished from R. filiozati by other means (e.g. R. pegma has oblique autozooidal apertures with wide japertural shelves, and R. scanica has autozooidal apertures ‘which occupy a significantly greater proportion of the frontal surface than in R. filiozati). According to Prof. E. Voigt, this is a rare species. It is known to occur with certainty only in the Coniacian of the Fécamp area of Seine-Maritime in northern France, although the Voigt Collection in Hamburg contains a possible example (un-numbered) from the Coniacian of Villedieu. DISTRIBUTION. Coniacian of Seine-Maritime, France. Figs 184, 209-216 MATERIAL. Holotype: EM RE 551.763.31.A745/1, Cenoma- nian, Essen, Westfalia, Germany. Paratypes: VH 10426, 10548 (8 fragments), Lower Cenomanian, Miilheim/Ruhr, Westfalia, Germany. Reptomultelea goldfussi sp. nov. NAME. In honour of Georg August Goldfuss (1782-1848), the first author to describe a melicerititid species, Ceriopora gracilis Goldfuss, 1827, later to become the type species of Meliceritites Roemer. DESCRIPTION. Colony (Figs 184, 211) unilamellar or multila- mellar, each layer about 0-25-0-35 mm thick, sometimes growing freely of underlying layers and incorporating second- ary substrata. Ancestrula not observed. Overgrowths origi- nate through intrazooecial fission, often from several closely- spaced parent zooids; pseudoancestrula (Figs 212, 213) an autozooid which may be occluded by a terminal diaphragm, initiating a secondary zone of astogenetic change of increas- ing zooid size. Organization fixed-walled. Zooidal apertures arranged in regular quincunx, disrupted close to overgrowth origins. Autozooids (Figs 209, 212) of moderate size, frontally elongate, about 1-5-2 x longer than wide, rhomboidal in outline, well-rounded distally; frontal wall occupying about half of frontal surface, slightly convex, with circular pseudopores; boundary wall salient, variably prominent. Aperture (Fig. 212) large, usually a little longer than wide, attaining maximum width just proximally of mid-length; apertural rim conspicuous but low, continuous with lateral zooecial boundary wall; apertural shelf rather narrow (less than 0-02 mm wide), tapering to nothing at widest point of aperture; hinge line with median bar and low teeth at either end. Operculum (Figs 210, 220) occasionally preserved in-situ, surface convex; pseudopores elongated parallel to growth direction, scattered across entire surface of opercu- lum. Terminal diaphragms seldom observed, pseudoporous. Intramurally budded autozooids uncommon. Kenozooids not observed. Eleozooids not observed, presumed absent. Gonozooids represented by one example in the holotype colony (Fig. 209); frontal surface slightly longer than wide, becoming triangular in outline soon after emergence of the distal frontal wall from the maternal aperture; margins of distal frontal wall irregular, indented by adjacent autozooids. Ooeciopore very large, about the same width as an autozooid aperture but less than half the length. MEASUREMENTS. autozooids (10 zooids with in-situ opercula from holotype EM RE 551.763.31.A745/1) frontal length: mean = 0-55 mm; SD = 0-051 mm; CV = 9-2; range = 0-48-0-63 mm mean = 0-32 mm; SD = 0-018 mm; CV = 5-5; range = 0-30-0-35 mm frontal width: P. D. TAYLOR Figs 209-210 Reptomultelea goldfussi sp. nov., EM RE 551.763.31.A745/1, holotype, Cenomanian, Essen, Westfalia, Germany; 209, gonozooid and autozooids, x 30; 210, autozooidal operculum, x 150. mean = 0:26 mm; SD = 0-022 mm; CV = 8:8; range = 0-23-0-30 mm mean = 0-25 mm; SD = 0-018 mm; CV = 7:2; range = 0:23-0:29 mm apertural length: apertural width: gonozooids (1 zooid from holotype EM RE 551.763.31.A745/1) frontal length: 2:03 mm distal frontal wall 1-80 mm length: frontal width: 1-80 mm ooeciopore length: 0-11 mm ooeciopore width: 0-24 mm REMARKS. The large and well-rounded autozooidal aper- tures distinguish Reptomultelea goldfussi from other species in the genus. As noted below (p. 91), R. goldfussi resembles R. reedi sp. nov. but is distinguished most convincingly by the occurrence of pseudopores across the entire surface of the operculum rather than in a crescentic row. The holotype (Figs 209, 210) differs from the paratypes (Figs 211-216) in having somewhat taller and larger autozooidal apertures. These differences may be astogenetic in origin because the paratypes comprise zooids close to overgrowth originations and probably within secondary zones of astogenetic change. DISTRIBUTION. Lower Cenomanian of Westfalia, Germany. Reptomultelea levinseni sp. nov. Figs 217-223 MATERIAL. Holotype: BMNH D58930 (2 fragments), Conia- cian or basal Santonian, decipiens Chalk, Vattetot, Seine Maritime, France, Taylor & Hammond Collection. Paratypes: BMNH D58931 (sample), D59248, D59249, same horizon and locality as holotype. ZMC Levinsen Colln M56, [?Coniacian], Fécamp, Seine Maritime, France; labelled by Levinsen ‘Clausimultelea sp.n.?’. NAME. For G.M.R. Levinsen, author of the most important work on melicerititid cyclostomes. DESCRIPTION. Colony generally multilamellar, each layer about 0-26 mm thick and occasionally growing free of under- lying layers and with an undulose basal lamina. Ancestrula not observed. Overgrowths originate by intrazooecial fission, often from several clustered parental zooids depressed beneath general level of colony surface; pseudoancestrula an autozooid or a kenozooid budding a ring of daughter zooids, sometimes including a gonozooid (Fig. 221), forming the initial part of a secondary zone of astogenetic change through which zooid size increases. Organization fixed-walled. Zooids arranged roughly in quincunx. Autozooids (Figs 217, 218) of large size, frontally elongate, about 2-2-5 x longer than wide, generally hexagonal to diamond-shaped in outline, pointed distally; frontal wall occupying more than half of the frontal surface, slightly convex, with circular pseudopores set in elongate depres- sions; boundary wall well-defined, salient. Aperture (Fig. 219) of moderate size, approximately 1-5 x longer than wide, gothic arch-shaped, pointed distally, attaining maximum width at a point between the hinge line and mid-length; apertural rim well-developed, pointed and raised distally; apertural shelf narrow; hinge line short, with low teeth at either end of a median bar. Operculum (Fig. 220) often preserved in-situ, convex, pseudopores present but number uncertain. terminal diaphragms may be present beneath level of operculum. Intramural buds not observed. Eleozooids (Figs 217, 218, 223) abundant, scattered; frontal surface about 3-4 x longer than wide, considerably longer than the autozooids but approximately the same width; frontal wall occupying more than half of the frontal surface, slightly convex and with a similar density of pseudopores to the autozooids. Aperture elongate, 2-3-5 x longer than wide, widest a little distally of the hinge line, with a long, narrow rostral area becoming parallel-sided or even very slightly spatulate; rostrum depressed distally; hinge line with a median bar proximal to which is a depressed area; aperture width constricted proximally by wall thickenings which join distally to form the rostral shelf. Operculum (Fig. 223) sometimes preserved in-situ, surface convex, especially distally. Terminal diaphragms and intramural buds not observed. 211 212 Figs 211-216 Reptomultelea goldfussi sp. nov., VH 10426, Lower Cenomanian, Miilheim/Ruhr, Westfalia, Germany; 211, small colony encrusting a shell fragment, x 14; 212, autozooids radiating from two closely-spaced pseudoancestrulae with apertures closed by terminal diaphragms, x 45; 213, detail of occluded pseudoancestrulae, x 130; 214, autozooidal operculum, x 200; 215, autozooidal aperture, x 200; 216, growing edge showing new buds with porous interior walls, x 97. Figs 217-222 Reptomultelea levinseni sp. nov. 217, ZMC Levinsen Collection M56, [?Coniacian], Fécamp, Seine Maritime, France; autozooids and two eleozooids with rostra resting on kenozooids, the right-hand eleozooid fouled by a small stomatoporid cyclostome; x 60. 218-222, Coniacian or basal Santonian, decipiens Chalk, Vattetot, Seine-Maritime, France. 218-220, BMNH D58930, holotype; 218, partly exfoliated specimen with operculate autozooids and eleozooids, - aa; a °.. ae = i ei ok ee ie | v vies irl ‘. ew 4 oma By ; —_ aes Py eaene | ' 3 | vi if ‘i or ee UL me aan tain iets " : a : { ' 2 om acy eo =<. lie te \ a ; a oo (pa atid “at i ee, My = Tin leks meal dad _- : ’ : ot J =) 7: : — = — or bahar = 7 on =e. 3 ¢ qmaglts- iy) GS me ogee), pe onl le ' ny er Vo See el Merl cai 4 eae ies _ ; (Me A. cor qd] “er ‘ae = ee = v ; o { ; ’ a ee 5S seme fe 5,8 2 Wiese Asam gah amyl, afl’ Ply | . : sy ae sae \ one Seervh ee exe's % r > ~~ as ry aii 4 ee ee — —— r amin = i - | a i vedniy : ' Aa i. at 008) re Wu a al pay ge a = - ~ ‘ Ly sy . alt oad AM als heel Cee Li SM at ; Nas i) | =e iF) - hears ai a ai am: @ eo citys fl i , \ ibe “ ; SD salt cine ae oe ; “ ’ : - i i = os 7 3 = = ; - | as ® s ino - %. 7 i ~ = ° ‘ 7 2 ate > o-~ i sans _ : d ; — = so : = ti : ~~, 4 oe ‘ » . - ; v4 - ‘ -» = — ‘stain & / J hers ot t a : > 20 Sear Pokey 3 oS See “=v ara ‘ ouepi 7 ar ; le eee core 7 ta 7 i = i ys, eee Td a Se 2 eee © «eps pe he edge - aa raha meme “Sw ay wiy 1D OT iran i Guee < ee oa ay ee Oe (ies «mt a ae ae ee aS ean ee ee ues ts Stee “pee euler area By fake a - a a » ples : ae Abe ~ ; nn ’ A co 7 fa ©. % i 1 a ~~ > = Pe 7 = } , t 7 oz a ey ee tire he j . oy i heie ——_ Gara ve o= i . j ete) @ ce ry ¢t < 0 taf o- = 7 ri — ea Cpa | " e4 ; — Gatlin Ge Teaabhe ann to wo ul} 7 ® *) ba : wo y : , , ’ 4 a re a is ° ; es ¢ we om.4 ey 7 : a » = then Syhades, Wher # i ; 7. —— wit) =.ar}gsosebede 3ponogsseoccpNceBbooeos00 snqeersasagheadoc: 148 Suyiseiianviky mae earaleal WHEN) Sccoccneatoscqadceusacuseen doooctacdes sadunadd ake oat> sonido dseocabedasnodsonocuLCRncesdsoso: 150 Qing tetera Cia AVOUIIG ISIS. geodcooeecdcane coocteeeobe 600-000¢ cerbusbbsbScb ocuccemEpooUjon socneascadeeboocnepEceauoCo> 150 QTE CZ AUIE 5). cocdeacedecdecodeGacheeJaddee eos 000 CDS 9B EER RRB 96 DaGccd .gdebaGdeuac con coud SoCUuCLCCQUUE aanpsa deo TOC acecceeeohbat 150 ©) The Natural History Museum, 1994 106 M. A. PARKES REUSCH EM AD Spr stern seats eseain ge astce islisis ati Souiels eee sites oleae lds nates a ano tet sass ese RC Rie RESORT see eR Cee 150 RY/[12} 1178S Omermenatte penned! seteccrsar starter dn aeer cane cero mmaenec nce. meee corereeeecn SoomeeteconesarCeeponeedeace semtecaeec csece 152 Saukrodictyaich ino | at | oo PORTRANE 22) Fe Uy, dy] [oui Rp 167 | 168 [isavti7o fee | 181 a KILDARE INLIER Leinster Granite e Portlaoise (Wicklow RATHDRUMe Leinster Granite 6 g Gorey Q URTOW BALLYKALE ° COORTIONN Kilkenny CLOLOGE, GREENVILLE SS e Enniscorthy® y RAHEEN, Clonmel e = glare Te oF ecane * Dungarvan c he 1 A, index map of Geological Survey of Ireland 1-inch sheets examined in whole or part for revision of fossil localities. B, location map for the main sections shown on Fig. 14 (p. 126), and other localities in the Leinster terrane and adjoining terranes. 109 110 Table 1 and 149, with present status. Localities from G.S.I. Memoir examined in Sheets 148 Loc. ‘'/4 sheet County & Situation & geological no. of6’map townland formation & 1” map Wexford Sheet 148 1 16/1 Clologe, Upper From debris, in field close to road, from Norris Mount to Milltown, Imile S of Camolin; light grey & brown slates, & tuffose rock. See later section. 2 20/1 Killabeg Quarry on bank of R. Bann, | mile S of Clone Wood & 3 miles S of Ferns; black slates. Graptolite loc. — not examined. 3) 20/2 Ballydonegan On road from The Harrow to Tinnacross, 1 mile SW of The Harrow; grey shales. No exposure found here. Wexford Sheet 149 4 7/4 Kildermot See Courtown localities. 5 7/4 é : 6 7/4 me 4 7 7/4 Ballymoney, Lr. 8 7/4 Seafield ‘ 9 11/2 Ballykale See later section. 10 11/2 Coolnaveagh About 2 miles S of Gorey a little W of Ballinatray Bridge; black slates. Not located. 11 11/3 Ballydaniel One mile W of Balloughter; bluish grey compact altered rock. No fossils found here. 12 11/4 Clogh and A little SE and SW of Clogh. Frankfort See later section. 13 12/1 Ballinatray, Lr. See Courtown localities. 14 12/1 i sf 15 12/1 ih M2 16 12/1 Seamount ‘ 17 12/2 Duffcarrick "4 18 12/3 Coolnahinch A little SW of Ballywalter House, 2-5 miles S of Gorey. Not traced. 19 12/4 Seamount Graptolite locality in Ribband Group — not examined. by a religious grotto about 100m from the bridge at Rathdrum. However, no fauna was found in the very slaty rock there. (ii) Wicklow 30/3 one mile on road from Rathdrum to Redcross. This was considered too obscure and the supposed specimens too poor even in 1869 and no attempt was made to trace the locality in this study. (iv) An old road cutting in the townland of Slieveroe, Co. Wicklow 30/1. This is discussed in brief below. M. A. PARKES Table 2 Localities from the G.S.I. Memoir examined in Sheet 158, with present status. Situation & geological formation & 1” map Loc. 1/4 sheet no. of 6” map County & townland Wexford Sheet 158 1 19/4 Kiltrea Graptolite loc. See Brenchley et al. (1967). Quarry near road, a little S of Moyne House, 1-5 miles N of Enniscorthy; dark grey slates. Quarry now infilled — no exposure at all. See Brenchley et al. (1977) for faunal lists from collecting in that revision. A little NE of Moyne House, 2 miles N of Enniscorthy; light bluish shales, weathering brown. No exposure at present. Old quarry, 1 mile N of Enniscorthy; dark grey shales. Fauna collected here in large blocks from newly ploughed strawberry field immediately adjacent to quarry. See later section. See later section. Between Ballynabarny House and White’s Bridge, 1-5 miles NE of Enniscorthy; dark grey slates. This site was not examined in this study or by Brenchley et al. (1977). A little N of Ballybrennan House, 1-5 miles W of 2 20/3 Moyne Upper 3 20/1 Moyne Lower 4 20/3 Greenville & Moyne Upper 5) 20/3 6 20/3 Greenville Clonhasten & Ballynacarny boundary 7 31/2 Ballybrennan Clonmore, & about 6 miles | | | | SW of Enniscorthy; grey shales and grits. Shannon (1979a) recorded fossils here but no exposure is now available. Near Chapel Village, 6 miles SW of Enniscorthy; So ivi | light grey shales & grits. | Raheen See later section. Sheet 129 The memoir for this sheet (Mitchell 1884) has one palaeonto- logical note by W.H. Baily concerning the only known locality at Ballintaggart, Co. Kildare. This was examined by Briick (1971) in his examination of fossil localities west of the Leinster granite, and is not included here. Sheets 138 and 139 Hull (1888), author of the G.S.I. memoir, mentions the only fossil locality in the text (p.8). A graptolite locality with poorly preserved forms ‘allied to, or identical with Grap- tolithus Sedgwickii’ was known a short distance NW of Arklow. The author has recently traced and curated these BRACHIOPODS OF DUNCANNON GROUP OF SE IRELAND | Table 3 Localties from the G.S.I. Memoir examined in Sheet 168, with present status. County & townland Situation & geological formation & 1” map Loc. '/4 sheet no. of 6” map Waterford Sheet 168 111 Table 4 Localities from the G.S.I. Memoir examined in Sheet 169, with present status. Loc. ‘'/4 sheet no. of 6” map County & townland Situation & geological formation & 1” map Wexford Sheet 169 } 1 9/4 Gibbet Hill New road cutting on S side of R. Suir, a little NW of Waterford; black argillaceous slates. Graptolite locality, not examined. Old quarry at the back of Farm House, 3 miles S of Waterford, on the road to Clohernagh Bridge; brown calcareous impure . limestone. G.S.I. 6” fieldsheets were not accessed for this area and it was not traced. Rocks on shore, a little N of Newtown Head, Waterford Harbour; dark grey concretionary shales. | Owen et al. (1986) have revised the trilobite fauna. D.A.T. Harper is revising the brachiopod assemblage from here. 1217/4 Killnre Raheen specimens in the G.S.I. collections. They have been identi- fied by Dr. A. Rushton, and indicate an early Ordovician, possibly Arenig age (John Morris G.S.I., pers. comm., 1994). ‘Sheets 148 and 149 This memoir (Hardman 1887) has palaeontological notes by W.H. Baily, who tabulated the localities. These are repro- ‘duced here with relevant information from this revision ‘(Table 1). | Sheets 158 and 159 W.H. Baily contributed palaeontological information io this ‘memoir (Kinahan 1882), again tabulating the fossil localities ‘and listing the fossils collected. These are listed here with applicable information from this work (Table 2). All are from |Sheet 158; none were known from Sheet 159. Sheets 167, 168, 178 and 179 This memoir (Du Noyer 1865) also has palaeontological notes by W.H. Baily, who listed the fossils collected and tabulated the localities on Sheets 168, 178 and 179 (none were known \from Sheet 167). The area of Sheet 167 is largely Caradoc or Silurian and has been studied by Penney (1980), who dis- ‘cussed recent faunal dating (p.319), including a Caradoc |graptolite record from the Ross Formation. Sheet 178 to the west of the main Tramore volcanics is also outside the scope of this research as discussed in the Introduction. Similarly, Sheet 179 is all outside the confines of the present project and the subject of revision by D.A.T. Harper, H. Carlisle and Oo n 2 = & iS) About 1-5 miles SE of Ballynabola; grey shales. See later section. A little N of Newbawn, 1-5 miles SE of preceding locality; grey shales. No fossiliferous exposure found here. A new quarry exposure with fossils was recently reported to me (pers. comm. M. Allen) but not visited in this research. A little S of Ballygarvan Bridge, 3 miles SW of preceding locality; grey shales. See later section. Rocks on shore a little W of Ballymadder Point; dark grey shales. Reconnaisance visit only made here. No fossils located. Dan Tietzsch-Tyler (pers. comm. 1986) did not find significant fossils while mapping the area in detail. Rocks on shore W of preceding locality; grey micaceous and argillaceous shales. Same comment as loc. 4 applies. Carrigadaggan Newbawn 3} 40/1 Ballygarvan 4 45/4 Ballymadder 5 45/4 Loftusacre myself. The area is described by Carlisle (1979). However, the Kilbride locality discussed below is within the confines of Sheet 179 and adjacent to localities 13-16. It is worth noting the confusion caused in the past by the similarity of the names of Newtown Head in Waterford Harbour (locality 3 — Raheen, but not the Raheen near Enniscorthy) and that of Newtown Cove, Great Newtown Head and Newtown Glen, all on the west side of Tramore Bay, fossilif- erous strata being found at all locations. Sheet 168 contains three localities, shown in Table 3. Sheets 169, 170, 180 and 181 Kinahan (1879) wrote the memoir to the four sheets covering southeast Co. Wexford, with W.H. Baily again contributing palaeontological information in tabulated form. Sheet 169 is the one relevant to this study. The table of ‘Lower Silurian’ localities is reproduced here with updated information (Table 4). Sheet 170 has two listed localities which are part of the Rosslare terrane and whose faunas were described by Brenchley et al. (1967b). These are currently being reassessed by Harper & Bates (in prep). Sheet 180 has only one graptolite locality, visited but not yielding any specimens; recent workers have not found any trace of them and suggest the deformation is too strong to preserve fossils (Gardiner 112 1967: 6). Sheet 181 has no fossil localities at all. A further significant point to note in connection with the composite list of fossils collected from these sheets is that in the sections on ‘Lower Silurian’ trilobites, brachiopods and graptolites many species are recorded from Locality 12. This is definitely a Carboniferous locality. According to the G.S.1. Map Curator, A.G. Sleeman (personal communication, 1988), these records are actually from Localities 16 and 17, which are both in the Tagoat area of the Rosslare terrane. He detailed many further complexities resulting from ‘some rather sloppy curating going on in the 19th century’. These errors clearly show the need for caution in utilizing the existing faunal lists alone in modern interpretative work. DETAILED LOCALITY INFORMATION The Kildare Inlier, Co. Kildare (1:126720 — SHEET 16, N724175 — Horizons 1 & 2, N724179 — Grange Hill Cottage) This inlier is described in detail elsewhere (Parkes and Palmer 1994). Only two main horizons and one minor one have been sampled extensively for the present work. All are of Caradoc age and on the flanks of the andesites of Grange Hill. The three localities sampled are shown in Fig. 2. The older fauna at the back of the ruined farm cottage, called Grange Hill House Cottage herein, is on the northern side of Grange Hill, on the edge of the common land. Grange Hill Horizons 1 and 2 are located only a few metres apart on the lowest slopes of Grange Hill in and just above an old obsolete field boundary. The oldest reference to the inlier seems to be M’Coy (1846), who listed many of his species from ‘the Chair of Kildare’ (in the townland of Carrickanearla). This has led to much subsequent confusion since the townland includes rocks of Caradoc and Ashgill age and fossils from different horizons were treated together. In 1858 the G.S.I. memoir to Sheet 119 (35 NE) was published (Jukes et al. 1858). A separate list was given for fossils found at Grange Hill House Cottage, but the identifi- cations were not indicative of significant differences between that locality and the combination list for the Kildare Lime- stone and Grange Hill. These fossils have never been described although Williams et al. (1972) stated it was a Soudleyan fauna. Wright (1970) published a study of the inarticulate brachiopod Orthisocrania divaricata, which is found only in the Caradoc siltstones dated as Longvillian on the basis of faunal similarity with the Gelli-grin Group of Bala in North Wales. Wright’s list is the only modern reference to the Grange Hill (Horizons 1 & 2) fauna, which is described fully herein. Kilbride and adjacent localities, Co. Waterford (1:126720 — SHEET 23, S578050) The Kilbride locality is not a G.S.I. nor a Griffith locality but is one of the few recently discovered fossil sites in the area. It was found and collected first by Hilary Carlisle in the course of her doctoral research. Carlisle (1979) described the stratig- raphy of the Tramore area, Co. Waterford, and listed the genera from Kilbride. Although Carlisle’s work was never M. A. PARKES completed, the material she collected was largely saved by Dr D.A.T. Harper with the intention that they should jointly complete the taxonomic study of the Tramore faunas. Although the Tramore Limestone Formation collection is currently under study by Harper, Carlisle and myself, the Kilbride material was lost. Extensive re-collection of this stratigraphically significant locality in the Upper Tramore Volcanic Formation was a high priority in the present study. The locality is shown in Fig. 3 and a detailed survey of the quarry is shown in Fig. 4, which pinpoints the position from — which the re-collection was made. In Du Noyer (1865:18) four localities are listed by Baily which are adjacent to Kilbride on 1” Sheet 179. These are Towergare (13 & 16), Munmahoge and Lisduggan (14) and Munmahoge (15), all townlands half a mile to the north of Kilbride. For all four localities the memoir descriptions are vague. Recourse to the 6” G.S.I. fieldsheets was necessary since the localities are not even indicated on the 1” Sheet 179. The original fieldsheets have been replaced by photographic copies in the G.S.I. and only limited information could be obtained. Munmahoge (15) was examined closely but the | other localities were not traced. No fossiliferous exposures were found. However, the faunal lists are short with only three or four species other than the ubiquitous bryozoan ‘Stenopora fibrosa’, although at Munmahoge (15) abundant specimens are indicated. Ballykale, Co. Wexford (1:126720 —- SHEET 19, 1147570) This locality was recorded as a G.S.I. locality (No. 9) in the Sheet 149 memoir (Hardman 1887). It was noted as being | ‘one mile and a half south of Gorey; tuffose rock’. The | quarter Sheet 11/2 of the G.S.I. fieldsheets showed the locality, although it is possible to confuse it with any of several adjacent localities such as Coolnaveagh (No. 10), Coolnahinch (No.18), or a number of other sites indicated by asterisks on the 1” Sheet 149, which are not easily correlated — with the memoir table. Kelly (1860) noted the locality as being ‘2 miles south of Gorey, on the east side of the road. The locality is nearly surrounded by a felspathic protrusion of yellow rock, such as is frequent thereabouts’. The actual position of the collection made by me is shown in Fig. 5. The locality has received no attention since the G.S.I. memoir. Field investigation did not reveal any exposure, but one large block in the base of the wall bounding the farm road was found to be packed with fossils, a nearly monospecific assemblage of Bimuria cf. dyfiensis Lockley (p.155). Although not in situ it is believed to be of local origin. The rock is a very tuffaceous mudstone and has been relatively strongly deformed, but the flattened fossil moulds are clearly identifiable. Clologe Upper, Co. Wexford (1:126720 - SHEET 19, TO51509) As locality 1 in the memoir for Sheet 148 (Hardman 1887), this site was also identified clearly on the G.S.I. 6" field- sheets, on quarter Sheet 16/1 of Co. Wexford. The memoir records that fossils came ‘from debris, in a field close to the road from Norris Mount to Milltown, one mile south of Camolin; light gray and brown slates, and tuffose rock’. Re-investigation of the area, shown on Fig. 6, failed to locate BRACHIOPODS OF DUNCANNON GROUP OF SE IRELAND i} Etrrri2=t=; gaa cee fossil GRANGE HILL 4 743 Or Be fossil localit . y Grange Hill Horizont Chair of Kildare , Wize Earl's Wel! on track - electricity ~ a rough 40°” contour transmission ~ ~/ grazing - line cable | fo) | river or 2" buildings | road or FY stream ve boundary [P,4/woodtand , Fig. 2 Locality map of the three fossil localities in the Kildare inlier; Grange Hill House Cottage, Grange Hill Horizon | and Horizon 2. 113 114 M. A. PARKES Fig. 3 Locality map of the Kilbride locality. Key as in Fig. 2. any in situ exposure. However, collecting amongst loose blocks in the steep wooded slope within a restricted area yielded a large fauna comparable with that listed by Baily (in Hardman 1887). The fossils were found most frequently in blocks of tuff but others came from slaty siltstones and mudstones. The preservation is generally poor and identifica- tion proved difficult beyond generic level. The similarity of the fauna and lithology to that described in the memoir, and the limited section of slope where fossiliferous blocks were found, suggest that they are from the outcrop below the surface drift. Carrigadaggan, Co. Wexford (1:126720 - SHEET 23, $313240) Despite being one of the most fossiliferous localities in the Duncannon Group this site has received scant attention in the last 100 years. Its exact position is shown on Fig. 7. M’Coy (1846) gives this locality for many of his species — an impressive list of 16, mainly brachiopods and trilobites, but including the rhombiferan cystoid Echinosphaerites granula- tus aS an ‘extremely common’ element of the fauna. The faunal list given in the G.S.I. memoir (Kinahan 1879) is even more comprehensive. Thirty-eight genera are listed, again mainly brachiopods and trilobites but also several bivalve and gastropod species, as well as bryozoans, orthocones, conu- lariids and Echinosphaerites aurantium (as a very abundant species). Forbes (1848) also dealt with the locality in connec- tion with Echinosphaerites. Kelly (1860) included the locality in his Wexford list and, unusually, named some species found there. Paul (1973) mentioned it, as one of 14 major cystoid localities, although no details were given. Williams ef al. (1972:57) noted the place as a shelly locality in the Caradoc rocks of Wexford and Waterford. Ballygarvan Bridge, Co. Wexford (1:126720 — SHEET 23, S792188) As one of the few fossil localities in south Wexford, Ballyga- rvan Bridge is often mentioned in connection with Carrigada- ggan, but similarly until now no systematic re-collection of the faunas has been attempted since the late 19th century. M’Coy (1846) listed 9 species; the G.S.I. memoir (Kinahan 1879) has an increased diversity of 11, but with some different species. The exact locality from which previous collections were made is in some doubt. Although the description in the memoir is characteristically imprecise, the G.S.I. 6” field- sheets have a precisely located asterisk indicating the locality. However, the geological boundaries adjacent to the bridge itself are complex and unclear, and since there is no exposure at the indicated spot the possibility of a cartographer’s mistake must be kept in mind. The present exposure is very /BRACHIOPODS OF DUNCANNON GROUP OF SE IRELAND 10m WOODEN FENCE 1 ClO $i ROCK EXPOSURE CLIFF TOP (ROCK) VEGETATED SCARP CONTACT WIT Oy ERY ING RHYOL OSED ne ene Spe m BUSHES & TREES \30/270 = & Lt a 7 - 63 > 1 ot 7 y¥FROM BASE OF ROCK 7” 1 EXPOSURE IN TUFFS 1 AND SILTSTONES Pal YFOSSILS COLLECTED »+++< Ly 3 |Fig.4 Detailed plan of the Kilbride locality. | limited, but a collection was made from hard grey slates in the |wooded banks of the river (see Fig. 8), approximately 20m away along the strike from the supposed locality. The fauna is sparse, although M’Coy (1846) described most elements of his list as common. All three collections were probably made from slightly different horizons in close proximity. Frankfort and Clogh, Co. Wexford (1:126720 — SHEET 19, T109550 (Frankfort), T123555 | (Clough)) These are listed by Baily (in Hardman 1887) as one locality. In fact the 1” sheet 149 and the G.S.I. 6” fieldsheets show | three separate localities, all of which were investigated in the |present work. No fossiliferous exposure was found at any |site, although a few fossils were found in loose blocks at |Clogh. The localities are the main source of fossils used to date the Ballymoney Formation of Hiller (1971) and Mitchell et al. (1972), although Hiller was only able to find wall blocks of fossiliferous rock, the original G.S.I. localities being | infilled or overgrown. The localities are shown in Fig. 9. Raheen, Co. Wexford (1:126720 — SHEET 23, _$891326) | The original G.S.I. locality listed by Baily (in Kinahan 1882) | for 1” Sheet 158 is an old quarry now extremely overgrown, with almost no exposure. Fortunately, during this study the local farmer had excavated a new pit roughly along the strike about 5m away from the old pit, for hardcore. On my discovery, it was already half filled with domestic refuse and / ie ea ; N Oo RUINED COTTAGE ra (CAT TLESHED) a ROAD TO TRAMORE ——s is probably by now completely filled. This temporary expo- sure allowed collection of a large fauna. Fig. 10 shows the exact position of Raheen. The locality was not known to M’Coy (1846) or Kelly (1860), and after collection by the G.S.I. no attention was paid to it until the 1970s. Shannon (1979a: 46) recorded a fossil assemblage indicative of a Caradoc age. Brenchley ef al. (1977) mentioned it in connec- tion with specimens of Plaesiomys, presumably from existing collections. It is an important locality, therefore, in that it provides data along strike from better-known sections at Enniscorthy, and between there and Carrigadaggan to the southwest. It is also important in that the newly collected fauna differs somewhat from previously listed assemblages. Greenville, Enniscorthy, Co. Wexford (1:126720 — SHEET 23, S962412) This locality is the most important of several known from the environs of Enniscorthy. M’Coy (1846) listed many species from here, as did the G.S.I. memoir (Kinahan 1882 — locality 5). The locality was reviewed in detail by Brenchley ef al. (1977), but it was revisited in this study, new material being collected for the sake of completeness, and with the specific aim of comparing elements of the brachiopod fauna with other sampled localities. It was also hoped to collect topo- typic material of the poorly known agnostid trilobite Trino- dus agnostiformis M’Coy, the type specimen of which was redescribed by Whittington (1950: 533). The site location is shown in Fig. 11. The rock is a very fractured buff coloured mudstone, occasionally tuffaceous. New material was exca- vated from shallow depth in the old farmyard, between the 116 Ballyminaun Hill ae PO me 0) Kn Fig. 5 Locality map of the Ballykale locality. Key as in Fig. 2. lowest doorway of the ruined farmhouse and the new gateway to the northwest. Greenville and Moyne Upper Boundary, Enniscorthy, Co. Wexford (1:126720 - SHEET 23, $967420) This locality (Fig. 11) should not be confused with Greenville itself. G.S.I. locality 4 in Kinahan (1882) is Greenville and Moyne Upper Boundary, where a fauna was re-collected from many large blocks exposed as a result of very recent ploughing. These were immediately adjacent to the hedge bounding the original locality, an old quarry now slurry filled. No fossiliferous horizon was located within the exposure in the mainly volcanic quarry. Courtown localities, Co. Wexford (1:126720 — SHEET 19, 1187566 (Ballinatray) Numerous localities in the area of Courtown are listed in an earlier section, Table 1 (p.110). These were reviewed by Crimes & Crossley (1968), Brenchley & Treagus (1970) and Mitchell et al. (1972). Further examination of all these localities failed, with one exception, to yield anything new or significant; fossils found were poorly preserved gastropods, a few external moulds of Glyptorthis and crinoid ossicles from M. A. PARKES Gorcy Saw ~ the Courtown Formation. No new material was recovered from the Ballymoney Formation. The only exception was a collection made from calcareous slates in the Ballinatray Formation, about 160 m west of Ballinatray Bridge (Fig. 12). The present bridge is probably more recent than that named in the G.S.I. memoir (Hardman 1887), since the Courtown to Gorey road has been re-aligned since the 6” mapping. There is considerable confusion about the exact position of localities in this area, since both the 1” and 6” G.S.I. maps have a profusion of fossil locality asterisks, not all of which can be | related to the named localities in the G.S.I. memoir. How- ever, both from the described position and from the fauna present it appears that this might be locality 14 of Hardman (1887), where ‘J/laenus Bowmanni’, ‘Leptaena sericea’ and ‘Orthis calligramma’ were recorded in some abundance. Slieveroe, Rathdrum, Co. Wicklow (1:126720 — SHEET 16, 7211890) This highly fossiliferous locality was known to Griffith’s collectors (M’Coy 1846) and to the G.S.I. mapping team (Jukes & Du Noyer 1869). It was re-collected after excava- tion by Brenchley ef al. (1977) and a mixed brachiopod- trilobite fauna recorded from there. Although it was re-collected by me, the combination of very strong deforma- tion in the fragile slates and shortage of time meant that no | | BRACHIOPODS OF DUNCANNON GROUP OF SE IRELAND S lo fe. ‘S A | WZ, HF fossil lgcali CLOLOGE UPPER | Fig. 6 Locality map of the Clologe Upper locality. Key as in Fig. 2. | further work was done on the material. The previous faunal lists are used in discussion of correlations. Other localities In the course of tracing the localities from the memoirs, several other records of fossils were noted. On Sheet 158 (Kinahan 1882: 26) a locality called Kellystown Bridge was described a little south of Raheen, but no details were given in the palaeontological notes and it was not traced on the G.S.I. 6" fieldsheets nor in reconnaissance fieldwork. Kelly (1860) gave details of various localities which were not all included on the G.S.I. 6” sheets or in the memoirs, including ‘Ballyminaun Hill and Carriganeagh, both south of Gorey. ‘These do not now have any fossiliferous exposure. Fossils were recorded by Shannon (1979a) from Raheen and Bally- brennan, both G.S.I. localities, and also from Wilton Castle southwest of Enniscorthy, but the area is now heavily for- ested and no fossils were found in the restricted exposure available. All other sites examined as a result of personal ommunications, or unnamed sites indicated by asterisks on 1” and 6" geological maps, proved to be either unfossiliferous : not now exposed. 117 A \W oS O ALS PY gn METHODOLOGY AND TECHNIQUES Sampling methods In all sampled localities, an initial collection was made by identification of the fossiliferous lithology and on-site collec- tion of specimens. Subsequently, the main collection of specimens was achieved by removing large volumes (between 10 kg and 80 kg, occasionally more) of the fossiliferous lithology, to be broken up and examined in a laboratory. The validity of this method was verified by the fact that successive seasons’ collections served to increase the numbers of speci- mens, but not the diversity of the fauna. The one exception to this was the Kilbride locality near Tramore, Co. Waterford (p.112). This debris flow appears to have ‘sampled’ various benthic associations in its downslope movement, and subse- quent collections made here increased the diversity of the fauna significantly from the initial sampling. 118 fossil locality . Fig. 7 Locality map of the Carrigadaggan locality. Key as in Fig. 2. Preparation techniques Standard preparation methods were used. The measurements of all material (in millimetres) were made using vernier scale calipers or a micrometric graduated microscope eyepiece, both accurate to 0-1 mm. The combination of camera, lenses and extension rings (Table 5) gave a range of magnifications of up to X5. The specimens were whitened with ammonium chloride sublimate before being photographed. Taxonomy and statistical analysis In this study both multivariate and bivariate analyses have been utilized where the measurement data were adequate. Table 5 Magnifications obtained by different lens and extension ring combinations. AF Micro-Nikkor 55 mm f/28 x1 AF Micro-Nikkor 55 mm f/2-8, PK11A, PK12, PK13 < ee 123 > | > | > yd ey nic Formations of Shannon (1978). It is also equivalent to the Lower Tramore Volcanic Formation and all formations above (except the intrusive rhyolites in the Upper Tramore Volcanic Formation of Carlisle (1979)), the Duncannon, Arthurstown, Ballyhack and Campile formations of Gardiner (1974), and the Ballymoney and Gorey Rhyolite formations of Brenchley & Treagus (1970), as well as the Upper Caradoc Raheen Formation of Owen et al. (1986). Tietzsch-Tyler’s Carrighalia Formation (equivalent to the Ross and Loftusa- cre Formations) with its Tramore Limestone Member is also the equivalent of the Courtown and Ballinatray formations of Brenchley & Treagus (1970). AGE OF THE ASSEMBLAGES Kildare, Grange Hill, Horizons 1 and 2 Wright (1970) claimed that these were Longvillian in age, on the grounds that their composition is similar to assemblages from the Gelli-grin Formation of the Bala district of Wales, and in particular in relation to the presence of the calcareous inarticulate Orthisocrania divaricata, whose occurrence could only be substantiated in Longvillian rocks in Britain. Whilst there are some differences noted (Tables 8, 9, p.128) between the assemblage described herein and that listed by Wright (1970), in essence the fauna still shows a strong affinity with the Welsh Bala faunas described by Williams (1963). Many of the genera are long-ranging in the Caradoc, or even the Ordovician, and where samples are inadequate to determine the species, then the overall generic composition has been used. However, at Grange Hill both 124 Orthisocrania divaricata and Cremnorthis parva occur, and these are at present only known from Longvillian rocks in Britain. Leptestiina oepiki is also recorded first in Longvillian Welsh localities (Williams 1963). Although Cremnorthis (Hints 1968, Llandeilo) and Orthisocrania (see p.136) are known from older Baltic successions and may have migrated to Leinster at a time different from their Welsh Basin arrival, on balance, with the total assemblage affinities with the Gelli-grin Formation, a Longvillian age is most likely for this assemblage. Given the close relationship of Grange Hill Horizon 2 to Horizon 1 with a substantially similar but reduced diversity assemblage, a similar Longvillian age is suggested for this horizon also. The position of the various localities at Kildare in the stratigraphical succession of the inlier is schematically illustrated in Fig. 13. Kildare, Grange Hill House Cottage The only modern reference to the faunal assemblage from this locality is Williams ef al. (1972). A Soudleyan age is claimed for the fauna, presumably on the basis of compari- sons with the faunal list published by Reynolds & Gardiner (1896). The recollection of the fauna, as described herein, confirms that suggested age. The palaeoecology of this assemblage differs substantially from other Duncannon Group faunas described here (Table 10, p.129), but the brachiopod species show a strong similarity with a fauna from Herbertstown, in the northernmost part of the Leinster green sandstones SILURIAN | _red mudst. | black shales HIRN. mudstones Kildare Limestone Fm. GRANGE HILL HOUSE mudstones COTTAGE & sandstones eo) = Lu a) z < -! =) z oc > Zz < a _ ARENIG Fig. 13 Schematic section of the succession in the Kildare inlier. M. A. PARKES terrane, described by Harper et al. (1985). The combination of Plaesiomys multiplicata and Oanduporella cf. reticulata is unknown elsewhere in the Leinster terrane (except possibly at Clologe Upper). Indeed, the enteletoidean Oanduporella was unknown outside the Baltic before the Herbertstown record. The assemblage at Herbertstown was noted as being unlike any other fauna recorded in eastern Ireland (Harper et al. 1985: 289), but the new Kildare sample shows strong resemblances. On balance, the Herbertstown fauna was restricted to the Caradoc, with a most probable Soudleyan age. Similarly at Kildare, the fauna cannot be younger than Longvillian since it lies below the andesites of Grange Hill, which themselves are below a Longvillian assemblage. On the whole a Soudleyan age is most likely since the common brachiopod Plaesiomys multiplicata is found in the Soudleyan of Glyn Ceiriog of North Wales (Bancroft 1945) in associa- tion with Rafinesquina, which is present at Kildare too. The relationship of this horizon to others at Kildare is shown in leg, I}. Kilbride At this locality Carlisle (1979) listed a fauna with some forms congeneric with those in the fauna described here (Table 11, p.129). There are several differences, but the principal gen- era are all indicative of a Caradoc age. Carlisle (1979: 552) recorded Decordinaspis, a trinucleid trilobite previously only found in the Harnagian/Soudleyan of Grangegeeth (Harper & Romano 1967). Thus she suggested a Soudleyan age for the base of the Upper Tramore Volcanic Formation, in which Kilbride is located, with an unknown upper limit but possibly extending into the Upper Caradoc. This author did not identify Decordinaspis here, but, based on the co-occurrence of Cremnorthis parva, Leptestiina oepiki and Sowerbyella sericea, a Longvillian age is postulated for the assemblage. This accords well with Carlisle’s interpretation. If the Orthis- ocrania she recorded proves to be O. divaricata, this would further enhance the reliability of the age assigned, since the locality was said (Carlisle 1979: 551) to be 200 m above the base of the formation. Ballykale This assemblage is relatively low in both specimen numbers and diversity, being dominated by Bimuria cf. dyfiensis (81%, see Table 12, p.130). This species has only been described previously from the Gelli-grin Formation of north Wales (Lockley 1980:215), which is of Longvillian age. In the other localities dealt with here, Bimuria sp. definitely occurs only at Kilbride, in a form probably conspecific or at least very close to B. cf. dyfiensis. This locality too is of probable Longvillian age and this seems the most reasonable estimate for the age of the Ballykale assemblage also. The few other brachiopods are all congeneric with Kilbride forms, notably the rare Ptychoglyptus. Clologe Upper Since the poor preservation of fossils from here permitted only a generic level identification at best, the precision of correlation can only be crude. Overall, the generic composi- tion indicates a Caradoc age and some strong similarities with the other assemblages described herein. However, the pos- sible occurrence of both Oanduporella and Plaesiomys, both == ag — axe . BRACHIOPODS OF DUNCANNON GROUP OF SE IRELAND reminiscent of the Soudleyan fauna at Grange Hill House Cottage, Kildare and the Soudleyan Herbertstown assem- blage (Harper et al. 1985), together with the apparent | absence of Cremnorthis and Orthisocrania, suggest that the | assemblage is older than Longvillian. Therefore, a probable Soudleyan, or perhaps Harnagian, age is postulated. Carrigadaggan | The brachiopod fauna contains numerous elements which _ allow confident assignment of the assemblage to the Longvil- lian, and enhance the strong inter-locality correlation. Lep- testiina oepiki, Cremnorthis parva, Orthisocrania divaricata, | Sowerbyella sericea, Skenidioides costatus and Kullervo aff. hibernica are all found in the Longvillian of Grange Hill, Kildare, as are many of the genera such as Nicolella, Platystrophia and ‘Orthambonites’. The species listed above, except Orthisocrania and Kullervo, are here recorded at Kilbride also (Table 14, p.131); Carlisle (1979) also listed _ Orthisocrania from there. The presence of Saukrodictya sp. is noted here, which although rare, also occurs at Kilbride, | Ballykale and Greenville-Moyne, all of which are probably of Longvillian age. The cystoid Echinosphaerites ct. granulatus is most common here, but was also found at Kilbride and | Ballygarvan Bridge. Although not stratigraphically useful, its occurrence in these three geographically close localities is | supportive of the brachiopod correlation. _Ballygarvan Bridge The sparse brachiopods from this locality allow little certainty _about correlation of the assemblage, neither does the restricted trilobite material. The generic composition and abundance of gastropods (Table 15, p.131) is notably similar to that of Grange Hill (Table 8, p.128) and Grange Hill House Cottage (Table 9, p.128) at Kildare. This may be | merely ecological control. However, as the large inarticulate Lingulella ovata occurs elsewhere in Leinster only in the | Longvillian of Grange Hill, Kildare, a possible Longvillian age is suggested. In spite of the low numbers of specimens, the congeneric forms enhance the consistency of inter-locality correlation in the Duncannon Group. | ‘F rankfort and Clogh The examination of these localities was unproductive for new _assemblages. However, the collections made by N. Hiller (for ‘a B.Sc. thesis, 1971) and subsequent publication (Mitchell er jal. 1972) correlating the Courtown area with the Tramore region, and now housed in the Ulster Museum in Belfast, | were examined for comparison with other collections made in the present study. The list of genera published by Mitchell et al. (1972) is quite long but does not reflect the proportions within the assemblage. The strong deformation and poor preservation in the tuffaceous rocks, collected from walls in \the area of the G.S.I. locality, made unequivocal identifica- |tions difficult. Many genera are represented by one question- able specimen only. The abundant genera are Cremnorthis, ‘Orthambonites’, Leptestiina and dalmanelloids. Less com- ‘mon but positively present are Kullervo, Platystrophia, Anisopleurella, Skenidioides, Nicolella and Orthisocrania. The apparent absence of some of the other listed genera could be due to incomplete donation of the collection, rather than absence from the assemblage, since not all specimens 125 were clearly labelled. The particular elements not seen in the material at the Ulster Museum were Diambonia, Ptychoglyp- tus, Eoplectodonta, Pseudolingula and Actinomena? The Christiania recorded is a somewhat equivocal single specimen which is highly strained. Thus, if the definitely present and abundant elements alone are considered, the assemblage clearly takes on a strong resemblance to the described assemblages from Grange Hill, Carrigadaggan, Greenville-Moyne and other localities in Wexford. The re-examination of this assemblage, in conjunc- tion with a revision of the Duncannon Group localities throughout Wexford and elsewhere in the Leinster terrane, reinforces the Upper Soudleyan — Longvillian age assigned to the Ballymoney Formation by Carlisle (1979: 552; fig. 3). On the basis of the existing brachiopod collections this will not be refined further, but new material or the current examination of the trilobite collections in the G.S.I. may yield better information. Raheen The assemblage described here (Table 16, p.131) is clearly inadequate for a confident correlation and is probably facies controlled, rather than of different age from other localities in the Duncannon Group. When the trilobite identification is complete a better constraint on the age might be made, but in the absence of better material the G.S.I. lists (Kinahan 1882) give a good indication of the age of adjacent beds in the succession. There are no species known to be restricted temporally, but typical Longvillian genera are absent. Some elements are reminiscent both of Greenville and the Brick- works Quarry Shale Formation of Grangegeeth (Romano 1980a), which are believed to be Harnagian or Soudleyan. Therefore, an early Caradoc age, possibly Harnagian or Soudleyan, is thought to be the best estimate. Greenville The collections made here (Table 17, p.132) are somewhat reduced by comparison with those of Brenchley ef al. (1977). These authors suggested a Harnagian age, and no significant evidence has been found to dispute that in the assemblage described here. Only the probable occurrence of Oandupor- ella, present in the Soudleyan of Herbertstown (Harper et al. 1985) and in the Soudleyan assemblage at Grange Hill House Cottage, Kildare, suggests a possible Soudleyan age. Brench- ley et al. (1977) attached some significance to the presence of ‘an early species of Sericoidea together with Anisopleurella aff. multiseptata Williams’. As discussed in the systematic description of Chonetoidea (the senior synonym of Seri- coidea), the variation amongst the population is considerable and assignment to C. aff. abdita Williams would now only suggest a Lower Caradoc age. Anisopleurella multiseptata, described from the Costonian Derfel Limestone Formation (Whittington & Williams 1955), is known also from the Longvillian Gelli-grin Formation of Bala, North Wales (Lockley 1980). Greenville-Moyne Several elements, such as Cremnorthis parva, Leptestiina oepiki and Skenidioides costatus, are conspecific with samples from Longvillian assemblages at Kilbride, Carrigadaggan and Grange Hill Horizon 1 at Kildare. Other forms only identifi- “6L61 AIS teD ‘Apnqs SITY} :WOWPIT /IPUg|Iy “6461 IIS4BD :‘uouUPIUNG “6861 JayA |. -YOSZ1 91, “Apnis SIU] :BI1VW SSO MON/URSSepesIeS “Apnis SIy) “O61 snseoly, mY Aayyousig ‘AYWOISIUUT/I|IAUDAIH 06] SNBvo1L 2? A2|Yousig “6L6] ISWIVD :uMoNOD “ApMys styi *Z7L6T 7Y 7 SWNT SPP “6L6T 7? J2 YOM :MOPPIM AS/wmupyey “L867 Aydin :uessiuqieg 196] Aydin ‘ZL6] (/Y Ja SWeI[IAA SUIPNoUT snoueA :auRIIOg/ARQUIL'T :S2d4NOG ‘pUR]>I] AS JO SULIID} 19}SUIIT] YI UIYIM SatTeoo] Pa}da]as 1OJ WeYO UONR]as109 JeoydessHens py “sty uolyeuwo04 eyeus asowed) uoNeuuio4 9u0}Sewr] esowed) “WW DIUBDIOA, esowel seddy uO lyeUO4 Keg Aojoog dnoiy pueqaqiy uo !yPUO4 umojseuuy dnoi5 pueqaqiy uolyeU0 4 8U0}]SELIT uMmoyNnoD UOI}EULIO4 uOolWeUO4 Aesyeuyeg uMO}SeuuYy uO }}eUIO4 AeuowA\yeg UONEULIOF Ayy Aeio5 dnoiy pueqaiy uo |VeUuLIO4 UMO}SUR|UOD uolyeuWO 4 aIUeDIO/A MO)MOIM ISPS UOIVEUO4 (S8}1Sepue ) uO!}eLUIO4 dweopg uo lyeULIO4 @U0}SeWI] 212 PII ~~ seysepuy Aeque eouenbes OIUBD|O/A AUBILOd sajeus 4oeIgé SNPHT, juosjyounut snjnosnjese} poe suepnyjnu juebuyjo syeaul] snjeuejduoo SN|eUIPsOB XE SINSYV ueIPURNIUM NYIANVT1 ueluo}soD ueibuewey ueAg|pnos uei||!ABu07 URIUO SOOM. ueIyOOsqusueyy ueluo}oy ueAayneog ueAeuymey uenuewiy, oTnaqnvT NVIDIAOGHO o90d0vVYVo THSHSV [__ seuojspny _| 7 snjdjnosied SYONVYL /20Ws VWauv SSOH MSN AHLYOOSINNA NVDDVaVOYVS /STIHANASYS MO1XODIM 3S /WONYOHLVY ‘S| AVEWYV1 /ANVYLEOd NONNVONNG NMOLYENOD SYvalIm NVSSWe Iva ANOZ 3LNOLdVYS BRACHIOPODS OF DUNCANNON GROUP OF SE IRELAND ' able generically or questionably assigned, e.g. Nicolella sp., _‘Orthambonites’ sp., Orthisocrania?, Rhactorthis? and Sower- | byella?, are all reminiscent of those assemblages, as is the ' rare occurrence of Saukrodictya sp. Kullervo aff. hibernica is ‘also present at Greenville-Moyne, Carrigadaggan and Grange Hill Horizon 1, all localities thought to be Longvillian in age, but it also occurs in the Knockerk House Sandstone Member (Romano 1980a) of the Grangegeeth Group (equivalent to the Upper Tuffs and Shales of Harper, 1952), _which is believed to be Costonian in age (Brenchley et al. 1977). However, the consistency of the inter-locality correla- | tions suggest that a Longvillian age is most probable for the _Greenville-Moyne assemblage. Ballinatray Although the assemblage from here is too deformed and broken to identify the brachiopods precisely, the age of the horizon is constrained by previous work. The age of the older Courtown Limestone Formation is believed to be mainly Llandeilo ranging up to the Costonian Stage of the Caradoc on the correlation with the Tramore Limestone Formation (Carlisle 1979). The succeeding Ballymoney Formation is believed to be of probable Longvillian age, and in any case gracilis Biozone graptolites have been recovered from the Ballinatray Formation (Mitchell et a/. 1972). An early Cara- ‘doc, at least partly Costonian age is suggested for the \formation. In the light of the age constraints clarified by a revision of these Duncannon Group faunas a revised correlation chart has been produced (Fig. 14), with sections additional to previously published correlations e.g. Williams er al. (1972) and Briick et al. (1979). PALAEOECOLOGY AND BIOGEOGRAPHY | | | | | | In some of the localities very little ecological data can be derived because the exposures were small, resricted and isolated from a clearly defined succession, or assemblages were not from in situ rocks. Equally, small assemblages like those from Ballygarvan Bridge (Table 15) are inadequate for detailed interpretation. Even with large assemblages such as Grange Hill Horizon 1 (N=646) some reservations are held about interpreting too much from them. | One evident point from this work is that assemblages often differ from previously recorded collections, especially those of M’Coy (1846) and the G.S.I. Memoirs. Many differences are simply taxonomic artefacts reflecting better systematic differentiation of this important phylum. Where the outstand- ing differences are not addressed directly below, the best explanation is that in the volcano-sedimentary setting of these ocalities the character of faunas changed as a result of tuff falls etc. and slightly different horizons have been sampled. This would apply to Ballygarvan Bridge, Raheen, Greenville and possibly Greenville-Moyne for example. For each locality a table of the total collected fauna is resented. Table 7, however, is a summary chart of presence/ absence data for each locality, highlighting the inter-locality affinities. The tables (8-19) include total counts of the brachiopod valves and various animals for the comparison of assemblages with recognised palaeocommunities discussed pelow. Following Lockley (1980: 171) this number has been 127 Table 7 Summary chart of fauna collected at each locality. Localties: A — Kildare, Grange Hill Horizon 1. B — Kildare, Grange Hill Horizon 2. C — Kildare, Grange Hill House Cottage. D — Kilbride. E — Ballykale. F— Clologe Upper. G - Carrigadaggan. H — Ballygarvan Bridge. I— Raheen. J - Greenville. K — Greenville-Moyne. L — Ballinatray. MJ} CJD I FOG isi tl Y IK Ie BRACHIOPODS Acanthocrania? OS tS 70] 5S 6 ee Anisopleurella cf. multi- septata = Cy e e - Bicuspina? Bimuria cf. dyfiensis ~ e = Bimuria sp. Chonetoidea abdita Cremnorthis parva dalmanellids indet. Dolerothis? - - Glyptorthis a Hedstroemina sp. Ce SE nS ee Hesperorthis sp. ere es lee eee Oe sy Hibernodonta? Sh Oe i te ee fae — Kjaerina? e = Kjerulfina? e - Kullervo aff. hibernica == = = Ser = Se = Leptaena sp. e Leptellina cf. llandeiloensis — Leptestiina oepiki e--eee Leptestiina oepikiampla — Lingulella ovata ee - Nicolella cf. actoniae e Oanduporella cf. reticulata — ‘Orthambonites’ sp. e Orthisocrania divaricata @ Oxoplecia? Petrocrania harperi Philhedra sp. Plaesiomys multiplicata —- -— @ — — Platystrophia sp. O46 =. @ Te = geneh= = ee Porambonites sp. - - e Ptychoglyptus sp. =—— -—_— 10) (0) =) Se Rafinesquina sp. - - @®@------ - Rhactorthis sp. e- - Salopia sp. - @ - Saukrodictya cf. sp. A -- - Schizotreta sp. - Skenidioides costatus e Sowerbyella sericea e Strophomena? e TRILOBITES Ampyx austinii - - @®----- @®e- - Arthrorhachis e Atractopyge =SSseae es OS sas = Autoloxolichas e - - Calyptaulax e----- @---- = Deacybele - e Encrinuroides - = - = = = = = = Flexicalymene e e Homalopteon sp. - = Remopleurides -------- - Tretaspis ----- @-- @®- - - ‘Trinodus agnostiformis’ — — — — e Xylabion e - - - - = OTHER ELEMENTS Echinosphaerites granulatus cheirocrinid cystoid crinoid ossicles gastropods ramose bryozoans prasoporid bryozoans tentaculitids orthocones bivalves conulariids ostracods eee! il | leoe0ee | e | eee!!! eeoec0c!| e e eeo0e eee! ee! | | | @eee7eee | | e @eeo0ee @ e@eeee! | | ee! | ee! e | | e | eeeeeoeeee i! e | | | | | eee! | | ee! | | 128 calculated using the formula: N=A + 0-51 + P (if P>B), or N = A + 0-SI + B (if B>P) where: A = no. of articulated valves, B = no. of brachial valves, P = no. of pedicle valves, I = no. of indeterminate valves. The problems of counting different groups of animals was discussed by Lockley (1980: 171-2) for Welsh Caradoc faunal associations and by Jaannusson (1984) for Baltoscandian Ordovician sequences. In the case of groups other than brachiopods absolute numbers are given. However, for some organisms such as fragmentary bryozoans, cystoid plates and crinoid ossicles, absolute numbers are given in brackets and an equivalent number of individuals given, generally one. An arbitrary correction factor is applied to bivalves for Kildare Grange Hill House Cottage, by halving the total, on the assumption that the valves are simply disarticulated in equal proportions. This is also applied to ostracods. For trilobites the largest value of pygidia or cephala is arbitrarily taken as Table 8 Total assemblage collected from Kildare, Grange Hill Horizon 1. Internal External Total Pw Bw © i PY C i wo, YW BRACHIOPODS Acanthocrania? | tee = 1 Sa It @eils) Chon. toidea sp. 1 <= 18=9.]— le si 2035 Cremnorthis parva St WH 4 = 1OAl SF = 5 37 dalmanellids, indet. 23) = 3S 8) 2 hy ANP) Hedstroemina sp. SSS eS 3 = 3 OS Kjerulfina? PSS Sa = = = i Pil Kullervo aff. hibernica 28-<=8=8 1 = =PS9 2203 Leptaena sp. Goi oa 35 =] 6 i tks Leptestiina oepiki 7 === = = isa elO Lingulella ovata - -=-- =-=-- 3 3 055 Nicolella cf. actoniae = 1 = SST = S05 ‘Orthambonites’ sp. 256 =8—) 20 = = = aorn0:9 Orthisocrania divaricata 38 23 —- 68 -— - - 108 92 14-2 Petrocrania harperi = 89 = ==—30 = = 89) 13:8 Philhedra sp. =8§=8—8=) — Ieee eles Platystrophia sp. Sl EG = We iyi 1 Sy Ss Rhactorthis sp. 1 <3 ty 1 27 SSO: 8 Skenidioides costatus Wo 32 i = Wilh 3 = Ils ils Sowerbyella sericea A Wy tl = IPI it @ 38 Sell Strophomena? 28=-s = 1S = = 2803 TRILOBITES ANH MUOTOUWGITS CHONG soccoscaenncadqnnnsne: oonnandABSnDS0on0006 3) eS) Cally ares? OSGI cooecoasansaqncotnoocosbon0onDooang0500090000 iL sis) Hlexicaly mene cephalonwerrce--eeerer eee eeeeeeee errr crete ee raee i pil OTHER ELEMENTS ClatsXorralvoxsyq io tox (7) gaepaeatoondpabEdnacaocotsdagedeenoswapedoqcebe iL (Opil5) PASOPOUS ee coer mrceckiecses. dee iaeeeemeeasccoeesaciewecsineet S30) Fed TAMOSE DLV OZOANS meme eree eeeece eee eee eee eeeeeac eer ere 4 0-6 prasoponidibiyOZOanSmen settee eee eee eee ee eee eee eee eee ile tentaculitids: .eecmaceesmesserecac een chisa tclarcosecenelekee sites 29 =4-5 OnthOCONES resesice chee socGeedeereh se eeeee alee meee ener 4 0-6 BIVAlVES: Aiciccsecgemetee de egetcinevseueseneerntecsee soak seeliemtaeee act 2:3) COND ALIGS’. «ca. estan gee neh acteesigs soeeme eee aeeepecseesseceaeses i pil SPONGESPICUIE He ctewesceee am atradeossemeresee ts -eecles ceeds. il. OpitS) Total 646 99-85 PV = pedicle valves, BV = brachial valves, C = conjoined valves, I = indeterminate valves, NO. = equivalent number of total animals. (N.B. Numbers in brackets are total fragments of colonial or fragmented individuals — see discussion). See discussion on p.127. M. A. PARKES the commonest animals, errors involved in counting other — animals are considered as negligible. The diverse assemblage from Kildare Grange Hill Horizon 1 is listed in Table 8. All the material consists of moulds from | decalcified mudstone/siltstone. The ratio of ventral to dorsal | valves is generally equal. Differences may be accounted for — by the problems of identifying fragments produced by the | sampling method. There is a large range of size variation | between and within species, and there is no suggestion of current winnowing. The shells are disseminated through the sediment, not collected together as in a coquina. For these reasons the assemblage is treated as a palaeocommunity. Grange Hill Horizon 2 (Table 9) is very similar but has reduced frequency of fossils and the addition of common — Lingulella ovata. The difference is partly because the rock is not decalcified. This biased the identifications in favour of larger distinctive species such as L. ovata, O. divaricata and Platystrophia. Another factor is that at least one thin tuff horizon is found between the two horizons, which may have subtly changed the assemblage. Some differences between Table 8 and the faunal list of | Wright (1970) should be noted. Cyclospira and Bicuspina were not collected by me, whilst taxa not recorded by Wright | but collected by me include Acanthocrania?, Chonetoidea, Hedstroemina, Kjerulfina, Kullervo aff. hibernica, Leptestiina | oepiki, Lingulella ovata, Nicolella ct. actoniae, Petrocrania | } | q | | | the number of individuals. Since brachiopods are generally | | harperi and Philhedra. All these were present as 1% or less of the assemblage, except for Petrocrania which alone consti- tuted 13-8% of the assemblage. Wright (1970) also recorded Eoplectodonta, but this genus is essentially indistinguishable | from Sowerbyella except for the presence of denticles along | the hinge line. No specimens in this study showed evidence of | such denticles. The older fauna from Grange Hill House Cottage (Table 10) is somewhat anomalous in comparison with the other Table 9 Total assemblage collected from Kildare, Grange Hill Horizon 2. ee Internal External Total VIB (CU WAY EAC It no. % BRACHIOPODS dalmanellids, indet. 2n3x=)= os thin meee Sail Hedstroemina sp. 7 --- Li==.5, 1 11-9 Leptaena? Sa Le 3-3 Lingulella ovata - -1- ---16 9 153 Orthisocrania divaricata erage ey a) 8:5 Petrocrania harperi 2 4 - 4 6-8 Platystrophia sp. S)ieov=y Sh2e= silgas B)| Salopia? Spliey a—) 1 17 Skenidioides costatus 3 --- --- - 3 Sal Sowerbyella sericea 2 - - 2 333 OTHER ELEMENTS @) EU NVOYHO) 1S Gagan goadcnan ance sodecadoogase sueagaRaDoIadacccooGaaZb0ue 3 5-1 [DHAO) KOVAC! lOVAYOVAOHIN scceosndancsoepsoobe ssbocodenapsecopeesconan 3) Sal DivalVe: sce dioscenee cht. teats seat ea- te netbe Steels ances epee CER 1 1-7 (ACHR) SXOTEES, densesuedassondsne sasaccbrcsoee cose dgubbonecBosoncSsBCHsc5 13 22:0 Total 59 100-0 SEE PV = pedicle valves, BV = brachial valves, C = conjoined valves, I = indeterminate valves, NO. = equivalent number of total animals. (N.B. Numbers in brackets are total fragments of colonial or fragmented individuals — see discussion). See discussion on p.127. / BRACHIOPODS OF DUNCANNON GROUP OF SE IRELAND Table 10 Total assemblage collected from Kildare, Grange Hill House Cottage. Internal External Total PVBVC I PVBVC I no. % ~BRACHIOPODS dalmanellids, indet. = ate SW IS 2 0-4 Dolerorthis? SS SS 1 0-2 _ Hibernodonta? = 3 == 2 = 3 4 0-8 _ Leangella? 1 = 1 0-2 Leptaena sp. fe= = Sse s 1 0-2 lingulid, indet. ==S=s5 =+=+-+4 4 0-8 Oanduporella cf. reticulata 13 10 — 9 6-- 13 2-5 | Petrocrania? el 2 0-4 | Plaesiomys multiplicata 2613 —- — 12 20 — - 26 5-0 Rafinesquina sp. § V= 2 10 Ss = 4 12 2-3 Skenidioides sp. 3. = 5) Sages 3 0-6 TRILOBITES _ Flexicalymene QDs [OWGRGTIAY Gobasedesaneseconseubeannce neepeas000 2 0-4 | Flexicalymene sp. cephala .............cssserceceesenseneeees 4 0-8 PEO COUGHOSIGEpNAlal .2...o.. sacle sae sire sscijeoane se series i 3 0-6 EIPIGSP: PYSIGIUM 22.222. c. 2... see eeceeeeeenecceereecmenees 1 0-2 OTHER ELEMENTS PRIS NSPECIES | (106) veces cme cwwemce ew eneween seca aammseaneennns 5310-1 MMDUMEDNNSIIC CICS a 2-c 08. 2225. Secwideaniulesesseesunenaeeveneeunes 246 47-2 Wastracods (Tetradella?) (132) .........ccccsveesccsseseeeeeceee 66) 12-7 PRETONCROSSIGIES ICL) orcrscicca can wncwasienenaaeeerescenewermen dormeets 1 0-2 immocone fragments (15)... .....eeseseee cece eene reese ene 15 2-9 MPRMETe TOS erernicen es Arcee rattaevereecretras esenancccaasoneste i @2 | SEONG ~ peécecdnn 008885 SAH CBaSE EBS ABBE ABRe es eae niece neecon couse 45 8-6 - SORRGT'Y oSdeSeOos doe sbBospbacbhenee oABEL Sapp badge Haare udoesee 15 BSS) Total 521- 100-2 PY = pedicle valves, BV = brachial valves, C = conjoined valves, I = indeterminate valves, NO. = equivalent number of total animals. (N.B. Numbers in brackets are total fragments of colonial or fragmented individuals — see discussion). See discussion on p.127. Leinster faunas. It is molluscan not brachiopod dominated, with different brachiopod genera present. Five of the 11 genera are not known from the other localities and make up 10-8% of the fauna, of which only 13-4% comprises brachio- pods. The fossils were excavated from muddy and tuffaceous siltstones and fine sandstones at the back of the cottage. |About 2-3 metres below the fossiliferous section are fine /quartz pebble conglomerates with fragments of the underly- ing slates. _ The fossils are interpreted as having lived in a shallow water but low energy environment, perhaps close to the shore of the emergent volcanic cone of Grange Hill. Supporting evidence includes the very close proximity of lavas inter- preted as being subaerial, and the abundance of an ostracod | (Tetradella?) which elsewhere is commonly indicative of ‘intertidal or shallow subtidal conditions. | Kilbride, within the Upper Tramore Volcanic Formation, iby contrast, represents a different setting. In the small exposure there are contorted streaks and lenses of tuff in the \dark siltstones, with very obscure bedding. The faunal evi- \dence from this work and the limited field evidence supports \Carlisle’s (1979: 551) interpretation of slumpedvolcaniclas- ‘tics, and that the lithology is a debris flow. The high diversity jand low frequency of any one species suggests that faunas from different areas and substrates were ‘sampled’ in the downslope movement of the sediment mass. Although Carl- isle (1979: 552) only listed certain genera as common and 129 others as ‘also in the fauna’, the assemblage listed in Table 11 shows significant differences, reflecting a different sample. In the fauna are numerous (22) fragmentary plates of a cheiro- crinid cystoid, of which 5 may be oral plates. These may belong (C.R.C. Paul, personal communication 1987) to Hadrocystis or Acantholepis (whose sole species A. jamesi M’Coy, 1846 was described from Raheen, Waterford Har- bour). The preservation of the cheirocrinids (Plate 16, figs 12-18), which are fractured across plates and not along boundaries, also reflects the disturbed nature of this horizon. There are also 9 plates of Echinosphaerites, some of which are broken. Comparable examples of debris flows include that described by Lockley (1984) from Builth, central Wales and Kilbucho in Scotland (Clarkson et al. 1992). Although it is reasonable to treat them as ‘in situ’ faunas Table 11 Total assemblage collected from Kilbride. Internal External Total PV BVC I PV BVC I no. % BRACHIOPODS Acanthocrania? 1 1 0-6 Anisopleurella? =- 1 -- = 1 1 0-6 Bicuspina? 2 1 2 1-1 Bimuria sp. 27 - -- 1--- Piss () Chonetoidea cf. abdita Id! @ = = Y 2 - 14 78 Cremnorthis parva OP 7 tS Ss Hee = 8 4-5 dalmanellids, indet. ee eee ee Dee Me 3} 1-7 Hesperorthis sp. Se t= eda 3 17 Leptestiina oepiki 6 -- 12-- 6 3)93) lingulid, indet. - 1 1 0-6 Nicolella cf. actoniae WM ==> § © == 10 5-6 ‘Orthambonites’ sp. Sal ee eo 14 qi Platystrophia sp. 2 ft gJoee= i lee 3 1-7 Porambonites sp. 2--- =-=-=-+ 2 1-1 Salopia? 1 1 0-6 Saukrodictya sp. i--- 1--- 1 0-6 Schizotreta sp. 1 1 - 1 0-6 Skenidioides costatus Ae Age ew 7 3-9) Sowerbyella sericea 18 7. — — 138 10,— — 18 10-0 sowerbyellid, indet. 2 - 2 1-1 TRILOBITES ey belinidicephal ong ence-cereese reenter ssasaeeer resect ners 1 0-6 Ne) nv ali(erejo) 1A oN" Sec eaeedagocermades dean sdosondadaceonansesopecoonodss 1 0-6 VAT NTONACHISI avsce ecto c seas tb = aot ee PER e eee eee eee eee 1 0-6 cybelinid librigenae (5) Mi eerviol? 2/9) NANO . soenadadecodeedacdons sobéooanoosaocdobodnandecas 1 0-6 2GHAHOEY |DYAENUIN cooacoogscocnnooadnuacooghoscsseecoeCooasaculs 1 0-6 OTHER ELEMENTS SONS arcoscedadaeadeddéssandeSoadsansdeonadoédasoosboscbDbDoddadoEee 1 0-6 PERSIA) OGL, _-sausseecooencdonesacseobhnoadedoooouddecobanpoapuaeopccace 6 3h) GSI Tcf0\c Soros ance penncecunBaster caneon be sessdecansuasbnonenaccodeotcigc 3) 1-7 (NTI OLKONEY "ss sceeodonacadonneadseanatndsos oc eoosdescsosadaconc0eOsD0c00 1 0-6 ConulantidS iaedas eat tasteep eee tapes oaacasd obegieetowower tines aoe 2 1-1 PLasOporidibryOzZOaNS e.eeeeeeteresseetesese erreeeater ere eccee 16 8-9 TOSS LOA OZORINS (27) aeaceanccpscosenccadcdorannodsorcsedosbtionse 1 0-6 COVA reece once mer iscece cece aniuectee cescsleyscmtictae tc sce slieeastiress 1 0-6 CLINOLAVOSSICIESI(T7)T oecaccc sane tc serce tone tecce settee matress 1 0-6 Cheinocninidlicystoid) erases. -cereasseeeeeceeretensce eer tteer iN 9-4 cheirocrinid cystoid oral plates? (5) JEG UI AON NUHAAHES [OENIES (6) copspdeeassococrenoononnboaecrogs.0ceC 1 0-6 Total 179 100-8 PV = pedicle valves, BV = brachial valves, C = conjoined valves, I = indeterminate valves, NO. = equivalent number of total animals. (N.B. Numbers in brackets are total fragments of colonial or fragmented individuals — see discussion). See discussion on p.127. 130 for stratigraphical correlation, the loose block nature of assemblages from Ballykale (Table 12) and Clologe (Table 13) offers little information, although the fauna from Clologe is from two distinct lithologies, one shaly and one tuffaceous. Hiller’s (1971) collections from Frankfort and Clogh also came from loose blocks. Carrigadaggan, although yielding a diverse large assemblage (Table 14), gave little information about palaeoecology, since exposure was poor and much material came from broken blocks not completely in situ. The coarse volcaniclastic lithology preserved details of the fossils only poorly. The presence of Echinosphaerites cf. granulatus M’Coy (Plate 16. figs 1-7,9-10) in some abundance at Carrigadag- gan, is worthy of note. It was listed as E. aurantium in the G.S.I. Memoir (Kinahan 1879). This species is found as almost complete individuals (6 specimens), suggesting rapid burial as in a coarse tuff fall, but there are also numerous isolated (15) or several associated plates (27) of the cystoid. Forbes (1848) monographed the British and Irish cystoid fauna, and Paul (1973, 1984) has produced a modern revision of some diploporite and rhombiferan cystoids. This material, which has been confirmed as Echinosphaerites (personal communication 1987) will be dealt with in the third part of Paul’s monograph. Bockelie (1981) has recently reviewed the functional morphology and evolution of the genus, from many different lithologies in Russia and Scandinavia. In spite of the fact that there are several nearly complete thecae, the fact that few if any preserve evidence of a stem, cover plates of the gonopore, brachioles or oral cover plates, suggests that they were not covered by sediment immediately after death. It is difficult to be unequivocal about these taphonomic factors, or the loss of periproctal plates (which occurred in less than 5% of specimens reviewed by Bockelie, 1981: 191), since the Carrigadaggan lithology is so coarse-grained that fine detail of the plates is not always preserved. Two speci- mens show projections, however, one of which is believed to be an oral projection, the other possibly the basal plates to which the stolon was attached (see Plate 16, figs 7, 9, 10). Possibly the most likely explanation is that Echinosphaerites and the cheirocrinid cystoids, of which there are 9 fragmen- tary plates similar to the Kilbride material (Plate 16, figs 8,11), were buried rapidly by volcaniclastic material but were subject to current action in shallow waters rather than inundated by distal tuff falls. Ballygarvan Bridge (Table 15) has volcanic rocks adjacent, but exposure is too poor and the sample is too limited to infer much. As previously noted, volcanic events may best account for the differences between the new collections from Raheen and the G.S.I. records (Baily in Kinahan 1882: 38-39). A diverse fauna of 25 species including 9 trilobites in abundance was recorded, whilst new collections (Table 16) are over- whelmingly dominated by a dalmanellid (probably Howel- lites, but remaining indeterminate because of the strong deformation) with only a few trilobites and generally low diversity. Tuff horizons (tens of cm thick) are common in the new section that was exposed and almost certainly account for apparent differences. The Greenville assemblage (Table 17) also differs from earlier collections. Since this was collected by digging beneath a thick farmyard deposit it is most likely that different horizons were sampled, but the cause of the changes in assemblages from horizons in close succession remains unknown. In contrast, the collections from Greenville-Moyne (Table 18) probably came from the same horizon as that M. A. PARKES | Table 12 Total assemblage collected from Ballykale. | Internal External mE: | PVABV CI SENS BVAG= aI % Bimuria cf. dyfiensis 5910 —-—- 16 5 = = 59 81-08 Chonetoidea 1 - - 1 1-3) dalmanellids, indet. 2) SS oa a eee | Leptestiina sp. D, - 2 pe Platystrophia - 1-- --- 1 2. 2a Ptychoglyptus i= Ht. = 2a | Saukrodictya sp. 1 = 1 1-3 Sowerbyella? 2D - 2 2-7} strophomenid, indet. 1 1 1-3 Total 73, , 90ne | OTHER ELEMENTS { ramose bryozoans (5) trilobite genal spines? (3) trilobite thoracic segments (2) PV = pedicle valves, BV = brachial valves, C = conjoined valves, | = indeterminate valves, NO. = equivalent number of total animals. (N.B. Numbers in brackets are total fragments of colonial or fragmented individuals — see discussion). See discussion on p.127. Table 13 Total assemblage collected from Clologe Upper. | Internal External Total PVBVEG a RVeB aCe no. % BRACHIOPODS Anisopleurella? 1 1 - 1 1-4 ) dalmanellids, indet. 24 Ne= 16. Sea 7 10-0 Dolerorthis? toi == 1 - -- i 1-4 Glyptorthis - --- ---1 1 1-4 Kjaerina? - --1 - --1 1 1-4 Leptestiina sp. 12.1- ---=+=- 3 4:3 | Nicolella sp. DOs NS = 2 2:9) Oanduporella? 2: 2 2-9 Plaesiomys? 1 1 1-4 | Ptychoglyptus sp. 1 1 1-4 Salopia? ---- -1-- i 1-4 Skenidioides sp. 1 --- 1 - - 1 1-4 Sowerbyella sp. 2 ? pes) | strophomenid, indet. 1 2 1 1-4 TRILOBITES tretaspid:cephiala: or.. s.as.ncetepeeber ees cesneacere -aereeee eee . 18 255i] tretaspid cephalon fragments (14) cybelinid librigenae (2) AGMOSTIG Mee hata eee See ce 1 1-4 lichid cephalonqcsiinc.cosguaedh seteasceseee ceases eee eee: 1 1-4] cheinunidicephalont a. ..-ee-seeeee-eeecees cass ease eee ee eee es 1 1-4 OTHER ELEMENTS | mamose bryozoansy(4)) fice. aceececsesso cece. ose se eee Reece 1 1-4 prasopontdibnyOzoans pee seeeeeet creer seeteeee reese seeeeeeeraee 1 1-4 OMMNOCOME tee eaes sean secs cee peat ee 7 1-8 Deacy beleicephalonwressserceseeeeee seca eee eee eee 1 1-35 ee 7 SP ; i - - 24 = 11 2:8 OTHER ELEMENTS ) Ort Pet waricata ae 4 = ; me DANO POM! TAWOZOBS — coccnssprocensuonopaoanuosccasseooessancce 20 27-0 Oxoplecia? | rs aie z Lsn Ors PATNOSS DINICZOLIS (BD) novcassocooeeseseacvocoovbnpassaecooossese 1 1-35 Platystrophia sp. See Wl 11- 9 D3 gastropods 37 50:0 Be eeriacdanjindet, = = == 2.- - 5 0:5 Eason vag Clee Rea din cing aniae's age seem dauasioete eh ees musta mee Thisatsipee | Salopia? a x 1 0:3 one i vere: cut asbaccondsastrocdbesspstebeseebobbesboze Me irodictya cf. sp. A Cee Mee oe a 5 13 OMMOSHACATTAD WNECA sonecsssacsocveanoacenondoccesondncdéeont 1 135 Skenidioides costatus By ae a 3 0-8 Total 74 99-95 Sowerbyella sericea 2 = eee 4 1-0 triplesiid indet. 1 --1- +--+ -- 1 0-3 triplesiid indet. 2 2) ee Oe 2 0-5 PV = pedicle valves, BV = brachial valves, C = conjoined valves, I = indeterminate valves, NO. = equivalent number of total animals. (N.B. TRILOBITES Numbers in brackets are total fragments of colonial or fragmented individuals trinucleid pygidia (1) — see discussion). See discussion on p.127. PPC LERGKCE PNA ay oct cuc sls sesesct scene sciiicscnssmeceoe messes 5 1-3 Atractopyge Peidia BA ce OS A Ie LR ee SAAR 18 4-6 Table 16 Total assemblage collected from Raheen. | Atractopyge cephala (9) ea (15) 5 0-5 Internal External Total oa stcseeenenucvtvecnson it OS ES a AC hei OOS PRPIGUIGX ICEPNAION a scccsterescseseescaserssesecesaceecdeuees 1 0-3 OTHER ELEMENTS Cen ral z ; a RPSONOIC DTV OZOANS) .<.ceececa-aececqecceceanesaecerceeteaneense 75 19-4 ee eeeeetein © al BeTS CHT OZ OAS) fa. eae es. Soe Sysco oautiamanens seateedse 42 10-8 pomere eves”) Oe ae ale Bh 820 PESIROPOGISPECIES, .2o0..2.0.eeeeeaeneeeenecernatsesooevenesemenece 29, «7-5 nats ae ad Lape A > 5, GunZ ; oe >“ IEAGIIGIS - coc eqicenbsouoldadacteceentacenee ceed te eee ener aeeeeee 2 10 I ee Cable sab anthas a Sen S op Ee DiVAIVeS ......... 2s. eeeeee cece este testes e teeter te tteetetteees 2 0-5 TRILOBITES = CRSICIES (AO) ceansoccadesdooseesesteeemnpaeaacobppacnaecrcs 1 ae ATIDYE PYCICA atte ek hte eene tera eee 2 1-5 ort peones te ee ee eLel ele dessins dRisica vieivslieinicininic vnrialen 11 2:8 firelaspistcephalonier pete eee 1 0-7 coral’ eaiisi wees eegeearseeseeseecacsuneneeeeeeaeeaeeteneenreereeees 1 0-3 iretaspidice phialond see Cee See. SO. inane 1 0-7 Echinosphaerites single plates (15) tretaspid cephalic fragments (2) Echinosphaerites attached plates) (27) iwressereeeeseeec eee: 5 1-3 tretaspid pygidium (1) PIRMIMOSPNGCrIESANECAE 22.2... -cccecencecrseeeseseeneesenecoees 5 ils3} MMMM AIC YSLOIG Meee cece ocjs sere ceasicch Ek ceeseer cee seeeceemenese 9 23 OTHER ELEMENTS (7g 1010) (0S (3) i chase csen ana SA cea aac aba aadddcdocnclsasneaspanannncbb 1 0-7 Total 387-1001 seastiopodsisn’ weenie: ase een meee 36-265 GORE ISCO) Scpesner popradenctocdocObecods aac coohpnachaccdadosocuspadayce 2 IES) 56 2 : PE clealves BY = brachial valves, C = conjoined valves, 1 pais BSS PREC oder CCD aC ceRRERRCt ore aecoreeceecodo tec nrorceacd g fe indeterminate valves, NO. = equivalent number of total animals. (N.B. Me OS ie BEDE ARS AINE E OE ES LS a ¥ Numbers in brackets are total fragments of colonial or fragmented individuals Total 136 100-0 —see discussion). See discussion on p.127. recorded by Brenchley et al. (1977) and slightly increase the genera known from here. Ballinatray (Table 19) preserves a rare occurrence of shelly fossils in a normally graptolitic facies. Coastal sections of the Ballinatray Formation north of Courtown have hitherto “yielded only graptolites (Crimes & Crossley 1968, Brenchley \% Treagus 1970). Shells are found in thin, densely packed | bands in calcareous slates. Restricted exposure in the bed and banks of the Owenavorragh River and strong deformation make assessment difficult, but the lack of size variation, restricted diversity and dense packing suggest that the fauna PV = pedicle valves, BV = brachial valves, C = conjoined valves, I = indeterminate valves, NO. = equivalent number of total animals. (N.B. Numbers in brackets are total fragments of colonial or fragmented individuals — see discussion). See discussion on p.127. was a coquina. Whether long distance transport or local winnowing was involved is not clear, but all specimens are small including the rare genera, elsewhere of a normaily larger size. 132 Table 17 Total assemblage collected from Greenville. M. A. PARKES Table 18 Total assemblage collected from Greenville-Moyne. Internal External Total Internal External Total JAY JERAV(C Il JAY AYE I no. % PY BAY EL JAY BY (CII no. % BRACHIOPODS BRACHIOPODS Chonetoidea abdita ES sy = i Bil 4 = D7 Il atlas Anisopleurella sp. 8 2-- 25 -- 8 6:0 dalmanellids, indet. - --- 41- - 4 Dei], Cremnorthis parva 8 2 8 6-0 Oanduporella? le =e= 9 i= = 3 2-0 dalmanellids, indet. 39 I © = IO i = = 45 = 33g Dolerorthis? bh 1 St - --- ly 0% TRILOBITES Kullervo aff. hibernica 3 -- - 3 = 2) 4 3:0 RATA NAUMAN Jos CIOVBVE) soonsaocesosonccseacenooacoscaovsoace 5 3-4 Leptestiina oepiki a Or = y Oe Bu D0) A ORAQHGOD SO. |OKBMUW soonneescnecnass0sc0nceontenaenconucs 1 0-7 lingulid, indet. 3 11 =0-748 Ampyx austinti cephala (8) Nicolella cf. actoniae Bee je A = 8 6-0 Ampyx austinii glabellar spines (7) ‘Orthambonites’? 5 ae i lke Siena ANAM ONEE CUISUI OH PONAGVE 5 ococcaccocsunacdacosonaccspon0qquosoGnn00 16 =10-9 Onthinoerania? | 1 - 0-78 Wirinoduswagnostiformisipy eld lalaueerereeeeeceeee raat eee eee 8 34; plectambonitacean, indet. 1 0-78 Miretaspis Cephalon) .aacshiseacsnis: ashe seetmanesemaseeet cere cate 1 0-7 Rhactorthis? 1 af 1 1 (0 thinucleidicephalatee. cesceneassecercer esters eect reece Rene cereee 4 2-7 Salopia sp. S432) 2h eee trinucleid pygidia @) Saukrodictya cf. sp. A ---- ---2 1 “07g trinucleid thoraces (2) Skenidioides costatus fet 2S. 9 hee 1 0-75 trinucleid genal spines (2) Sowerbyella? Bh Lees Deis me 340020) OTHER ELEMENTS strophomenid, indet. 1 = 1 0-75 ENOTOGTING! GVA! sacoccassnascasanassacooosconccesacnadoocuaeN 1 0-7 TRILOBITES gastropods wid au Gahoiad a Mepeaame cio ueensen ane oteecastnepeetoes meter 38 ©25:9 cybelinid? librigenae (8) bivalve _begoonoeoana9 noe sandansoconnauTaaDdooObODsUsHDOoROOAECCOLOIORC 1 0-7 indet. pygidia (1) (OWE EIGN: Se acsascosasaedsdocaapcacode race sbecucadascnososaaaoosoas 4 2-7 indet. cephala (2) Sn ae eee 2 1-5 Total 147 100-0 OTHER ELEMENTS erinoidiossicles: (14) steers fees eee 1 0-75 i ans 22 : PV = pedicle valves, BV = brachial valves, C = conjoined valves, I = Bee BEYOZOADS 3-25-6200 052s soreness aaa a 4 x indeterminate valves, NO. = equivalent number of total animals. (N.B. OSS WW OZ0ans, So aa ec oe aa aaa aia sare us OOO O5D ‘ Numbers in brackets are total fragments of colonial or fragmented individuals Cheirocrimid Cystoids ©.-....-.r.... one. eee nne eee dene een gee 3 22 —see discussion). See discussion on p.127. (OLY H (C60) 0 LS ate aaemaenoe oaoepeetnton santadbododdaserchaddiedchsonnas8 3 2:2 Total 134 100-0 Comparisons and contrasts with coeval associations A prerequisite to recognition of faunal provinciality in bra- chiopod assemblages is the determination of whether appar- ently different faunas are simply facies controlled. This was undertaken as an integral part of this project and is briefly summarised here. Due to the problems noted before, only some localities yielded assemblages which could be treated as palaeocommunities. These were Grange Hill Horizon 1, Grange Hill House Cottage, Carrigadaggan, Raheen, Green- ville and Greenville-Moyne. In fact, the assemblages are clearly dissimilar to platform sequence faunas from Laurentia, and to the marginal Scoto- Appalachian faunas from Ireland and Britain. Few assem- blages of similar age from the Lake District are known and there are no modern quantitative studies for direct compari- son. Essentially, only comparisons with Anglo-Welsh associa- tions were applicable. Principal Component Analysis (PCA) using Palstat (Harper & Ryan 1987) is an effective way of discriminating and illustrating the elements which contribute most to the variation between samples. One example is given to illustrate the comparitive analysis undertaken. Figure 15 shows a plot of vector 1 against vector 3 for the Grange Hill Horizon 1 association and 11 samples of the Nicolella Asso- ciation from the Gelli-grin Formation from Lockley (1980). Plots of vector 1 against 2 discriminated abundance trends of dalmanellids. The plot in Fig. 15 clearly differentiated the Grange Hill elements as contributing substantially to the variation on vector 3. Whilst not a precise test, it shows important differences which do not permit the association to be considered as an equivalent to the Nicolella Association, PV = pedicle valves, BV = brachial valves, C = conjoined valves, | = indeterminate valves, NO. = equivalent number of total animals. (N.B. Numbers in brackets are total fragments of colonial or fragmented individuals — see discussion). See discussion on p.127. Table 19 Total assemblage collected from Ballinatray. Internal External Total PVABWVAC SIE VABWVaGaal no. % BRACHIOPODS | Chonetoidea sp. Lj = = Sy = l 1-8 dalmanellids, indet. 38 26 -- 810 - 14 45 78-9 Dolerorthis? 12-- --- = 2 33 Leptestiina sp. 11-- --- = i 18 plectambonitacean, indet. 1 2-- - —- — 3 4 7-0 Sowerbyella? 1 --- - -- - 1 i TRILOBITES TPO MT OWES? FNFBCWEL -onceadaad soaosvabesnaeses08se8eG0 2 7300802 2 3-3 OTHERS CHINOIG OSSICIO Ss: pcieowsasatnede hess caboasete figs 1—5 MATERIAL AND LOCALITIES. Kildare, Grange Hill, Horizon 1: 6 internal moulds of brachial valves, 2 internal moulds and 2 external moulds of pedicle valves, both incomplete. Kil- bride: 8 internal and 1 external moulds of pedicle valves, 14 internal and 3 external moulds of brachial valves. MEASUREMENTS (mm) X1 xe P1.3, fig. 1. 10:5 12-6 P1.3, fig. 2 4-0 4-6 P1.3, figs 3, 4 6-6 9-0 P13, fig. 5 5-2 73 DISCUSSION. The small samples are inadequate to justify assignment to any species and in any case the taxonomy of ‘Orthambonites’ is in need of revision. Subfamily PRODUCTORTHINAE Schuchert & Cooper, 1931 Genus NICOLELLA Reed, 1917 Nicolella cf. actoniae (J. de C. Sowerby, 1839) Pl. 3, figs 6-9, 11 MATERIAL AND LOCALITIES. Kilbride: 10 internal and 8 external moulds of pedicle valves, 6 external moulds of brachial valves; all material is fragmentary. Carrigadaggan: 3 internal and 8 external moulds of pedicle valves, 1 internal and 5 external moulds of brachial valves; material is mostly fragmentary. Greenville-Moyne: 3 internal and 7 external moulds of brachial valves, 3 internal and 4 external moulds of pedicle valves and 1 conjoined internal mould. Kildare, Grange Hill, Horizon 1: 1 internal and 1 external mould of a brachial valve. DISCUSSION. Williams (1974: 58) commented on the mor- phological stability of Nicolella actoniae from mid Caradoc to early Ashgill times. All the samples here are probably very close to N. actoniae, or the subspecies N. actoniae obesa Williams (1963) from Bala, North Wales, a much deeper form than the nominate subspecies. All the samples collected here are poorly preserved and mostly broken, as well as relatively small in numbers; hence measurements of variation and counts of rib numbers are not possible. Thus assignment to one species, or several, is unrealistic until larger samples are available to assess the variation in morphology. The Greenville-Moyne sample is apparently not as deep in the ventral valve as that from Carrigadaggan or Kilbride, but this may be a preservational effect, owing to disparity in compaction between the fine-grained mudstones _ of Greenville-Moyne, and the coarse volcaniclastics of Carri- gadaggan or the tuffaceous siltstones of Kilbride. The latter M. A. PARKES | 140 | BRACHIOPODS OF DUNCANNON GROUP OF SE IRELAND two samples are strongly convex in the pedicle valve and | appear most similar to N. actoniae obesa, from the Gelli-grin ‘Formation of the Bala district. The broken nature of the samples, particularly of the Kilbride one (as are most species from there), generally precluded counts of rib numbers, but ‘three brachial valve external moulds from Kilbride had 10, 12 /and 15 ribs, and one pedicle valve had 11 in total. __ Type specimens of N. interplicata (M’Coy, 1846), from the /Kildare Limestone Formation, were examined (NMI- ‘lectotype F4565, paralectotypes F5564 and F11604), but they are inadequate for formal comparison. Indeed the latter specimen is probably not a Nicolella at all. As noted by Cocks (1978) the evaluation of topotypes from the Kildare Lime- stone Formation is necessary to assess this species. Whether | Nicolella calcarata M’Coy (1846), cited as rare in the slates of ‘Greenville and very common in the slates of Slieveroe, Rathdrum, is simply a deformed version of N. cf. actoniae or a separate species is not clear, since no Greenville topotypes were recovered. The lectotype (F4567) is broken and did not appear similar to any of the samples, being more transverse ‘and coarsely costate. A paralectotype (F5509) from Slieveroe jis also deformed and broken but appears similar to N. cf. actoniae. Larger collections of better-preserved material would be ‘desirable to enable both a biometric comparison with N. cf. actoniae and described subspecies as well as pre-Longvillian species such as N. humilis (Whittington & Williams 1955) and N. cf. strasburgensis (Williams 1962). Provisionally, the present material is compared to N. actoniae which, as pres- ently understood, encompasses a wide range of variation and a lengthy time span (mid Caradoc — mid Ashgill). \Nicolella ? sp. Pieseticele MATERIAL AND LOCALITY. Kilbride: 1 internal and external {mould of a brachial valve. DiscussION. This single concave brachial valve is question- ably assigned to Nicolella, but is different from the other samples of the genus, including the Kilbride material, princi- pally in being large and having a much greater ribbing frequency (at least 26 ribs). The brachial interior has the erect cardinal process, short divergent brachiophores and heavy deposits of secondary shell ankylosing the brachiophores to a short median ridge which are characteristic of Nicolella. It may possibly be compared to N. asteroidea Reed, which has more ribs than N. actoniae, but the preservation of the specimen is inadequate to assess the branching pattern of costellae. | PLATE 3 L 12661, internal mould of pedicle valve, x 4. 141 Family DOLERORTHIDAE Opik, 1934 Subfamily HESPERORTHINAE Schuchert & Cooper, 1931 Genus HESPERORTHIS Schuchert & Cooper, 1931 Hesperorthis sp. Else ie 10 MATERIAL AND LOCALITY. Kilbride: 3 internal moulds of pedicle valves. DIscussION. The small numbers of the genus are inadequate for specific determination, but the long apsacline interarea and overall shape suggest the specimens belong to Hesperor- this. The genus is known from the Caradoc rocks of Grange- geeth, as the Estonian species H. inostranfzefi, but the Kilbride species is dissimilar and much smaller. Two species are known from Girvan (Williams 1962) and one from the Llanvirn of Wales (Lockley & Williams 1981), but formal comparison requires a better preserved and larger sample from Kilbride. Family PLAESIOMIIDAE Schuchert, 1913 Subfamily PLAESIOMIINAE Schuchert, 1913 Genus PLAESIOMYS Hall & Clarke, 1892 Plaesiomys multiplicata Bancroft, 1945 ; Pl. 3, figs 13-16; Pl. 4, figs 1-6; Pl. 7, fig. 12 21896 Orthis flabellulum Sowerby; Reynolds & Gardiner: 589. 1945 Dinorthis (Plaesiomys) multiplicata Bancroft: 244; pl. 35, figs 4-6; pl. 36, figs 1-3. 1968 Dinorthis multiplicata Bancroft; Romano: 47; pl. 5, fig. M. 1978 Plaesiomys multifida (Salter); Cocks: 50 (pars). 1978 Dinorthis multiplicata Bancroft; Brenchley: 160. 1980b Lordorthis sp.; Mitchell, in Romano: 206. cf. 1985 Plaesiomys cf. multiplicata Bancroft; Harper et al.: 291; figs 6-24. Diggens & MATERIAL AND LOCALITY. Kildare, Grange Hill House Cot- tage: 26 internal and 12 external moulds of pedicle valves; 13 internal and 20 external moulds of brachial valves. DESCRIPTION. Exterior. Large, dorsibiconvex to convexi- plane valves of rounded subquadrate outline, with maximum width just anterior of hinge line, and about two-thirds as long as wide. Evenly convex anterior and lateral profiles in brachial valve, but pedicle valve flat to weakly concave except for swollen posterior axial surface. Ventral interarea flat and apsacline with open delthyrium. Dorsal interarea flat and orthocline, with open notothyrium. Radial ornament of evenly rounded costae and costellae and interspaces, and Figs 1-5 ‘Orthambonites’ spp. 1, 34 Kildare, Grange Hill Horizon 1. 1, BC 12658, internal mould of brachial valve, x 4. 3, 4, BC 12660, internal mould of brachial valve, latex cast and mould, x 4. Figs 2, 5 Kilbride. 2, BC 12659, internal mould of brachial valve, x 4. 5, BC igs 6-9, 11 Nicolella cf. actoniae (J. de C. Sowerby). 6-7, 9 Kilbride. 6, 7, BC 12662, internal mould of pedicle valve, latex cast and mould, x 4. 9, BC 12664a, internal mould of pedicle valve, x 2. Figs 8, 11 Greenville-Moyne. 8, BC 12663b, external mould of pedicle valve, x 2. 11, BC 12663a, internal mould of pedicle valve, counterpart of Fig. 8, x 2. Fig. 10 Hesperorthis sp. Kilbride. BC 12666, internal mould of pedicle valve, x 10. fe 12 Nicolella? sp. Kilbride. BC 12665b, external mould of brachial valve, x 2. igs 13-16 Plaesiomys multiplicata Bancroft. Kildare, Grange Hill House Cottage. 13, 16, BC 12667, internal mould of brachial valve, and | latex cast, x 4. 14, 15, BC 12668, external mould of pedicle valve, and latex cast, x 4. | f I | / M. A. PARKES 142 | BRACHIOPODS OF DUNCANNON GROUP OF SE IRELAND very fine concentric growth lines. Counts of 18-25 costae and 'costellae are present on 2, 0, 0, 1, 1,1, 0 and 1 valve exteriors ‘at the 5 mm growth stage and counts of 27 and 31 costae and ‘costellae on 2 and 1 valve exteriors at the 10 mm growth stage. | Ventral interior. Stout, small teeth directed dorsilaterally from anterior margins of wide delthyrium are supported by ‘strong receding dental plates. External ornament strongly ‘impressed, particularly near anterior margins. Dorsal interior. Simple, linear cardinal process is slightly ‘thickened posteriorly, situated on notothyrial platform which extends anteriorly for one-third of valve length as a low broad ridge. Blade-like, divergent brachiophores supported by \stout bases which, with the hinge line, define deep sockets. | ‘Discussion. Although large, the sample cannot easily be ‘statistically compared to the existing descriptions of the ‘species because many of the specimens are broken or poorly preserved. However, sufficient distinctive material is \described to assign it confidently to the species illustrated by ‘Bancroft (1945) from the Soudleyan of Glyn Ceiriog, North Wales, and in particular the sample described by Harper & \Mitchell (Harper et al., 1985) from the Clashford House Formation of Co. Meath. Re-collection of more specimens will allow a better assessment of the variability in the species, !particularly in external ornament, since the available material |shows a few specimens more like the closely related genus Dinorthis, than like Plaesiomys. The relationship of these two genera is in need of reassessment. However, the record of Orthis flabellulum from here (Reynolds & Gardiner 1896) is probably the present species. Lamont (1953) noted that it was ‘apparently a late variety of J. de C. Sowerby’s species with ‘bifurcation and trifurcation of ribs’. The specimens collected by Lamont are now held in the National Museum of Ireland, but were apparently never figured or described. They are labelled as ‘Dinorthis peplos’ on Lamont’s labels, but no publication of this name is known. | Family PLECTORTHIDAE Schuchert & Le Vene, 1929 Subfamily PLATYSTROPHINAE Schuchert & Le Vene, 1929 Genus PLATYSTROPHIA King, 1850 Platystrophia sp. 1 Pl. 4, figs 7-17; Pl. 5, figs 1-3 MATERIAL AND LOCALITIES. Kildare, Grange Hill, Horizon 1: 30 internal and 17 external moulds of pedicle valves; 33 internal and 13 external moulds of brachial valves; 6 internal and 10 external conjoined moulds; 12 external fragments. (Most of the material assigned to Platystrophia is incomplete. Kildare, Grange Hill, Horizon 2: 1 conjoined shell and one rachial valve. 143 Discussion. Although the large sample of specimens from Grange Hill, Kildare, was clearly identifiable as Platystrophia in both internal and external moulds, the material is almost all fragmentary or partially broken, precluding valid mea- surements. A statistical assessment was not possible. Since Platystrophia is a ubiquitous genus in Middle and Upper Ordovician rocks in Europe and America, with little variation amongst the many described species, it is necessary to reiter- ate the need for a complete species revision. Many authors such as Williams (1962: 126; 1963: 371), Wright (1964: 206), Cocks (1978: 55) and Hiller (1980: 143) have discussed the artificial nature of Cumings’ (1903) scheme of species group- ings, elaborated further by McEwan (1920) and modified in terminology by Schuchert & Cooper (1932: 67). In this scheme, the present material is all placed in the bicostate group, with 2 costae in the ventral sulcus and 3 on the dorsal fold. The counts on suitable material showed there were 1, 5, 3 and 2 pedicle valves with 4, 5, 6 and 7 costae respectively, on each flank. The Kildare population is finely pustulose, but distinguish- ing it from other species is a concentric ornamentation inviting comparisons with P. caelata Williams from the Soud- leyan of Shelve, Shropshire (Williams 1974: 76-77; pl. 12, figs 13, 14, 16-19). The ornament is of differentially developed lamellae, but further investigation would be needed to assess whether the lamellae are of the distinctive P. caelata type or merely accentuated growth lines. In his description Williams stated that P. caelata is uniplicate, but later noted all speci- mens are biplicate (bicostate); this would appear to be correct from the figures. Platystrophia sp. 2 PI. 5, figs 4, 5 MATERIAL AND LOCALITY. Kilbride: 1 external and 3 inter- nal moulds of brachial valves; 1 internal and 1 external moulds of pedicle valves, all incomplete. DISCUSSION. The present material is inadequate for specific determination but appears to differ from Platystrophia sp. 1 from Kildare, Grange Hill in having 7 ribs on the ventral flanks, although being of smaller average size. It also lacks the strongly developed overlapping lamellae, although in other respects it is similar, belonging to the bicostate group. Subfamily RHACTORTHINAE Williams, 1963 Genus RHACTORTHIS Williams, 1963 PI. 5, figs 6-11 MATERIAL AND LOCALITY. Kildare, Grange Hill, Horizon 1: 4 internal and 2 external moulds of brachial valves, 2 external moulds of pedicle valves and 1 conjoined internal and exter- nal mould. Rhactorthis sp. LATE 4 igs 1-6 Plaesiomys multiplicata Bancroft. Kildare, Grange Hill House Cottage. 1, 2, BC 12669, internal mould of pedicle valve, and latex cast, x 4. 3, 4 BC 12670, external mould of brachial valve, latex cast and mould, x 2. See also PI. 7, fig. 12. 5, 6, BC 12671, internal mould of brachial valve, latex cast and mould, x 4. Figs 7-17 Platystrophia sp. 1. Kildare, Grange Hill Horizon 1. 7, BC 12672, internal mould of pedicle valve, x 2. 8, BC 12673, latex cast of external mould of brachial valve, x 4. 9, BC 12674, latex cast of external mould of pedicle valve, x 4. 10, BC 12675, latex cast of external 4. mould of pedicle valve, x 2. See also Pl. 5, fig. 2. 11, 17, BC 12676, internal mould and latex cast of brachial valve, x 4. 12, 15, BC 12677, external mould and latex cast of brachial valve, x 4. See also PI. 5, fig. 1. 13, BC 12678a, internal mould of pedicle valve, x 2. 14, BC 12679, latex cast of external mould of brachial valve, with a Petrocrania attached, x 2. 16, BC 12680a, internal mould of brachial valve, x M. A. PARKES 144 id ) BRACHIOPODS OF DUNCANNON GROUP OF SE IRELAND MEASUREMENTS (mm) | XX. XS PI.5, figs 8,9 8-1 10-4 3-1 (conjoined internal mould) PLS, fig.10 8-2 11-1 4-1 (brachial valve) PI.5, figs 6,11 9-8 9-5 — (strained brachial valve) | Discussion. The sparse numbers of this genus from Kildare are inadequate to justify assignment, either to the type species R. crassa Williams from the Longvillian of Bala, or to 2ither of the other Caradoc species actoniae and grandis erected by Hurst (1979a), from the type Caradoc of Shrop- shire. Its presence, however, serves to emphasize the similari- ties of the Kildare fauna to that of the Bala district of North Wales. Family CREMNORTHIDAE Williams, 1963 Subfamily CREMNORTHIDAE Williams, 1963 Genus CREMNORTHIS Williams, 1963 i Cremnorthis parva Williams, 1963 | Pl. 5, figs 12-16; Pl. 6, figs 1-7 1963 Cremnorthis parva Williams: 379; pl. 4, figs 15-23; text-fig. 9. MATERIAL AND LOCALITIES. Kildare, Grange Hill, Horizon |: 38 internal moulds of brachial valves, 31 internal moulds of yedicle valves; 14 external moulds of brachial valves, 7 external moulds of pedicle valves. Carrigadaggan: 3 internal ind 1 external moulds of pedicle valves, 3 internal moulds of yrachial valves. Kilbride: 9 internal and 1 external moulds of pedicle valves, 6 internal moulds of brachial valves. sreenville-Moyne: 8 internal moulds of pedicle valves and 2 nternal moulds of brachial valves. DESCRIPTION. Exterior. Subcircular to semicular outline, noderately biconvex, with both valves about three-tenths as leep as long. Maximum width occurring at less than one-third he length of the shell. Length of brachial valve about even-tenths of the width. Pedicle valve length about four- ifths of width. Brachial valve gently sulcate, with flatly onvex lateral profile. Dorsal interarea short and anacline, entral interarea apsacline and about a quarter the length of he valve. Radial ornamentation costellate with angular cos- ae and costellae about 5 per mm at 2 mm anterior of imbones. Shell impunctate. | Ventral interior. Short teeth connected to shell floor by »LATE 5 pedicle valve (probably a juvenile specimen), < 10. 145 thickened deposits, and long apsacline interarea, which together bound a deep umbonal cavity. Subtriangular muscle scar does not extend much beyond cavity anteriorly. Muscle field composed of wide median adductor scars flanked by pair of narrow diductor scars which are lobate anteriorly. Sagittal length of muscle field nearly two-fifths length of valve. Dorsal interior. Large cardinal process, standing above hinge line, is continuous with blade-like median septum which extends four-fifths of the valve length, and is highest at about mid-length. Brachiophores short and slightly diver- gent, continuous with bases which curve posterolaterally as fulcral plates to define subtriangular sockets. Brachiophore bases are situated at about a quarter of the valve length. MEASUREMENTS Pedicle valve internal moulds (Kildare, Horizon 1): Variates X11 X2 X3 X4 X5 X6 X9 X10 Means 3:31 3-39 1:22 0-96 2-84 0-96 1-33 1-08 Sample size29 30 30 29 IM I A DW Variance- covariance 0:50 0-39 0-21 0-10 Matrix 0:57 0-14 0-08 0-18 0-05 0-05 0-23 0-36 0-09 0-06 0-39 0-04 0-04 0-03 0-01 0-03 0-02 0-20 0-14 0-13 0-04 0-08 0-03 0-14 0-14 0-14 0-08 0-03 0-08 0-02 0-08 0-07 Brachial valve internal moulds (Kildare, Horizon 1): Variates XX XGA SK 4 Means 3:07 3-44 1-13 0-87 0-77 1-28 2-54 Sample size i) Su. «SS 3S) SB) BE Variance- covariance 0:46 0:38 0:14 0:08 0-08 0-10 0-33 Matrix 0:76 0:09 0:10 0:08 0-15 0:26 0-10 0-01 0-03 0-02 0-01 0-04 0-01 0-03 0-06 0-03 0:03 0-06 0-06 0-08 0.30 DISCUSSION. The numerical data in the description are based only on the mean values for the large sample from Grange Hill, Kildare, where this species is a very common element of the fauna. The poor preservation of the samples from Kil- igs 1-3 Platystrophia sp. 1. Kildare, Grange Hill Horizon 1. 1, BC 12677, detail of ornament of latex cast, x 10. See also Pl. 4, figs 12, 15. 2, BC 12675, detail of ornament of latex cast showing the accentuated lamellae, x 10. See also PI. 4, fig. 10. 3, BC 12681, internal mould of igs 4-5 Platystrophia sp. 2. Carrigadaggan. BC 12682, internal mould of brachial valve and latex cast, x 4. ‘igs 6-11 Rhactorthis sp. Kildare, Grange Hill Horizon 1. 6, BC 12683a, internal mould of brachial valve, x 4. 7, BC 12684, external mould 16, x 4 x 10. | of pedicle valve, x 4. 8, 9, BC 12685a, ventral and dorsal views of a conjoined internal mould, x 4. 10, BC 12685b, latex cast of external mould of conjoined valves, counterpart of Figs 8-9, dorsal view, x 4. 11, BC 12683b, external mould of brachial valve, counterpart of Fig. igs 12-16 Cremnorthis parva Williams. Kildare, Grange Hill Horizon 1. 12, BC 12686, latex cast of external mould of brachial valve, x 10. 13, 14, BC 12687, external mould and latex cast of brachial valve, x 10. 15, 16, BC 12688, external mould and latex cast of brachial valve, 146 LAURENT UA OCEAN 1APETUS Avalonia (after Parkes, 1992). bride and Carrigadaggan made measurement of all variates difficult, but a principal component analysis (PCA) of all three samples shows that in plots of the first four eigen- vectors (see Fig. 17) the two small samples fall within the same region as the Kildare sample. It is possible that analysis of a larger topotype sample of the species would show significant differences between the Welsh and Irish forms, but the original description was based on fewer than ten NS ANGLO-WELSH GONDWANALAND i Fig. 16 Palaeogeographic reconstruction of the Iapetus region in mid-Caradoc times (c. 448 Ma) showing the main continental masses of Laurentia, Baltica, Gondwana and Eastern and Western Avalonia. The main brachiopod provinces of Scoto-Appalachian, Baltic and Mediterranean affinity are also shown. At this time, an Anglo-Welsh Province, including these Leinster faunas, was centred on Eastern SCOTO-APPALACHIAN PROVINCE _ ‘BALTIC PROVINCE EAST AVALONIA valves. The mean percentage length of the ventral muscle field relative to the valve length is significantly longer in the Irish form (Kildare — 39% compared to 35%, p < 0-01). This | is considered inadequate to justify erection of even a new subspecies, since the Bala sample consisted of only 5 speci- mens. A larger, better preserved sample would probably encompass the same variation seen in the Kildare form. f PLATE 6 Figs 1-7 Cremnorthis parva Williams. Figs 1-5, Kildare, Grange Hill Horizon 1. 1, BC 12689, internal mould of brachial valve, x 5. 2, 3, BC 12690, internal mould of pedicle valve, latex cast and mould, x 15. 4, BC 12691, internal mould of brachial valve, and BC 12692, internal mould of pedicle valve, both x 6. 5, BC 12693, internal mould of brachial valve, x 3. Figs 6, 7, Carrigadaggan. 6, BC 12694, ventral view of conjoined internal mould, x 10. 7, BC 12695, internal mould of pedicle valve, x 10. Figs 8-16 Skenidioides costatus Cooper. Kildare, Grange Hill Horizon 1. 8, 12, BC 12696, dorsal and ventral views of conjoined internal mould, x 10. 9, BC 12697, internal mould of brachial valve, x 10. 10, BC 12698, internal mould of brachial valve, x 10. 11, BC 12699, ventral view of conjoined internal mould, x 10. 13, BC 12700, internal mould of pedicle valve, x 10. 14, BC 12701, internal mould of pedicle valve, x 10. 15, BC 12702, internal mould of pedicle valve, x 10. 16, BC 12703, internal mould of pedicle valve, x 10. 3RACHIOPODS OF DUNCANNON GROUP OF SE IRELAND 147 148 Family SKENIDIIDAE Kozlowski, 1929 Genus SKENIDIOIDES Schuchert & Cooper, 1931 Skenidioides costatus Cooper, 1956 Pl. 6, figs 8-16; Pl. 7, figs 1-5 1956 Skenidioides costatus Cooper: 493; pl. 97, figs 38-48. aff. 1962 Skenidioides aff. costatus Cooper; Williams: 126; pl. 11, figs 24-27, 52. cf. 1963 Skenidioides cf. costatus Cooper; Williams: 375-377; pl. 4, figs 7-14. cf. 1974 Skenidioides cf. costatus Cooper; Williams: 82-83; pl. 13, figs 14-16; pl. 14, figs 1-3. cf. 1979a Skenidioides cf. costatus Cooper; Hurst: 242; figs 145-159. MATERIAL AND LOCALITIES. Kilbride: 2 internal and 3 exter- nal moulds of pedicle valves; 3 internal and 6 external moulds of brachial valves. Kildare, Grange Hill, Horizon 1: 10 internal and 10 external moulds of brachial valves; 19 internal and 3 external moulds of pedicle valves. Kildare, Grange Hill House Cottage: 2 internal and 1 external moulds of pedicle valves; 1 internal and 2 external moulds of brachial valves. Carrigadaggan: 1 internal mould of a pedicle valve. Kildare, Grange Hill, Horizon 2: 3 internal moulds of pedicle valves. Greenville-Moyne: 1 internal and 1 external moulds of a pedicle valve. DESCRIPTION. Exterior. Ventribiconvex, subpyramidal Sken- idioides with pedicle valve length about seven-tenths of the length, and about 40% as deep as long. Brachial valve gently convex with distinct median sulcus, about 50-70% as long as wide. Ornament of radial costellae, about 2-5, commonly 3, ribs per mm 2mm anteromedially of umbo, in both valves. Commonly a wider median rib on the pedicle valve, with total rib counts of between 12 and 20 with 15-17 the most common frequency. Ventral interarea high, catacline to apsacline with open delthyrium. Dorsal interarea shorter, anacline. Ventral interior. Generally unsupported spondylium about a quarter as long as the valve, and about 94% as long as wide. Some shells have a median thickened ridge of shell support- ing the spondylium. Dorsal interior. Thin median septum, continuous anteriorly from shaft-like cardinal process, extending about 90% of valve length. Slender brachiophores with bases convergent onto median septum defining a diamond shaped cruralium about a third as long as valve. DISCUSSION. Previously described samples compared to S. costatus (Cooper 1956) differ in some proportions from each other and from the material described here, but the differ- ences are not considered important enough to warrant taxo- nomic recognition. Principal component analysis of all the PLATE 7 M. A. PARKES || MEASUREMENTS Pedicle valve internal moulds (Kildare, Horizon 1): Variates Xl X2 x4 XG eel 4 Means 3:37 4-64 1:29 0-81 0-84 Sample size 19 19 I) 18 18 Variance-covariance 0-46 0-42 0:06 0-13 0-06 Matrix 0:77 0-05 0-10 0-08 0-08 0-01 0-01 0-06 0-03 0-02 ° Brachial valve internal moulds: Variates X1 X2 X15 X16 X24 Means 3:00 4-11 0-98 0-83 2-76 Sample size 10 10 10 10 10 Variance-covariance 0:25 0-24 0:06 0-02 0-26 Matrix 0:78 0-02 0-10 0-14 0-03 0.0 0.06 0-02 0-0 0-32 Brachial valve external moulds: Variates Xl X2 x4 Means 2:73 5-32 10-47 Sample size 10 10 7 Varlance-covariance 0-43 0-63 ~=0-05 Matrix 1A S014) 0-12 material shows no differentiation on the first four vectors, as shown in Fig. 18. As noted by Mitchell (1977: 50) and Hiller (1980: 146), internal features of many Skenidioides species show no significant differences, the species being separated on the basis of the density of external ornament. The rib counts for this Irish material is comparable to previously described populations of S. costatus of similar size range. One aspect which apparently requires further investigation is the branching mode of the ribs of the species. Williams (1974: 83) discussed the differences between Shelve and Bala stocks, with new costellae arising only from the ventral median rib in the former and branching freely from lateral costae in the latter. In the Bala stocks, the majority of costae branched externally (Williams 1963: 377). However, Hurst (1979a: 242) states that the Shropshire stock only branched Figs 1-5 Skenidioides costatus Cooper. Kildare, Grange Hill Horizon 1. 1, 3, BC 12704, external mould of brachial valve and interareas, x 6, and enlarged oblique posterior view of latex cast of interareas, showing hinge line and open delthyrium and notothyrium, Xx 10. 2, BC 12809, latex cast of external mould of pedicle valve, x 6. 4, 5, BC 12705, external mould of brachial valve and interareas, and latex cast showing open delthyrium, x 10. Figs 6-11 Oanduporella cf. reticulata Hints. Kildare, Grange Hill House Cottage. 6, 9, BC 12706, external mould of pedicle valve, latex cast and mould, x 10. 7, 8, BC 12707a (upper) and BC 12708a (lower); internal moulds and latex cast of 2 pedicle valves, x 4. 10, BC 12711b, external mould of pedicle valve, x 4. 11, BC 12711a, internal mould of pedicle valve, counterpart of Fig. 10, x 4. Fig. 12 View of slab, containing BC 12670, external mould of Plaesiomys multiplicata Bancroft (P; see also Pl. 4, figs 3-4), to illustrate the typical assemblage at Kildare, Grange Hill House Cottage; with Oanduporella cf. reticulata Hints (A), Rafinesquina sp. (B), and numerically dominant gastropods (C). x 1. ; p 4sRACHIOPODS OF DUNCANNON GROUP OF SE IRELAND 149 150 V-2 -5.1 -3.3 -1.5 0.3 2.1 3.9 V-1 C24V2 0.9 0.5 0.1 V-4 -0.3 -0.7 -0.5 0.3 1.1 1.9 V-3 C24V2 $2511” Fibs) Fig. 17 Principal component analysis of samples of Cremnorthis parva, internal moulds of pedicle valves only. Top, vector 1 against vector 2. Below, vector 3 against vector 4. VW = Carrigadaggan, * = Kilbride, 0 = Kildare, Grange Hill Horizon il. internally. The present material has few external moulds with costellae, but those that do, show both internal and external branching. M. A. PARKES | Superfamily ENTELETOIDEA Waagen, 1884 Family DALMANELLIDAE Schuchert, 1913 Genus OANDUPORELLA Hints, 1975 | Oanduporella cf. reticulata Hints, 1975 Pl. 7, figs 6-12; Pl. 8, figs 1-7 | cf. 1975 Oanduporella reticulata Hints: 19, 105; pl. 1, figs 1-15; pl. 2, figs 1-5. | 1980a? Ravozetinal/Onnizetina; Mitchell, in Romano: 206. 1985 Oanduporella cf. reticulata Hints; Harper & Mitch- ell, in Harper et al.: 295, figs 25-37. | MATERIAL AND LOCALITY. Kildare, Grange Hill House Cot- | tage: 10 internal and 6 external moulds of brachial valves, 13 | internal and 9 external moulds of pedicle valves. DISCUSSION. Harper & Mitchell gave a full description | (Harper et al., 1985) of material they compared to Hints’ species from the east Baltic, with which the present material - from Kildare accords well. The Herbertstown material, from ' the Clashford House Formation, was the first record of the | genus from Britain or Ireland and the present sample repre- | sents the second known occurrence from these areas. It serves to emphasize the similarities of the Kildare fauna to | that from Herbertstown with two conspecific forms, Plaesi- omys multiplicata Bancroft and Oanduporella cf. reticulata | Hints, present, with a possible third, Hibernodonta? Harper | & Mitchell (in Harper et al. 1985). There are few suitable — specimens but similar rib counts (5-6 per 2 mm at 5 mm_ sagittally) are seen in the Kildare specimens, although the | microsculpture is well developed on most specimens. Oanduporella sp. (Not figured) . MATERIAL AND LOCALITY. Greenville: 3 external and 1 internal moulds of pedicle valves, 1 external mould of a) brachial valve. . DISCUSSION. The poor preservation, and deformation in this mudstone lithology made it impossible to compare this mate- rial to the specimens from Kildare, Grange Hill House | Cottage. Nevertheless, this small sample shows the character-_ istic pitted microsculpture of the genus and extends the known geographical range. | Genus REUSCHELLA Bancroft, 1928 Reuschella ? sp. Pl. 11, fig. 16 | | | | MATERIAL AND LOCALITY. Kilbride: A single internal mould | of a pedicle valve. DISCUSSION. The single poorly preserved mould is assigned | to Reuschella on the basis of the sharp median carina, curved long apsacline interarea, the ventral muscle scar and massive teeth. The specimen represents the sole occurrence of the genus within the southeast Ireland Caradoc. It is known from the Actonian of Shropshire (Hurst 1979a), the Soudleyan of Shelve, Shropshire (Williams 1974) and Bala (Williams 1963) as well as from Girvan (Williams 1962). e ; 3RACHIOPODS OF DUNCANNON GROUP OF SE IRELAND 151 0.9 0.5 | 0.1 . V4 | -0.3 -0.7 -1.1 -214 -1.1 ft) 09 19 2.9 V-3 SKENPV B 2.9 1.9 0.9 v2 —2.1 One W073 1.5 Pet eye) -21 -0.9 0.3 1.5 27 ~3t9 V-1 SKBVEX V-1 SKBVEX Ea 199 087 0.5 Ai” 72:9 -3.1 -19 -0.7 0.5 i729 | V-1 SKEEXT V-1 SKEEXT lig. 18 Principal component analysis of Skenidioides costatus, from Kilbride (@) and Kildare, Grange Hill House Cottage (*) and Horizon 1 , (Q). A, pedicle valve internal moulds; B, brachial valve external moulds; C, pedicle valve external moulds. 152 Family LINOPORELLIDAE Schuchert & Cooper, 1931 Genus SALOPIA Williams, in Whittington & Williams 1955 Pl. 11, figs 18-20 MATERIAL AND LOCALITIES. Kildare, Grange Hill, Horizon 2: a single internal mould of a pedicle valve. Greenville- Moyne: 4 internal and 1 external moulds of brachial valves. Salopia sp. DESCRIPTION. Ventral valve. Interior, strongly convex mould, slightly sulcate, over half as deep as long, and slightly wider than long. Maximum width just anterior to hinge line, giving a subcircular outline to shell with high apsacline, but nearly catacline interarea. Short teeth are supported by divergent dental plates. Muscle field extends beyond dental- plates to about one-third of valve length. External ornament only seen where it is impressed around margins of shell interior. Dorsal valve. Interior, gently convex in mould form, with low notothyrial platform between divergent brachiophores carrying a thin linear shaft which is continuous with a low median septum, extending to about mid-length. External ornament impressed slightly around margins of interior. MEASUREMENTS (mm). BC 12758: X1 = 11-5, X2 = 12, X4 = 6, X9 = 4-5 (Pl. 11, fig. 19). DiscussION. Salopia is known from the Llandeilo (Lockley & Williams 1981: 51) and Lower Caradoc (Williams 1963, 1974; Whittington & Williams 1955) of Wales and Shrop- shire. The single pedicle valve from Kildare is inadequate for formal comparison, but is apparently significantly deeper than described species. The sample from Greenville-Moyne is indistinctly preserved, and is lacking in pedicle valves, so cannot be directly compared to the Kildare specimen. The closest obvious comparison is with Salopia salteri (Davidson, 1869). Family SAUKRODICTYIDAE Wright, 1964 Genus SAUKRODICTYA Wright, 1964 Saukrodictya cf. sp. A of Hints Pl. 8, figs 8-15 cf. 1979 Saukrodictya sp. A, Hints: 57; pl. 2, fig. 10; pl. 4, figs 15-22. MATERIAL AND LOCALITIES. Carrigadaggan: 1 internal and 3 external moulds of pedicle valves, 4 external moulds of indeterminate valves. Greenville-Moyne: 2 indeterminate external moulds. DESCRIPTION. Exterior. Typical ornament of exopuncta, up to 6 radial rows, closely spaced, in the interspaces between narrow ribs. Interspaces are relatively wide and rounded. Ventral valve gently convex, wider than long. Ventral interior. Nearly 50% wider than long. Gently convex profile, rectimarginate commissure. (Dorsal valve unknown). M. A. PARKES jh sh DIscussION. The mould material, although very poor, 1s | assigned to Saukrodictya rather than Salacorthis because of | the typical pitted ornament found only in the interspaces and not on the thin ribs. The frequency of ribs is also greater than in Salacorthis costellata Williams (1974), the only known species, and the pedicle valves described here are not sulcate, as are those of Salacorthis. There are a number of described species of Saukrodictya, but in all cases they are based on limited material and are not well known. The present material does not permit a detailed comparison with described species, but the illustrations of Saukrodictya sp. A by Hints (1979) from the Idavere and Johvi Stages in Estonia (L. Caradoc — multidens Biozone) are most similar to this material. The species apparently lacks the strong fold of S. reticula Vinassa, 1927 (Villas, 1985). It has a — lower frequency of ribs than S. rotundopora Hints or S. | oblongatopora Hints, both of which also have a fold. Simi- | larly, S. porosa is sulcate and also has a greater rib frequency, though its general outline is similar (Havlicek 1977). The type | species S. hibernica Wright (Wright 1964; see also Hiller, | 1980) is strongly sulcate. However, Villas (1985) has sug- gested that S. hibernica may be conspecific with S. reticula (Vinassa, 1927), ‘but there are too many gaps in the knowl- edge of ‘British‘ and Sardinian Saukrodictyae’. The present — sample unfortunately does nothing to clarify the definition of species, but is stratigraphically and biogeographically signifi- — cant. These are the oldest known occurrences of Saukrodictya in Ireland. S. rotundopora Hints (1979: 53) and S. oblongato- pora Hints (1979: 55) are from approximately contemporane- ous stages in the Middle Caradoc of Estonia. Other occurrences are Ashgill in age, including the type species S. hibernica from Portrane (Wright 1964: 216) and Wales (Hiller 1980: 165), S. wrighti from Belgium (Sheehan 1987) or from | the Llandovery (S. sp. from Wales (in Temple, 1970: 32); S. . | H | sp. B from Estonia (Hints, 1979: 58)). According to Havlicek (1977) the oldest occurrence of species of Saukrodictya are S. porosa from the Liben and Letna Formations of Bohemia | (Middle Llandeilo to Costonian) and in the Costonian/ | Harnagian of Portugal (Mitchell 1974). It would thus appear to have a Gondwanan origin and to have migrated north: | ward, reaching Ireland by the Longvillian or earlier. Superfamily GONAMBONITOIDEA Schuchert & Cooper, ) 1931 Family KULLERVOIDEA Opik, 1934 | Genus KULLERVO Opik, 1934 Kullervo aff. hibernica Harper, 1952 1218 aff. 1952 Kullervo hibernica Harper: 100; pl. 6, figs 6-8. aff. 1977 Kullervo aff. hibernica Harper; Brenchley et al.: | 70. | PI. 9, figs 1-10, PLATE 8 Figs 1-7 Oanduporella cf. reticulata Hints. Kildare, Grange Hill House Cottage. 1, 2, BC 12709, external mould of brachial valve, and latex cast, X 10. 3, 4, BC 12710a, internal mould of brachial valve, and latex cast, x 10. 5, 6, BC 12710b, external mould of brachial valve, counterpart of Figs 3—4, latex cast, x 10. 7, BC 12712, internal mould of brachial valve, x 72. Figs 8-15 Saukrodictya cf. sp. A of Hints. Carrigadaggan. 8, 11, BC 12713, external mould, latex cast and mould, x 10. 9, 10, BC 12714, external mould and latex cast, x 10. 12, BC 12715a, internal mould of pedicle valve, x 8. 13, BC 12716b, latex cast of external mould, x 10. 14, 15, BC 12715b, external mould of pedicle valve, counterpart of Fig. 12, latex cast and mould, x 10. } a oa) w 3RACHIOPODS OF DUNCANNON GROUP OF SE IRELAND M. A. PARKES 154 i ) BRACHIOPODS OF DUNCANNON GROUP OF SE IRELAND MATERIAL AND LOCALITIES. Carrigadaggan: 3 internal moulds and 5 external moulds of pedicle valves, 2 internal and 3 external moulds of brachial valves and 8 indeterminate ‘external moulds. Kildare, Grange Hill, Horizon 1: 1 external mould of a pedicle valve and2 fragments of internal moulds of pedicle valves. Greenville-Moyne: 3 internal and 3 external moulds of pedicle valves, 2 external moulds of brachial valves, and 2 indeterminate external moulds. DESCRIPTION. Exterior. Strongly ventribiconvex Kullervo with pyramidal pedicle valve and maximum width along hinge line. Concave posterolateral flanks on extended hinge line, anterior slope evenly convex. Shallow median sulcus in brachial valve, but essentially rectimarginate commissure. Ventral interarea high, curved, apsacline near umbo, cata- cline near hinge. Dorsal interarea short, anacline. Ornament of distinctive reticulate pattern, of strongly developed con- centric lamellae and regular radial ribs. Radial ornament absent on posterolateral flanks. | Ventral interior. Spondylium with hemisyrinx supported by well-developed median septum extending nearly to mid length. External ribs impressed on anterior margins of valve interior. | Dorsal interior. Cardinalia of thick divergent socket ridges about three times as long as wide, extending less than half valve width. Thin cardinal process in narrow space between their ends. Socket ridges merge anteromedially with thick notothyrial platform, itself passing into thick median ridge anteriorly. At about mid-length of valve the ridge tapers to a thin, low median septum, separating very poorly impressed adductor scars. Discussion. Harper’s (1952) original description of Kullervo hibernica was based on limited material. In respect of the external ornament, overall shape and ventral interior the described material is comparable to the paratype material of Kullervo hibernica Harper from Grangegeeth (NMING: F14035, 14036). However, a single well-preserved brachial valve interior from Carrigadaggan (PI. 9, figs 12, 13) shows some differences from the holotype of hibernica (NMING: \F14034; Pl. 9, fig. 18), which is itself broken and missing posteriorly; in the hibernica holotype the impression of external ribs is more regular, stronger and abruptly and evenly terminated. In the Carrigadaggan specimen the ribs fare variably impressed, also more irregular and longer. However, an additional broken, poor specimen does show a more reg 10F SUIPUD}X9 puke JUoIINSap Ayyensn uldAplyy se = = 4 = AL - puod JO aseq edu }dd9xa siyoes 0} pasny Ajjenied Ayjewuou ore yey) sajnuulg Z ‘ € ( % tfc b/c (-) aeuurdipenb 10 (¢) ayeuutdiy *(Z) ajeuurdiq soysueig euuid Arewutig = = = = = = 4 puoiy jo jaed 19MO] Ul sajnuuld pliajdojoAd Wo}1uas 10 Ie|NdIQIO 4 = = = tL 4b - doRJINS [RIXEGR UO SOWOYOI] AP]NYJION|NYY i = + aL ; i i - aoRjins jerxege uo aviideg AD ({)uy (61d wy /AD Le yArs| ig/juy uy (wry) sn Ad0joAoryduue JO (AD) aAd0]949 (1g) on Aoviedéyoeig ‘(uy) sAd0WOUR PROS 4 = dk ae ae ae ~ sjjem [euljonue (—) ou Aypenq4ia JO (+) Su01}s YIM JJIINNd jIexeqy 4 = de +4 = de - Jayjour duo Woy (—) Ie IWS 10 (+) JUdIAIJIP ddRyINs [BIXepe UO S][99 [PISOSIOJUT pur [e]SOD + - - = ~ = = (+) soorjins y10q 10 (—) 990RJ.IMs [eIxege A[UO UO e}PUIOIS siajdojuopoinan sual ding Stajdoyjajvanan sidajdyjv20Anan S11a]d0Anauosrvy si4ajdoinan. si4ajdoauiaav T SE ee ea ee Se ae ‘(1 9192.1 *Z66T) 21NYS 2 Jea[D Woy popuedxg -1aded sty} ul pod eIDUIS-WIOF dy] DsOUSEIP 0} pasn siajJoRIeYO Ie[NONO pue [RoIsojoydiow ssois oy], Tage. 10 — cS CHINES INS TPL Wel) SIONS SS Cee ora, we Za nefeman' GSS Fig. 5 Laveineopteris rarinervis (Bunbury) Cleal et al. Copy of Carpentier (1930: fig. 1), showing orbicular cyclopterids attached near the dichotomy of the primary rachis. Here reproduced at x 0-7 life size. with the Trigonocarpales (Oestry-Stidd 1979). Little is known of the fructifications, other than that large ovules were attached directly to the frond, probably at the end of ultimate pinnae (Kidston 1904). Form-genus MACRONEUROPTERIS Cleal, Shute & Zodrow (1990: 488) Type. M. macrophylla (Brongniart) Cleal, Shute & Zodrow COMMENTS. Most neuropteroid fronds have a dichotomy of the primary rachis producing tripinnate or occasionally quad- ripinnate branches. In some species, however, the dichotomy of the primary rachis produces less-divided, essentially bipin- nate, primary rachis branches (Figs 6-7). These species also have a number of distinctive epidermal characteristics, such as brachyparacytic or cyclocytic stomata (Fig. 17E,F). It was for this distinctive group of species that Cleal et al. (1991) proposed the form-genus Macroneuropteris. The fronds of Macroneuropteris are very similar to Neurop- teris sensu stricto, except that they are less divided. Of particular significance is the presence in at least one macro- neuropterid species (M. scheuchzeri) of so-called ‘Odontopt- C.J. CLEAL AND C.H. SHUTE | ~ eris lindleyana’- type pinnules (e.g. Crookall 1959: pl. 57, fig. 1), which can be compared with laciniate pinnules in the lower part of true neuropterid fronds (e.g. Stockmans 1933: pl. 11 fig. 1; pl. 12 fig. 2; Zodrow & Cleal 1988: pl. 4 fig. 3). j There is no evidence of the orbicular cyclopterid pinnules of | Laveineopteris or Margaritopteris. Nothing is known of the fructifications. Beeler (1983) claimed that the rachis anatomy is of a type typical of the Trigonocarpales. However, this was based purely on evi- dence of association; she could find no such rachides with macroneuropterid pinnules directly attached. Form-genus MARGARITOPTERIS Gothan (1913: 168) Type. M. coemansii (Andra) Gothan COMMENTS. Most species included in this form-genus have broadly attached and/or lobed pinnules, and prior to Goth- an’s protologue were assigned to Odontopteris (Brongniart) ~— ial ee i i H Sternberg, 1825 or Sphenopteris (Brongniart) Sternberg, 1825 | (see Laveine et al. 1977). However, one species, originally | included in Neuropteris also belongs here (‘N.’ multivenosa | Purkynova). Laveine et al. (1977) have shown that it is almost | certainly the ancestral form of Callipteridium. Nothing 1s| known of the fructifications or stem/rachis anatomy. Form-genus NEURALETHOPTERIS Cremer ex Laveine (1967: 97) TyPE. N. schlehanii (Stur) Laveine COMMENTS. This form-genus is used for alethopterid-like fronds, in which the pinnules have a constricted base (Fig. 8). Most of its component species were originally described as neuropterids, but they in fact have little to do with that form-genus in its currently defined sense. The taxonomy of the form-genus has been thoroughly discussed by Laveine (1967), and need not be repeated. Our only disagreement with his analysis concerns the authorship of the taxon. Laveine quotes Cremer (1893), but this is a] thesis that was not effectively published. Wagner (1963, 1965) suggested that the name should be resurrected, but provided neither a diagnosis nor type. The first validly published diagnosis is in fact in Laveine’s study, who must therefore be taken as the author of the genus. The architecture of the frond has been established with reasonable certainty by Laveine et al. (1992). As with most of the trigonocarpaleans, the frond had a dichotomy of the primary rachis producing two tripinnate primary rachis branches. Most significantly, there appear to be no interca- lated elements on the primary rachis branches between the secondary pinnae. The lack of this feature separates Neur- alethopteris from most of the other neuropteroid fronds and helps confirm that its affinities lies closest with the aletho- | pterids. There have been a number of reports of sporangial organs attached or closely associated with neuralethopterid fronds (Dix 1932, 1933; Arnold 1949; Jongmans 1954; Stockmans & Williére 1961; Laveine 1967). Dix and Arnold both referred them to the form-genus Aulacotheca, but Jongmans identified them as Whittleseya. By studying a range of specimens from a single locality, Stockmans & Williére concluded that this apparent taxonomic difference in fact reflected infraspecific ' NEUROPTEROID FOLIAGE FROM CARBONIFEROUS AND LOWER PERMIAN 11 } ‘ig. 6 = Macroneuropteris macrophylla (Brongniart) Cleal et al. Specimen showing lower part of frond. V.2970. Westphalian D, Radstock, Somerset, UK. Natural size. | 12 ‘WNEUROPTEROID FOLIAGE FROM CARBONIFEROUS AND LOWER PERMIAN 23 ‘REASON FOR GENERIC ASSIGNMENT. Epidermal structure ‘(Barthel 1962, Cleal 1985). Also, pinnate fragments of this \species are almost invariably associated with orbicular cyclop- terid pinnules. (COMMENTS. This species has been widely reported from the ‘Iberian Peninsula. However, Cleal (1981) analysed these records and showed that they were based either on specimens lof Neuropteris resobae Cleal (q.v.), or on unidentifiable ‘fragments; L. tenuifolia would seem to be absent from this area. ‘OCCURRENCE. South-West UK (Lan-WeD), Pennines (Lan- Bol), Franco-Belgian Basin (Lan-WeD), S. Limburg (Lan), NW Germany (Lan-WeD), Lublin (Lan-Bol), Intra-Sudetic Basin (Duc-Bol), U. Silesia (Lan-Bol), Saar-Lorraine (Bol), Svoge (Duc-Bol), Donets (Duc-WeD). | Form-genus MACRONEUROPTERIS Cleal, Shute & Zodrow Macroneuropteris britannica (Gutbier) Cleal, Shute & _ Zodrow * 1835 Odontopteris britannica Gutbier: 68, pl. 9, figs 8-11. * 1990 Macroneuropteris britannica (Gutbier) Cleal, Shute & Zodrow: 488. REASON FOR GENERIC ASSIGNMENT. Epidermal structure (Barthel 1962). OCCURRENCE. Zwickau-Oelsnitz (WeD). Vacroneuropteris macrophylla (Brongniart) Cleal, Shute & Zodrow Figs 6-7 17E-F 1831 Nevropteris macrophylla Brongniart: 235; pl. 65, fig. | . * 1990 Macroneuropteris macrophylla (Brongniart) Cleal, . Shute & Zodrow: 488. k REASON FOR GENERIC ASSIGNMENT. Type species. COMMENTS. Cuticles have been described by Cleal & Zod- “ow (1989). The frond architecture is currently under review py Cleal, Laveine & Shute. | The specimens from North Caucasus illustrated by Anisi- nova (1979) as this species are clearly misidentified. They are solated pinnules which resemble those of Paripteris (e.g. P. seudogigantea), although they would seem to have origi- nated from rather a high stratigraphical position (WeD) for hat form-genus. ICCURRENCE. South-West UK (WeD-Can) Macroneuropteris scheuchzeri (Hoffmann) Cleal, Shute & Zodrow 1827 Neuropteris scheuchzeri Hoffmann: 157; pl. 1b, figs 144. 1990 Macroneuropteris scheuchzeri (Hoffmann) Cleal, Shute & Zodrow: 488. | REASON FOR GENERIC ASSIGNMENT. Epidermal structure ‘Barthel 1961, Cleal & Zodrow 1989). The frond architecture S currently under review by Cleal & Laveine. i= Havlena (1953: pl. 5, fig. 3) figured a fragment | reputedly from the Stephanian C of the Intra-Sudetic Basin as Neuropteris cordata. He claimed similar material also occurred in the Autunian of this region. As pointed out by Laveine (1967), however, the figured specimen is almost certainly M. scheuchzeri. If it does belong there and its stated provenance is correct, this is by far the highest stratigraphical occurrence of this species in Europe. OCCURRENCE. South-West UK (Duc-Can), Pennines (Lan- Bol), Franco-Belgian Basin (Duc-WeD), S. Limburg (Duc), NW Germany (Duc-WeD), NE Germany (Duc-Bol), Lublin (Bol), Intra-Sudetic Basin (Duc-WeD, ?StC-Aut), U. Silesia (Lan-Bol), Saar-Lorraine (Bol), Alps (Can), Svoge (Bol), Donets (Duc-WeD), N. Caucasus (WeD), Turkey (WeD), N. Portugal (WeD), NW Spain (WeD-Can), S. Portugal (WeD). Macroneuropteris subauriculata (Sterzel) Cleal, Shute & Zodrow T 1855 Neuropteris auriculata Brongniart: Geinitz: pl. 27, figs 4-7, 9. 1901 Neuropteris subauriculata Sterzel: 100. 1990 Macroneuropteris subauriculata (Sterzel) Cleal, Shute & Zodrow: 488. * § REASON FOR GENERIC ASSIGNMENT. Epidermal structure (Barthel 1962) and possibly frond architecture (Daber 1957). COMMENTS. This species appears to be endemic to the Zwickau-Oelsnitz Basin. The record from North Caucasus by Anisimova (1979) is based on extremely poorly preserved material and is unconvincing. Laveine (1989: pl. 60, fig. 1) has figured a single fragment of this species from the upper Westphalian D (or possibly basal Cantabrian) of Saar-Lorraine. It bears a close similarity to the fragmentary types of Neuropteris germeri de Jong, from slightly older strata in Saar-Lorraine, and which we have provisionally assigned to N. ovata. Cuticles from this Saar-Lorraine material could help resolve their taxonomic position. OCCURRENCE. Zwickau-Oelsnitz (WeD). Form-genus MARGARITOPTERIS Gothan Margaritopteris multivenosa (Purkynova) Cleal & Shute, comb. nov. * 1970 Neuropteris multivenosa Purkynova: 223-224; pl. 45, fig. 1, pl. 46, fig. 1. REASON FOR GENERIC ASSIGNMENT. The prominent midvein and the size and texture of the pinnules (see Laveine et al. 1977). Also the manner of lobing of the pinnules. COMMENTS. Laveine et al. (1977) were clearly of the opinion that this species belongs to Margaritopteris, and was the precurssor of the more familiar Westphalian species, although no formal proposal of transference was made. The factors mentioned above, particularly well shown in Laveine et al. (1977, pl. 19, fig. 3), make it unnecessary to postpone the proposal of transference. OCCURRENCE. U. Silesia (Alp). 24 Form-genus NEURALETHOPTERIS Cremer ex Laveine Neuralethopteris densifolia Josten * 1983 Neuralethopteris densifolia Josten: 144; pl. 53, fig. 1; pl. 54, fig. 1. REASON FOR GENERIC ASSIGNMENT. The alethopteroid nature of the venation. OCCURRENCE. NW Germany (Kin- Yea). Neuralethopteris doubravica (Purkynova) Cleal & Shute, comb. nov. * 1971 Neuropteris doubravica Purkynova: 165-166; pls 6-9. REASON FOR GENERIC ASSIGNMENT. The cordate base of the pinnules and the prominent midvein. COMMENTS. The generic position of this species is far from certain. At least some of the specimens figured in the protologue, particularly those with smaller pinnules, approach Neuropteris in venation and pinnule shape (e.g. pl. 8, fig. 1a). As pointed out by Purkynova, however, the larger pinnules share many characters with Neuralethopteris, espe- cially N. jongmansii, and so we propose to transfer the species there. OCCURRENCE. U. Silesia (Lan). Neuralethopteris jongmansii Laveine * 1967 Neuralethopteris jongmansii Laveine: 107; pls 2-4. REASON FOR GENERIC ASSIGNMENT. The typically alethop- terid nature of venation (Laveine 1967). Also, Whittleseya sporangial structures and Trigonocarpus ovules have been linked by Jongmans (1954) to foliage identified as N. jong- mansii by Laveine (1967). OCCURRENCE. South-West UK (Lan), Pennines (Lan), Franco-Belgian Basin (Lan), S. Limburg (Lan), NW Ger- many (Lan), U. Silesia (Lan). Neuralethopteris larischii (Susta) Laveine ‘ 1930 Neuropteris Larischi Susta: 5, pl.1. ’ 1967 Neuralethopteris larischi (Susta) Laveine: 102; pl. 1. REASONS FOR GENERIC ASSIGNMENT. Typically alethopterid nature of venation (Laveine 1967). OCCURRENCE. Franco-Belgian Basin (Arn-Lan), NW Ger- many (Arn-Lan), U. Silesia (Kin-Lan), N. Caucasus (?Kin- ?Yea), Turkey (Yea), NW Spain (Lan). Neuralethopteris neuropteroides (Susta) Josten * 1927 Alethopteris neuropteroides Susta: 4; pl. 1, fig. 2. * 1983 Neuralethopteris neuropteroides (Susta) Josten: 138; pl. 50, fig. 1. REASON FOR GENERIC ASSIGNMENT. The close similarity in pinnule shape to N. schlehanii, and the tendency of the pinnules to be fused to the rachis at the base. COMMENTS. This species seems to occupy a position interme- diate between Neuralethopteris and true Alethopteris. C.J. CLEAL AND C.H. SHUTE OCCURRENCE. Franco-Belgian Basin (Kin-Lan), NW Ger- many (Kin-Lan), U. Silesia (Yea-Lan). Neuralethopteris rectinervis (Kidston) Laveine *v 1888 T 1959 Neuropteris rectinervis Kidston: 314; pl.1, figs 2-4. Neuropteris Schlehani forma rectinervis (Kidston) Crookall: 145-147; pl. 35, figs 6-8. Neuralethopteris rectinervis (Kidston) Laveine: 120; pl. 9. 8 1967 REASON FOR GENERIC ASSIGNMENT. typically alethopterid nature of venation (Laveine 1967). OCCURRENCE. South-West UK (Lan), Pennines (Lan), Franco-Belgian Basin (Lan), NW Germany (Lan), U. Silesia (Lan), Donets (Lan). Neuralethopteris schlehanii (Stur) Laveine NST 1953 Neuropteris Schlehani Stur: 289; pl.28, figs 7-8. Neuropteris schlehanioides Stockmans & Williére: 233; pl. 31, figs 3, 7; pl. 36, fig. 2. Neuropteris loriformis Stockmans & Williére: 234; pl. 16, fig. 2. Neuropteris rectinervis forma obtusa Tenéov: 59-60; pl. 20, figs 3-4. Neuropteris lata Tenéov: 60; pl. 21, figs 2-3. Neuropteris longifolia Tenéov: 61; pl. 21, figs 4-9. oP IPSS) 1977 1977 1977 REASON FOR GENERIC ASSIGNMENT. Type species. COMMENTS. Epidermal structures have been described by Cleal & Shute (1992). Also, Aulacotheca sporangial struc- tures were reported in close association with it by Laveine (1967: pl.5, fig.3). The types of N. schlehanioides clearly represent fragments | from high in the pinna of N. schlehanii and can be compared with parts of Laveine (1967: pl. 6, figs 2-3). The type of N. | loriformis is less typical, having extremely long, slender pinnules. As pointed out by Stockmans & Williére (1953), however, it occurs in association with N. schlehanii and has a comparable venation pattern. Neuropteris longifolia and N. lata were erected for speci- mens from the Svidnaya Formation of the Svoge Basin. The pinnules are rather large (up to 30 mm long), but are otherwise very similar to N. schlehanii, with which they are closely associated. As Laveine (1967: pl. 8) has figured specimens of N. schlehanii with pinnules approaching these in size, there seems little reason for separating these species. Tenéov (1977) described the types of N. rectinervis forma obtusa as having a venation nearer to that of N. rectinervis than N. schlehanii. However, the veining is in fact quite compatible with N. schlehanii, being broadly arched (cf. Laveine 1967: pl. 7 fig. 1; pl. 8 fig. 4). OCCURRENCE. South-West UK (Lan), Pennines (Lan), Franco-Belgian Basin (Pen-Lan), S. Limburg (Lan), NW Germany (Mrd-Lan), NE Germany (Kin-Lan), (?)Lublin (Kin-Yea), Intra-Sudetic Basin (Lan), U. Silesia (Alp-Lan), Svoge (Yea-Lan), Donets (Pen-Lan), N. Caucasus (?Kin- Lan), Turkey (Lan), Alps (Lan), Pyrenees (Kin), NW Spain (Lan). Fig. 8 | NEUROPTEROID FOLIAGE FROM CARBONIFEROUS AND LOWER PERMIAN 25 Form-genus NEUROCALLIPTERIS Sterzel Neurocallipteris gallica (Zeiller) Cleal & Shute, comb. nov. * 1888a Neuropteris gallica Zeiller: 248; pl. 29, figs 1-3. REASON FOR GENERIC ASSIGNMENT. Close similarity to N. planchardii in both pinnule shape and venation. _ COMMENTS. This species is not well documented, and it is not entirely certain that it is distinct from N. planchardii. As pointed out by Wagner (1963), many of the differences _ claimed by Zeiller may be merely a function of the position of _the pinnules within a frond. The most obvious difference is _ the presence of hairs near the midvein of N. gallica, but this could be influenced by taphonomic factors. There have, however, been a number of records of the species in recent years, and so it has provisionally taken to be ‘good’. OCCURRENCE. Saar-Lorraine (Bar), Massif Central (StC), NW Spain (StB), S. Spain (StC), N. Portugal (StC). ' Neurocallipteris neuropteroides (G6ppert) Cleal, Shute & Zodrow Figs 9-10 : -* 1836 Gleichenites neuropteroides Goppert: 186; pls 4-5. * 1990 Neurocallipteris neuropteroides (Géppert) Cleal, Shute & Zodrow: 489. | REASON FOR GENERIC ASSIGNMENT. Type species. COMMENTS. Epidermal structure (Barthel 1962, 1976) and frond architecture (Barthel 1976, Setlik 1980) have been | documented for this species. OCCURRENCE. Saxony (Aut), Intra-Sudetic Basin (StB- | Aut), Massif Central (StC), Pyrenees (StC), NW Spain (StC), 'N. Portugal (StC-Aut). | Neurocallipteris planchardii (Zeiller) Cleal, Shute & Zodrow '* 1888a Neuropteris planchardii Zeiller: 246; pl. 28, figs 8-9. * 1990 Neurocallipteris planchardii (Zeiller) Cleal, Shute & Zodrow: 489. IREASON FOR GENERIC ASSIGNMENT. Similarity of epidermal structures to N. neuropteroides (see Reichel & Barthel 1964, Barthel 1976) OCCURRENCE. Saxony (Aut), Intra-Sudetic Basin (Aut), N. Caucasus (StC), Alps (?Can), Massif Central (StB-Aut), Pyrenees (StC), NW Spain (WeD-StB), N. Portugal (StC- Aut). Form-genus NEURODONTOPTERIS Potonié | } Veurodontopteris auriculata (Brongniart) Potonié Figs 11-12 ? 1830 Nevropteris auriculata Brongniart: pl. 36. 1831 Nevropteris auriculata Brongniart: 236. 1831 Nevropteris dufrenoyi Brongniart: 246. _ 1893 Neurodontopteris auriculata (Brongniart) Potonié: 124. .(?)1937 Neuropteris densinervosa (Grigoriev) Zalessky: 183; fig. 31. REASON FOR GENERIC ASSIGNMENT. Type species. COMMENTS. There have been no descriptions of cuticles under this species name. However, Z. Simunek (pers. comm. 1992) has shown that cuticles described by Barthel (1976) as Neuropteris cordata in fact belong to Neurodontopteris auriculata. In contrast to typical N. cordata from the Massif Central, the specimens which yielded the cuticles have smaller pinnules (<60 mm long) with a rounder apex, and a denser venation (30 veins per cm on the pinnule margin). Siminek has prepared very similar cuticles from specimens of N. auriculata from the Intra-Sudetic Basin, examples of which are shown in Fig. 11. The inclusion of N. densinervosa here must be regarded as tentative, since only two specimens have been illustrated in the literature and one of those (the holotype) only as a drawing. However, the large pinnules and high stratigraph- ical position (Gzhelian) of the specimens would seem com- patible with N. auriculata. OCCURRENCE. Saar-Lorraine (Aut), Saxony (Aut), Intra- Sudetic Basin (StB-Aut), NW Spain (StB), S. Spain (StC- Aut). Form-genus NEUROPTERIS (Brongniart) Sternberg Neuropteris antecedens Stur * 1875 Neuropteris antecedens Stur: 53; pl.15, figs 1-6. 1953 Neuropteris mathieui Stockmans & Williére: 227. 1955 Neuropteris condrusiana Stockmans & Williére: 12; pl. 6, figs 1-7, 9-15; pl. 9, figs 1-8. 1955 Neuropteris papilioniformis Stockmans & Williére: 13; pl. 2, fig. 3. 1955 Neuropteris pseudozamites Stockmans & Williére: 13, pl. 2, fig. 1. REASON FOR GENERIC ASSIGNMENT. The tendency of the pinnules to be broadly attached to the rachis and on the venation sometimes being flexuous (Crookall 1959). COMMENTS. The type of N. mathieui was found associated with typical specimens of N. antecedens, and it is difficult to see why it is not merely the small-pinnuled form of that species. The types of N. condrusiana, N. papilioniformis and N. pseudozamites all originated from the same horizon and locality, and were associated with a specimen which Stock- mans & Williére (1955) identified as their N. mathieui. All of this material clearly belongs to a single species, and bears quite a striking similarity to the types of N. antecedens; it in fact represents one of the best documentations in the litera- ture of the morphological variability of that species. OCCURRENCE. South-West UK (Arn), Pennines (Asb-Arn), Franco-Belgian Basin (Pnd-Arn), U. Silesia (Asb-Arn), Svoge (?Arn). Neuropteris bohdanowiczii (Zalessky) Gothan * 1907 Sphenopteris bohdanowiczi Zalessky: 33, 65; pl. 2, fig. 2. 26 8 1913 Neuropteris bohdanowiczi (Zalessky) Gothan: 210; pl. 44, fig. 4; pl. 53, fig. 1. REASON FOR GENERIC ASSIGNMENT. The relatively weakly developed midvein, the slightly flexuous lateral veins, and the tendency of the pinnules to be fused to the rachis. COMMENTS. This is a relatively poorly known species, the best documented records being by Gothan (1913) and Kota- sowa (1968). Zalessky (1907) suggested that it should be placed in a new subgenus, Sphenopteris subg. Neurospheno- pteris. However, it is doubtful if the latter is validly published according to the ICBN (Danzé 1956). OCCURRENCE. U. Silesia (Alp). Neuropteris ervedosensis (Teixeira) Wagner * 1942 Mixoneura ervedosensis Teixeira: 8; pl. 1. S 1963 Neuropteris ervedosensis (Teixeira) Wagner: 27. REASON FOR GENERIC ASSIGNMENT. The apparent occurence of enlarged, ‘forma impar’-like pinnules in the lower part of the frond. Also, the veining, although rather less dense, has some resemblance to that of N. ovata and N. flexuosa. COMMENTS. Opinion is divided as to the affinities of this species. De Jong (1974) regards it as being closely related to N. ovata, while Wagner & Sousa (1983) state that there is probably no relation with this species or N. flexuosa. As stated above, its affinities seem to lean towards Neuropteris, particularly as it probably has enlarged pinnules low in the frond, but this must be taken as provisional, at least until better information on the frond architecture becomes avail- able. OCCURRENCE. N. Portugal (WeD). Neuropteris flexuosa Sternberg T 1823 Osmunda gigantea, var. B Sternberg: pl.32, fig.2. * 1825 Neuropteris flexuosa Sternberg: xvi. T 1959 Neuropteris ovata Hoffmann, forma flexuosa (Stern- berg) Crookall: text-fig. 52. REASON FOR GENERIC ASSIGNMENT. Epidermal structures (Cleal & Zodrow 1989) and close similarity of pinnule form to N. ovata. COMMENTS. The European records of this species, other than those in Britain, are misidentifications of species such as Laveineopteris tenuifolia (e.g. Novik 1952, 1954) and L. jongmansii (e.g. Corsin 1932). OCCURRENCE. South-West UK (WeD-Can), S Portugal (WeD). Neuropteris ghayei Stockmans & Williére T 1933 Neuropteris grangeri Brongniart; Stockmans: pl.12, fig.3. * 1954 Neuropteris ghayei Stockmans & Williére in Pastiels & Williére: 59. REASON FOR GENERIC ASSIGNMENT. Underlying similarity in pinnule form with N. heterophylla and N. obliqua. Also, on the presence of large ‘impar’-type pinnules (Laveine 1967: pl.49, fig.5), similar to those found in the lower part of the N. obliqua frond. § C.J. CLEAL AND C.H. SHUTE OCCURRENCE. Franco-Belgian Basin (Lan), NW Germany (Lan) Neuropteris heterophylla (Brongniart) Sternberg 15-16 * 1822 Filicites (Nevropteris) heterophyllus Brongniart: 239; pl. 2, fig. 6. 1825 Neuropteris heterophylla (Brongniart) Sternberg: XVi. T 1831 Nevropteris (sic) heterophylla Brongniart: pl. 71 (neotype — vide Laveine, 1967). T 1967 Neuropteris heterophylla Brongniart (sic); Laveine: | pl. A. | Figs | REASON FOR GENERIC ASSIGNMENT. Type species. COMMENTS. Both epidermal structures and frond architec- | ture have been documented by Cleal & Shute (1991a). Specimens of this species have sometimes been recorded as _ Neuropteris grangeri Brongniart (see comments by Laveine |. 1967). The specimens from the Donets that have been illustrated | in the literature as N. heterophylla appear to be misidentified | specimens of Laveineopteris loshii (Novik 1952: pl. 61, figs 14; 1954: pl. 20, figs 5-6). However, Fissunenko and Laveine (1984) claim that true N. heterophylla occurs here and, in view of Laveine’s familiarity with the type specimens, | the record has been accepted. OCCURRENCE. Pennines (Duc), Franco-Belgian Basin (Lan- Bol), NW Germany (Lan-Bol), U. Silesia (Lan-Duc), Donets (Yea-WeD). Neuropteris obliqua (Brongniart) Zeiller Figs 13A—B, 14 T 1833 Pecopteris obliqua Brongniart: pl. 96, figs 14. * 1834 Pecopteris obliqua Brongniart: 320-321. ’ 1888b Neuropteris obliqua (Brongniart) Zeiller: 284-289. 1953a Neuropteris marginenervis Jongmans: 15; pl. 5, figs 29-30; pl. 6, figs 31-32 (vide Laveine, 1967). .v 1959 Neuropteris lanarkiana Kidston ex Crookall: 174; pl. 50, figs 1-2. T 1967 Neuropteris obliqua (Brongniart) Zeiller; Laveine: pl. E, figs 1-2. REASON FOR GENERIC ASSIGNMENT. Epidermal structure (Cleal & Shute 1992) and frond architecture (Fig. 13; Gotha 1953: fig. 7; Scott 1978: pl. 27, fig. 1). COMMENTS. The frond architecture of this species has still t be fully documented, but the available evidence suggests tha it was essentially similar to that of N. heterophylla, excep that the pinnules at the base of the frond were rather large (the so-called ‘forma impar’-type — e.g. Crookall (1959: pl.47 fig.4). Crookall (1959) distinguished N. lanarkiana from N. obli qua by its more slender, triangular and widely-spaced pin nules, and less dense and straighter veins. However, suc pinnules are not atypical in the N. obliqua fronds, and can b compared with a specimen figured by Laveine (1967: pl. 50 fig. la). Stockmans & Williére (1965: pl. 3, fig. 8; pl. 5, figs 1-2 document specimens from NW Spain as this species, an Laveine (1967) has agreed with the identity of at least some 0 NEUROPTEROID FOLIAGE FROM CARBONIFEROUS AND LOWER PERMIAN 27 them. However, the specimens are very fragmentary and, on their own, are inadequate for identification. There are no better documented specimens of this species from the Iberian Peninsula (the records by Wagner & Bowman 1983 and Alvarez- Vazquez in Wagner 1990 are unillustrated). OCCURRENCE. South-West UK (Lan-Duc), Pennines (Lan- Bol), Franco-Belgian Basin (Mrd-Bol), S. Limburg (Lan- Duc), NW Germany (Mrd-Bol), NE Germany (Yea-Bol), Lublin (Lan-Bol), U. Silesia (Yea-Bol), Intra-Sudetic Basin (Lan), Svoge (Lan), Donets (Yea-Bol), N. Caucasus (?Kin- Lan), Turkey (Yea-Duc), Alps (Lan-Alps), S. Spain (Duc). Neuropteris ovata Hoffmann Fig. 17A-B * 1826 Neuropteris ovata Hoffmann: 266. T 1827 Neuropteris ovata Hoffmann: pl. 1b, fig. 6 (vide Satzwedel, 1969). 1888a Nevropteris stipulata Zeiller: 255; pl. 29 fig. 5 (tenta- tively suggested by Wagner & Alvarez-Vazquez, 1991). ._p 1960 Neuropteris valdensis (Heer) Jongmans: 57; pl. 18, in. JOFe 'T 1969 Imparipteris ovata (Hoffmann) Gothan; Saltzwedel: pl. 24 figs 1-2. -.y 1973 Mixoneura polyneura Doubinger & Germer: 50-51; pl. 1, fig. 2. .) 1974 Neuropteris germeri de Jong: 58; pls 21-22. _.v 1975a Neuropteris pilosa Doubinger & Germer: 18; pl. 7, fig. 1. REASON FOR GENERIC ASSIGNMENT. Epidermal structures | (Barthel 1962, Cleal 1985, Cleal & Zodrow 1989) and frond architecture (Zodrow & Cleal 1988). COMMENTS. Cleal & Zodrow (1990) recognize varieties of this species, based partly on differences in epidermal struc- ture. Also, Wagner (1963) assigns most of the Stephanian ‘examples of the species to a separate variety (var. grandeuryi ' Wagner), based on the lateral veins being denser and less ‘oblique to the pinnule margin. While recognizing that these | Varieties almost certainly have some biological validity, they will not be separated in this analysis. The identity of upper Stephanian C specimens from north Portugal, described by Wagner & Sousa (1983) as Neuropteris ovata var. pseudovata Gothan & Sze, centers on one of the most contentious issues concerning the taxonomy of this species, viz. the difference between it and Neurocallipteris neuropteroides. There have been many analyses of this prob- ‘lem, the most detailed being by Zalessky (1909), Barthel (1976), Setlik (1980) and Wagner & Sousa (1983). Setlik and Wagner & Sousa have shown there are certain very subtle differences in pinnule form, but without the evidence of cuticles it is far from certain that they would be regarded as Sufficient justification for separating them as species, let alone in different form-genera. It would seem that two quite _\separate groups of trigonocarpaleans have developed analo- gously similar pinnule morphologies, perhaps in response to similar environmental pressures. Consequently, identifying specimens with this type of pinnule in the upper Stephanian, f epidermal characters are unknown, is very difficult, if not mpossible. Wagner & Sousa’s Portugese specimens have some of the characters (somewhat subtriangular pinnules, veins slightly oblique to pinnule margin) that tend to be commoner in N. neuropteroides than N. ovata. We have therefore provisionally transferred their specimens to N. neuropteroides. However, we recognize that this needs to be confirmed by epidermal evidence or, if this is impractical, by larger specimens showing the form of the intercalated pin- nules (cf. Setlik 1980). M. polyneura and N. pilosa were separated from N. ovata on minor characters of venation and surface detail, and their distinction cannot be maintained (Cleal 1985). N. germeri, which was described mainly on just two specimens, has a veining pattern and pinnule shape compatible with the larger forms of N. ovata, especially those of the var. sarana as described by Cleal & Zodrow (1989). The veining density is a little lower than is typical, but can probably be accommo- dated within the lower end of the range of variation. As the types of N. germeri are associated with more typical frag- ments of N. ovata var. sarana (Bertrand) Cleal & Zodrow, there seems little reason for distinguishing them taxonom- ically. The type of N. stipulata is strikingly similar to N. ovata in pinnule shape, being relatively squat, having a basiscopic auricle and a short midvein. If Zeiller’s drawing of the type of N. stipulata is accurate, the vein density is c.40 veins per cm, which is compatible with the form of N. ovata, normally found in the Stephanian, and known as var. grandeuryi Wagner. OCCURRENCE. South-West UK (WeD-Can), Pennines (WeD), Franco-Belgian Basin, (WeD), NW Germany (WeD), U. Silesia (WeD), Saar-Lorraine (WeD), Donets (?Bol-?Bar), N. Caucasus (WeD, StB), Turkey (WeD), Alps (WeD-StC), Massif Central (Bar-StB, ?StC), NW Spain (WeD-StB), S. Spain (StC). Neuropteris parvifolia Stockmans * 1933 Neuropteris parvifolia Stockmans: 28-29, pl. 8, figs 1-5. REASON FOR GENERIC ASSIGNMENT. Similarity of pinnule morphology and underlying venation pattern to N. obliqua (see Laveine 1967). OCCURRENCE. Franco-Belgian Basin (Duc-Bol), NW Ger- many (Duc-Bol), NE Germany (Duc-Bol), Lublin (Duc- Bol), Alps (Bol). Neuropteris plicata Sternberg * 1833 Neuropteris plicata Sternberg: 70; pl. 19, figs 1,3. REASON FOR GENERIC ASSIGNMENT. Close similarity of the pinnules to N. ovata. COMMENTS. This is so similar to N. ovata that it is far from certain that it is a distinct species. Although Setlik (1921) attempted an analysis of the morphological variation of the pinnules, the results were equivocal on this point. If they are the same species, then Sternberg’s species would take prior- ity. In view of the important palaeobotanical and biostrati- graphical role played by N. ovata, a more thorough analysis of N. plicata should be undertaken before any nomenclatural changes are proposed. OCCURRENCE. Intra-Sudetic Basin (WeD). 28 Neuropteris praeovata (Némejc) Cleal & Shute, comb. nov. * 1949 Mixoneura praeovata Némejc: 17-18; text fig. 2; pl. 4, figs 1-7. REASON FOR GENERIC ASSIGNMENT. Similarity of pinnule shape and venation to N. ovata. COMMENTS. As pointed out by Némeje, this clearly belongs to the group of neuropterids allied to N. ovata, from which it can be reliably distinguished only by the more prominent midvein. Also, it is the only known member of this group of species to occur below the Westphalian D. OCCURRENCE. Intra-Sudetic Basin (Bol). Neuropteris resobae Cleal * 1981 Neuropteris resobae Cleal: 79, pls 1-2. REASON FOR GENERIC ASSIGNMENT. Dense venation with relatively weak midvein. Pinnule limb often partly fused to rachis, and with basiscopic auricle. COMMENTS. Little is known of the frond architecture, other than that the ultimate pinnae are terminated by a single apical pinnule, and nothing of the epidermal structures. Its reten- tion in this form-genus is thus based entirely on pinnule morphology and venation, which clearly cannot be conclu- sive. The only other (albeit negative) piece of evidence is that, despite the size of the collection on which the species was described, not a single cyclopterid pinnule was found in association. OCCURRENCE. NW Spain (Duc-WeD). Neuropteris schaeferi Doubinger & Germer * 1975a Neuropteris schaeferi Doubinger & Germer: 10-11; pl. 4, fig. 1. REASON FOR GENERIC ASSIGNMENT. Similarity of epidermal structure to N. ovata (see Saltzwedel 1968). COMMENTS. This species is not really adequately docu- mented, as only one small specimen has been described in the literature. However, in view of the epidermal evidence presented by Saltzwedel (1968), it has been included in the present analysis. OCCURRENCE. Saar-Lorraine (StB). Neuropteris semireticulata Josten * 1962 Neuropteris semireticulata Josten: 39-40; pl.3, figs 2-5. REASON FOR GENERIC ASSIGNMENT. Similarity of pinnule morphology and underlying venation pattern to N. obliqua (see Josten 1962). OCCURRENCE. South-West UK (Duc-Bol), Pennines (Duc), Franco-Belgian Basin (Duc-Bol), NW Germany (Duc-Bol), Lublin (Bol). i Neuropteris willierei Laveine * 1967 Neuropteris willierei Laveine: 224-227; pl.62. C.J. CLEAL AND C.H. SHUTE | REASON FOR GENERIC ASSIGNMENT. It is almost indistin- guishable from N. parvifolia, differing only in having smaller, more broadly attached pinnules, and occurring stratigraphi- cally lower (Laveine 1967). OCCURRENCE. Franco-Belgian Basin (Lan-Duc), NW Ger- many (Lan-Duc). Form-genus PARIPTERIS Gothan Paripteris gigantea (Sternberg) Gothan * 1821 Osmunda gigantea Sternberg: 33; pl.22. ‘1941 Paripteris gigantea (Sternberg) Gothan: 427. T 1953 Neuropteris gigantea Sternberg; Havlena: pl. 4; pl. 5), he, 2. .v 1959 Neuropteris maltbyensis Crookall: 164; pl. 33, figs 7-8. ? 1965 Paripteris veeni Stockmans & Williére: pl. 2, figs 4-7 (vide Wagner & Bowman 1983). REASON FOR GENERIC ASSIGNMENT. Type species. OCCURRENCE. South-West UK (Lan-Duc), Pennines (Lan- Duc), Franco-Belgian Basin, (Mrd-Duc), NW Germany (Mrd-Bol), NE Germany (Kin-Lan), Lublin (Kin-Duc), Intra-Sudetic Basin (Lan-Bol), U. Silesia (Mrd-Bol), Svoge (Yea-Lan), Donets (Kin-Duc), N. Caucasus (?Kin-Lan), Turkey (Lan), Alps (Lan), Pyrenees (Kin), NW Spain (Mrd, Lan), S. Spain (Lan). Paripteris linguaefolia (Bertrand) Laveine * 1930 Neuropteris linguaefolia Bertrand: 31-32; pl. 15. ’ 1967 Paripteris linguaefolia (Bertrand) Laveine: 266-267; pls 77-78. REASON FOR GENERIC ASSIGNMENT. Frond architecture. OCCURRENCE. Franco-Belgian Basin (Duc-Bol), NW Ger- many, (Bol), Lublin (Duc-Bol), Intra-Sudetic Basin (Duc- Bol), Saar-Lorraine (Duc-Bol), Donets (Duc-WeD), Turkey (Duc), Alps (Duc-Bol), NW Spain (Duc), S. Spain (Duc). Paripteris linguaenova (Bertrand) Cleal & Shute, comb. nov. * 1930 Neuropteris linguaenova Bertrand: 29; pls 13-14. eee — me tad ie er fe, ee , REASON FOR GENERIC ASSIGNMENT. Similarity of pinnule shape and venation to P. linguaefolia. COMMENTS. Laveine (1967) assigned the types of this species to P. pseudogigantea. However, they are significantly larger (many are longer than 4 cm), have a thinner midvein, and lateral veins that diverge from the midvein at a narrower angle. More problematic is its distinction from P. linguae- folia. Bertrand (1930) separated them because P. linguaefolia has virtually no midvein, and ‘la disposition et l’aspect des nervures sont trés différents. . .". As the types of P. ling- uaenova occur within the stratigraphical range of P. linguae- folia, these differences really need to be re-examined more critically. For the time being, however, the separation has been retained. OCCURRENCE. Saar-Lorraine (Bol). SS NEUROPTEROID FOLIAGE FROM CARBONIFEROUS AND LOWER PERMIAN 29 Paripteris pseudogigantea (Potonié) Gothan Fig. 13C-D * 1897 Neuropteris pseudogigantea Potonié: 113; text fig. 102. 1941 Neuropteris scheuchzeri forma minor Novik: 457; pl. Dips 5 7h * 1953 Paripteris pseudogigantea (Potonié) Gothan: 63-64; pl. 37, figs 1-4. _ REASON FOR GENERIC ASSIGNMENT. Frond architecture (Laveine 1967: pls 73-76). : / OCCURRENCE. South-West UK (Duc-Bol), Pennines (Duc- _ Bol), Franco-Belgian Basin (Duc-Bol), NW Germany (Duc- Bol), NE Germany (Duc-Bol), Lublin (Duc-Bol), Saar- | Lorraine (Duc-Bol), Donets (Duc-Bol), NW Spain (Duc). Paripteris schuetzei (Potonié) Daber : | * 1903 Neuropteris schiitzei Potonié: 399. T 1912 Neuropteris schiitzei Potonié: 122; fig. 84. _* 1963b Paripteris schiitzei (Potonié) Daber: 1212, fig. 2. . | REASON FOR GENERIC ASSIGNMENT. Frond architecture and » general aspect of pinnules. } OCCURRENCE. S. Limburg (Lan), U. Silesia (Duc). Form-genus SPHENONEUROPTERIS Shchegolev * Sphenoneuropteris brongniartii Shchegolev ) 1979 Sphenoneuropteris brongniartii Shchegolev: 159; pl. 53, fig. 2. | REASON FOR GENERIC ASSIGNMENT. General similarity in | -pinnule morphology and venation with type species. _ COMMENTS. The distinction between this and S. elegans is far ) from clear, and there must be a strong likelihood that they . are synonyms. | OCCURRENCE. N. Caucasus (StC). | Sphenoneuropteris dimorpha (Lesquereux) Cleal & Shute, comb. nov. IT 1879 Pseudopecopteris dimorpha Lesquereux: pl. 35, figs 1-6. 1880 Pseudopecopteris dimorpha Lesquereux: 201. 1978 Neuropteris dimorpha (Lesquereux) Boersma: 59; pl 8, tig. 3; pl. 12; figs 1-6. * REASON FOR GENERIC ASSIGNMENT. Mainly the venation (widely forking veins, oblique to pinnule margin, producing low vein density), and the large, lax-limbed pinnules. COMMENTS. Little is known of the frond architecture and nothing of the epidermal structure of this species. However, the pinnules show a remarkable similarity, especially in their venation (e.g. Doubinger & Germer 1975b, pl. 4), to the ypes of Sphenoneuropteris. This species is often thought to have characteristically leeply-lobed pinnules (e.g. Wagner 1958). However, 30ersma (1978) showed that this was at least partially a onsequence of the thin limb of the pinnules, which rarely lay lat in the matrix, and would undulate in and out of the plane long which the fossil was split. It is unlikely that this can explain all specimens with undulate margins, but the remnant examples may simply be from the distal regions of pinnae, where pinnules are in transition to ultimate pinnae. This species was initially assigned to Pseudopecopteris Lesquereux, 1880. However, this form-genus was not typified and included within it was a variety of disperate types of frond; it is thus a nomen dubium, and cannot be used as an alternative name for Sphenoneuropteris. OCCURRENCE. Saar-Lorraine (Bol-StB). Sphenoneuropteris elegans Shchegolev * 1979 Sphenoneuropteris elegans Shchegolev: 158; pl. 54, figs 1,2. REASON FOR GENERIC ASSIGNMENT. Type species. OCCURRENCE. N. Caucasus (StC). Sphenoneuropteris nemejciana (Purkynova) Cleal & Shute, comb. nov. * 1971 Neuropteris nemejciana Purkynova: 168; pls 10-11. 1971 Neuropteris venceslai Purkynova: 171; pl. 12. REASON FOR GENERIC ASSIGNMENT. Similarity of pinnule form and venation to S. dimorpha. COMMENTS. Purkynova’s specimens occur stratigraphically lower than any of the other species included in Spheno- neuropteris. However, it has many of the characteristic gross morphological features of that form-genus, including large pinnules (30-35 mm long) with a lax limb and somewhat undulate margin, and a low vein density (16 veins per cm on pinnule margin). The type and only known specimen of N. venceslai ori- ginated from the same locality and horizon as the types of S. nemejciana. It has similarly large, relatively thin-limbed pin- nules, thin midvein, and low vein density; compare for instance the specimen figured by Purkyfova on her pl. 11, fig. 2. The pinnules have a more acute apex, and are marginally larger, but not execssively so; the largest recorded pinnule of S. nemejciana is 5.5 cm long, as opposed to 7.5 cm in N. venceslat. All in all, there seems little reason to regard these as separate species. OCCURRENCE. U. Silesia (Lan). Sphenoneuropteris praedentata (Gothan) Cleal & Shute, comb. nov. * 1909 Neuropteris praedentata Gothan: figs 1,2. REASON FOR GENERIC ASSIGNMENT. Based mainly on vena- tion (widely forking veins, oblique to pinnule margin, pro- ducing low vein density), and the relatively large, subtriangular pinnules. COMMENTS. The general aspect of the pinnules, particularly the venation, seems to exclude this from Neuropteris as it is interpreted in this work. The venation seems to fit in far better with that given in the diagnosis of Sphenoneuropteris given by Shchegolev (1979). It is recognized that this is far from a satisfactory basis for recognizing ‘natural’ form- genera. However, until cuticle and frond architecture data become available, Shchegolev’s form-genus provides a conve- nient repository for this species. 30 The only large specimens of this species to have been published are in Zeiller (1888a: pl. 26) and Zeiller (1906: pl. 26), both under the name Neuropteris crenulata Brongniart. They both show bipinnate frond fragments, with intercalated pinnules on the penultimate rachis. Laveine (1967: text-fig. 6d) interprets the 1906 specimen as essentially a pinnate frond. However, the penultimate rachis in the 1888 specimen is noticeably curved, suggesting that it might be from a bipartite frond, similar to that present in many of the other neuropteroid form-genera. The numerous records of this species from the Iberian Peninsula have been analysed by Knight (1983). He has concluded that, although they show some similarity to S. praedentata, they differ in having smaller, thinner-limbed pinnules with weaker crenulations on the margin, and thinner veins. They have since been transferred to a separate species, S. wagneri (see below). Significantly, Knight also observed that the Spanish material shared some features in common with S. dimorpha, providing some support for the idea that S. dimorpha, S. praedentata and S. wagneri cluster together to form a reasonably natural form-genus. OCCURRENCE. (?)Saar-Lorraine (Bar), Massif Central (Bar- St@)? Sphenoneuropteris wagneri (Lorenzo) Cleal & Shute, comb. nov. * 1980 Mixoneura wagneri Lorenzo: 11-13; pl. 1. REASON FOR GENERIC ASSIGNMENT. The large, relatively lax pinnules with a wide venation. COMMENTS. This species was established for the Spanish specimens that were traditionally assigned to “Neuropteris’ praedentata (see comments on previous species). OCCURRENCE. NW Spain (Bar-Aut). Species of uncertain taxonomic position Included here are those species which, although clearly circumscribed and thus ‘good’, cannot be readily assigned to any of the above form-genera. Cuticular evidence is lacking, and their pinnule and pinna morphologies do not provide any obvious comparison with one or other of the more completely known species. Neuropteris bourozii Laveine * 1967 Neuropteris bourozii Laveine: 152; pls 23-25. COMMENTS. Some of the pinnules of this species show simil- arities to Laveineopteris (Laveine 1967: pl.24, fig.5), while others are of a more typical neuropterid-type (Ibid. pl.23, fig.5). Laveine (1967) assigned specimens from the Pennines Basin figured by Bolton (1926: pl. 6) to this species, but they almost certainly belong to L. tenuifolia. OCCURRENCE. Franco-Belgian Basin (Duc), NW Germany (Duc). Neuropteris cordata Brongniart * 1831 Neuropteris cordata Brongniart: 229; pl. 64. 1890 Nevropteris Raymondii Zeiller: 147; pl. 9a, fig. 4. C.J. CLEAL AND C.H. SHUTE 1893 Neuropteris pseudoblissii Potonié: 137. 1964 Mixoneura raymondii (Zeiller) Wagner: 9. COMMENTS. This species cannot readily be fitted into any of the other form-genera. The general aspect of the pinnules suggests affinities with Neurocallipteris or possibly even Neu- ropteris, but what little is known of the frond architecture (e.g. Langiaux 1984: fig. 111) would seem to separate it from both genera. The species is in clear need of a revision. The type and only known specimen of N. raymondii Zeiller (Mixoneura raymondii (Zeiller) Wagner) was figured photo- graphically by Doubinger (1956: pl. 12, fig. 3; pl. 13, fig. 1). It originated from the Mont Pel Formation in the Autun-Epinac Basin, and occurs together with specimens of N. cordata. The pinnules are rather smaller (c.16 mm long) than is typical for N. cordata but the venation is very similar. Doubinger (1956) claims that the veining density is higher in N. raymondii, but the measured value of 22 veins per cm on the pinnule margin | is quite compatible with some of the smaller forms of N. cordata (cf. Zeiller 1906: pl. 27, fig. 3). In view of the | evidence of association and of the similarity of the venation, it seems reasonable to assume that N. raymondii is merely a | small-pinnuled form of N. cordata, possibly from the more distal regions of the frond. N. pseudoblissii is still being recorded in the modern literature for specimens from the upper Stephanian with very elongate pinnules, but which are otherwise very close to N. cordata. Zeiller (1888a), who figured the types of Potonié’s species under the incorrect name Neuropteris blissii, noted that isolated fragments would be difficult to distinguish, and it is also significant that the two species almost invariably occur together (e.g. see records in Doubinger 1956). There thus seems little justification for separating the two species. The single specimen from the Duckmantian of the Pen- nines figured by Crookall (1959: pl. 41, fig. 5) as N. pseudo- blissii is an indeterminable fragment, possibly of a mariopterid. OCCURRENCE. Massif Central (StB-StC), Pyrenees (StC), Alps (StC), N. Portugal (StC). Neuropteris duprei Laveine * 1967 Neuropteris duprei Laveine: 164, pl.29, figs 14. COMMENTS. This is a very distinctive species with elongate, often asymmetrical pinnules and very oblique lateral veins. The only other similar material reported from Europe are th specimens described by Némejc (1949, pl.1, figs 1-8) “4 Odontopteris stradonicensis Andra. There is also a recor from the Langsettian of NW Spain (Wagner & bona 1983), but it is not illustrated. OCCURRENCE. Franco-Belgian Basin (Lan-Duc). Neuropteris dussartii Laveine * 1967 Neuropteris dussartii Laveine: 191; pl.48. COMMENTS. Laveine argued that this species shared a num ber of features in common with laveineopterid species such a L. loshii and L. rarinervis. However, Laveine also pointe out certain similarities with Neuropteris ovata, such as th presence of a basiscopic auricle on some of the pinnules. OCCURRENCE. South-West UK (Bol-WeD), Franco-Belgia Basin (WeD), NW Germany (WeD). NEUROPTEROID FOLIAGE FROM CARBONIFEROUS AND LOWER PERMIAN 31 Neuropteris teberdensis Shchegolev * 1979 Neuropteris teberdensis Shchegolev: 163; pl. 51; nl. Sa ike, WS pol, S3i5 104, Il, COMMENTS. The pinnules of this very late species show a _ marked resemblance to Neuropteris ovata, suggesting that it is a true neuropterid. However, one of the specimens (Shche- golev 1979: pl. 52, fig. 1) suggests that the frond might have been only bipinnately divided, with intercalated pinnules on the primary rachis branches. This fact, together with its high Stratigraphical position, suggests that the species may instead - belong to Neurodontopteris. _ OCCURRENCE. N. Caucasus (StC). Neuropteris zeilleri de Lima _T 1864 Neuropteris cordata Brongniart; Géppert: 100; pl. 11, figs 1-2. * 1890 Neuropteris zeilleri de Lima: 140. COMMENTS. This species has been widely quoted in the literature as occurring in the upper Stephanian of Europe (e.g. Havlena 1953, Doubinger 1956, Wagner 1963, Vetter 1968, Wagner & Sousa 1983). As pointed out by Zeiller /(1906) and Vetter (1968), however, there are problems with ithe typification of the species; that quoted above is the one ‘normally accepted, but it is far from clear if de Lima regarded Goppert’s specimens or his own Portugese specimens as types. The distinction from Neuropteris cordata is also far ‘from clear and according to Zeiller is based mainly on the fact that there is not a single midvein, but a number of separate, fine veins lying along the long axis of the pinnules. This distinction has never been properly documented and there must be a strong suspicion that it is purely taphonomic. Whatever the outcome, however, there can be little doubt hat N. zeilleri will end up in the same form-genus as N. vordata, whatever that will prove to be (see above). JCCURRENCE. (?) Intra-Sudetic Basin (Aut), Massif Central 2StB, StC-Aut), NW Spain (StB, ?StC), N. Portugal (StC- Aut). Nomina dubia he first group of species included here were initially escribed on just one or two fragments and additional jaterial has not been published. There is thus insufficient vidence of morphological variation to be able to recognize ie species reliably, or of features such as frond architecture ‘epidermal structure, by which their generic position could = assertained. They are listed below without further com- ent. "paripteris flabellinervis Gothan, 1953: 59; pl. 9, figs 2-3; pl. 28, fig. 2; pl. 30, fig. 6. europteris asturiana Jongmans MS ex Wagner, 1962: 757 {nomen nudum]. 2uropteris beveridgei Crookall, 1959: 189, pl. 40, fig. 4. puropteris bulupalganensis Zalessky in Zalessky & Chirkova, 1933: 9; fig. 1. ?uropteris(?) delasii Zeiller, 1892: 45; pl. 8, fig. 6. "uropteris dispar Zeiller, 1888a: 253; pl. 29, fig. 6. ‘uropteris horrida Zeiller, 1888a: 251; pl. 32, figs 1-2. | | Neuropteris jugosa Kidston ex Crookall, 1959: 164; pl. 41, fig. 3 Neuropteris matheronii Zeiller, 1888a: 245; pl. 28, fig. 7. Neuropteris pseudoimpar Stockmans & Williére, 1953: 235: pl. 44, fig. 2; pl. 50, fig. 12. Neuropteris squarrosaeformis Kidston ex Crookall, 1959: 163: pl.50, fig.6. Neuropteris subsessilis Stockmans & Williére, 1955: 14; pl. 8, fig. 1. Neuropteris waltonii Stockmans & Williére, 1953: 227-228. In addition to the above, there are a number of other species, for which more specimens are known, but which are still impossible at present to identify reliably. These require further comment. Mixoneura muensterifolia Némejc * 1949 Mixoneura muensterifolia Némejc: 15-16; pl. 3, figs 10-14. 1949 Mixoneura grandifolia Némejc: 18-20; text fig. 4. COMMENTS. Némejc established this species for a number of fragments from the middle Westphalian, that were claimed to have a pinnule shape similar to Neuropteris obliqua, but with more flexuous veins. These are similar to the characters used to define N. semireticulata, of which it would be an earlier synonym. However, the illustrations used by Némeje are poor and the specimens fragmentary. It would thus be unwise to give it priority over N. semireticulata, at least until Ném- ejc’s species is better documented. Némejec reported larger pinnules in close association with M. muensterifolia, and used them as the types of another new species, M. grandifolia. However, the figured specimens would seem to correspond with forma impar-type pinnules found in the proximal parts of the fronds of the N. obliqua group. It is thus almost certain that they are conspecific with the specimens that he assigned to M. muensterifolia. Neuropteris arberi Crookall * 1959 Neuropteris arberi Crookall: 148; pl. 50, fig. 7; pl. 51, figs 2-4. COMMENTS. Based on three fragments, none of which show details of the apical pinnules or the pattern of lobing. Their affinities may be more mariopterid than neuropterid. Remy & Remy (1975) attempted to use this species for German specimens, but it is difficult to see how this can be justified in the light of the extremely imperfect types. Neuropteris kosmannii Potonié * 1903 Neuropteris kosmanni Potonié: 399. T 1913 Neuropteris kosmanni Potonié; Gothan: pl. 47, fig. 3; pl. 50, figs 14. COMMENTS. Although this species periodically re-appears in the literature (e.g. Kotasowa 1968), it has only ever been described from small fragments. They all show vaulted, extremely thick-limbed pinnules, often with a somewhat undulate margin, quite atypical for any of the neuropteroid form-genera, with the possible exception of Margaritopteris. In the absence of more complete material, it is impossible either to give it a useful circumscription as a species, or to determine their generic position. 32 Neuropteris lubnensis Havlena * 1953 Neuropteris lubnensis Havlena: 153-154; pl. 6, figs 1-2. COMMENTS. This is based on forty-eight specimens preserved in a sandstone, although only two were figured. They are undoubtedly unusual, having very large pinnules (up to 4 cm long and 2 cm wide), and do not fit into any previously described species. However, the arenaceous matrix is far from perfect for preserving this type of fossil, and the number of specimens illustrated is inadequate to determine the range of morphological variation, let alone frond architecture. Much better material needs to be documented before any- thing can be done with this species. Neuropteris montana Heer * 1879 Neuropteris montana Heer: 22; pl. 6, figs 22, 23. T 1960 Neuropteris montana Heer; Jongmans: pl. 21, fig. 121. COMMENTS. From the form of the distal part of the pinna, the type clearly belongs to a paripinnate frond, presumably of the Potonieaceae. It is reputed to originate from the Cantabrian or lower Barruelian, which is far higher stratigraphically than Paripteris normally occurs. The venation is very poorly preserved, but may be anastomosed with very elongate vein-meshes. If so, then it may belong to Linopteris neu- ropteroides (Gutbier) Potonié, 1899, which sometimes occurs as high as Barruelian. However, the material is really inad- equate to give an unequivocal statement on this. ROBUSTNESS OF GENERIC TAXONOMY The statistics of this taxonomic analysis are summarized in Table 2 and Fig. 18. A total of 101 neuropteroid species have been recorded from Europe over the last half century, of which nearly a half (43-5%) are either unsatisfactory because they are based on insufficient material, or are later synonyms of other species. Of the remaining fifty-seven ‘good’ species, all but six (10-5%) can be assigned to one or other of the 9 form-genera summarized in the early part of this paper. Total good species (0.6) Inedequately circumscribed (0.2) Synonyms (0.3) Fig. 18 The robustness of the taxonomy of neuropteroid fronds. (a) The proportions of synonyms, inadequately described and ‘good’ species among all those neuropteroids used since 1940. (b) The proportion of the ‘good’ species belonging to each of the form-genera. Neurodontopteris (0.0) C.J. CLEAL AND C.H. SHUTE The six species that cannot yet be placed in our generic classification fall into three groups. 1. N. cordata and N. zeilleri appear closely related to each other and it is far from certain that they are not in fact conspecific. Although widely recorded from the Stephanian and Autunian of France and the Iberian Peninsula, little is known of the frond architecture and nothing of the cuticles (the cuticles assigned to N. cordata by Barthel, 1976, in fact belong to Neurodontopteris auriculata — see above). 2. N. duprei has unusual, asymmetrical pinnules unlike | any of the other species included in this analysis; in fact they | are different from any type of foliage previously assigned to | the trigonocarpaleans. It may well belong to a new form- genus, but details of the frond architecture and/or cuticles will be needed before any decision on this can be made. 3. From the general aspect of the pinnules, it is likely that N. bourozii, N. dussartii and N. teberdensis belong to either | Neuropteris, Laveineopteris or Neurocallipteris. Again, evi- | dence of frond architecture and/or cuticles will be needy | | | before a decision can be made on their classification. In conclusion, the analysis has allowed us to see where the main gaps are in our knowledge of these fossil fronds. In particular, the 6 species that cannot currently be assigned need to be further investigated. Nevertheless, we believe that | the results support the essential robustness of our generic} classification of neuropteroid fronds, and points to it being a| potentially useful tool for understanding more clearly the distribution of these plants. | DIVERSITY ANALYSIS Diversity of the neuropteroids as a whole As a by-product of this study, whose original goal was merely to ascertain the robustness of the generic classification of neuropteroid fronds, we have built up a database of the stratigraphical and geographical distribution of species within Europe. This would appear to invite further analysis of diversity variations. Diversity analysis has become a popular pursuit in recent years, but can be prone to serious problems| Paripteris (0.1) Sphenoneuropteris (0.1) | Neuropteris (0 3) Uncertain affinities (0.1) Laveineopteris (0.2) Neurocallipteris (0.1) Macroneuropteris (0.1) Margaritopteris (0.0) Neuralethopteris (0.1) NEUROPTEROID FOLIAGE FROM CARBONIFEROUS AND LOWER PERMIAN 33 Table 2 Statistics of neuropteroid taxonomy No. of Form-genera species % (Total) % (Good) Laveineopteris 9 8-91% 15-79% Macroneuropteris 4 3-96% 7:02% Margaritopteris 1 0-99% 1-75% Neuralethopteris 7 6:93% 12:28% Neurocallipteris 8) 2:97% 5:26% Neurodontopteris 1 0:99% 1:75% Neuropteris 15 14-85% 26-32% Paripteris 5 4-95% 8:77% Sphenoneuropteris 6 5-94% 10-53% - Uncertain affinities 6 5-94% 10-53% Inadequately circumscribed 17 16-83% Synonyms 27 26-73% Total ‘good’ species / 56-44% ' Total ‘bad’ species 44 43-56% Grand total 101 100-00% ) due at least in part to the tendency to use data trawled ' uncritically from the literature (cf. comments by Cleal 1988). Our database, although based only on a small range of species, at least has the merit of having been critically . compiled. | To this end, a tabulated set of statistics has been compiled to represent diversity, first-appearances and extinctions for each stage (Table 3). This has been done separately for each ‘of the form-genera, as well as for the group as a whole | (including those species unassignable to any of the form- genera). | The diversity of the group as a whole follows a fairly simple _pattern, showing a marked peak in the Westphalian, followed )by a rapid decline and then a subsidiary peak in the upper Stephanian (Fig. 19A). The Westphalian peak would seem to be confirmed by observations made by Boulter et al. (1988) on diversity changes in the wider plant adpression record for ithe palaeoequatorial belt. It almost certainly reflects varia- tions in the available non-marine strata in Europe; Niklas et al. (1980, p. 29) demonstrated that 98:5% of plant fossil diversity (at least between the Carboniferous and Jurassic) can be accounted for by this single factor. Numerical data on the available strata in each stage are not available for Europe. However, our observations would seem to confirm the gen- eral impression that delta-plain, fluvio-lacustrine deposits, which presumably reflect the habitats favoured by the plants yroducing these fronds, are at a maximum in the Westphalian ind upper Stephanian, with a low in the Cantabrian and, to {in extent, the Baruellian. | Figs 19B and 19C show the patterns of appearances and pxtinctions per stage, both corrected for variation in the ‘ength of the stage. These show curves with a similar double- eaked form to the diversity curve. A broad correlation ‘etween species turn-over and diversity is not surprising. fowever, the species profit/loss curve (Fig. 19D) shows a hore interesting pattern. Up to the Kinderscoutian the \ituation is relatively stable, but at higher stratigraphical pvels there are major fluctuations. Peaks occur in the sinderscoutian, Langsettian and Baruellian/Stephanian B. ‘he first of these can be correlated with the first appearance f large-scale deltas across northern Europe; the second the | | proliferation of coal-swamp conditions on the delta-tops; and the third the expansion of intra-montane basins in central and southern Europe. The trough in the Cantabrian presumably reflects the change-over from predominantly paralic to pre- dominantly intra-montane conditions over much of Europe. So, the diversity of the neuropteroids as a whole is merely a function of the general diversity of the tropical swamp vegetation. If the form-genera outlined earlier in this paper have any basis in the genetic relationships of the parent plants, diversity patterns of the individual form-genera may tell a different story. Fig. 20 shows the diversity curves of six of the most abundant of the form-genera plotted separately. This clearly shows that the story is far more complex. However, the style of analysis dealt with so far in this paper is not really suitable for uncovering the more detailed distribu- tional patterns. For this, we need to look at the detailed variations in diversity of the species within each of the form-genera. The problem here is the limited amount of suitable data available. There have been studies documenting the quantita- tive stratigraphical variations of different species, such as by Davies (1929). However, such work is mostly old, largely unillustrated and uses unreliable taxonomy. Also, as pointed out by Scott (1985), there are serious weaknesses with the sampling that was usually employed. Scott himself suggested that quadrat analysis, similar to that sometimes used to study living plant ecology, could produce more reliable results. However, while quadrat analysis might prove valuable in the detailed relationship between facies and plant fossils at a specific locality, it would need a considerable number of such studies before it would reveal any meaningful stratigraphical patterns of plant fossil distribution. We have instead adopted an alternative approach, by looking at the numbers of localities from where a species is recorded at different stratigraphical levels. To do this, it was decided to restrict the analysis to one particular area, which would help minimize potential palaeolatitudinal variations. The area should have numerous records spread over a reasonably long stratigraphical range. The data should also preferably be based on identifications made by a single authoritative palaeobotanist, thus minimizing the potential for subjective variations in identification. In fact, only one area was found to have all these virtues, namely the Franco- Belgian Basin, through the monographic study by Laveine (1967). Species diversity analysis (Franco-Belgian Basin) Laveine’s (1967) monograph provides a taxonomically reli- able record of most of the neuropteroid species found in the paralic belt between the Kinderscoutian and Westphalian D. For each species, he individually lists the localities where they are found in the Nord-Pas-de-Calais Coalfield, divided strati- graphically into lower, middle and upper divisions of the formations there. Using this data, we have plotted the diversity curves for each species of four of the form-genera (Figs 21-23). Neuropteris (Fig. 21). These 7 species appear to fall into two groups. The early group consists of N. obliqua, N. heterophylla, N. ghayei and N. willieri, which occur predomi- nantly in the Langsettian and basal Duckmantian (in Bel- gium, N. obliqua is reported to extend down to the Marsdenian, but the French records on which the present C.J. CLEAL AND C.H. SHUTE 34 ‘a8U}s YoUa UI Soldads JO SSO|/J1IJOId (p) ‘o8U}S YOUS Ul SIVSA UOl|[lul IsU SuOTSUT;XS Serosas| ———— Jo Joquinu (9) ‘asv}s yore ul sivak vont sod Suumeadde sarsads yo saquinu (q) :e8v)s yore ul saioads jo Jaquinu (v) :2]oym v sv soloeds prosraidoinsu yo dnoss ay} 10} sono AusIOAIG 61 “BIA sabes sebels iny OIS GIS Jeg UeEDGeM 10g ING Ue BeA PIW UI diy oud Wy Pud Wg QSV iny 91S gIS 22g UBD GEM 10g 9NG Ue BEA PIW UIM diy CUD Wy PUd 1G ASV g- Lor 10) 9- L = m Woe reed (o} Ss =a ee € 5 a Ss One o ©, 5a o = me] ° rd ® 3 o s‘s yo 7) ® 9 9 0) g /L. sabes sabes inv OIS ais Jeg UeDGaM I0G ING Ue BBA PW UH div oud Wy Pud Ya aAsSY ny OIS gIS Jeg UEDGEM 10g 2NG UBT BEA PJW Uy diy oud Wy Pud WG ASV fo) 0 c = S 2 = 14 » oO 9 =} °o Ce) > o Ol WwW 9 o 3 = 9. = g ° 8 5 < Si 7) OL . "q Vv 02 N = 35 NEUROPTEROID FOLIAGE FROM CARBONIFEROUS AND LOWER PERMIAN “98R]S UI SUONOUNXA JO JaquINN= J “983s ul Sulivadde ysiy satoads jo (s1e9X uoriW ut) uOnRINp asRIDAY = AL ‘asvys ul saouvsieodde jsmyj jouoquny = 4 “a8e3s ul juasaid saisads yo 1aqunN = Ny 9 9 I I G G 9 +£:0 € II & CO) € # I I I € I Pe ¢ 6 G if c:0 Te a1 Gps of Ss if o-9 E ¢ Le So Coan OPS I € V2 iL I I C I OL 6:1 6 LI I CG kee AS o/h (OH) IE O ine 8 LI Ore i i € a0:Ca ah aS iS Gel jE 8 GG 9 9] é WS € 6 GSE Gal BY, 8 VAG WAL Ol! eG: (ee (ME I € I Cl C UW 9 80 9 if Ll I 9 I O-S I! I iS 0-7 I ¢ I OHO yt U € 8S € 7 OH | il 0°S € c c-0 C € I S0) Lot I I I I 0-S I C I I 0-01 if I I I I (PEI It I OMe LU I BNL gl IN Uo oath gl WN WU sh gt IN T HL dN IT dl an T dL dN DIO]. stuajdodnauouads suajduvg Siiajdoinan Si4ajdyjo20Anan stuajdoyjajoanan NV oS ais 1eg ue) d2M log ond uey BOX PAN uly] div oy) uly pud lg gsv NN [oe) — ise) ae tN AOA Ga) VANS) sI \o =) COUGNIE CN i lool a) T dL AN a8e1¢ 14a] dOANIUOAIDIN Ue wel al INI SlajdoauiaavT en Pa Se ee ee ee Pe eee “e1ouasd plorajdoinony ul suoyeuea AjIsIoAIq ¢ aIquy, top) (oa) [e>) No. of species present ine) a = o——+— Asb Bri Pnd Arn Cho Alp Kin Mrd Yea Lan Duc Bol WeDCan Bar StB StC Aut Stages > Laveineopteris + Neuralethopteris ~™* Neurocallipteris -* Neuropteris % Paripteris 4 Sphenoneuropteris Fig. 20 Stratigraphical diversity of six of the more abundant neuropteroid form-genera, showing complexity hidden by generalized graph in Figure 19a. analysis is based only show it as far back as the Langsettian). It then undergoes a significant decline in the lower Duckman- tian. Only one of the species extends much beyond the middle Duckmantian, and that is what is referred to in the chart as the N. obliqua group. This pattern seems to be in general agreement with what is seen in areas other than the Franco-Belgian Basin. As with the other neuropterids, N. obliqua sensu stricto undergoes a marked decline in the lower Duckmantian. However, in the upper Duckmantian there is the start of a progressive change in the venation, which becomes more flexuous and eventually culminates in the anastomosed- veined form known as Reticulopteris Gothan. Details of this gradual change in venation through the Duckmantian and Bolsovian have been documented by Josten (1962), and the possible adaptive advantage of this style of venation is discussed by Zodrow & Cleal (1993). This morphological change is accompanied by a proliferation of the group of species, especially in the Bolsovian. The abundance of the N. obliqua group then undergoes a dramatic collapse in the topmost Bolsovian and it eventually becomes extinct in the upper Westphalian D. This collapse in abundance coincides approximately with the appearance of the second group of neuropterids at the base of the Westphal- ian D. In the Franco-Belgian Basin, this consists of just one species, namely N. ovata, but elsewhere in Europe there are other, very similar species which come in at about the same level (N. flexuosa, N. plicata and N. ervedosensis). This is near the top of the Upper Carboniferous succession in this basin and so provides no direct evidence of the diversity of these neuropterids at higher levels. However, in other areas such as South Wales (Cleal 1978) and NW Spain (Wagner et al. 1983, Wagner & Alvarez-Vazquez 1991) it is clear that the group continues to be abundant at least through the Westphal- ian D and Cantabrian, and in some cases beyond. There is no direct evidence from the Franco-Belgian Basin of the phylogenetic origins of the N. ovata group. There is a possible precursor in the Bolsovian of the Intra-Sudetic Basin (N. praeovata), but this throws little light on potential ances- C.J. CLEAL AND C.H. SHUTE \—"DRIER INTERVAL” = “| i N. ovata ; nN N. willieres 0 ' i ron N. ghayei ! ! y_ VW N. heterophylla N_ obliqua group WeD = Fig. 21 Detailed abundance variations of Neuropteris species, plotted against stages (using abbreviations shown in Fig. 2). In the graph of the Neuropteris obliqua group, black represents N. obliqua, fine stippling N. parvifolia, diagonal hatching N. semireticulata, and coarse stippling Reticulopteris muenstert. Based on data from Laveine (1967), determined from the Franco-Belgian Basin. The shaded expansion of the N. ovata curve reflects its proliferation in other areas. tors. Almost certainly, the group evolved in an extra-basinal habitat, possibly from a N. heterophylla-like ancestor. Laveineopteris (Fig. 22). The laveineopterids may b divided into two main groups: those with larger pinnules (th L. tenuifolia/loshii group) and those with smaller pinnule: (the L. rarinervis group). The larger pinnuled-group firs appears in the Langsettian with L. loshii, which reaches it acme in the upper Langsettian. At about the Langsettian, Duckmantian boundary, however, it undergoes a significan decline in abundance, and is replaced by a number of specie with more elongate pinnules (L. tenuifolia, L. hollandica, L jongmansii, L. morinii). These species, especially L. tenut folia, remain abundant and characteristic elements of the Duckmantian and Bolsovian, but then towards the top of th Bolsovian decline sharply to become extinct in the lowe Westphalian D. The L. tenuifolia group shows a reduction in abundance about the Duckmantian-Bolsovian boundary, but is otherwis an important and characteristic element found in thos stages. However, towards the top of the Bolsovian it unde goes a second and this time terminal decline, finally becomin extinct in the basal Westphalian D. 1 NEUROPTEROID FOLIAGE FROM CARBONIFEROUS AND LOWER PERMIAN 37 r — "DRIER INTERVAL” —— L. rarinervis L. morinii L. jongmansil ~ hollandica ~ tenuifolia ~ loshii WeD H 1 ' = uU c oO [e} 2 mo wv Malden fa L onlacs! | ) (Fig. 22 Detailed abundance variations of Laveineopteris species, | plotted against stages (using abbreviations shown in Fig. 2). | Based on data from Laveine (1967), determined from the Franco-Belgian Basin. The shaded expansion of the L. rarinervis | curve reflects its proliferation in other areas. | The L. rarinervis group of very small-pinnuled species shows a somewhat different distributional pattern. In the Franco-Belgian Basin it starts in the Bolsovian, having possi- bly originated from the slightly older L. nicolausiana. It proliferates during the Bolsovian. In the lower Westphalian D it appears to decline in the Franco-Belgian Basin, but this is symptomatic of it being at the top of the Upper Carbonifer- Dus succession here; elsewhere in the paralic belt of 20alfields, it continues to be abundant through into the ‘Cantabrian. Veuralethopteris (Fig. 23). It is well known that this form- yenus is restricted to the Namurian and Langsettian, a point vhich is borne out by the Franco-Belgian data. The only ther point of possible significance is that, compared with many of the other neuropteroid taxa whose extinctions are sormally marked by a gradual decline in abundance, the xtinctions of most of the neuralethopterids is characterized ly a sudden proliferation followed by a sudden decline. aripteris (Fig. 23). According to Laveine et al. (1989), the jlant that bore paripterid fronds migrated from China to < ted NW Spain Fig. 25 Cluster analyses using geographical distributions of neuropteroid species (see text for details of methods of analysis); (a) Chokierian to Yeadonian; (b) Langsettian; (c) Duckmantian to Bolsovian. NEUROPTEROID FOLIAGE FROM CARBONIFEROUS AND LOWER PERMIAN 43 3 2 Ss S SJ S by iS Buip Sonepat dane oe alee (d) RRS Sees iS) Sees eNS2 8 e 3 § & SSS S FR Ss 8 Ss § ea Ss 8 8 Ss § 25883 § N 100 90 80 70 60 50 40 30 20 10 0 PVPues=sezeRe Vey Nishi: Pennines Westphalian D - x x x Turkey _ Cantabrian i Me RS Donets CS Ie Sy Tes NW Germany x x x xX France-Belgium tat a U. Silesia x x Intra-Sudetic x x N. Caucasus x x x x K NW Spain Pa ES ES OS i SW UK x * Saar-Lorraine x < Alps x os a S. Portugal = N. Portugal x x Zwickau St < S 38 8 3 Seca! Gere suey (ies (e) aS. § Ge Ss SS) RRS SePsNsssryse &§ 8 § sss § g § $3 100 90 80 70 60 50 40 30 20 10 0 a Foe See RRS TS es SS SS eS Ee ee ae eee eee, RS) rs ears Re Ec aie Gh x x x x Intra-Sudetic Barruelian - palate! adi toes ipa nine NW Spain Autunian x MOS RS se x Massif Central * x x x x*K N. Portugal Saar-Lorraine S. Spain Donets x N. Caucasus Pig. 25 cont studied, at least in recent years. There are some exceptions, such as Darrah’s (1969) monograph on the Mazon Creek lant fossils, and some useful records by Gillespie et al. 1975), Gillespie & Pfefferkorn (1976), Gillespie & Crawford 1985) and Gillespie & Rheams (1985). These indeed suggest close similarity to the European assemblages. However, on heir own they are not really sufficient to allow a comprehen- ‘ive assessment of the North American records, which is why hey were not incorporated into the analysis presented in the resent paper (for a further review of the North American ecords, see Pfefferkorn & Gillespie 1980). | The western part of North America in the Carboniferous \as been assigned to two separate phytochoria, which may be eferred to as the Cordillera Palaeoarea of the mid-west ates and the Oregon Palaeoarea of the Pacific coastal area Pfefferkorn & Gillespie 1980, Cleal & Thomas in Cleal 991). No neuropteroids have been reported from the regon Palaeoarea. The Cordillera Palaeoarea is very poorly ‘ocumented, with the sole exception of the plant fossils from | | (d) Westphalian D to Cantabrian; (e) Barruelian to Autunian. the Manning Canyon Shale (Tidwell 1967). Of Tidwell’s records, the most significant is of Neuropteris cf. pocahontas White, which is undoubtedly a neuralethopterid similar to N. schlehanii. However, his record of ‘Neuropteris’ gigantea is undoubtedly incorrect (at least one pinnule in the figured specimen has a basiscopic lobe — it may in fact be an elongate neuralethopterid) and his ‘Neuropteris’ ampelina Tidwell is a Eusphenopteris. Gondwana. There are no neuropteroids recorded from the Carboniferous of the middle and high palaeolatitudes of Gondwana (the so-called pre-Glossopteris and early Glosso- pteris floras — reviewed by Wagner et al. 1985). However, the palaeoequatorial parts of Gondwana, such as the Mérida Andes of Venezuela, the Djerada Basin of Morocco and the Sud-Oronais region of Algeria, yield typical Euramerian-type assemblages. The published records include species of Neu- ropteris, Laveineopteris, Macroneuropteris, Paripteris and Neurocailipteris (Jongmans & Deleau 1951, Jongmans 1952b, 44 Pfefferkorn 1977, Migier 1982). However, these are either unillustrated records, or just show small fragments, which are difficult to assess; their generic affinities are probably correct, but any further statement will have to await a more complete documentation. Cathaysia. Although in very similar palaeolatitudes to Europe during the Carboniferous, only a few neuropteroids are found in China. The most significant from an evolution- ary point of view is Paripteris, which seems to have first evolved in China in the late Visean (possibly Brigantian) and only later migrated west to Europe in the Namurian (Laveine et al. 1989, 1992). The Chinese specimens have traditionally been referred to as Paripteris gigantea (e.g. Li et al. 1974, Yang et al. in Wagner et al. 1983). However, recent work by Zhang et al. (1992) and Laveine et al. (1992) has shown that, although similar, the Chinese material is not conspecific with that from Europe and it awaits a new name. There is also some evidence that Neuropteris may occur in China. There are numerous records from the Upper Carbon- iferous (thought to be approximately equivalent to the Stephanian in the Heerlen Classification) of North China of Neuropteris ovata. However, their veining is denser and the pinnules more broadly attached to the rachis than the typical Westphalian D specimens of this species from Europe, and Gothan & Sze (1933) referred them to a separate species, N. pseudovata. Wagner (1963) went further, to suggest that there is a close similarity between these Chinese fossils and the species which is now referred to as Neurocallipteris neuropteroides. This clearly raises a difficulty as to the status of the Chinese fossils, as there is no published evidence of their cuticles to prove whether they are neuropterid or neurocallipterid. In view of their relative high stratigraphical occurrence, these Chinese fossils are in clear need of revision. Li et al. (1974) described some fragmentary specimens from the Namurian of China as Lopinopteris intercalata Sze. Laveine et al. (1987) have argued that they may be very closely related to Neuropteris obliqua. However, there will have to be a more complete documentation of the Chinese material before its taxonomic position can be confirmed. Angara. There have been a number of records of Neuropt- eris from this palaeokingdom (e.g. Neuburg 1948, Gorelova et al. 1973). Among the more completely known species are ‘N.” pulchra Neuburg and ‘N.’ izylensis (Chirkova) Neuburg. Although only a few specimens of these species have been documented in the literature, and the illustrations of these are mostly poor, they demonstrate certain significant features of frond architecture: they have ultimate pinnae terminated by a pair of pinnules and intercalated pinnules on the penultimate racheis. These are characteristic features of the form-genus Paripteris, although the pinnule form and vena- tion is rather different from any of the European or Chinese species. Also of possible paripterid affinity is ‘N. “ dichotoma Neuburg, although this observation is based on the similarity of its pinnules and venation to the European species P. gigantea; little of its frond architecture has been documented. Two species with very large pinnules (up to 70 mm long) have been described under the names ‘N.’ siberiana Zalessky and ‘N.’ balachonskiensis Gorelova. One specimen of the former, figured by Neuburg (1948: pl. 31, fig. 1), shows pinnules apparently with two basal lobes or incipient pinnules (again, the quality of the illustrations make their interpreta- tion difficult). A comparison with Macroneuropteris is thus hinted at, but far more material needs to be examined before this could be confirmed. A rather unusual-looking species has been described as ‘N.’ ignotus Gorelova in Gorelova et al. (1973). It has very tapered, subfalcate pinnules, spaced widely along a very wide rachis, and is quite different from anything that has been previously assigned to the neuropteroid group. A comparison with the once-pinnate peltasperm frond Compsopteris is possible, although without more complete material, prefer- ably including cuticles, this affinity would be difficult to confirm. Most of the other Angaran species that have been assigned to Neuropteris (e.g. “N.’ tomiensis (Zalessky) Radchenko, ‘N.’ orientalis Radchenko) are all too small and poorly illustrated to assess. As far as it is possible to make out, other | than some possible paripterids, no good examples of neu- | ropteroid fronds have been described from these floras. C.J. CLEAL AND C.H. SHUTE | | | | Kazakhstan. ‘The Carboniferous plant assemblages found here are intermediate in composition between those typical of | Eurameria and Angara (Meyen 1987). According to both Vakhrameev ef al. (1978) and Cleal & Thomas in Cleal (1991), about half of both species and form-genera in the Middle Carboniferous (in the Russian chronostratigraphy, equivalent approximately to the Namurian and Westphalian of the Heerlen Classification) of Kazakhstan are also found in Europe, and include some neuropteroids. | The best documented records of Carboniferous plant fos- sils from here are by Radchenko (1954, 1985) and Oshurkova | (1967). Other than some large, isolated pinnules from the Upper Carboniferous (in the Russian sense, i.e. approxi- mately Stephanian), identified as the Angaran species “Neu- ropteris’ dichotoma Neuburg (see above), most neuropteroid- like material originates from the upper Visean and Namurian. The latter are all characterized by relatively small, vaulted, lateral pinnules with a weakly developed midve and a distinctive, round apical pinnule. The lateral pinnules| vary to an extent in shape, from round to oval to subrectang- ular with a round apex, and have been assigned to various species including Neuropteris antecedens Radchenko non Stur, N. heterophylla Oshurkova non Brongniart, N. pseudoheterophylla Radchenko, N. bulupalganensis Rad- chenko non Zalessky and N. karagandensis Borsuk. How-| ever, these morphological variants are frequently found associated together, and they almost certainly belong to one and the same species. Goganova et al. (1992) have recently described some remarkably complete examples of this species and found that it is fundamentally different from Neuropteris. They propose that the correct name is Cardioneuropteris asiatica (Radchenko) Goganova et al. Although the fronds are bipartite, producing tripinnate primary rachis branches, there are no intercalated elements between the secondary pinnae. Also, in close association were numerous Aulacotheca-like sporangial clusters, which in Europe ar normally associated with the frond form-genus Alethopteris. It is clear that Cardioneuropteris is fundamentally differen from any of the neuropteroid form-genera found in Europe. Mention should be made of specimens recorded b Oshurkova (1967) from somewhat higher (probably West phalian equivalent) strata under the name Neuropteris e qua. Unfortunately, only one extremely small fragment wa illustrated (Ibid.: pl. 15 fig. 8), which is totally inadequate fo taxonomic assessment. It seems that, other than the possible paripterid ‘N. NEUROPTEROID FOLIAGE FROM CARBONIFEROUS AND LOWER PERMIAN 45 dichotoma and the inadequately documented N. obliqua, no unequivocal neuropteroid form-genera (at least in the Euro- pean sense) have been recorded from Kazakhstan. CONCLUDING REMARKS We are minded at the end of our study to quote from the preface to John Woodward’s (1729) pioneering palaeonto- logical study; Now, that I have been for some time engaged in Mineral Studyes, with no small Application, ‘tis a Pleasure to me to find that it has not been wholly without Fruit. When we first started out on our project we intended it purely as a means of testing the robustness of the taxonomic scheme proposed by Cleal et al. (1990). However, we have ended up on a much longer journey into the realms of palaeogeogra- phy, biostratigraphy, palaeoclimatology and population dynamics. Trying to improve the taxonomy of a group of organisms, whether living or extinct, has its own internal _ logic, but we discovered that is has also provided an improved tool for understanding the pattern of the temporal and spacial distributions of the species. The distributions of the indi- - vidual species were of course mostly already known, but the _ more general patterns were obscured by the wholly artificial » generic taxonomy traditionally employed. Grouping the spe- - cies into what seem to be more natural form-genera provided » a context for at last seeing more clearly these more general patterns; we have been able to see the trees for the wood! This demonstration of its geological utility of course also | adds further support for the essential ‘naturalness’ of the _Tevised taxonomic scheme. That a group of species responds | in the same way to environmental pressures does not prove | that they are closely related. However, if the species are also ' morphologically very similar at both the macroscopic (frond architecture) and microscopic (cuticles) levels, there must iclearly be a strong likelihood that they are a genetically homogeneous group. There will always be the potential for convergent evolution to confuse the issue, especially with | organs such as leaves, but by using as many morphological | characters as possible it should be possible to detect this. A case in point is the close gross-morphological similarity between the mainly Westphalian D to Barruelian Neuropteris ovata and the mainly Stephanian C to Autunian Neurocalli- |pteris neuropteroides. Some authors have gone as far as to ‘suggest that the latter is a descendant of the former (e.g. Wagner 1963). However, their epidermal structures are very different, as are their apparent responses to environmental changes within the forests, and it is almost certain that the ‘similarity in gross morphology merely represents convergent evolution. Our study provides clear evidence of the long-known but often forgotten fact, that there is a close symbiotic relation- _|ship between the study of plant fossils and geology; the fossils cannot be properly understood without an understanding of the geological (sedimentological, stratigraphical, palaeogeo- graphical) context in which they are found. Equally, the plant fossils provide invaluable palaeoecological, biostratigraphical and palaeophytogeographical data for improving our under- standing of the geology. This information can then be re-cycled back to improve our understanding of the original vegetation (Cleal 1991: 223). As our study has demonstrated, this iterative process is dependent on the availability of a robust taxonomy, not only at the rank of species but also of form-genus. Obviously, a form-genus cannot be the exact equivalent of a whole-plant genus, being based only on a single plant organ. Nevertheless, the aim should be to make a form-genus as near as possible to a phylogenetically coherent concept (Cleal 1986), and this can only be achieved by detailed morphological and taxonomic study of the fossils. ACKNOWLEDGEMENTS. We are deeply indebted to the following for providing information on some of the species referred to in the study, and with which we were not previously familiar: Professor J.-P. Laveine (Université Science et Techniques, Lille), Professor R.H. Wagner (Jardin Botanico, Cordoba), Professor M. Barthel (Museum fiir Naturkunde, Berlin), Professor J.H.F. Kerp (Westfalische Wilhelms-Universitat, Munster), Dr. H.W.J. van Amerom (Geolo- gisch Bureau, Heerlen) and Dr. Z. Siminek (Ustiedni tstav geolog- icky, Prague). Professor Barthel and Dr Simtinck, together with Dr. A.C. Scott (Royal Holloway and Bedford New College, London), are particularly thanked for providing some of the photographs illustrated in this paper. The remaining photographs were produced by the Photographic Unit of The Natural History Museum, to whom we are grateful. We would like to thank Professor Barry Thomas (National Museum of Wales) for permission to reproduce the recon- struction shown in Figure 1. For assistance with statistical proce- dures, we would like to thank Dr. A.B. Smith and Mr. C. Montcrieff (The Natural History Museum, London), Professor M.C. Boulter (N.E. London Polytechnic) and Dr. W.L. Kovach (University of Aberystwyth). We gratefully recognize the contributions of Dr E.L. Zodrow (University College of Cape Breton, Sydney), particularly in the early phases, in the development of the taxonomy used in this paper, and for useful discussions on this subject in general. Finally, we would like to thank Dr. J. McEvoy (Newbury) for help with computer facilities, and Mr. J.A. Cleal (Solihull) and Dr T.J. Ferrero (The Natural History Museum) for producing the illustrated charts. 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Review of Palaeobotany and Palynology, Amsterdam, 30: 57-66. | i a ® & | NEUROPTEROID FOLIAGE FROM CARBONIFEROUS AND LOWER PERMIAN 51 INDEX OF GENERA AND SPECIES 0 eee ee EEE a ee ee ee eee ee eee This is an index of the systematic section, not the whole paper. Species which are regarded as ‘good’ in the sense used in this paper are shown in bold Roman type, while earlier synonyms and combinations, and species based on inadequate type specimens, are in italics. The archaic spelling variant Nevropteris is not distinguished in the index, and its entries are to be found under Neuropteris. Alethopteris neuropteroides 24 ' antecedens, Neuropteris 25 arberi, Neuropteris 31 . asturiana, Neuropteris 31 auriculata, Neurodontopteris 25, 32 Neuropteris 23, 25 beveridgei, Neuropteris 31 ) blissii, Neuropteris 30 bohdanowiczii, Neuropteris 25 Sphenopteris 25 - bourozii, Neuropteris 30, 32 britannica, Macroneuropteris 23 Odontopteris 23 | brongniartii, Sphenoneuropteris 29 » bulupalganensis, Neuropteris 31 ! chalardii, Neuropteris 20 _condrusiana, Neuropteris 25 cordata, Neuropteris 23, 25, 30, 31, 32 ; :delasti, Neuropteris 31 densifolia, Neuralethopteris 24 ‘densinervosa, Neuropteris 25 )dimorpha, Neuropteris 29 Pseudopecopteris 29 Sphenoneuropteris 29, 30 ‘dispar, Neuropteris 31 idoubravica, Neuralethopteris 24 Neuropteris 24 dufrenoyi, Neuropteris 25 )duprei, Neuropteris 30, 32 \dussartii, Neuropteris 30, 32 elegans, Sphenoneuropteris 29 \ervedosensis, Mixoneura 26 ervedosensis, Neuropteris 26 Filicites (Nevropteris) heterophyllus 26 | tenuifolius 22 Mabellinervis, Imparipteris 31 ‘Texuosa, Neuropteris 26 ‘ormosa, Neuropteris 20 gallica, Neurocallipteris 25 | Neuropteris 25 hermeri, Neuropteris 23, 27 Neuralethopteris 24 Neuropteris 20 jugosa, Neuropteris 31 kosmannii, Neuropteris 31 lanarkiana, Neuropteris 26 larischii, Neuralethopteris 24 Neuropteris 24 lata, Neuropteris 24 Laveineopteris guadiatensis 20 hollandica 20 jongmansii 20, 26 loshii 20, 22, 26, 30 morinii 22 nicolausiana 22 piesbergensis 22 rarinervis 22, 30 tenuifolia 22, 23, 26, 30 linguaefolia, Neuropteris 28 Paripteris 28 linguaenova, Neuropteris 28 Paripteris 28 longifolia, Neuropteris 24 loriformis, Neuropteris 24 loshii, Laveineopteris 20, 22, 26, 30 Neuropteris 20 lubnensis, Neuropteris 32 Macroneuropteris britannica 23 macrophylla 23 scheuchzeri 23 subauriculata 23 macrophylla, Macroneuropteris 23 Neuropteris 23 maltbyensis, Neuropteris 28 Margaritopteris multivenosa 23 marginenervis, Neuropteris 26 matheronii, Neuropteris 31 mathieui, Neuropteris 25 Mixoneura ervedosensis 26 grandifolia 31 muensterifolia 31 polyneura 27 praeovata 27 raymondii 30 wagneri 30 auriculata 23, 25 beveridgei 31 blissii 30 bohdanowiczii 25 bourozii 30, 32 bulupalganensis 31 chalardii 20 condrusiana 25 cordata 23, 25, 30, 31, 32 delasii 31 densinervosa 25 dimorpha 29 dispar 31 doubravica 24 dufrenoyi 25 duprei 30, 32 dussartii 30, 32 ervedosensis 26 flexuosa 20 formosa 20 gallica 25 germeri 23, 27 ghayei 26 gigantea 28 grangeri 26 guadiatensis 20 hemingwayi 20 heterophylla 26 hollandica 20 horrida 31 jongmansii 20 Jugosa 31 kosmannii 31 lanarkiana 26 larischii 24 lata 24 linguaefolia 28 linguaenova 28 longifolia 24 loriformis 24 loshii 20 lubnensis 32 macrophylla 23 maltbyensis 28 marginenervis 26 matheronii 31 mathieui 25 shayei, Neuropteris 26 montana, Neuropteris 32 montana 32 sigantea, Neuropteris 28 morinii, Laveineopteris 22 morinii 22 Osmunda 28 Neuropteris 22 multivenosa 23 Paripteris 28 muensterifolia, Mixoneura 31 nemejciana 29 var. 8, Osmunda 26 multivenosa, Margaritopteris 23 nicolausiana 22 sleichenites neuropteroides 25 Neuropteris 23 obliqua 22, 26, 27, 28, 31 /randifolia Mixoneura 31 obliqua forma impar 22, 26 rangeri Neuropteris 26 nemejciana, Neuropteris 29 ovata 23, 26, 27, 28, 30 juadiatensis, Laveineopteris 20 Sphenoneuropteris 29 ovata forma flexuosa 26 _ Neuropteris 20 Neuralethopteris densifolia 24 ovata var. grandeuryi 27 doubravica 24 var. pseudovata 27 emingwayi, Neuropteris 20 eterophylla, Neuropteris 26 eterophyllus, Filicites (Nevropteris) 26 ollandica, Laveineopteris 20 Neuropteris 20 orrida, Neuropteris 31 nparipteris flabellinervis 31 ovata 27 _ plesbergensis 22 ‘ngmansii, Laveineopteris 20, 26 | jongmansii 24 larischii 24 neuropteroides 24 rectinervis 24 schlehanii 24 Neurocallipteris gallica 25 neuropteroides 25, 27 planchardii 25 Neurodontopteris auriculata 25, 32 Neuropteris antecedens 25 arberi 31 asturiana 31 var. sarana 27 papilioniformis 25 parvifolia 27, 28 pilosa 27 planchardii 25 plicata 20, 27 praedentata 29 praeovata 27 pseudoblissii 30 pseudogigantea 28 pseudoimpar 31 pseudozamites 25 52 rarinervis 22 raymondit 30 rectinervis 24 rectinervis forma obtusa 24 resobae 23, 28 rytoniana 20 schaeferi 28 scheuchzeri 23 scheuchzeri forma minor 28 schlehanii 24 schlehanii forma rectinervis 24 schlehanioides 24 schiitzei 29 semireticulata 28, 31 squarrosaeformis 31 stipulata 27 subauriculata 23 subplicata 20 subsessilis 31 teberdensis 31, 32 tenuifolia 22 valdensis 27 venceslai 29 waltonii 31 willierei 28 zeilleri 31, 32 neuropteroides, Alethopteris 48 Gleichenites 25 Neuralethopteris 24 Neurocallipteris 25, 27 nicolausiana, Laveineopteris 22 Neuropteris 22 obliqua, Neuropteris 22, 26, 27, 28, 31 Pecopteris 26 forma impar, Neuropteris 22, 26 Odontopteris britannica 23 stradonicensis 30 Osmunda gigantea 28 gigantea var. 3 26 ovata, /mparipteris 27 Neuropteris 23, 26, 27, 28, 30 forma flexuosa, Neuropteris 26 var. grandeuryi, Neuropteris 27 var. pseudovata, Neuropteris 27 var. sarana, Neuropteris 27 papilioniformis, Neuropteris 25 Paripteris gigantea 28 linguaefolia 28 linguaenova 28 pseudogigantea 23, 28 schuetzei 29 veenti 28 parvifolia, Neuropteris 27, 28 Pecopteris obliqua 26 piesbergensis, /mparipteris 22 Laveineopteris 22 pilosa, Neuropteris 27 planchardii, Neurocallipteris 25 Neuropteris 25 plicata, Neuropteris 20, 27 polyneura, Mixoneura 27 praedentata, Neuropteris 29 Sphenoneuropteris 29 praeovata, Mixoneura 27 Neuropteris 27 pseudoblissii, Newropteris 30 pseudogigantea, Neuropteris 28 Paripteris 23, 28 pseudoimpar, Neuropteris 31 Pseudopecopteris dimorpha 29 pseudozamites, Neuropteris 25 rarinervis, Laveineopteris 22, 30 Neuropteris 22 raymondii, Mixoneura 30 Neuropteris 30 rectinervis, Neuralethopteris 24 Neuropteris 24 forma obtusa, Neuropteris 24 resobae, Neuropteris 23, 28 rytoniana, Neuropteris 20 C.J. CLEAL AND C.H. SHUTE schaeferi, Neuropteris 28 scheuchzeri, Macroneuropteris 23 Neuropteris 23 forma minor, Neuropteris 28 schlehanii, Neuralethopteris 24 Neuropteris 24 forma rectinervis, Neuropteris 24 schlehanioides, Neuropteris 24 schuetzei, Paripteris 29 schiitzei, Neuropteris 29 semireticulata, Neuropteris 28, 31 Sphenoneuropteris brongniartii 29 dimorpha 29, 30 elegans 29 nemejciana 29 praedentata 29 wagneri 30 Sphenopteris bohdanowiczii 25 squarrosaeformis, Neuropteris 31 stipulata, Neuropteris 27 stradonicensis, Odontopteris 30 subauriculata, Macroneuropteris 23 Neuropteris 23 subplicata, Neuropteris 20 subsessilis, Neuropteris 31 teberdensis, Neuropteris 31, 32 tenuifolia, Laveineopteris 22, 23, 26, 30 Neuropteris 22 tenuifolius, Filicites 22 valdensis, Neuropteris 27 veenii, Paripteris 28 venceslai, Neuropteris 20 wagneri, Mixoneura 30 Sphenoneuropteris 30 waltonii, Neuropteris 31 willierei, Neuropteris 28 zeilleri, Neuropteris 31, 32 Bull. nat. Hist. Mus. Lond. (Geol.) 51(1): 53-72 Issued 29 June 1995 The Upper Cretaceous ammonite Pseudaspidoceras Hyatt, 1903, in north-eastern Nigeria P. M. P. ZABORSKI bk €1%¢¢60 Department of Geology and Mining, University of Jos, P.M.B. 2084, Jos, Nigeria CONTENTS | VaUHROYG LLL) er sepa be pSE RE SSBe SERUM cite Neen MCE BE ee PCa sccreassomicenmarconabeadasscee aceccnit en re 53 NikeeAshalkavandseinci gs aysecuions atssrcatertsesoeeer- 5 ee tea eee ee ee ns SY SLE IMAC CESCLIPUONS mee seetr tree neretnrm mee atte oc Na ccs ahincienecre cee tea et eT eee, eh tears BESS Pavnavily AN CANNITUSKOXCSIRVANTDIDYANE, GRORSOINS ~cooacbponnsbadecasbesasenéoonssocedecabepdpesbesueoarenaédercnbobGscccoeebeces 55 SublamilysEU ONMPE ALO GER AGWINAEN Cooper tc--ee ites saan eee ee eee SS GonustESHUDASEID O CRRA SWclyattimenee names sere ics eee sae eee Eee eee EE ee 55 SEudas pid Ocerasipseudonodosorresl(@hoitat) mse steetee ee eee ee 57 BS EUMAS DIG OGErGSs OOLEAUI (StOMCZKa) meeereee eee eee eee ee ee 59 [ESL DILOCAES DURING REST soooancecooosets see se ocaoneneadedeone seaosod ocopposbopeeooeboaamncuasmnencusedeose: 63 LESAN AOCT ES LEDS RONAN cogecionasudcoooseeaeoee osc eeeesbeoorcn decd ioconobersascposodeenaseoeeeon sceshon aber 63 GenngBURROCERAS CobbanwHookséa Kennedy, -:es:5s5-ssss4.0 ce eee eee 67 UIA ROTEAERY GD), conn Sp oS eeEeECe OCG: PREC EERE DECOTH So neRERR RPE Peer RRRSRCER i cia eee ee om nei ne ioe 69 Sitiasi rap CALdS CUSSION aa a asset oer. sees ee oo oes eeGoe se cero sala dace ese meee es cate eeeROGeE eee 69 IRGIEINENESS | cenisoddangooodde sob eO aaa nee en eee CP CSgE CE ERE E EE Oe ENR ETE er Weert mune ie uli mares ee Ait flat ewe I ited 71 Synopsis. The following species of Pseudaspidoceras Hyatt occur in north-eastern Nigeria, from oldest to youngest: P. pseudonodosoides (Choffat), P. footeanum (Stoliczka), P. paganum Reyment and P. flexuosum Powell. P. pseudonodosoides is represented by paedomorphic populations. P. flexuosum marks the basal Turonian of the region. The remaining species are all of Late Cenomanian age. P. pseudonodosoides occurs in beds correlatable with the Neocardioceras juddii Zone in south-western New Mexico but the Nigerian sequence is expanded in comparison; no equivalents of the horizons with P. footeanum and P.paganum are known in the former region. The Nigerian material demonstrates the great potential of Pseudaspidoceras in detailed correlation over the Cenomanian- Turonian boundary. Below the beds with Pseudaspidoceras, specimens probably referrable to Burroceras Cobban, Hook & Kennedy occur. INTRODUCTION In recent years renewed attention has been paid to the Cenomanian-Turonian ammonite faunas of north-eastern Nigeria (Wozny & Kogbe 1983, Popoff et al. 1986, Meister 1989, Zaborski 1990, Courville et al. 1991, Courville, 1992). Early descriptions were provided by Woods (1911), Reyment 1954a, 1954b, 1955) and, especially, Barber (1957, 1960), dut these works generally lacked information concerning the wecise stratigraphical occurrences of the material involved. (he impetus for fresh studies has largely been provided by he opening of the Ashaka Cement Company’s quarry some 00 km north of the town of Gombe. Here, unusually for the egion, large ammonite collections can be made within a clear tratigraphical context. Coupled with information gathered rom scattered localities elsewhere in north-eastern Nigeria, otably the Pindiga stream section, data from Ashaka have nabled a regional ammonite biostratigraphy to be worked jut (Zaborski 1990). North-eastern Nigeria is emerging as a ' ) The Natural History Museum, 1995 | region of key importance in understanding the ammonite succession across the Cenomanian-Turonian boundary. The expanded sequences are dominated by vascoceratid faunas, but also contain acanthoceratid ammonites of wider geo- graphical distribution, allowing correlations to be made with zonal schemes from north-western Europe and North America. One of the genera having such value is Pseudaspi- doceras Hyatt, 1903, which is represented by four species, in ascending stratigraphical order: P. pseudonodosoides (Chof- fat), P. footeanum (Stoliczka), P. paganum Reyment and P. flexuosum Powell. In addition, forms probably referrable to the closely related genus Burroceras Cobban, Hook & Kennedy, 1989 occur lower in the sequence. The purpose of this contribution is to document the Nigerian material and to highlight its stratigraphical value. The Ashaka and Pindiga sections Although Pseudaspidoceras occurs widely in north-eastern Nigeria, only at Ashaka and Pindiga have more than one 54 species been collected in an observed stratigraphical sequence. The locations of these sections and other localities mentioned herein were shown by Zaborski (1990), while a general account of the geology of the region was given by Carter et al. (1963). A continuous section made up of ammonite-bearing lime- stones and interbedded barren shales of Late Cenomanian and Early Turonian age is exposed at Ashaka (Fig. 1). The Pindiga stream section is discontinuous and partly subject to burial and re-exposure during the summer rains. Figure 1 shows those parts of the section that have been logged over the period 1986 to 1992. The Ashaka section is as follows (see Fig. 1; numbers in parentheses are the equivalent horizons in Meister (1989): Bed Metres Alternating shales and sandstones, the lat- ter glauconitic, feldspathic and calcite- cemented below, quartzose above and with a 15 cm shelly, sandy limestone towards the top Glauconitic, gypsum-bearing clay Pale grey, nodular limestone with clay and gypsum laminae. Vascoceras harttit (Hyatt), V. obscurum Barber, Pseudotis- sotia nigeriensis (Woods), Eotissotia sim- plex Barber Blue-grey shales with gypsum Yellow, nodular limestone with Thalassi- noides burrows on upper _ surface. Pseudotissotia nigeriensis, Eotissotia sim- plex Blue-grey shales with gypsum Yellow, nodular limestone with gypsum. Thalassinoides burrows on upper surface. Pseudotissotia nigeriensis, Eotissotia sim- plex, Wrightoceras munieri (Pervin- quiere) Blue-grey shales with gypsum Greenish-grey nodular limestone, weather- ing to creamy yellow, with clay and gyp- sum stringers. Vascoceras sp., Pseudotissotia nigeriensis, Eotissotia sim- plex Blue-grey shales with gypsum Glauconitic, calcareous clay with black phosphate pebbles. Pseudaspidoceras flexuosum Powell, Watinoceras aff. colo- radoense (Henderson), Vascoceras pro- prium proprium (Reyment), V. obscurum Barber, Thomasites gongilensis (Woods), Pseudotissotia nigeriensis, Wrightoceras munieri and Choffaticeras sp. Cream-brown limestone with Pseudotissotia nigeriensis and Vascoceras proprium pro- prium (30) Blue-grey shales with gypsum ................. Cream-brown calcareous concretions, forming a continuous layer in places Dark grey to blue-grey shales Glauconitic, calcareous clay X (34) W (34) V (34) U (32) T2 (30) T1 (30) S (28) 12-00 P.M.P. ZABORSKI Hard, pale grey limestone. Pseudaspi- doceras paganum Reyment, Vascoceras proprium globosum (Reyment), Thoma- sites gongilensis, Pseudotissotia nigerien- sis Calcareous, glauconitic, shelly clay, with shale partings forming a more distinct shale unit up to 50 cm thick in some places Massive, hard, grey-green limestone. Vas- coceras sp., Thomasites gongilensis Dark grey shales with gypsum; a 14 cm horizon crowded with small thin-shelled bivalves, bone fragments and small phos- phatic pebbles occurs 15 cm below the top in some places 1p Clay, with gypsum and scattered white cal- careous nodules Dark grey shales with gypsum O (21-22) Hard, grey-green, nodular limestone in top 8-15 cm, rubbly limestone with shale part- ings below. Pseudaspidoceras footeanum (Stoliczka), Vascoceras proprium costatum (Reyment), V. proprium globosum, V. bullatum (Schneegans), V. nigeriense Woods, V. cauvini Chudeau, Thomasites gongilensis, ammonite gen. et sp. nov. R (26) Q (24) N (20) M (19) Hard, grey, massive crystalline limestone .. Rubbly, impure limestone with shale part- ings. Pseudaspidoceras pseudonodosoides (Choffat), Vascoceras sp. nov. aff. gamai Choffat, V. cauvini Hard, pale cream-grey, massive limestone . Rubbly, impure limestone with shale part- ings. Pseudaspidoceras pseudonodosoides, Vascoceras sp. nov. aff. gamai, V. cau- vini L (18) K (17) J (16) places 1 (14-15) Hard, grey limestone, cross-bedded in places H (12-13) Hard, grey limestone, cross-bedded in places. Vascoceras cauvini G(1i1) Massive, pale to dark grey limestone, pass- ing laterally into bioturbated grey-green limestone. Vascoceras cauvini Rubbly limestone with interbedded harder limestones. Burroceras? sp., Vascoceras cauvini F (10) E (10) Rubbly, grey-green to dark grey limestone with clay and gypsum stringers. Vasco- ceras cauvini in upper part D Massive, grey-green, glauconitic, quartzose limestone. Exogyra-rich. Nigericeras gad- eni (Chudeau) Brown-yellow, calcareous sandstone Ferruginous sandstone Grey-green, fine-grained, poorly consoli- dated sandstone becoming calcareous in its upper part >wWoO ~ mel THE UPPER CRETACEOUS AMMONITE PSEUDASPIDOCERAS HYATT, 1903, IN NORTH-EASTERN NIGERIA The Pindiga section, or more precisely that part of it which has been seen, is as follows, from top to bottom, see Fig. 1 (numbers in parentheses are the equivalent horizons in Popoff et al. (1986)): Bed Metres Blue-grey shales with gypsum (base of Pin- diga Formation shale member) V (22) Earthy, yellow-brown, glauconitic, calcare- ous clay, with gypsum and white calcare- ous nodules. Pseudotissotia nigeriensis, ISCISXOTEL DO QUE’ csccectoaccog CEE RO EERCCA ORDO 0-15 Blue-grey shales with gypsum ................. 1-00 U(21) Pale grey-green, poorly bedded, impure limestone. Pseudotissotia nigeriensis, JE DYUBCOME OTD DIAS cococucseadangodenorecdeoqee0e 0-70 T (21) Brown-grey, poorly bedded, calcareous shale, with gypsum and shale laminae. Pseudotissotia nigeriensis, Eotissotia sim- [DUB o accee tr Gamearno dssosccdcococa eee eA eRe ena 0:27 Blue-grey shales with gypsum ................. 0-15 18 (21) Cream-grey, irregularly bedded marl. Pseudotissotia nigeriensis, Eotissotia sim- [DUBE cer eapepeheap atise4c0¢ oc 08e0 aac een Rees 0-38 ‘R(17) Light grey, poorly bedded limestone with shale partings. Pseudotissotia nigeriensis. 7 cm hard, pale grey, nodular limestone | CAPPS see acdeeteee eee te triniss oo isteslucuies tea 0-40 KO) Pale yellow, nodular limestone ............... 0-06 | Blae-oney Shales! qeecseeeeeeetere ses -icseceise cose. 0-04 | Hard, fine-grained grey limestone ........... 0-08 P Porous grey limestone. Thomasites gong- ilensis, Pseudotissotia nigeriensis ........... 0-61 ILENE ONS. TAO) scaoaadooondadedquaconnaoaeeees 0-12 Hard, fine-grained grey limestone ........... 0-08 | Blue oneysshales|mesecserreeeece aecacecaceee seen 2-10 10) Intensely hard, grey, orange-weathering, shelly limestone. ?Pseudaspidoceras paganum, Vascoceras proprium globo- sum, Thomasites gongilensis, Pseudotisso- IL GATLLRCVLCTIS Stren ene erie: Ae DE ae an eR ia 91 INTHROGMEHIOD .s8stebBococodebtina san eaeleenbecsb Cosson cael eke coEc Co oeEe ETD nee mBEn Annee ars a ni Ceti Lae pee 91 SIMON? BREE pec cS Sood dbAbdogbROuSgHgERS EE CEE ER ORDoceIaCOOSONE NOPE REE MPH eTe rete eres: is Nanette la nee nen 93 ithotaciecrancbaunalkassem laces a aceedeeeeccete sss ohascce ei sneect ee eener eeaehccoeteeceee icc en eS 95 General palaccenvironmentalisynthesismepestessmererecetes.-< ceeeseeet etcetera cee ee 107 BOs Ural PTAPICAlMESUL ts merece etcunees sete teehee ec mee cece ov ceiscucsutease METRE Sree eave ce cuttin to wee. ls 107 Synopsis. The stratigraphy and macrofauna of the Qahlah and Simsima Formations is described for 14 sections along the western margins of the North Oman Mountains between Al Ain and Al Dhaid. These are the earliest autochthonous deposits following emplacement of the Semail ophiolite complex and are dated, on the basis of ammonites, inoceramid bivalves and echinoids, as latest Campanian to Maastrichtian in age. Deposition over a deeply weathered surface of ultrabasic rocks commenced with nearshore conglomerates, grits and cross-bedded sands which, in places, have an in situ bivalve fauna. Clastic input was abruptly terminated by a sea-level rise and replaced by carbonate deposition in the early Upper Maastrichtian. The basal carbonate bed is composed of coarse shoreface reefal debris formed during rapid transgression. This is overlain by a highly fossiliferous series of muddy carbonate sands deposited in shallow water around wave-base. There is indirect evidence for algal stands from the associated macrofauna, and level-bottom thickets of corals/rudists are present. Upper beds are well-washed carbonate sands rich in larger benthic foraminifers but with few macrofossils that were deposited in broad subtidal flats. In places these are overlain by deeper shelf-basinal marls. Bed by bed faunal lists for each section are provided. INTRODUCTION The Oman Mountains form a prominent arcuate range along the northeastern corner of the Arabian Peninsula. Their geological history is complex, but work by Glennie et al. (1974), Glennie (1977), Hughes-Clarke (1988) and contribut- ers to Robertson et al. (1990) has done much to improve our understanding of this region. The rocks forming the Oman Mountains can be divided into seven major geological units, ranging in age from late Proterozoic to early Tertiary. Of direct interest here are the Hawasina Group, the Semail Nappe and the Maastrichtian to early Tertiary autochthonous marine clastics and carbonates. The Hawasina Group is formed of tectonic slices of Permian to mid-Cretaceous sedimentary sequences deposited over the ocean floor and continental slope of the Arabian continent. They were dbducted onto the continental margin of the Arabian plat- orm during the Upper Cretaceous. The Semail Nappe repre- ‘ents a massive slice of former oceanic crust, generated by ) The Natural History Museum, 1995 subduction-related spreading during the Cenomanian- Turonian and emplaced before the Upper Campanian. Both are overlain by marine sediments of Maastrichtian to Palae- ocene age. Initially these beds were deposited around the newly emergent margins of the Arabian shield, following the obduction event. Later, a broad carbonate platform formed over the region as it subsided. Only remnants of this once widespread succession now remain, forming small jebels along the western margins of the Oman Mountains. The late Cretaceous to early Palaeocene rocks are conven- tionally divided into three units (Skelton et al. 1990), which are, from bottom to top: 1. The Qahlah Formation — a marine clastic sequence of sands and conglomerates of late Campanian or early Maastrichtian age. 2. The Simsima Formation — a platform carbonate sequence of Maastrichtian age. 3. The Pabdeh Group — a basal limestone conglomerate of reworked Simsima Formation with an erosive base, of 92 presumed early Palaeocene age, overlain by thin-bedded basinal marls of late Palaeocene age. Until recently little was known of the fauna of the late Cretaceous rocks of this region. The first published account of Cretaceous fossils was that of Carter (1852), who described late Cretaceous sediments of the Hadramaut region of south- east Oman. Parts of the fauna collected by Carter were described by Duncan (1865) and are Cenomanian in age. Lees (1928) gave the first authoritative account of the geology of the Oman Mountains and provided much new palaeonto- logical data. He described late Cretaceous faunas from sev- eral localities, including Jebel Bu Milh (‘Jabal al Milah in Wadi Sharm’), where he recorded and described 39 taxa of gastropods (‘the product of an hour’s collecting’). Clegg (1933) also described a small number of late Cretaceous species from Oman, but without stratigraphic details. In the past few years there has been renewed interest in the geology and palaeontology of the late Cretaceous deposits around the fringes of the Oman Mountains. The regional setting of these deposits was outlined by Alsharham & Nairn (1990, 1994), who also described lithofacies and listed micro- fauna for type sections. The late Cretaceous stratigraphy and faunas from the south western Oman Mountains (Dhofar region, Oman) were studied by Platel & Roger (1989), Roger et al. (1989) and Roman et al. (1989). Late Cretaceous ammonites, echinoids, foraminifera and calcareous algae were described from the central Oman Mountains (Smith er al. 1990, Kennedy & Simmons 1991). To the north, along the United Arab Emirates-Oman border region, Skelton er al. Fig. 1 Map of the Arabian Peninsula showing the area of study. A.B. SMITH, N.J. MORRIS, W.J. KENNEDY AND A.S. GALE (1990) studied key late Cretaceous sections from both a sedimentological and palaeontological standpoint. They pub- lished lithostratigraphic sections and faunal lists for these sections, with particular emphasis on the rudist bivalve fau- nas, from which they were able to provide the first detailed assessment of late Cretaceous palaeoenvironments for the region, demonstrating that the carbonate succession recorded a variety of facies ranging from intertidal to shelf basinal settings. Finally, there are a series of papers documenting part of the diverse echinoid fauna (Ali 1989, 1992a,b) and a few of the molluscs and corals from this region (Ghalib, 1989, 1990; Metwally 1992). Our interest in the faunas of this region began in 1984, when Dr. S. Nolan (then at the University of Swansea) and Dr. P.W. Skelton (Open University) brought back their collection of late Cretaceous fossils for identification. This, together with material brought to the Natural History Museum for identification by amateur collectors (notably, Mrs Valerie Chalmers), alerted us to the importance of the Maastrichtian faunas of the western fringes of the Oman Mountains. In April, 1991, A.B. Smith carried out a prelimi- nary survey of some late Cretaceous fossil localities along the Oman-United Arab Emirates border. The echinoid fauna proved to be exceedingly rich and also remarkably well- preserved, and without doubt represents the most important Maastrichtian Tethyan echinoid fauna yet known. During this initial survey it became rapidly apparent that the molluscan and coral faunas were also exceedingly rich and likely to be of equal importance, both in terms of new taxa and their significance for understanding late Cretaceous CUA E: Arabian Peninsula | E | } | t | LATE CRETACEOUS CARBONATE PLATFORM FAUNAS benthic community structure. Indeed, the small collections brought back to London for examination here by relevant experts generated considerable interest. Consequently a sec- ond expedition was mounted to study the entire macrofauna from these important sections and to investigate the lithofa- cies and palaeoenvironments in more detail. In January, 1992 three of us (ABS, NJM, ASG) spent two weeks exploring, logging and collecting from the late Cretaceous outcrops that form the western margins of the Oman Mountains along the United Arab Emirates Oman borders region. It is the results of these two bouts of fieldwork that form the subject of the following papers. STUDY AREA Our study area lies along the border between the United Arab Emirates and the Sultanate of Oman (Fig. 1). Outcrops in this region comprise small, generally outlying jebels along the western fringes of the Oman Mountains. Collection and logging was carried out at eight sites ranging from Jebel Huwayyah in the south to Jebel Faiyah in the north (Fig. 2). Sections were measured at each jebel and macrofauna col- lected or noted bed by bed. Details of the outcrops studied are as follows: 1. Jebel Huwayyah (Figs 2, 3). Two sections were logged and ' their faunas collected systematically. Jebel Huwayyah, section 1. Southeastern corner of the | U-shaped Jebel immediately to the north of the dirt track and _ about 3 km SE of the northwestern corner of the jebel, 10 km NE of Al Ain, United Arab Emirates. Map reference: Buraymi Sheet 1:100,000 NG—40-140; grid reference | 842,878. ' Jebel Huwayyah, section 2. Western face of the eastern limb of the jebel approximately 2 km SSE of the northwestern corner of the jebel, 10 km NE of Al Ain, United Arab | Emirates. Map Reference: Buraymi Sheet 1:100,000, _ NG-40-140; grid reference 823,877. | 2. Jebel Bu Milh (Fig. 3). Two sections were studied in detail. Jebel Bu Milh, section 1. Western face of a prominent ridge at | the northwestern tip of the jebel, 10 km NW of the village of -Mabdah, Oman. Map reference: Buraymi Sheet 1:100,000 NG+0-140; grid reference 895,075. Jebel Bu Milh, section 2. Southeastern corner of a prominent knoll, isolated from the main jebel at its northern end, some 10 km NW of Mabdah, Oman. Map Reference: Buraymi Sheet 1:100,000 NG-40-140; grid reference 906,075. 3. Jebel Rawdah (Figs 2, 4B). Six sections were examined in detail and a further two briefly investigated but not logged or systematically sampled. Logged sections are as follows: lebel Rawdah, section 1. Slope and cliff at the eastern end of “he northern flank of a valley some 50 m east of the head of he valley, 3 km east of quarry weigh-bridge and site office, jebel Rawdah, east of Al Madam, United Arab Emirates. dumayni sheet NG-40-14A 1:100,000; grid ref. 925,528. ebel Rawdah, section 2. Slope and cliff on northern side of ‘ebel Rawdah, about 2 km east of quarry weigh-bridge and ite office, Jebel Rawdah, east of Al Madam, United Arab ‘mirates. Sumayni sheet NG—40-14A 1:100,000; grid ref. 93 Al Hair Jepel Huwayyah Fig. 2. Locality map showing the position of the four major jebels studied (asterisked) where there are important outcrops of late Cretaceous sediments. For regional placement see Fig. 1. 913,544. Jebel Rawdah, section 3. Slope and cliff on south side of valley from 200 to 600 m east of the head of the valley, about 3 km east of quarry weigh-bridge and site office, Jebel Rawdah, east of Al Madam, United Arab Emirates. There were three measured sections, labelled from west to east a, b, and c. Sumayni sheet NG—40-14A 1:100,000; grid refs 932,528 (section 3a): 928,527 (section 3b) and 927,527 (sec- tion 3c). Jebel Rawdah, section 4. Slope and cliff at the eastern end of the northern flank of a valley, 600 m east of the head of that valley, 2.5 km east of quarry weigh-bridge and site office, Jebel Rawdah, east of Al Madam, United Arab Emirates. Sumayni sheet NG-40-14A 1:100,000; grid ref. 928,531. 4. Jebel Buhays (Figs 2, 4A). Two sections were logged and their faunas systematically collected. A third section (2), exposing the lowest beds of the sequence was impossible to 94 oe \4Jebel Qatar }:/4 % : | Fig. 3 Detailed locality map for Jebel Huwayyah and Jebel Bu Milh. 1, 2 = studied sections. For regional placement see Fig. 2. log, but collections were made from the scree slope. Jebel Buhays, section 1. East face of the most northerly hill forming Jebel Buhays, 4 km north of Al Madam, United Arab Emirates. Dhayd Sheet 1:100,000, NG—40-107; 780,681. A second section 300 m to the south and forming another small hill was also examined and collected from, but no detailed section was made. Jebel Buhays, section 2. Scree slope at the southwestern corner of Jebel Buhays, 4 km north of Al Madam, United Arab Emirates. Dhayd Sheet 1:100,000, NG—40-107; grid ref. 779,668. No measured section could be made, but the beds are almost vertical here and the scree material is all derived from the lowest few metres of the sequence. Jebel Buhays, section 3. Northeastern corner of Jebel Buhays, just southeast of a television mast, 4 km north of Al Madam, United Arab Emirates. Dhayd Sheet 1:100,000, NG—40-107; grid ref. 788,670. A.B. SMITH, N.J. MORRIS, W.J. KENNEDY AND A.S. GALE U U ‘ 5. Jebel Thanais (Fig. 4A). Northeastern side of the jebel, about 4 km north of Al Madam, United Arab Emirates. | Dhayd Sheet 1:100,000, NG—40-107; grid ref. 783,699. 6. Jebel Aqabah (Fig. 4A). Southwestern face of jebel, forming a steep cliff about 200 m northeast of the tip of Jebel Thanais and about 4 km north of Al Madam, United Arab Emirates. Dhayd Sheet 1:100,000, NG—40-107; grid ref. 785,698. | 7. Jebel Faiyah (Figs 2, 4A). One section was logged in detail. A further three were explored but found to be unprofitable: collections were made from only one. Jebel Faiyah, section 1. Eastern scarp face from the southern, nose of the jebel northwards for 500 m, 5.5 km north of Al Madam, United Arab Emirates. Detailed logs were made a the northern end of the section (section 1a) and approxi L Jebel Buhays } aA LATE CRETACEOUS CARBONATE PLATFORM FAUNAS mately midway along (section 1b). Dhayd Sheet 1:100,000, NG—~40-107; grid ref. 800,697 to 802,702. Jebel Faiyah, section 2. Eastern scarp face of the jebel approximately 3 km NNE of the southern tip of the jebel, and 8 km N of Al Madam, United Arab Emirates. Dhayd Sheet 1:100,000, NG—40-107; grid. ref. 806,722. All but the lowest 2 metres of outcrop was highly indurated and thus fossils collected come from only these basal beds. 8. Qarn Murrah (Fig. 4A). Northeastern slope of the garn, some 8 km west of the northern tip of Jebel Faiyah and 15 km north of Al Madam, United Arab Emirates. Dhayd Sheet 1:100,000, NG—40-107; grid ref. (approx.) 760,795. The section here was small and the rocks hard and, well-lithified, making collecting difficult. No section was logged. LITHOFACIES AND FAUNAL ASSEMBLAGES A total of 14 sections were logged and their macrofauna recorded. Lithological samples were collected systematically from the thickest sequence, (Jebel Rawdah, section 2), and sectioned for petrographic analysis. The lithological descrip- tions given for other sections are based solely on field observations and are consequently of a preliminary nature. The results of this work are summarized in the measured sections (Figs 5-11) and in the faunal lists of the Appendix. As the successions in the various jebels differ significantly and correlation between jebels was not initially obvious, each succession is documented in turn before attempting a synthe- sis. Qarn Murrah N 1 ar wal Qarn ee Mulayh tbe a 2S 3, 4 = studied sections. For regional placement see Fig. 2. Jebel Rawdah (Figs 5-7) 1. Sedimentary lithofacies at section 2. From field logging and petrographic analysis we recognize the following major litho- facies: Facies 1. The succession commences with an ultrabasic clast conglomerate (bed 1). This can reach more than 10 m in thickness in places, and comprises well-rounded clasts of mean size 10-20 cm. There are rare rounded fragments of rudist and occasional acteonellid gastropod shells. Similar sediments in the region have been interpreted as beach deposits by Skelton e¢ al. (1990), a view supported in our interpretation. Facies 2. There then follows a mixed clastic-bioclastic coarse sand, with relatively well-rounded coarse sand to gravel-sized clasts, ca. 1 m thick (beds 2-4). This includes up to 20% ultrabasic sand. The high sphericity and excellent sorting of the clasts indicates these sediments accumulated in a very high-energy environment; either intertidal or very shallow subtidal. We interpret these as shoreface sands. Facies 3/4. The majority of lower beds in the succession are poorly sorted mollusc-coral-foraminiferal packstones that contain components of three size classes: (i) clay and silt- grade carbonate, now replaced by a weak ferroan calcite microsparite cement; (ii) well-rounded, fine sand-grade par- ticles, with a sizeable component of ultrabasic grains in the lower beds, as, well as rolled and bored mollusc and echino- derm debris; (iii) diverse larger bioclasts which vary in composition from bed to bed. There are two broad facies distinguishable in hand specimen on the basis of the major clast component. Facies 3 is a poorly sorted calcarenite rich in mollusc clasts (especially rudist clasts) as well as obvious ultramafic sand- Detailed locality map for Qarn Murrah and Jebels Buhays, Thanais, Aqabah and Faiyah (map A), and Jebel Rawdah (map B). 1, 2 96 Figures 5 - 11. Measured sections through the late Cretaceous Qalah and Simsima Formations. Locality details for each named section are given in Figures 2 - 4 and in the text. Logs are drawn to a scale of 5 mm = 1 m for Figs 5, 6,9 - 11 and 4 mm = 1 m for Figs 7 and 8. Lithological data are based on field observations except for Jebel Rawdah, section 2. Faunal assemblages noted in the right-hand column are also largely based on field observations; full faunal lists for each section are given in Appendix 1. Symbols used are as follows:- Bioturbation Poorly-sorted bioclastic packstones with major grade clasts. This makes up the lower beds (beds 5-10) and includes various thin coquina lenses and horizons of coarse coral and rudist debris. Facies 4 is characterized by an abundance of orbitoid foraminiferan clasts, with fragments of the rudist Dictyopty- chus common in lower beds (beds 11-12) and of corals in upper beds (beds 15-19). Furthermore, in these higher beds there are also layered blebs of mud-grade carbonate, incorpo- rating sand-grade bioclasts, which are storm-reworked pellets of mud-grade sediment. Skelton er al. (1990) interpreted facies 3 and 4 as deposits formed in a point bar and tidal channel system. However, the original presence of significant quantities of mud-grade car- bonate (probably algal aragonite) in this facies, together with its overall poor sorting is taken as evidence of deposition below normal wave-base. The sand-grade component, which Siliciclastic sands & conglomerates Well-sorted bioclastic A.B. SMITH, N.J. MORRIS, W.J. KENNEDY AND A.S. GALE Coarse, poorly sorted bioclastic wackestones Red-weathering marly seams and wackestones orbitoid foraminifer packstones | content | Weathered serpentenite Rudists ©o@ Rudists ; (Semail complex) ANI (Dictyoptychus) ( Hippurites) Rudists Infaunal bivalves = Bivalves | (large Vaccinites) © (Scabrotrigonia) ‘S~ (undifferentiated) Gastropods ae Gastropods Gastropods : (Campanile) (Acteonella) @ (undifferentiated) | “, 4 | Corals CS Stromatoporoids SOF Echinoids Loftusia (benthic LL ASE NCCES foraminiferan) | is well-rounded and was originally well-sorted, was probably | washed in from shallower water (intertidal to shallow sub- tidal) by storm activity. Metre- to several metre-scale bedding cycles (most obvi- ously expressed in banded rhodolite development) are appar- ent in this facies. The origin of such cyclicity is enigmatic, but the deeper-water setting proposed above militates against their being tidal channel in origin. Facies 5. The upper part of the succession (comprising beds 19-28) is composed of well-sorted, well-rounded sand-grade bioclasts, grain supported and originally containing little or no calcite mud. There are rare larger (up to 1 cm) bioclasts. The foraminiferal component is dominant (50-60%) and includes both broken and rolled orbitoids, entire small rota- | lines and miliolines. Up to 10% dasycladacean algae also occurs. This facies was deposited significantly above wave- LATE CRETACEOUS CARBONATE PLATFORM FAUNAS Jebel Rawdah 2 (lower part) Massive well-sorted packstones with scattered orbitoids Prominent parting Massive well-sorted packstones with scattered orbitoids Nodular well-sorted bioclastic sand Red marly parting Nodular weltsorted bicckstic sand Bioturbated orbitoid- rich bioclastic Packstone Orbitoid-rich wackestone Poorly sorted bioclastic wackestones rich in orbitoids and with rhodolite bands Red-weathering, flaggy bedded silty cakcarenite Poorly sorted orbitoline-rich biociastic wackestone Nodular, coarse bioclastic wackestone with orbitoids Orange parting Grey poorty- sorted bioclastic Ppackstone with siliciclastic grains Sandy calcarenite Boulder conglomerate Sandy calcarenite Boulder conglomerate Campanile 28 Occasional Hemipneustes Echinoid level - Hemipneustes, Pygurostoma, Petalobrissus, Noetlingaster; Barbatia, Pycnodonte, Campanile and gastropod moulds Pygurostoma, Noetlingaster, Hemipneustes Petalobrissus, occasional Globator, Faujasia; Large Astrogyra; Dictyoptychus, Tylostoma and other molluscs 27 Abundant Cunnolites and gastropods Loftusia common Bournonia Petalobrissus, occasional Hemipneustes; Loftusia in levels; gastropods common; Cunnolites moderately common Hardouinia, Hemipneustes, Petalobrissus 26 Alternating Dictyoptychus and Cunnolites beds, with Loftusia and Petalobrissus Giant Dictyoptychus Cucullaea, Pholadomya; Petalobrissus, 25 Globator Noetlingaster, Zuffardia, Petalobrissus; Cunnilites; Modiolus Abundant Petalobrissus; occasional mollusc debris 24 Echinoid-ich level with Porosoma Bournonia abundant Lots of broken moliuscan debris, rare Dictyoptychus and stromatoporoids; rare Faujasia, Goniopygus and Orthocyphus Molluscs, Faujasia, rare Goniopygus, 23 Orthocy phus Echinoid-rich level Molluscan debris Echinoid-rich level Rolled and bored stromatoporoid heads, ocassional compound corals and Dictyoptychus Molluscan debris; abundant Bournonia; Faujasia, Goniopygus, Porosoma, Salenia Rudist and other molluscan debris; Faujasia Jebel Rawdah 2 (upper part) Massive-bedded wel-sorted packstone with scattered orbitoids and other bioclastic debris Nodular, red-weathering silty calcarentte Massive well-sorted packstones with scattered orbitoids Fig.5 Measured section made at Jebel Rawdah, section 2 (see Fig. 4B for locality). 97 Large Dictyoptychus and other poorly preserved rudists abundant Occasional levels of echinoid and molluse debris seen in cross-section Petalobrissus, Hemipneustes, Noetlingaster, Pygurostoma, Faujasia; Campanile Noetlingaster Campanile 98 base in a high energy environment, as indicated by the high degree of sorting and lack of mud-grade carbonate. 2. Succession and faunal assemblages (Figs 5-7). Lateral variation is considerably greater here than at any other jebel. Of particular note is the marked attenuation of the entire lower part of the succession eastwards along Jebel Rawdah 3 (Fig. 7). However, although there is marked variation in bed development from one section to the next, the same general succession can be identified at them all. The basal pebble to boulder conglomerate varies tremen- dously in thickness, being best developed at Jebel Rawdah 3a and thinnest at Jebel Rawdah 4. There is very little in the way of sand and grit lenses developed, the entire sequence being exceedingly coarse. However, rare Acteonella are present in the upper part of the sequence at Jebel Rawdah 3. The coarse siliciclastics are succeeded by a thin transitional sandy calcarenite facies quickly followed by grey, shelly bioclastic limestones with obvious scattered sand-sized grains of ophiolite. These basal calcarenites are relatively coarse and are well-lithified. At several horizons the beds contain rolled and often bored heads of the compound coral Actina- cis, some up to 30 cm in diameter, as well as a variety of allochthonous compound corals and mollusc debris. The infaunal cassiduloid Faujasia eccentripora is the only obvious autochthonous element in these beds. The basal bed at Jebel Rawdah 4 is notable for the abundance of transported hippuritid and Durania rudists. Within the succession at section 2 there are two closely-spaced shell-rich partings with many echinoids, notably Goniopygus, Phymotaxis and Orthocyphus. Towards the top of this calcarenitic succession is a promi- nent, orange-weathering, silt-enriched parting (top of bed 10, section 2). This bed is thickest at Jebel Rawdah 2 and has the same echinoid fauna as found below. The overlying beds are still bioclastic calcareous sands but are more poorly sorted, and in places include significant amounts of sand-grade siliciclastics (e.g. section 1, bed 1). They contain the solitary discoidal coral Cunnolites and the infaunal bivalves Cucul- laea, Pholadomya and, in places, Scabrotrigonia. Orbitoid Foraminifera occur but only in minor abundance. Overlying these poorly sorted bioclastic limestones come orbitoid-rich packstones, with abundant specimens of the infaunal cassiduloid echinoid Petalobrissus. At the base there is a distinctive infaunal echinoid assemblage dominated by the cassiduloid Zuffardia and the epifaunal regular echinoid Noetlingaster. One to three metres above the base of orbitoid-rich limestones there is a prominent horizon of very large examples of the rudist Dictyoptychus, probably in life position. The succeeding 2-3 metres at Jebel Rawdah 2 consist of beds with Dictyoptychus alternating with beds rich in the solitary discoidal coral Cunnolites, the larger benthic foraminifer Loftusia, and the infaunal cassiduloid echinoid Petalobrissus. It is at about this level that the ammonite Brahmaites (Anabrahmaites) vishnu (Forbes, 1846) was found at Jebel Rawdah 1. A second major silt-enriched level occurs above this, and these flaggy beds yield a fauna of large infaunal cassiduloid and holasteroid echinoids (Hardouinia, Hemipneustes). There then follows a succession of rather fine, muddy, thoroughly bioturbated, bioclastic limestones with orbitoid Foraminifera and bands of rhodolite nodules. There are occasional infaunal holasteroid echinoids (Hemipneustes), but the predominant component of the fauna are the solitary A.B. SMITH, N.J. MORRIS, W.J. KENNEDY AND A.S. GALE discoidal corals (Cunnolites and Asteraea) and a variety of gastropods, together with the larger benthic foraminifer Loftusia. The succeeding bed (bed 19, section 2) contains large compound corals (Astrogyra) and rudist material. This bio- clastic limestone also has occasional large infaunal echinoids and a mixture of other molluscs. The next few metres are composed of well-sorted, red- weathering, silt-rich calcarenites. They have a significant fine siliciclastic content, particularly in the Hemipneustes beds (bed 21, section 2). There then follows a thick and fairly monotonous sequence of well-sorted calcarenites with sparse to moderate numbers of orbitoid Foraminifera. This facies has little in the way of macrobenthos, other than the occasional Dictyoptychus, Noetlingaster or Campanile. There is, however, one distinct nodular, red-weathering, silt-enriched calcarenite level which has a diverse fauna of infaunal cassiduloid and holasteroid echinoids towards the top. Large, poorly preserved rudists occur in abundance at the top of the sequence and the succession is terminated by an unconformity. 3. Palaeoenvironmental interpretation. It is clear that at Jebel Rawdah local sedimentation patterns were controlled by topographic variation of the underlying sea-floor. In particu- lar, the south-eastern corner appears to have been relatively starved of sediment compared, for example, to the northern flank of the jebel. In environmental terms deposition commenced with pebble conglomerates laid down in a near-shore high-energy environment around the newly emergent obduction complex. These were replaced rapidly, as the obduction complex subsided and the region became flooded, by coarse, poorly sorted carbonate sands. These contain significant amounts of reefal debris and represent immediately offshore sands accu- mulating below wave-base. The following sequence of mixed orbitoid-rich platform shoals alternating with more protected platform bioclastic sands with their gastropod and solitary coral fauna probably represent local variation between topo- logical highs and dips over a broad carbonate platform in probably no more than 10-20 m water depth. Towards the top of the section a regressive phase is marked by a brief period of patch-reef development and the influx of fine siliciclastics. The remainder of the succession is composed of shallow-water calcarenites formed within active wave-base and supporting only a sparse fauna. Jebel Huwayyah. 1. Lithological succession and faunal assemblage (Fig. 8). The general succession is similar throughout the jebel, though with some lateral variation in bed thicknesses. The lowest beds, best seen at section 1, consist of poorly sorted silts and sands with rare lenses of the oyster Acutostrea. The succes- sion passes up into pebble- and grit-sized conglomerates and cross-bedded siliciclastic sandstones, the pebbles being pre- dominantly igneous in origin and well-rounded. Bed 7 is of particular interest because it contains broken fragments of thick-shelled rudists, and occasional pebbles that are encrusted by Acutostrea or, more rarely, by small compound corals. After a small gap in exposure there follows a succession of highly bioturbated, poorly sorted, brown-weathering, silt-rich Jebel Rawdah 1 More massive-bedded , packstones; well-sorted Nodular boturbated orbitoxd-rich bwclastic packstone Orange-weathering rubbty parting Nodular-bedded, poorty- and rhodolite bands Reddish parting Brown-weathering, grains with siliciclastics sorted, coarse bioclastic packstone with orbitoids Orbitoid - rich, bioclastic packstone, poorly sorted, bioturbated, poorly sorted packstone with siliciclastic Poorly sorted packstone LATE CRETACEOUS CARBONATE PLATFORM FAUNAS Coenholectypus ?Proraster (in cross-section) Agerostrea, Pycnodonte Rare Dktyoptychus Scattered gastropods, occasional large coral heads, Spondylus Hattopsis, Hemipneustes, Noetlingaster; discoidal corals; occasional Loftus and Dictyoptychus Pycnodonte, Neithea, Acteonella, large infaunal bivalves, lots of gastropods and discoidal corals; Loftusia; ammonites Large Dictyoptychus; Globator, Petalobrissus, Orthopsis Faujasia-rich level Abundant Scabrotrigonia, Cucullaea and Cunnolites Abundant Pycnodonte Fig.6 Measured sections made at Jebel Rawdah, sections 1 and 4 (see Fig. 4B for locality). Jebel Rawdah 4 Prominent red marl Parting Prominent red marl parting More massive bedded Hemipneustes Faujasia, Coenholectypus Dictyoptychus Globator, Pygurostoma, Coenholectypus bioclastic packstones with scattered orbitoids Rubbly, coarse brown-weathering sandy wackestone Rubbly, coarse, poorly-sorted packstone More massive orbitoid-rich bioclastic packstones Nodular orbitoid-rich biockastic packstone Soft, bioturbated marl seam Orbitoid-rich bioclastic packstone Hard nodular rhodolite- rich bed Red marly calcarenite Rubbly, poorty-sorted bioclastic packstone with orbitoids Rubbly, poorty-sorted biociastic packstone with orbitoids Red sandy marl Rubbly, coarse bioctastic limestone with siliciclastics Hard sandy calcarenite packstone Orthopsis, Salenia, Globator Scabrotrigonia, Pycnodonte Pycnodonte Scabrotrigonia, Pycnodonte Globator, Orthopsis Dictyoptychus Dictyoptychus Dictyoptychus, Globator, Hemiaster Noetlingaster common, Globator, Petalobrissus, Coenholectypus Lophids and Campanile Dictyoptychus Globator Phymechinus, Globator, Orthocyphus, Petalobrissus Globator, Petalobrissus, Faujasia, Glyphopneustes Globator Orthocyphus, Faujasia, Globator, Petalobrissus Bournonia, oysters, gastropods Abundant rudists (Hippurites, Dictyoptychus), gastropod and bivalve debris 99 100 A.B. SMITH, N.J. MORRIS, W.J. KENNEDY AND A.S. GALE Jebel Rawdah 3c Jebel Rawdah 3b 12 Massive, well-sorted orbitoid Packstone with chert nodules Jebel Rawdah 3a "1 Massive red-weathering Packstone with orbitoids Thin-bedded, red-weathenng, silt-rich wackestone Relatively thick-bedded orbitoid-rich packstone More massive orbitoid-nch packstone Red weathering parting Nodular, bioturbated, poorly sorted packstone with scattered orbitoids and thodolite bands Coarse, poorty sorted packstone == —_— — —_ —_— Ss Coarse conglomerate with rare lenses of sands and grits Fig. 7 Measured sections made at Jebel Rawdah, section 3 (see Fig. 4B for locality). Cucullaea, Tylostoma Agerostrea, Amphidonte, Barbatia Acteonella Mecaster, Actinophyma, Barbatia, Modiolus, Pycnodonte, Amphidonte Diptyoptychus, Agerostrea, Pycnodonte Clavagella Eligmid bivalves, sparse Dictyoptychus Loftusia, solitary discoxdal corals, Pycnodonte; Hemipneustes, Mecaster Strombid gastropods, Barbatia, mytiloids; Rolled and bored compound coral heads; Dictyoptychus and other rudists; occasional Acteonella, pycnodontids Occasional Acteonella fragments Jebel Huwayyah 1 Agerostrea and Pycnodonte Fine-grained carbonate sands/silts with chert nodules and scattered orbitoids Proraster Thallasinoides burrow systems Agerostrea and Pycnodonte; occasional solitary corals Campanile Oyster debris More massive, orbitoid-rich, packstone Bioturbated poorty-sorted bioclastic packstone with orbitoids Diverse fauna, including abundant gastropods including Campanile; Globator Pygurostoma, Orthopsis;. corals, bivalves Coarse, poorty sorted bioclastic packstone Pebble conglomerate Silty bioclastic wackestone with calcareous nodules Bioclastic packstone Scattered Loftusia Abundant Loftusia; Faujasia; Vaccinites, Spondylus solitary discoidal corals; rare compound corals, ammonites Silty bioclastic wackestone Rubbly bedded sandy bicclastic wackestone Abundant Loftusia Pebble conglomerate Cross-bedded, coarse sand with strings of grit Rudist fragments; some pebbles oyster encrusted (Acutostrea); bryozoans Pebble conglomerate Cross-bedded, coarse sand with strings of grit Poorly sorted conglomerates with cross-bedded sand lenses Fine carbonate sand Sparse Acutostrea with scattered clasts Poorly sorted conglomerates with sand lenses Small Acutostrea and Gastrochondrites Sand with scattered clasts le. 8 Measured sections made at Jebel Huwayyah, sections 1 and 2 (see Fig. 3 for locality). | | Jebel Huwayyah 2 Bioclastic packstone Rubbly bedded sandy bioclastic wackestone Rubbly bedded sandy bioclastic wackestone Rubbly bedded sandy bioclastic wackestone Compound corals; Dictyoptychus and Vaccinites; rare ammonites Abundant Loftusia and discoidal solitary corals; diverse molluscs; Hemiaster Abundant Loftusia and discoidal solitary corals; diverse mollusc fauna Abundant Loftusia and discoidal solitary corals; diverse mollusc fauna; Coptodiscus, Glyphopneustes 101 102 bioclastic limestones with significant amounts of dark-green, sand-sized, igneous grains. These beds contain abundant specimens of the benthic foraminifer Loftusia as well as scattered molluscan debris and occasional infaunal spatan- goid echinoids. The large solitary button coral Cunnolites is also abundant. Within this facies are two prominent marker limestones that are thicker and better developed on the western limb of the jebel. They contain a rich fauna of compound corals, the hippuritid rudist Vaccanites and the occasional large, tall-spired gastropod Campanile. None of the rudists are found in life position and the beds appear to represent allochthonous accumulations. However, the base of the upper of these beds is typically rich in Spondylus and Plicatula that are clearly not transported, as most are pre- served with both valves connected. Ammonites, some quite large, are occasionally found at this level, presumably trans- ported into this environment. Above the Loftusia-rich beds comes a sequence of less fossiliferous silty bioclastic carbonate sands, which are in turn succeeded by a thick pebble conglomerate. This conglomer- ate is composed of very well-rounded clasts, and represents such an anomalous lithofacies change that it must have formed through exposure and reworking of earlier conglom- eratic beds. Above the conglomerate siliciclastics abruptly disappear and are replaced by relatively clean bioclastic limestones of the Simsima Formation. This starts with an initial shell-rich bioclastic limestone, followed by 2 to 3 metres of highly bioturbated and poorly sorted bioclastic limestones with sparse sand-grade siliciclastics. These sedi- ments are very fossiliferous, the fauna dominated by shallow infaunal cassiduloids and holectypoids (Pygurostoma and Globator), infaunal naticids and other gastropods, including Campanile. Occasional pectinid bivalves are also present. Higher beds pass into less fossiliferous and much more massive and well-sorted, orbitoid-rich carbonate sands and silts. There is at least one level of thallasinoid burrows. Rare Dictyoptychus are found here. The orbitoid content of the sediments decreases upwards so that higher beds, which contain the epifaunal bivalves Agerostrea, Pycnodonte and the infaunal spatangoid Proraster, are finer-grained carbon- ate sands or silts with relatively few orbitoid clasts. 2. Palaeoenvironmental interpretation. The succession com- mences with unconsolidated subtidal sand and cobble beds deposited around the margins of the newly emergent obduc- tion complex. The sequence shallows upwards into the shore- face facies of bed 7, with increasing cross-bedding and coarser clastic content. The clastic supply then sharply diminishes, presumably marking the final flooding of nearby ophiolite islands, and the initiation of carbonate platform develop- ment. The poorly sorted Loftusia-rich muddy carbonate sands represent extremely shallow-water, back bar or lagoonal deposits formed in a protected environment below wave-base. Coral-hippuritid thickets were able to develop locally and occasional open-water ammonites were washed in. The thin, well-sorted calcarenitic sands with Faujasia may represent protected beach-face sands. A brief regression caused by local uplift led to the exposure of Qahlah Formation conglomerates, or their source rock, and pebble conglomerates were once more briefly deposited in a high-energy shallow marine environment (bed 13). This was short-lived and carbonate platform conditions returned once again, bringing about the more or less complete elimina- tion of siliciclastic components. This presumably marks a A.B. SMITH, N.J. MORRIS, W.J. KENNEDY AND A.S. GALE renewed phase of subsidence with transgressive seas once more flooding the region and cutting off siliciclastic input. The initial coarse bioclastic fossiliferous sands (bed 14), we interpret as deposits formed at or immediately below normal wave-base. They are replaced immediately upwards by thick beds of shallower-water orbitoid-rich carbonate sands with little macrobenthos, formed within wave-base. These are succeeded in turn by fine carbonate sands and silts suggestive of shelf-basinal conditions. Jebel Bu Milh 1. Lithological succession and faunal assemblages (Fig. 9). The succession begins, as in Jebel Huwayyah and elsewhere, with a rather thick series of well-rounded pebble conglomer- ates, grits and coarse clastic sands, with some small-scale cross-bedding. Although largely unfossiliferous, occasional clumps of the rudist Durania can be found, apparently in situ. Towards the top is a 1-2 m thick conglomeratic sequence, packed with well-sorted shells of the gastropod Acteonella, showing good current alignment of individual layers. Occa- sional specimens of the rudist Pseudosabinia also occur. The succeeding clastic sands are strongly cross-bedded and con- tain occasional Acteonella specimens, as well as large logs bored by lithophagid bivalves. There is then a sharp reduction in the siliciclastic content of the sediments and the succeeding 1.0-1.5 metres are com- posed of highly fossiliferous calcarenitic limestones with sparse sand-sized quartz grains. The basal bed is a fine, silty calcarenite that is highly nodular and extensively bioturbated. It contains rare nautiloids and ammonites and a rich gastro- pod fauna, with naticids and neogastropods dominating. An erect, branching sponge is a common element of this fauna, though none appear to be preserved in situ. Above this basal carbonate bed is a thin, red-weathering, silt-enriched parting, followed by a harder, less bioturbated shelly bioclastic lime- stone that contains at its base the large tall-spired gastropod Campanile in some abundance, together with Acteonella, Dictyoptychus and other large molluscs. Succeeding beds are well-sorted orbitoid-rich bioclastic limestones. These contain little in the way of macrofauna except for a very distinctive horizon of large hippuritid rudists in situ, some 4-5 m above the base of the limestone succes- sion. 2. Palaeoenvironmental interpretation. The succession at Jebel Bu Milh commences with quartz sands, gravels and conglomerates representing open-water, nearshore deposits around the newly emergent obduction complex. The succes- sion shallows upwards, with the Acteonella conglomerates representing open shore-face, shell-lag accumulations at wave base, and the overlying cross-bedded sands with beached | driftwood representing tidal sandbars. The clastic succession | is abruptly replaced by carbonate platform deposits, presum- ably as nearby islands became submerged. A thin deposit of silty bioclastic limestone rich in naticid gastropods represents shallow subtidal sands, possibly formed at wave base in a_ protected back-bar environment. The presence of the large neritid Lisocheilus indicates that shallow rocky substrates lay in the vicinity. The remaining carbonate succession, by comparison with beds at Jebel Rawdah, represent shallow subtidal carbonate sand flats formed above wave base. a) LATE CRETACEOUS CARBONATE PLATFORM FAUNAS Jebel Bu Milh 1 Well-sorted orbitoid- rich bioclastic packstone Rubbly level Orbitoid-rich bioclastic packstone Bioclastic packstone Sands with conglomerate and grit levels, occasional cross-bedding Shel-rich, occasional coral Scattered molluscan debris Dictyoptychus and Campanile Gastropods in cross-section Abundant Campanile and Tylostoma, rare Deltonautilus Abundant Dictyoptychus and Acteonella; rare Campanite and Durania Clumps of Durania 103 Jebel Bu Milh 2 Massive-bedded, well- sorted biockastic packstone with orbitoids Welksorted bioclastic Giant Vaccinites Packstone with orbitoids Bioclastic orbitoid- Sparse molluscan debris rich packstone Shelly bioclastic Campanile, Dictyoptychus, Acteonella packstone and other molluscs Nodular poorly-sorted iAbundantinaticd and thiid t d sandy wackestone lant naticid and cerithiid gastropods, Orange sand occasional Deltonautilus, sponges Strongly cross-bedded Occasional bored logs, some large; sands with grit partings Acteonella also occurs Cross-bedded coarse sands and gravels Acteonella abundant, forming shell beds, also some cerithiid gastropods and the rudist Colveraia Poorly sorted, cross- bedded sands and grits with conglomerate lenses Pebble conglomerate fig. 9 Measured sections made at Jebel Bu Milh, sections 1 and 2 (see Fig. 3 for locality). 104 Jebels Buhays, Thanais and Aqabah 1. Lithological succession and faunal assemblages (Fig. 10). These three jebels show virtually the same succession and can therefore be treated together. The initial siliciclastic succes- sion of pebble conglomerates, sands and gravels is thinner than at Jebel Rawdah or Jebel Bu Milh. At Jebel Aqabah the succession commences with a 50 cm well-sorted calcarenite resting directly on top of slightly weathered ophiolite. This basal bed is notable for the small, uniform size of its fossils; a small heteromorph ammonite, Glyptoxoceras sp., was recov- ered from here. Elsewhere the basal contact is not seen. Towards the top of the clastic sequence, at Jebel Buhays, section 1, come 3 to 4 metres of laminar and cross-bedded sands. These contain a fauna of infaunal venerid bivalves and small turreted gastropods. As elsewhere, there is a sudden elimination of siliciclastics coincident with the onset of biocalcarenite deposition, mark- ing the base of the Simsima Formation. The limestone succession begins with a hard, 80 cm thick, sandy bioclastic shell bed full of the gastropods Acteonella and Campanile, the echinoid Goniopygus and rudist fragments (Durania and hippuritid), as well as other molluscan debris. Occasional compound corals occur at this level, but apparently not in situ. The succeeding 3 to 4 metres consist of highly bioturbated, poorly sorted coarse- to medium-grained bioclastic lime- stones, with scattered orbitoid foraminifera that increase in abundance upwards. These beds are rich in infaunal echi- noids (cassiduloids, especially Pygurostoma, and the holecty- poid Globator) as well as small epifaunal regular echinoids. They are also rich in gastropods and bivalves, both infaunal and soft-bottom epifaunal forms. The only rudist to occur here in abundance is the small recumbent Glabrobournonia. Within this succession there is a major red-stained, silt- enriched layer that immediately overlies a thin bed of fine, ?dolomitized limestone. There is then a second major shell-lag deposit, full of large sponge-bored shells of Acteonella and other molluscs. Above this the bedding becomes much more massive and the lime- stones better sorted and cemented. Immediately overlying the Acteonella shell bed is an orbitoid-rich packstone with rhodolite bands and a low diversity fauna of the rudist Dictyoptychus and the gastropod Campanile. A major red-weathering silt-enriched horizon occurs towards the top of the sequence and is immediately overlain by a one metre thick brown-weathering silty limestone, rich in tall cylindriform hippuritids. Associated here is the echi- noid Codiopsis, indicative of rocky shore habitats. The silt- enriched bed records a marked increase in the clastic component at this level. It is overlain by massive-bedded, well-sorted carbonate sands/silts with orbitoids. 2. Palaeoenvironmental interpretation. The palaeoenviron- mental setting is similar to that seen elsewhere, with shallow marine sands, gravels and conglomerates deposited above active wave base. The basal calcarenite at Jebel Aqabah probably represents a shoreface sand, as suggested by the well-sorted clasts and fossils. These sands, gravels and conglomerates are replaced abruptly by coarse, poorly sorted bioclastic calcarenites con- taining a diverse molluscan and echinoid fauna. These beds are highly bioturbated and were deposited in a protected environment below wave-base, possibly in a shallow lagoonal A.B. SMITH, N.J. MORRIS, W.J. KENNEDY AND A.S. GALE setting. The occasional large compound coral near the base suggests that reefal patches developed nearby. Periodic minor adjustments in sea-level resulted in red-weathering, silt-enriched partings and beds, marking temporary influxes of fine iron-rich silt-grade clasts while thin dolomitic levels may signify temporary supratidal exposure. A subsequent regressive phase led to the deposition at wave base of the Acteonella shell-lag of bed 11 followed by orbitoid-rich carbonate sands with a low diversity macro- fauna. We interpret these well-sorted and well-lithified beds as shallow water platform shoals formed above active wave base. There was a brief event that created the upper iron-rich siltstone band (bed 14) which was followed by a bed with hippurited thickets and the shallow-water echinoid Codiopsis, indicative of nearshore conditions. Shallow-water carbonate shoals form the remainder of the succession. Jebel Faiyah 1. Lithological succession and faunal assemblages (Fig. 11). Here the sequence overlying weathered ophiolite begins with pebble conglomerates, with interspersed grit and sand lenses becoming more common towards the top. Occasional frag- ments of thick-shelled mollusc are present indicating a marine origin for the unit. As in other sections, there is an abrupt change to carbonate sedimentation, commencing with a 40-50 cm shell bed full of | Acteonella, together with lesser amounts of rudist fragments — and other thick-shelled molluscan material. The succeeding 3 metres consist of highly bioturbated, poorly-sorted bioclastic _ limestones, with rhodolites and orbitoid foraminifera, a diverse fauna of gastropods, small regular echinoids and Dictyoptychus bands. There is a major, 10 cm thick, red-coloured, silt-enriched bed which, as at Jebel Buhays, is interpreted as marking a — minor change in sea-level with increased erosion of lateritic | soils from the hinterland. It is succeeded by in situ coral thickets with associated hippuritid rudists (bed 6, section 1a). The majority of in situ colonies are erect, branching forms some 50 cm in diameter, with intermixed erect colonies of | cylindrical hippuritids. A variety of other colonial corals and rudists also occur. There is also a moderately diverse fauna of small regular echinoids, such as Glyphopneustes, and small gastropods at this level. Next come a few metres of thick-bedded, orbitoid-rich | : | | | ee ee es ee | ~ bioclastic limestones with occasional levels of Dictyoptychus and rhodolites but little else. There is then a return to | red-weathering, poorly sorted bioclastic limestones with | significant silt-grade component (beds 8-10). These beds appear to be relatively iron-rich. The fauna is diverse and dominated by the epifaunal bivalves, Pycnodonte and Agerostrea. Small discoidal corals (Cunnolites), occasional compound corals, the foraminifer Loftusia and rudist Glabro- bournonia all occur at this level. The succeeding thick sequence of rather massive, well lithified and well-sorted carbonate sands, rich in orbitoid] foraminifera, but with no observable macrofauna, is tun cated by an erosion surface. 2. Palaeoenvironmental interpretation. The succession begins with shallow-water sands and gravels deposited above active wave base. In places they may even become intertidal or supratidal (Skelton et al. (1990) reported the presence 0 sedimentary structures indicative of intertidal or supratida i f 4 vl) LATE CRETACEOUS CARBONATE PLATFORM FAUNAS Jebel Buhays 1 Massive-bedded orbitoid packstone Brown-weathering, silty fine calcarenite Red silty wackestone Massive-bedded, well- sorted orbitoid packstone with rhodolite bands Shelly cakcarenite Rubbly bedded, poorly sorted, orbitoid-rich bioclastic wackestone with rhodolite bands ?dolomitized Ist Rubbly orbitoid- rich wackestone Red silty wackestone Orange, bioturbated poorty-sorted bioclastic wackestone with some orbitoids and rhodolites Sandy bioclastic packstone Pebble conglomerate Calcareous sand with grit and pebble lenses Pebble conglomerate le. 10 Measured sections made at Jebel Buhays, Codiopsis; occasional Dictyoptychus Small hippuritids common; Globator rare Hemiaster Dictyoptychus Scattered Dictyoptychus Large Dictyoptychus, Campanile, Tylostoma and Neithea Acteonella shell lag with Neithea, Bournonia, lophids and Dictyoptychus Acteonella (small) and other molluscs; occasional Astrogyra; Globator common. Campanile, Plagiostoma, Neithea, Scabrotrigonia, rare Dictyoptychus, vermiform gastropod; Phymosoma, Globator Small Acteonellids and other gastropods; Pycnodonte, Bournona; Cunnolites; regular echinoids, Globator; decapod claws Lyria, strombid gastropods, eligmids, Granocardium, Scabrotrigonia Campanile, Acteonella, Crassatella, Pycnodonte, rare Durania fragments; Porosoma Mesocallista and small gastropods in lenses Jebel Aqabah Sandy packstone Coarse sand/gravel with conglomeratic lenses Boulder conglomerate Well-sorted shelly calcarenite packstone sections 1 and Jebel Aqabah (see Fig. 4A for locality). 105 Acteonella common with much rudist debris and other molluscan material Infaunal venerid bivalves common, with low diversity gastropod fauna Small shelly fossils including Faujasia and Petalobrissus: Nostoceras 11 Fig. 11 Jebel Faiyah la Thick-bedded welt sorted packstones with scattered orbitoids and bands of chert nodules Bioturbated, silty, poorly- sorted wackestone; red- weathering { Rhodolite band Poorly sorted bioclastic wackestone Thick-bedded, well-sorted orbitoid-rich packstone with rhodolite bands Well-sorted, thick-bedded bioclastic packstone with abundant leptorbitolines Highly bioturbated red mart-rich wackestone Orange poorly-sorted bicclastic packstone with scattered orbitoids and some siliciclastic grains Well cemented sandy bioclastic packstone with rhodolites Sands and pebble conglomerates Pycnodonte, Spondylus, Agerostrea, Amphidote, Dictyoptychus, Cunnolites, Noetlingaster, Loftusia Pycnodonte uncinella Rare Dictyoptychus Large Dictyoptychus Noetlingaster Branching coral thickets with hippuritid colonies. Dictyoptychus Dictyoptychus, small gastropods, Hattopsis Acteonella, small gastropods, Sparse rudists, turrilitids and Callianasid claws Jebel Faiyah 1b Bioclastic packstone with orbitoids Red silty wackestone Orbitoid-rich packstone with occasional rhodolite bands Bioclastic packstone with orbitoids Poorly sorted bioclastic packstone Reddish marl parting Fine carbonate sand Coarse poorly-sorted bioclastic packstone Shelly bioclastic packstone Pebble conglomerate with sand lenses Measured sections made at Jebel Faiyah, sections la and 1b (see Fig. 4A for locality). Loftusia, Pycnodonte, bivalves and gastropods; Mimiosalenia, Globator Occasional molluscan debris Dictyoptychus Occasional Dictyoptychus Branching coral thickets in situ; Hippurites colonies many gastropods; rare Noetlingaster Abundant Dictyoptychus and many gastropods and small echinoids Acteonella abundant at base LATE CRETACEOUS CARBONATE PLATFORM FAUNAS environments from these beds). Clastic input ceased sud- denly, marking a major transgressive event, and was replaced by carbonate deposition. The carbonate succession began with a transgressive shell-lag deposit rich in acteonellids, after which shallow lagoonal conditions were established for a short period. There was then a minor shift in sea-level, creating a silt-enriched band, which was immediately fol- lowed by the establishment of coral-rudist level bottom reefal thickets and peri-reefal bioclastic sands. These in turn were quickly replaced by shallower-water bioclastic carbonate sands with abundant orbitoid foraminifera. These we inter- pret as shallow-water platform shoals formed above active wave base. Later, a second transgressive phase, marked by an input of siliciclastic material, brought a brief return to deeper water conditions, below active wave base, and the establis- ment of a more diverse fauna once again. This, however, was short-lived and shallow-water shoal conditions quickly returned with the deposition of massive bedded carbonate sands and the virtual disappearance of benthic fauna. GENERAL PALAEOENVIRONMENTAL SYNTHESIS The autochthonous late Cretaceous succession was deposited over a deeply weathered surface of ultrabasic rocks, which must have been exposed subaerially for some time. Initial deposits were locally derived conglomerates and grits formed around the shores of the newly uplifted ophiolite massifs. Pebbles and boulders are well-rounded. Although rudist and acteonellid debris does form a component of these beds, they _ probably represent shell coquinas washed onshore from sub- tidal sand-flats rather than autochthonous fauna. There is evidence locally for more sheltered shoreface facies with infaunal bivalves, or for more stable cobble-bottom develop- ‘ment, with an in situ fauna of encrusting oysters and/or corals. In places, as Skelton et al. (1990) note, the rudist Durania can be found living in situ in cross-bedded sands. _ A marine transgression largely drowned these ophiolite islands and led to the onset of carbonate deposition. During this transgression, shoreface reefal debris, with mixed hippu- ritid and radiolitid rudists and massive compound corals, was deposited as a coarse lag at the base of the transgression. The ‘corals are, for the most part, encrusting forms and are accompanied by a shallow intertidal to immediately subtidal tegular echinoid fauna composed of species adapted for life on hard substrata within the zone of active wave surge. Acteonellid gastropods are a characteristic feature of such Shell-lag deposits. Faujasia is the only common infaunal echinoid in this environment and probably lived in nearshore or shore-face clean, well-washed sediments. Slightly more protected sand beds were colonized by non-siphonate infau- tal bivalves such as Scabrotrigonia and Cucullaea. This facies was succeeded, as sea-level rose, by a thick succession of rather muddy sands formed at or below active wave base. At some levels the large semi-recumbent rudist Dictyoptychus is common, together with bands of rhodolites, lense orbitoid foraminiferal aggregates and abundant small nfauna!l cassiduloid echinoids (Petalobrissus and Zuffardia). The sea bottom must have been loose and unconsolidated, ying at or just below active wave base. It regularly received torm-washed bioclastic material. 107 Elsewhere, solitary discoidal corals and the larger benthic foraminifer Loftusia occur in profusion together with pycn- odont oysters and a moderately diverse gastropod assemblage in what appears to have been a slightly muddier sand. Infaunal echinoids are more or less absent from this facies, though the small epifaunal regular echinoid Hattopsis is often locally abundant and may indicate the presence of algal stands. The molluscan fauna is dominated by gastropods (especially the ?algivore Acteonella borneensis and the filter- feeding Umboniidae gen. nov.) and the epifaunal ‘Pycn- odonte’ uncinella. We interpret these beds as stable, possibly algal-bound, sands lying below wave-base. Where this facies occurs close to reefal thickets there is a much higher diversity of regular echinoids accompanied by the holectypoid Globa- tor. Unconsolidated calcarenite shoals, deposited above active wave base, form a major part of the upper succession and contain the lowest faunal diversity. The absence of any large-scale cross-bedding suggests these formed as shallow, broad, subtidal expanses of sand. Rotaline and milioline foraminifera are the predominant bioclasts in this facies, along with dasycladacean algae. It is in this environment that the large selective deposit feeding irregular echinoids Hemip- neustes, Pygurostoma and Stigmatopygus are found. The only regular echinoid found to inhabit this environment was Noetlingaster. At Jebel Huwayyah, however, the highest beds appear rather finer grained and contain the infaunal spatangoid echinoid Proraster. These may be local basinal sediments deposited below wave base, and possibly represent the deep- est water sediments present in the sequence. The palaeoenvironmental reconstruction that emerges from our combined sedimentological and faunal evidence differs somewhat from that given by Skelton ef al. (1990). They interpreted the lower part of the Simsima Formation as tidal deposits, whereas we believe that this is inconsistent with both the sedimentological and faunal evidence. The petrography of the sediments indicates deposition below wave base. This is supported by the diverse echinoid fauna, which is also at variance with a tidal environment. The faunal associations that we recognize represent more- or-less contemporary assemblages that replace one another in vertical succession as a result of shifting sedimentological facies. We found very little stratigraphic variation within single lineages, except in species of the echinoids Hattopsis and Hemipneustes. Consequently, we believe that the differ- ences in the assemblages reflect variations in palaeoenviron- mental conditions over the carbonate platform and represent coexisting communities. BIOSTRATIGRAPHICAL RESULTS Maastrichtian biostratigraphic zonation. The Qahlah Forma- tion has been regarded as Campanian or early Maastrichtian in age, and the Simsima Formation as Maastrichtian (e.g. Skelton et al. 1990). The boundary between Campanian and Maastrichtian stages has been defined on the basis of a range of biostratigraphic criteria, involving belemnites, ammonites, planktonic foraminifera and coccoliths (amongst other groups). However, recent reviews (Burnett ef al. 1992, Kennedy et al. 1992, Hancock et al. 1992) have demonstrated 108 that these various data do not correspond. Indeed, Obradov- ich (1993) concluded that the interval between the widely adopted planktonic foraminiferal boundary datum (the extinction point of Globotruncanita calcarata) and the widely adopted belemnite definition (appearance of Belemnella lan- ceolata) is at least three million years. There is as yet no agreed definition of the boundary, and subdivisions of the stage into Lower/Upper or Lower/Middle/Upper vary widely between authors. Indeed, in the absence of a clear statement of which definitions of stage and substage limits are used, these terms are near meaningless. Burnett et al. (1992) were able to place the putative markers for the base of the Maastrichtian in sequence, as well as certain other key fossil occurrences. Their scheme is as follows: (Youngest) — last occurrence of the nannofossil Quadrum trifidum [or Tranolithus phacelosus (= orionatus)]|. — first occurrence of the ammonites Pachydiscus (P.) neu- bergicus and Acanthoscaphites tridens. — last occurrence of the nannofossil Broinsonia parca. — first occurrence of the ammonite Hoploscaphites constric- tus s.l. — first occurrence of the belemnite Belemnella (B.) lan- ceolata. — last occurrence of the ammonite Nostoceras (N.) hyatti. — last occurrence of the nannofossil Reinhardtites anthopho- rus (or Eifféllithus eximus). — first occurrence of the ammonite Nostoceras (N.) hyatti. — last occurrence of the planktonic foraminifer Globotrun- canita calcarata. — first occurrence of the planktonic foraminifer Globotrun- cana falsostuarti. — first occurrence of the nannofossil Reinhardtites levis. (oldest) As they note, the first occurrence of the belemnite Belemnella lanceolata is widely taken to indicate the base of the Maastrichtian stage, and this is the definition adopted in the present work. Birkelund ef al. (1984), in their review of the conclusions of the 1983 Copenhagen Meeting on Creta- SUBSTAGE Belemnella casimirovensis Upper Maastrichtian Belemnitella junior Belemnella occidentalis Lower Maastrichtian Belemnella obtusa Belemnella lanceolata correlation. BOREAL BELEMNITE ZONATION Belemnella fastigata Belemnella cimbrica Belemnella sumensis Belemnella pseudobtusa Belemnella lanceolata Fig. 12 Biostratigraphic zonation schemes for the Maastrichtian based on belemnites, ammonites and inoceramid bivalves, and their A.B. SMITH, N.J. MORRIS, W.J. KENNEDY AND A.S. GALE ceous stage boundaries, noted that it was ‘widely accepted to keep the base of the Maastrichtian close to the appearance of Belemnella lanceolata, as this datum is so well-defined and widely accepted in the Boreal realm. However, there is a strong need for finer correlation of this boundary level with the succession in the Tethyan Realm possibly by planktonic foraminifera or coccoliths’. Since the work of Burnett et a/. and the partial achievement of the needs expressed at the Copenhagen Meeting (see also Schonfeld & Burnett 1991, Kennedy et al. 1992, Hancock et al. 1992, Hancock and Kennedy 1992) further refinements in correlation of the base of the Maastrichtian in the classic Boreal sense with Tethyan sequences have been achieved through the work of Burnett, Kennedy & Ward (1992), Hancock et al. (1993), Hancock & Kennedy (1992) and Ward & Kennedy (1993). These results are summarized in Figure 12. Ward & Kennedy (1993) recognized a threefold ammo- nite zonation of the Maastrichtian, with a fourth zone of Pseudokossmaticeras tercense that was possibly in part Cam- panian, in part Maastrichtian. Taking the base of the Maas- trichtian at the first appearance of Belemnella lanceolata, the work of Hancock & Kennedy (1992) and Hancock et al. (1992) demonstrate that Pachydiscus (P.) epiplectus, Hoplos- caphites constrictus and P. (P.) neubergicus first occur within a very narrow interval, such that the base of the Boreal lanceolata Zone and Tethyan epiplectus Zone are coeval, within the current limits of biostratigraphic correlation. For subdivisions of the Maastrichtian we use Lower and Upper Substages, as is widely accepted by workers in the Boreal Realm, the base of the Upper Maastrichtian lying at the base of the Belemnitella junior Zone (Fig. 12). Anapachydiscus fresvillensis (index species of the second zone of the Maastrichtian of Ward & Kennedy 1993) first occurs in the junior Zone in the Netherlands (Kennedy 1987). The base of the Boreal junior Zone and Tethyan fresvillensis Zone are coeval, within current limits of biostratigraphic correlation. Anapachydiscus terminus, index of the highest Tethyan Maastrichtian ammonite zone, is now known from the Boreal casimirovensis Zone (as Anapachydiscus aff. fres- villensis of Birkelund 1993 in Denmark, and, subsequently, in the Netherlands (Jagt, in press)). For a more refined division AMMONITE ZONATION Fi Anapachydiscus terminus | Anapachydiscus fresvillensis Trochoceramus morgani Trochoceramus lanjonaensis Pachydiscus (P.) epiplectus Trochoceramus monticulae/radiosus INOCERAMID ZONATION kL LATE CRETACEOUS CARBONATE PLATFORM FAUNAS of the Lower Maastrichtian reference will be made to the zonation of Schulz (1979: fig. 12). Good palaeontological dating and correlation between the isolated jebels is very limited. Within each jebel there is evidence of sedimentation having taken place on an uneven sea-floor and relativily rapid changes of thickness and facies are sometimes visible by casual inspection. Larger benthonic foraminifera are often abundant but apparently controlled in their distribution by palaeoenvironment. Further careful col- lecting would be necessary before their stratigraphical poten- tial is realized. Therefore, here we use evidence from the ammonite, inoceramid bivalve and echinoid faunas, together with inferred sea-level changes. Ammonite biostratigraphy. The small number of late Creta- ceous ammonites from the United Arab Emirates-Oman borders region (Kennedy, this volume) provide us with some constraints on the age of the succession. More weight has been placed on the pachydiscids than the heteromorphs, largely because they are better preserved. The sections at Jebel Huwayyah include at least three ammonite horizons. The first and lowest contains an uniden- tified species of Pachydiscus, which was discovered at the top of the sand and conglomerate sequence of the Qahlah Forma- tion, just below the Loftusia-rich beds, and only indicates a late Upper Cretaceous age. The Loftusia-rich beds themselves have yielded the great- est number of ammonites, including Pachydiscus (Pachydis- cus) dossantoi (Maury, 1930), Lewyites ambindense (Collignon, 1971), Nostoceras (Nostoceras) major Kennedy & Cobban, 1993, Nostoceras (Nostoceras) sp., and a Libyco- ceras sp. (not described; photograph only seen). The implied ages of these ammonites are somewhat contradictory. Pachy- discus (P.) dossantoi (Maury, 1930) is imprecisely dated within the Maastrichtian but occurs in the highest of three ammonite horizons in the Nkporo Shale of south-eastern Nigeria (Zaborski, 1985). At that locality it is closely associ- ated with the inoceramids Trochoceramus ianjonaensis (Sor- nay) and Endocostea coxi (Reyment), which indicate a late Lower Maastrichtian to early Upper Maastrichtian age. Lewyites ambindense (Collignon, 1971) is known only from the lower Upper Maastrichtian, fresvillensis Zone. Nostoceras | (Nostoceras) major Kennedy & Cobban, 1993 is Upper, but not uppermost, Maastrichtian, with an inferred fresvillensis Zone age. The presence of Lewyites ambindense and Nosto- ceras (Nostoceras) major imply a lower Upper Maastrichtian, fresvillensis Zone age. However, there are also two species that suggest an earlier age for at least part of the unit; Nostoceras (Nostoceras) sp. is uppermost Campanian or lower Lower Maastrichtian in age, and a specimen of Libyco- ceras, now in the Arab Emirates University Museum, has a bifid outer saddle indicative of the earliest of the three Libycoceras horizons described by Zaborski (1982) from Nigeria, and a late Campanian age. The age of the Loftusia- beds could therefore be as old as late Campanian or as young as early Upper Maastrichtian. In the northwestern part of the outcrop at Jebel Huwayyah, close to where it is cut by the road, the lower part of the Simsima Formation yielded Pachydiscus (Pachydiscus) _neubergicus neubergicus (Hauer, 1858), and Lewyites ambin- | dense (Collignon, 1971). Pachydiscus (P.) neubergicus neu- | bergicus (Hauer, 1858) implies a lower Lower to lower Upper Maastrichtian, epiplectus and fresvillensis Zones age. As pointed out above, Lewyites ambindense (Collignon) is 109 known only from the lower Upper Maastrichtian. A specimen of Desmophyllites diphylloides (Forbes, 1846) was found in bed 21 at Jebel Rawdah, section 2, well up in the Simsima Formation. However, the species is not age diagnos- tic, and its known range is Santonian to Upper Maastrichtian, fresvillensis Zone, and possibly higher. However, from Jebel Rawdah section 1, from scree almost certainly derived from the top of bed 4, we found a specimen Brahmaites (Anabrah- maites) vishnu (Forbes, 1846). The few well-dated specimens of this species are Upper Maastrichtian, fresvillensis Zone, and possibly younger in age. In the basal transgressive calcarenite shell bed beneath the sands and conglomerates of the Qahlah Formation at Jebel Agabah a Glyptoxoceras sp. was recovered. However, the genus is not age diagnostic and ranges from Santonian to upper Upper Maastrichtian. At Jebel Buhays, section 2, a Libycoceras sp. was found in the basal conglomerate of the Qahlah Formation. Libycoceras species first appear in the Upper Campanian and may range into the Lower Maastrich- tian (Zaborski, 1982). The basal beds of the Simsima Formation at Jebel Buhays, section 1, yielded a specimen of Pachydiscus (P.) dossantoi (Maury, 1930) which suggests a ‘mid-Maastrichtian’ age, presumably not very different to the Loftusia-rich beds at Jebel Huwayyah. In conclusion then, the meagre ammonite data that we have suggests that the basal transgressive sands and conglom- erates are late Campanian to ‘mid-Maastrichtian’ in age, whereas the main carbonate deposits of the Simsima Forma- tion are of early Upper Maastrichtian, fresvillensis Zone, age. At Jebel Huwayyah the Loftusia-rich horizons may be as old as late Campanian or as young as early Upper Maastrichtian, fresvillensis Zone. We do not know whether or not sedimen- tation continued to the end of the Maastrichtian. Inoceramid biostratigraphy. Inoceramid biostratigraphy has been based on what appear to be evolving lineages, often expressed as the ranges of subspecies. These are usually defined as the timespan of particular ‘morphs’ within an evolving species, although short-term biological events, such as the widespread ‘flood’ of a particular species or subspecies, are also important. Both methods can provide useful bios- tratigraphical data. In recent years much progress has been made on integrat- ing biostratigraphic data derived from Upper Cretaceous inoceramids and other fossil groups with data from sequence stratigraphy. Of all the Upper Cretaceous stages, however, the inoceramid biostratigraphic scheme for the Maastrichtian is the least well-established. Species of the genus Trochoceramus hold the most promise for establishing a sound Maastrichtian biostratigraphy, since they apparently represent a relatively simple evolutionary lineage. Trochoceramus species have a somewhat rounded shell outline, with their umbones set back from the anterior, and have evenly spaced, rounded, comarginal ribs and dis- tinctive radial ribs. The earliest species is narrow, with a regularly curving shell, but later species have a more convex shell, which is narrow in the early stages, but then undergoes a significant change in coiling direction, markedly increasing the relative volume of the mantle cavity. A similar change is also observed in the earlier genus Cremnoceramus. Here we recognize three successive inoceramid faunas of Trochoceramus which can be used to zone the Maastrichtian. The lower division is characterized by the narrow, regularly 110 curved species Trochoceramus cf. monticulae (Fugger & Kastner) (?= Trochoceramus radiosus (Quaas)). These flat forms first occur in very late Campanian strata (e.g. in the Nacatoch Sand of Navarro County, Texas, U.S.A. (Stephen- son 1941, pl. 13, fig. 3), and from Ammoniten Berg in the western Egyptian desert, apparently co-occuring with Libyco- ceras ismaeli (Quaas, 1902). However, the species becomes much more widespread in the lower part of the Lower Maastrichtian (Dhondt, 1983). The middle division is characterized by a more convex species of Trochoceramus, T. ianjonaensis (Sornay), its con- vexity arising from a sharp change in shell curvature following an initial flattish stage. Such forms are known from Madagas- car, where they have been dated as ‘lower’ Maastrichtian (Besairie 1972), although the ammonites listed suggest levels well into the Upper Maastrichtian as defined here. They are also known both from the Calabar district of Nigeria, where they are ‘mid’-Maastrichtian, and the St. Lucia Formation of Zululand, Republic of South Africa, where they almost certainly extend into the Upper Maastrichtian. Finally, they are also reported from Libya, but apparently not associated with ammonites. The upper division is characterized by Trochoceramus morgani (Sornay). It is of Upper Maastrichtian age and, so far, has only been recognized from the Calcaire a Baculites of the Cotentin Peninsula, France. 7. morgani clearly differs from T. ianjonaesis by the smaller average size of the initial flat part of the shell (this average is 67% of the average size in T. ianjonaensis). Put another way, T. ianjonaensis includes forms with large and small initial shells, whereas 7. morgani includes only specimens with small initial shells. Because this reduction in size of the initial flat portion of the shell has not been recognized in Trochoceramus from the Republic of South Africa, it is uncertain whether this size reduction represents a genuine world-wide change in the lineage or simply a geographical variation. A tentative biostratigraphic scheme for the Maastrichtian based on inoceramid bivalves and compared with the cephalopod biochronology is pre- sented in Fig. 12. Although inoceramid bivalves are proving useful for the division of the Maastrichtian they occur only rarely in the Upper Cretaceous deposits of the U.A.E. and Oman. We have discovered them at only three horizons, and so far only the material from Jebel Rawdah is at all well-preserved. Furthermore, none of this material includes species of Tro- choceramus. The lowest horizon includes only a fragment indeterminate at generic level, which was recovered from the conglomerate series below the Loftusia-beds at Jebel Huwayyah. At the second horizon, in the basal transgressive shell bed of the Simsima Formation at Jebel Bu Milh, we found a single large fragmentary specimen of ‘Platyceramus’ sp. This genus occurs in some abundance in the Maastrichtian succession of the St Lucia Formation in the Republic of South Africa, where it is common in the Lower Maastrichtian, but also seems to be dominant in the uppermost inoceramid horizon of that formation. A rather poorly preserved fragment of Endocostea ct. bebahoaensis comes from the Loftusia-beds at Jebel Huwayyah. It is not sufficiently well preserved to be dated better than Campanian or Maastrichtian. The highest of the three levels, some way up the Simsima Formation at Jebel Rawdah, has yielded the most important inoceramid fauna. At Jebel Rawdah, section 1, just below the base of the more massive, well-cemented limestones (base of A.B. SMITH, N.J. MORRIS, W.J. KENNEDY AND A.S. GALE bed 5) Endocostea (Cataceramus) semaili sp. nov. occurs. This is most comparable with specimens from Nagoryany, Ukraine, and indicates a ‘mid-Maastrichtian’ horizon. Drs Skelton and Nolan also collected this species, together with Endocostea (Endocostea) cf. coxi, Endocostea sp. and ‘Endocostea’ bebahoaensis, from the south-western part of the jebel at the same locality and possibly the same horizon as the ammonite Pachydiscus (P.) neubergicus neubergicus. Endocostea coxi occurs with Trochoceramus ianjonaensis in Nigeria and Zululand, while ‘Endocostea bebahoaensis occurs with the same Trochoceramus in Madagascar, but is known to be long-ranging (it occurs in the Upper Maastrich- tian at Cotentin, France). The inoceramids from Jebel Raw- dah imply an early Upper Maastrichtian age, thus giving support to the age suggested by the ammonites from the same horizon. Echinoid biostratigraphy. Although useful biostratigraphi- cally for division of late Cretaceous—Palaeocene strata in the Boreal realm, echinoid distribution in Tethyan carbonate sequences is at present too poorly known to provide reliable dating of the succession. Furthermore, the environmental constraints of most species makes them highly restricted in their occurrence, and thus of limited value. However, our work has identified two biostratigraphically useful species lineages, Hemipneustes spp. and Hattopsis spp. There are three species of Hemipneustes which differ in the relative elevation of the test, sharpness of the anterior sulcus and position of the apical disc. At Jebel Rawdah the lowest beds contain the flattest species, H. persicus Cotteau and Gauthier, and this species has a relatively broad and shallow anterior sulcus. Above this level, and possibly co-occurring for a short interval, comes H. arabicus, a species that is equally flat in profile but with a narrower and more sharply defined anterior sulcus. Higher beds contain an elevated species of Hemipneustes, H. compressus, and the relative elevation of the test increases up the succession, so that in the upper half of the Simsima Formation, Hemipneustes compres- sus begins to develop a strong apical elevation and narrower frontal groove, resembling the boreal species H. striatoradi- atus. The oldest species, Hemipneustes persicus, occurs (as H. sardanyolae Vidal) in the late Campanian of Spain. H. arabicus is endemic to this area, but the highest species, H. compressus, is known from the Maastrichtian of the Mari Hills, West Pakistan, where, unfortunately, the precise dating of the beds remains uncertain. At Jebel Rawdah, the replace- ment of H. persicus by H. arabicus seems to take place within the Upper Maastrichtian, according to the evidence of the ammonites and inoceramids. The second biostratigraphically useful lineage comprises the two species of Hattopsis. At Jebel Faiyah, two species succeed one another, with H. paucituberculatus predating H. sphericus. This is useful for local correlation, since it is H. paucituberculatus that occurs at the base of the Simsima Formation at Jebel Buhays and H. sphericus that occurs near the base of the Simsima Formation at Jebel Rawdah. This then implies that the base of the Simsima Formation is not strictly contemporaneous between jebels. Finally, it is noteworthy that the basal bed in the Loftusia- rich unit at Jebel Huwayyah yields a distinct suite of echinoids seen at no other jebel, including the holectypoid Coptodiscus. Coptodiscus is known from the late Campanian of Arabia LATE CRETACEOUS CARBONATE PLATFORM FAUNAS 111 Rawdah 2 . roo Faiyah Poh [a0 cr rj See Inferred sea-level change = oo deep <> shallow ‘=o ‘aoe Rawdah 3 Humeayga Cot Coo J] = ecos fof =] 1S saa se roo =o aww as Buhays == = ro = ‘ot Siliciclastic sands, grits and conglomerates HE Poorly sorted bioclastic packstones Poorly sorted orbitoid-rich packstones and wackestones Well-sorted bioclastic packstones Red-weathering silt-rich wackestone Fig. 13 Tentative correlation between jebels with inferred sea-level curves. | (Kier 1972) and from a presumed similar ‘Senonian’ horizon / in southern Iran. Global Sea-level curves. In broad terms the stratigraphic | Successions in the five jebels described here follow the same pattern, but in detail, precise correlation remains difficult, due to rapid shifts in facies and bed thickness across even small areas (Figs 6, 13). Sedimentation upon the Semail _Nappe and Hawasina Group follows a typical transgressive pattern, often with massive boulder beds of serpentinite at the base overlain by arenitic and then calc-arenitic sediments, tall deposited in shallow or nearshore marine environments. The lower clastic facies of conglomerates, sands and gravels jare commonly included in the Qahlah Formation, whereas tiie sediments above, which are essentially bioclastic cal- Carenites, form the Simsima Formation. There is no reason to | suppose, however, that the change from boulder beds and essentially quartz clastics upwards into calcarenites took place at the same time in all the sections we have examined. The general deepening and shallowing cycles that can be recognized within individual sections allow a means of corre- lation. In all sections the initial coarse clastics of the Qahlah Formation are abruptly terminated, presumably marking the submersion of the ophiolitic complex within the region. This allowed carbonate production to dominate. For most of the Simsima Formation deposition appears to have more-or-less kept pace with subsidence so that sea-level remained around wave-base. Furthermore, at Jebel Buhays, Jebel Faiyah and Jebel Rawdah there are conspicuous reddened beds and partings that probably contain land-derived iron oxides, which are a common constituent of lateritic soils formed on exposed 72 ophiolite. These red partings and thin beds presumably formed when lateritic soils were eroded and periodically flushed out to sea in run-off after heavy rains. Some may mark minor fluctuations in sea-level. They are potentially traceable over a wide area and may be important for local correlation. The problem is that there are not the same number of iron-enriched bands currently recognized in the different sections, and it becomes problematic as to which of several alternative bands should be correlated. A tentative correlation is presented in Fig. 13. However, mineralogical and geochemical analyses of these bands is needed to estab- lish whether any have distinctive signatures. The Cretaceous part of the section at Jebel Rawdah is topped by a limestone conglomerate apparently made up of eroded fragments of the Simsima Formation and including large chunks of rudists. This is in turn overlain by Lower Tertiary limestones. The lack of rounding and poor sorting of the clasts in the limestone conglomerate are probably indica- tive of subaerial erosion and deposition but we have no way of more accurately dating this period of emergence. It lies somewhere between late Maastrichtian and early Tertiary. Flexer & Reyment (1989) identified two late Cretaceous transgressive events affecting the Arabo-Nubian shield: one in the late Campanian-early Maastrichtian, and the other in the late Maastrichtian. Although local tectonic events, associ- ated with the ophiolite emplacement, may have had a pro- found effect on the local sea-level signature, it is tempting to associate the initial submergence of the ophiolitic islands and the sands and conglomerates of the Qahlah Formation with the late Campanian-early Maastrichtian transgression, and the later flooding of these beds and initiation of carbonate platform deposition with the second of these major sea-level transgressions in the late Maastrichtian. The end of the Cretaceous saw a major drop in sea-level. Summary. Combining the evidence from ammonites, inoc- eramid bivalves, echinoids and global sea-level curves we conclude that the basal siliciclastic beds of the Qahlah Forma- tion are of latest Campanian age. At Jebel Huwayyah, the Loftusia-rich levels of the Qahlah Formation probably encompass Lower to early Upper Maastrichtian. Finally, the Simsima Formation appears to be early Upper Maastrichtian, fresvillensis Zone or later. 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Campanian and Maastrichtian sphenodiscid ammonites from southern Nigeria. Bulletin of the British Museum (Natural History), Geology Series, 36 (4): 303-332. APPENDIX Macrofossils collected in the United Arab Emirates/Oman border region. Echinoids identified by A.B. Smith, nautiloids, bivalves and gastropods by N.J. Morris, P.W. Skelton and R.J. Cleevely, ammonites by W.J. Kennedy, corals by J. Darrell and B.R. Rosen, bryozoans by P.D. Taylor and brachiopods by E.F. Owen. Locality details are given in the main text: bed numbers refer to those shown in figures 5-11. Numbers in square brackets after each name refer to the number of specimens collected. Jebel Buhays, Section 1 Top of Bed 4. Echinoids: Arnaudaster cylindriformis sp. nov. [1]; Circopeltis? emiratus sp. nov. [1]. Ammonite: Pachydiscus dossan- toi (Maury) [1]. Bed 10. Echinoids: Hemipneustes sp. [1]; Coenholectypus cf. baluchis- tanensis (Noetling) [1]. Bed 11. Bivalves: Dictyoptychus morgani (Douvillé) [2]; Gastropods: Acteonella crassa (Dujardin) [6]. Bed 12, base. Echinoids: Goniopygus arabicus sp. nov. [1]; Hattopsis paucituberculatus sp. nov. [1 fragment]. Coral: Hydnophoraraea sp. [1]. Bivalves: Scabrotrigonia sp. [3]. Bed 15. Echinoid: Codiopsis lehmannae sp. nov. [1]. Bivalve: Hippu- rites cf. cornucopiae Defrance [6]. Lowest part of Simsima Formation (beds 4-10) — mostly collected loose. Echinoids: Rhabdocidarid, gen. et sp. indet. [1]; Prionoci- daris morgani (Gauthier) [2]; cidarid spines [2]; Heterodiadema buhaysensis sp. nov. [2]; Orthopsis miliaris (d’ Archiac) [10]; Sale- nia nutrix Peron & Gauthier [5]; Goniopygus arabicus sp. nov. [22]; Glyphopneustes hattaensis Ali [41]; Hattopsis paucitubercula- tus sp. nov. [11]; Noetlingaster paucituberculatus (Noetling) [3]; Phymosoma hexoaporum Lambert [13]; Actinophyma spectabile Cotteau & Gauthier [4]; Plistophyma asiaticum Gauthier [3]; Circopeltis? emiratus sp. nov. [1]; Coenholectypus inflatus (Cotteau & Gauthier) [8]; Coenholectypus cf. baluchistanensis (Noetling) [3]; ‘Globator’ bleicheri (Thomas & Gauthier) [96]; Conulus dou- villei (Cotteau & Gauthier) [19]; Vologesia rawdahensis Ali [3]; Pygurostoma morgani Cotteau & Gauthier [26]; Petalobrissus cf. setifensis (Cotteau) [1]; Nucleopygus magnus sp. nov. [27]; Arnau- daster cylindriformis sp. nov. [5]; Hemipneustes sp. indet. (frag- ments) [3]; Hemiaster hattaensis Ali [4]. Bivalves: Arca sp. [1, bivalved]; Cucullaea sp. A [3, 2 bivalved]; ‘Modiolus’ cf. capitatus Zittel [3, bivalved]; Myzilus sp. nov? [1, bivalved]; Modiolus aff. typicus Forbes [3, bivalved]; Neithea regularis (Schlotheim) [33]; Spondylus sp. C [1]; Spondylus sp. D [1]; Ctenoides aff. scaberrima (Stolizcka) [14]; ?Osculopha sp. [1]; ?Gyropleura sp. [1]; ?Pycn- odonte uncinella (Leymerie) [8]; Amphidonte pyrenaicum (Leyme- tie) [2]; Agerostrea ungulata (Schlotheim) [4]; Eligmidae [6, bivalved]; ?Tellinella sp. [1]; Glabrobournonia arabica Morris & Skelton sp. nov. [12]; Tancrediidae cf. Tatella sp., sp. nov. [1]; Clavagella cf. semisulcata Forbes [2, bivalved]; Pholadomya sp. B [1, bivalved]. Gastropods: Bathrotomaria cf. verdachellensis (Forbes) [2]; Calliomphallus sp. [1]; Angaria sp. [1]; ‘Tectus’ ex. gr. rozeti (d’Archiac) [7]; Umboniinae nov. gen. [9]; ?Rhabdocon- cha sp. [1]; Turritellidae [2]; Pyrazus sp. [1]; Campanile sp. [8]; Tylostoma incerta (Forbes) [9]; Naticidae [1]; Strombidae, gen. nov., cf. crassicostatus (Noetling) [1]; Strombidae, gen. nov. [1]; Strombacea [3]; ?Ovula expansa dArchiac & Haime [4]; Cypraeidae [1]; ?Tonnacea [1]; ‘Volutilithes’ dubia Noetling [7]; ?Volutoderma sp. [4]; Caricella sp. [1]; Volutidae A [1]; Volutidae B [2]; Volutidae C [2]; Volutidae D [3]; Neogastropoda indet. [1]; Acteonella crassa (Dujardin) [4]; Acteonella caucasica Zekeli cf. styriaca Kollman [3]; Acteonella sp. cf. A. borneensis Nuttall & Leong [19]; Acteonella caucasica Zekeli subsp. nov. [7]; Acteonella crassa (Dujardin) [9]; Neocylindrites cf. minutus Sohl [3]; Neocylin- drites sp. [1]; opisthobranch [2]. Corals: Polytremacis sp. [2]; Aspidas- traea sp. [1]; Cunnolites sp. [4]; Diploctenium sp. [3]; Hydnophoraraea sp. [2]; Paraplacocoenia orbignyana (Reuss) [1]; cerioid colony [1]; phaceloid colony [1]; solitary form [1]. Stromatoporoid: massive stromatoporoid (bored by bivalves) [1]. Sponge: chaetetid [1]. Bryo- zoan: ‘Onychocella’ sp. [1]. Decapod crustaceans: Carcineretes sp. [3 carapaces, 5 limb segments]; crab carapace, species A [1]; crab carapace, species B [2]; claw, indet. [1]. Jebel Buhays Section la Small hill 150 m to south of Jebel Buhays 1. Collected from scree, in rock fall from lowest 4 m of Simsima Formation (beds are steeply dipping and all loose material must be derived from the basal beds here). Echinoids: Heterodiadema buhaysensis sp. nov. [1]; Orthopsis miliaris (d’Archiac) [7]; Salenia nutrix Peron & Gauthier [2]; Goniopygus arabicus sp. nov. [7]; Glyphopneustes hattaensis Ali [9]; Phymosoma hexoaporum Lambert [3]; Actinophyma spectabile Cot- teau & Gauthier [1]; Plistophyma asiaticum Gauthier [3]; Coenholec- typus inflatus (Cotteau & Gauthier) [3]; ‘Globator’ bleicheri (Thomas 114 & Gauthier) [occurs]; Conulus douvillei (Cotteau & Gauthier) [occurs]; Vologesia rawdahensis Ali [2]; Petalobrissus linguiformis (Peron & Gauthier) [1]; Nucleopygus magnus sp. nov. [9]; Hemip- neustes sp. [1]; Hemiaster hattaensis Ali [3]. Bivalves: Cucullaea sp. A [4]; Barbatia sp. [1]; Lyriochlamys ternatus (Minster) [6]; Neithea regularis (Schlotheim) [6]; Agerostrea ungulata (Schlotheim) [4]; ?Pycnodonte uncinella (Leymerie) [1, bivalved]; Scabrotrigonia sp. [8]; Dictyoptychus morgani Douvillé [1, lid]; Glabrobournonia ara- bica Morris & Skelton sp. nov. [2]; Clavagella cf. semisulcata Forbes [2]. Gastropods: Tylostoma incerta (Forbes) [2]; ?Strombidae, gen. nov. [1]; Pseudomelania [Trajanella] sp. cf. conica (Stolizcka) [2], Acteonella crassa (Dujardin) [13]; Acteonella cf. laevis laevis (Sow- erby) [6]; Acteonella cf. borneensis Nuttall and Leong [4]; Neocylin- drites sp. cf minutus Stolizcka [1]. Cephalopods: Cimomia aff. sowerbyana (d’Orbigny) [1]; Deltoidonautilus salisfilius sp. nov. [1]. Corals: Polytremacis sp. [1]; Favia sp. cf. ‘Diplocoenia’ klogsdorfen- sis Trauth [1]; Barysmilia irregularis (Reuss) [1]; Cunnolites sp. [1]; Diploctenium sp. [1]; cerioid colony [1]. Sponge [1]. Jebel Buhays 2 Collections made from the scree-slope. Here the bedding is very steep and material is derived from the lowest few metres of section only. Echinoids: Salenia nutrix Peron & Gauthier [1]; Goniopygus arabicus sp. nov. [4]; Coenholectypus inflatus (Cotteau & Gauthier) [1]; ‘Globator’ bleicheri (Thomas & Gauthier) [3]; Conulus douvillei (Cotteau & Gauthier) [3]; Pygurostoma morgani Cotteau & Gauthier [2]. Bivalves: Dictyoptychus morgani (Douvillé) [1]; Glabro- bournonia arabica Morris & Skelton, sp. nov. [16]; Ammonite: Libycoceras sp. [1]. Jebel Buhays 3 Basal limestone (lowest Im). Echinoids: Goniopygus arabicus sp. nov. [4]; Nucleopygus magnus sp. nov. [1]. Above first major red-weathering siltstone level. Echinoids: Circopel- tis emiratus sp. nov. [2]. Loose. Echinoids: Salenia nutrix Peron & Gauthier [2]; Hattopsis paucituberculatus sp. nov. [2]; Phymosoma hexoaporum Lambert [1]; ‘Globator’ bleicheri (Thomas & Gauthier) [1]. Bivalves: Hippu- rites cornucopiae Defrance [1]; ?Biradiolites aff. baylei Toucas [2]. Jebel Thanais Lowest 4 m of Simsima Formation. Echinoids: Heterodiadema buhay- sensis sp. nov. [1]; Orthopsis miliaris (d’Archiac) [1]; Salenia nutrix Peron & Gauthier [2]; Goniopygus arabicus sp. nov. [2]; Glyphop- neustes hattaensis Ali [6]; Hattopsis sp. [1]; Phymosoma hexoaporum Lambert [3]; Plistophyma asiaticum Gauthier [2]; Coenholectypus cf. baluchistanensis (Noetling) [2]; ‘Globator’ ble- icheri (Thomas & Gauthier) [5]; Conulus douvillei (Cotteau & Gauthier) [8]; Vologesia rawdahensis Ali [1]; Pygurostoma mor- gani Cotteau & Gauthier [1]; Nucleopygus magnus sp. nov. [4]; Hemiaster hattaensis Ali [1]. Bivalves: ?Barbatia sp. A, cf. B. morgani (Douvillé) [2, bivalved]; ‘Modiolus’ aff. typicus Forbes [1, bivalved]; Neithea regularis (Schlotheim) [2]; Pycnodonte vesicu- laris (Lamarck) [1, bivalved]. Gastropods: Acteonella sp. cf. A. borneensis Nuttall & Leong [1]; Acteonella caucasica Zekeli subsp. cf. styriaca Kollman [1]; Acteonella caucasica caucasica Zekeli [1]. Corals: Cunnolites sp. [1]; Diploctenium sp. [1]; Moltkia isis Steenstrup [1]; Paraplacocoenia orbignyana (Reuss) [1]; coarse meandroid colony [1]. Bryozoan: ‘Onychocella’ sp. [1]. Decapod crustacean: limb segments [3]. Upper beds (equivalent to bed 15 of Jebel Buhays 1). Bivalves: Hippurites cornucopiae Defrance [1]. Loose in scree. Bivalves: Cucullaea sp. A [2 ,1 bivalved]; ‘Modiolus’ A.B. SMITH, N.J. MORRIS, W.J. KENNEDY AND A.S. GALE aff. typicus Forbes [1, bivalved]; Chlamys dujardeni (Roemer) [1]; Neithea regularis (Schlotheim) [3]; Spondylus sp. E [1]; Ctenoides aff. scaberrima (Stolizcka) [2]; ?Amphidonte cf. pyrenaicum (Leymerie) [1]; ?Osculopha sp. [1]; Lucinidae gen. indet., sp. B [1, bivalved]; Dictyoptychus morgani (Douvillé) [1]; Glabrobournonia arabica Morris & Skelton, sp. nov. [1]; Colveraia variabilis Kling- hardt [1]; Vaccinites vesiculosus (Woodward) [4]; Lapeirousia sp. [2]; Pholadomya sp. B [1, bivalved]. Gastropods: Tylostoma incerta (Forbes) [1]; Acteonella crassa (Dujardin) [5]. Jebel Aqabah Excellent outcrop of the basal part of the marine sequence resting directly on serpentinized ultramafics. Bed 1. Echinoids: Hattopsis paucituberculatus sp. nov. [1]; ‘Globator’ bleicheri (Thomas & Gauthier) [2]; Faujasia eccentripora Lees [6 juveniles]. Ammonite: Glyptotoxoceras sp. [1]. Loose. Bivalves: ?Pycnodonte uncinella (Leymerie) [2]; Dictyopty- chus morgani (Douvillé) [1]; ?Biradiolites aff. baylei Toucas [3]. Gastropods: Acteonella crassa (Dujardin) [2]. Cephalopod: Cimo- mia aff. sowerbyana (d’Orbigny) [1]. Coral: Cunnolites sp. [1]. Jebel Huwayyah, Section 1 Bed 1. Bivalves: ?Acutostrea sp. [ca. 10]. Bed 3. Bivalves: ?Acutosrea sp. [abundant]. Bed 7. Stromatoporoid [1]. Bed 9. Echinoids: Faujasia eccentripora Lees [6]; Hemiaster sp. cf. H. hattaensis Ali [1]. Bivalves: Spondylus sp. A [1]; Vaccinites vesicu- losus (Woodward) [5]; ?Granocardium sp. [1]. Gastropods: ?Tec- tus sp. [1]; Campanile sp. [1]; Amauropsina aff. bulbiformis (J. de C. Sowerby) [1]; ‘Amputllina aff. splendida’ (Deshayes) [3]. Corals: Cunnolites sp. [4]; cerioid colony (with Lithophaga borings) [1]. Ammonite: Pachydiscus sp. [1]. Beds 10-11. Echinoids: Faujasia eccentripora Lees [33]; Hemiaster sp. cf. H. hattaensis Ali [1]. Bivalves: Endocostea cf. bebahoaensis (Sornay) [1]; Spondylus sp. A [2]; Plicatula hirsuta Coquand [4]; ?Acutostrea sp. [6]; Glabrobournonia arabica Morris & Skelton, sp. nov. [1]; Pholadomya? sp. B [1]. Gastropods: cf. Tylostoma incerta (Forbes) [1]. Corals: Cunnolites sp. [2]; Paraplacocoenia orbignyana (Reuss) [1]; large trochoid solitary [1]; large flabellate solitary [1]; cerioid colonies (5 genera) [7]. Ammonites: Pachydis- cus dossantoi (Maury) [1]; Lewyites ambindense (Collignon) [1]; Nostoceras (Nostoceras) major Kennedy & Cobban [1]. Beds 9-11 undifferentiated. Bivalves: Torreites sanchezi (Douvillé) [2] Durania form A [1]. Bed 13. Bivalves: Durania spp. [fragments]. Beds 14/15. Echinoids: Coenholectypus inflatus (Cotteau & Gauthier) [1]; ‘Globator’ bleicheri (Thomas & Gauthier) [54]; Pygurostoma morgani Cotteau & Gauthier [3]; Hemipneustes sp. [1]. Bivalves: Cucullaea sp. A [1]; Lyriochlamys ternatus (Minster) [7]; Neithea | regularis (Schlotheim) [8]; Spondylus sp. A. [2]; Dictyoptychus morgani (Douvillé) [1]. Gastropods: Execochirsus sp. [1]; Campa- | nile sp. [1]; Cerithiidae [1]; Tylostoma incerta (Forbes) [6]; Strom- bacea [1]; ?Aporrhaidae [1]; Neogastropods [2]. Ammonite: Pachydiscus dossantoi (Maury) [1]. Coral: Cunnolites sp. [1]. Bed 16. Bivalves: Pycnodonte vesicularis (Lamarck) [2]; Agerostrea ungulata (Schlotheim) [12, bivalved]. Bed 17. Bivalves: Lyriochlamys ternatus (Minster) [1]; Neithea regularis (Schlotheim) [2]; Scabrotrigonia sp. [4]; Durania sp. 2]. Gastropods: Bathrotomaria sp. [1]. Bed 18, towards top of section. Echinoids: Proraster geayi Cottreau {7]. Loose. Echinoid: Mecaster sp.? [1]. Bivalve: ?‘Inoceramus’ sp. [1]. ——————— - ———————— | | LATE CRETACEOUS CARBONATE PLATFORM FAUNAS Jebel Huwayyah, Section 2 Bed 1. Echinoids: G/yphopneustes hattaensis Ali [1]; Coptodiscus magniproctus sp. nov. [2]. Beds 3-5. Echinoids: cidarid spine [1]; Orthopsis sp. [1]; Salenia microprocta sp. nov. [1]; ?Hattopsis sp. [1]; Faujasia eccentripora Lees [1]; Hemiaster sp. cf. H. hattaensis Ali [3]; Mecaster sp. [4]; Proraster geayi Cottreau [2]. Bivalves: Dictyoptychus morgani (Douvillé) [1]; Corals: Cunnolites sp. [27]; Diploctenium sp. [4]; Placosmilia sp. [22]. Bryozoans: cf. Euritina lata Canu [1]; ‘Wilber- topora’ sp. [1]. Decapod crustacean: pincer [1]. Brachiopods: terebratulid gen. et sp. nov. [3]. Ammonite: Nostoceras sp. [1]. Bed 7. Bivalves: Vaccinites vesiculosus (Woodward); Corals: Actina- cis sp. [3]; Cladocora ?humilis (Michelin) [7]; ?Aspidastraea sp. [1]; Astraraea sp. [3]; Calamophylliopsis simonyi (Reuss) [1]; Parapla- cocoenia orbignyana (Reuss) [1]; cerioid colonies (3 genera) [4]; thamnasteroid colonies [4]; coarse meandroid colonies [5]; elongate-oval flat solitary corals [3]; flabellate solitary corals [2]; turbate solitary corals [10]; cylindrical solitary corals [3]. Ammo- nite: Neancycloceras sp. [1]. Loose in scree, but derived from Loftusia beds, beds 2—7 unless otherwise stated. Echinoids: Prionocidaris? emiratus sp. nov. [1]; Coenholectypus inflatus (Cotteau & Gauthier) [3] (probably derived from Simsima Formation above?). Bivalves: Endocostea cf. bebahoaensis (Sornay) [1]; Lyriochlamys ternatus (Minster) [8]; Neithea regularis (Schlotheim) [19]; Neithea (Neithella) notabilis (Munster) [1]; Spondylus sp. A [49, 19 bivalved]; Spondylus sp. B [12, 4 bivalved]; Plicatula hirsuta Coquand [23]; ?Osculopha sp. [4, 1 bivalved]; Amphidonte pyrenaicum (Leymerie) [10, 2 bivalved]; Agerostrea ungulata (Schlotheim) [1, bivalved]; ?Eligmidae indet. [1]; Ctenoides sp. [2]; Chama noetlingi [7]; Biradiolites aff. baylei Toucas [1]; Glabrobournonia arabica Morris & Skelton, sp. nov. [10]; Semailia smithi Morris & Skelton, sp. nov. [1]; Vaccinites vesiculosus (Woodward) [25]; Pholadomya sp. A, cf. indica Noet- ling [2]; Pholadomya sp. B [1, bivalved]. Gastropods: ?Angaria sp. [5]; Turbinidae sp. [5]; Turritellidae sp. [10]; ?Pyrazus sp. [3]; Cerithtidae sp. [1]; ?Perrisoptera sp. [7]; ?Naticidae sp. [7]; ?Fasci- olariidae sp. [1]; Volutidae sp. A [1]; Volutidae sp. B [1]; ?Mitridae sp. [1]; neogastropods [2]; Acteonella sp. [1]. | Jebel Rawdah, Section 1 Bed 2. Echinoid: Petalobrissus cf. setifensis (Cotteau) [1]. Bivalves: ?*Modiolus’ cf. capitatus Zittel [1, bivalved]; Cucullaea sp. A [11]; Lyriochlamys ternatus (Minster) [2]; Neithea regularis (Schlo- theim) [7]; Pycnodonte vesicularis (Lamarck) [1]; ?Pycnodonte uncinella (Leymerie) [7]; Agerostrea ungulata (Schlotheim) [1]; Indet. ribbed oyster [1]; Scabrotrigonia sp. [62, 7 bivalved]; Durania spp. [2]; Glabrobournonia arabica Morris & Skelton [4]; Tancrediidae, cf. Tatella, sp. nov. [2, bivalved]. Gastropods: Acteonella sp. cf. borneensis Nuttall & Leong [4]. Corals: Cunno- lites sp. [7]; Diploctenium sp. [1]. Bed 3 (lower part) and top of Bed 2. Echinoids: Orthopsis miliaris (@Archiac) [2]; Salenia nutrix Peron & Gauthier [2]; Hattopsis sphericus Ali [19]; Noetlingaster paucituberculatus (Noetling) [5, fragments]; Coenholectypus cf. baluchistanensis (Noetling) [1]; ‘Globator’ bleicheri (Thomas & Gauthier) [49]; Faujasia eccentri- pora Lees [5]; Zuffardia morgani (Cotteau & Gauthier) [4]; Petalobrissus cf. setifensis (Cotteau) [17]; Phymechinus? sp. [frag- ment]. Bivalves: ?Barbatia sp. A, cf. B. morgani Douvillé [1, bivalved]; Cucullaea sp. A [4, 3 bivalved]; ?‘Mytilus’ nitens Forbes [1, bivalved]; ‘Modiolus’ cf. capitatus Zittel [5, 4 bivalved]; Pinna _ sp. [3]; Endocostea (Selenoceramus) semaili sp. nov. [1]; Lyrio- chlamys ternatus (Minster) [3]; Neithea regularis (Schlotheim) [23]; Limidae [1]; Plicatula hirsuta Coquand [1]; Pycnodonte vesicularis (Lamarck) [6]; Pycnodonte vesicularis (Lamarck) var. hippopodium Nillson [1]; ?Pycnodonte uncinella (Leymerie) [10, 4 115 bivalved]; ?Amphidonte cf. pyrenaicum (Leymerie) [2]; Scabrot- rigonia sp. [6, 2 bivalved]; Lucinidae sp. [1]; ?Plagyoptychus sp. [1]; Dictyoptychus morgani (Douvillé) [10]; Glabrobournonia ara- bica Morris & Skelton, sp. nov. [22]; Durania spp. [6 fragments]; ?Trapeziidae [1]; ?Mesocallista sp. [5, bivalved]; Tellininae [1]; ?Tancrediidae, cf. Tatella, sp. nov. [2]; Clavagella cf. semisulcata Forbes [6]; ?Brechites cf. aspergilloides (Forbes) [1]; Pholadomya sp. B [1]. Gastropods: Calliomphalus sp. [4]; ?Angaria sp. [2]; ?Tectus sp. [3]; ?Pseudoliotina sp. [1]; ?Umboniinae, gen. nov. [64]; euomphalid gen. nov. [6]; ?Rhabdoconcha sp. [1]; ?Turritel- lidae [2]; Vermetidae or Siliquariidae [1]; Cerithiidae [7]; Cypraeidae, sp. [4]; ?Cypraeidae. [3]; ?Volutacea [1]; Ampullina aff. ‘splendida’ (Deshayes) [4]; Tylostoma incerta (Forbes) [2]; Mesogastropod [2]; ?Anchura sp. [3]; 7Aporrhaidae [1]; Hipponi- cidae [3]; ?Fasciolariidae [1]; ?Volutidae [16]; ?Buccinacea [7]; Lyria sp. [8]; ?Lyria sp. [2]; Acteonella sp. cf. borneensis Nuttall & Leong [30]; Acteonella laevis laevis (J. de C. Sowerby) [1]; Acteonella sp. [4]; Neocylindrites sp. [2]; oliviform opisthobranch [1]; ?Scaphander sp. [1]; bullaeform opisthobranchs [2]; elongate opisthobranch [1]. Cephalopods: Cimomia cf. sowerbyana (d’Orbigny) [1]; Brahmaites (Anabrahmaites) vishnu (Forbes) [1]. Corals: Cunnolites sp. [42]; Diploctenium sp. [9]; Aspidastraea sp. [2]. Decapod crustacean: pagurid claw [1]. Bed 3 (upper part). Echinoids: Hattopsis sphericus Ali [3]; phymo- somatid fragment [1]; Conulus douvillei (Cotteau & Gauthier) [1]; Zuffardia morgani (Cotteau & Gauthier) [1]; Hemipneustes sp. [3, fragments]. Bivalves: ?Chlamys sp. [1]; ?Ctenoides sp. [1]; Cardi- idae [3]; ?Fragum sp. [2]. Gastropods: ?Umboniinae, gen. nov. [6]; ?Rhabdoconcha sp. [2]; Campanile sp. [1]; ?Cerithiidae indet. [3]; ?‘Ampullina aff. splendida’ (Deshayes) [1]; ?Tylostoma incerta (Forbes) [4]; ?Naticidae sp. [2]; Cypreidae spp. [3]; Lyria sp. [2]; elongate, fusiform genus [1]. Bed 4. Echinoids: Glyphopneustes hattaensis Ali [1]; Hattopsis spheri- cus Ali [15]; Hemipneustes arabicus Ali [3]. Bivalves: Neithea sp. [1]; Durania form B [1]; ?Pycnodonte uncinella (Leymerie) [7]; cf. Fragum sp. [3]. Gastropods: ?Angaria sp. [1]; ?Trochacea, euomphalid gen. nov. [3]; ?Umboniinae, gen. nov. [4]; Turritel- lidae [1]; ?Cerithiidae, sp. [2]; Cypraeidae, sp. [1]; Caricella sp. [2]. Cephalopod: Cimonia aff. sowerbyana (d’Orbigny) [1]. Deca- pod crustacean: small spinose pincers [2]. Bed 4 (top). Echinoids: Salenia nutrix Peron & Gauthier [2]; Hattopsis sphericus Ali [29]; Noetlingaster emiratescus Ali [1]; Noetlingaster sp. [1, fragment]; Phymechinus perplexus sp. nov. [2]; ‘Globator’ bleicheri (Thomas & Gauthier) [15]; Faujasia eccentripora Lees [2]; Hemipneustes persicus Cotteau & Gauthier [1, fragment]. Bivalves: ?‘Mytilus’ nitens Forbes [1, bivalved]; ‘Modiolus’ aff. typicus Forbes [2, bivalved]; Endocostea (Selenoceramus) semaili sp. nov. [2]; Neithea sp. [6]; ?Pycnodonte uncinella (Leymerie) [16]; Sca- brotrigonia sp. [5, 1 bivalved]; Gyropleura sp. [4]; ?Tellinella sp. [1, bivalved]; ‘Quenstedtiidae’, gen. nov. [1]; Clavagellidae, gen. nov. [3]. Gastropods: ?Umboniinae, gen. nov. [27]; Turritellidae [1]; ?Rhabdoconcha sp. [2]; Cerithiidae [1]; Strombidae, gen. nov. [3]; Buccinacea sp. [1]; Lyria sp. [2]; ?Volutomorpha sp. [2]; Volutidae, cf. Melo [1]; ?Cancellariidae [1]; 7Neogastropods indet. [5]. Cephalopod: Cimonia cf. sowerbyana (d’Orbigny) [1]. Corals: Cunnolites sp. [4]. Collected in scree or from slipped blocks. Echinoids: Orthopsis miliaris (d’Archiac) [4]; Salenia nutrix Peron & Gauthier [1]; Noetlingaster sp. [1, fragment]; ‘Globator’ bleicheri (Thomas & Gauthier) [2]; Faujasia eccentripora Lees [6]; Petalobrissus sp. [2]; Hemipneustes compressus Noetling [1]; Hemipneustes sp. [2, frag- ments]. Bivalves: Cucullaea sp. A [2, 1 bivalved]; Neithea regularis (Schlotheim) [1]; Pycnodonte vesicularis (Lamarck) [2]; ?Pycn- odonte uncinella (Leymerie) [2]; Agerostrea ungulata (Schlotheim) [3]; Scabrotrigonia sp. [14]; Glabrobournonia arabica Morris & Skelton, sp. nov. [1]; Radiolitidae [1]; Tancrediidae, cf. Tatella, sp. nov. [1, bivalved]. Gastropods: ‘Tectus’ sp. [1]; Turbinidae [1]; 116 ?2Umboniinae, gen. nov. [1]; ?Turritellidae indet. [1]; ‘Ampullina aff. splendida’ (Deshayes) [2]; Tylostoma incerta (Forbes) [2]; 2Naticidae sp. [1]; ?Cimolithium sp. [1]; ?Aporrhaidae sp. [2]; Lyria sp. [3]; Acteonella sp. cf. borneensis Nuttall & Leong [8]; opisthobranch [1]. Corals: Cunnolites sp. [66]; Diploctenium sp. [1]. Ammonites: Brahmaites (Anabrahmaites) vishnu (Forbes) [1]. Bed 6. Echinoids: Coenholectypus inflatus (Cotteau & Gauthier) [8]; ‘Globator’ bleicheri (Thomas & Gauthier) [1]; Hemipneustes perst- cus Cotteau & Gauthier [2]; Hemiaster hattaensis Ali [3]. Rebel Rawdah, Section 2 Bed 4: 60-120 cm above basal conglomerate. Echinoids: Goniopygus arabicus sp. nov. [1]; Plistophyma asiaticum Gauthier [1]; Echino- tiara perebaskinei Lambert [6]; Faujasia eccentripora Lees [14]; Petalobrissus rawdahensis sp. nov. [3]. Bivalve: Durania [1]. Gas- tropod: ?‘Ampullina aff. splendida’ (Deshayes) {2]. Decapod crus- tacean: limb segments [4]. Bed 5. Echinoids: Faujasia eccentripora Lees [2]. Beds 6-8. Echinoids: Salenia nutrix Peron & Gauthier [10]; Goniopy- gus arabicus sp. nov. [6]; Codiopsis lehmanni sp. nov. [1]; Phy- mechinus perplexus sp. nov. [6]; ?Phymechinus sp. [1]; Echinotiara perebaskinei Lambert [58]; Coenholectypus inflatus (Cotteau & Gauthier) [1]; ‘Globator’ bleicheri (Thomas & Gauthier) [1]; Faujasia eccentripora Lees [92]; Petalobrissus rawdahensis sp. nov. [4]; Petalobrissus cf. setifensis (Cotteau) [9]; Hemipneustes arabicus Ali [1]. Bivalve: ?Ctenoides sp. [1]; Durania form B [4]. Coral: Cunnolites sp. [1]. Decapod crustacean: limb and claw segments [4]. Bed 10. Echinoids: Goniopygus arabicus sp. nov. [1]; Glyphopneustes hattaensis Ali [2]; Faujasia eccentripora Lees [1]; Petalobrissus sp. [2]. Bivalves: Durania cf. apula (Parona) [1]. Gastropod: Stromba- cea [1]. Bed 11: Cucullaea bed immediately overlying orange-weathering band. Echinoids: Noetlingaster paucituberculatus (Noetling) [1, fragment]. Bed 11 (middle, ca. 1 m above orange-weathering band). Echinoids: Goniopygus arabicus sp. nov. [1]; Faujasia eccentripora Lees [6]; Petalobrissus sp. [1]; Nucleopygus magnus sp. nov. [4]; Stigmatopy- gus pulchellus sp. nov. [2]; Hemipneustes arabicus Ali [4]. Gastro- pods: ?Umboniinae, gen. nov. [1]; Campanile sp. [1]; Pyrazus sp. [1]; Strombidae, gen. nov. [1]; Volutidae sp. [3]. Bed 11: 2.5 m above base. Echinoids: Echinotiara perebaskinet Lambert [18]; Coenholectypus cf. baluchistanensis (Noetling) [1]; Faujasia eccentripora Lees [49]; Petalobrissus sp. [40]. Gastropod: ‘Ampullina aff. splendida’ (Deshayes) [1]. Corals: Cunnolites sp. [3]; massive meandroid colony [1]. Bed 11 (unspecified). Echinoids: Salenia nutrix Peron & Gauthier [2]; Glyphopneustes emiratensis Ali [5]; Hattopsis sphericus Ali [2]; Noetlingaster paucituberculatus (Noetling) [4]; Phymechinus? per- plexus sp. nov. [1]; Echinotiara perebaskinei Lambert [3]; Coen- holectypus cf. baluchistanensis (Noetling) [8]; ‘“Globator’ bleicheri (Thomas & Gauthier) [10]; Faujasia eccentripora Lees [46]; Zuffar- dia morgani (Cotteau & Gauthier) [41]; Petalobrissus rawdahensis sp. nov. [247]; Petalobrissus cf. setifensis (Cotteau) [124]. Bivalves: ‘Modiolus’ aff. typicus Forbes [1, bivalved]; Cucullaea sp. A [8, bivalved]; Pholadomya sp. C. cf. P. connectans Forbes [22, bivalved]. Coral: Cunnolites sp. [1]. Bed 13. Echinoids: Salenia nutrix Peron & Gauthier [2]; Glyphop- neustes hattaensis Ali [1]; ‘Globator’ bleicheri (Thomas & Gauthier) [1]; Petalobrissus rawdahensis sp. nov. [230]; Petalobris- sus cf. setifensis (Cotteau) [4]; Nucleopygus magnus sp. nov. [3]. Bivalves: ‘Modiolus’ aff. typicus Forbes [1, bivalved]; Dictyopty- chus morgani (Douvillé) [common]; ?Biradiolites aff. baylei Tou- cas [1]; ?Fragum sp. [1]; ?Clavagellidae indet. [1]. Gastropods: ?Umboniinae, gen. nov. [6]; Turritellidae, sp. [1]; ?Rhabdoconcha sp. [1]; ?}Cimolithium sp. [2]; ‘Ampullina aff. splendida’ (Deshayes) A.B. SMITH, N.J. MORRIS, W.J. KENNEDY AND A.S. GALE Bed 14. Echinoids: Noetlingaster paucituberculatus (Noetling) [5 plus Bed 15. Echinoids: Conulus douvillei (Cotteau & Gauthier) [2]; Beds 16-19. Bivalve: ‘Modiolus’ aff. typicus Forbes [1, bivalved]. Bed 16. Echinoids: ‘Globator’ bleicherei (Thomas & Gauthier) [3]; Bed 18. Echinoid: Pygurostoma morgani Cotteau & Gauthier [1]. Bed 19. Echinoids: Noetlingaster paucituberculatus (Noetling) [4]; Bed 20. Echinoids: ‘Globator’ bleicheri (Thomas & Gauthier) [2]; Bed 21 (base). Echinoids: Noetlingaster paucituberculatus (Noetling [4]; Cypraeidae, sp. [1]; cf. ?Columbellidae [2]; ?Neogastropod indet. [1]. Corals: Cunnolites sp. [15]; Diploctenium sp. [2]; Aspidastraea sp. {1}. fragments]; Phymechinus ? sp. [1]; ‘Globator’ bleicheri (Thomas & Gauthier) [7]; Conulus douvillei (Cotteau & Gauthier) [1]; Fauja- sia eccentripora Lees [7]; Pygurostoma morgani Cotteau & Gauthier [3]; Arnaudaster cylindriformis sp. nov. [4]; Petalobrissus rawdahensis sp. nov. [17]; Petalobrissus cf. setifensis (Cotteau) [15]; Petalobrissus linguiformis (Peron & Gauthier) [4]; ?Stigmato- pygus pulchellus sp. nov. [6]; Hemipneustes arabicus Ali [1]; H. compressus Noetling [2]. Gastropods: Campanile sp. [2]; ?Campa- nile sp. [1]; ‘Ampullina aff. splendida’ (Deshayes) [2]; ? Pseudocas- sis sp. [1]; Strombidae, gen. nov. [2]; Fasciculariidae, sp. [1]; ?Volutidae indet. [1]. Corals: Aspidastraea sp. [1]. Petalobrissus rawdahensis sp. nov. [25]; Hemipneustes compressus Noetling [2]. Bivalves: ?Granocardium sp. [3]; rudist [1]. Gastro- pods: Pyrazus sp. [2]; Cerithiidae [1]; ?Naticidae, sp. [1]; Strom- bidae gen. nov. [3]. Corals: Cunnolites sp. [21]; Aspidastraea sp. [9]; massive meandroid colony [1]. Corals: Cunnolites sp. [2]; Aspidastraea sp. (2]. Faujasia eccentripora Lees [3]; Petalobrissus rawdahensis sp. nov. [5]. Bivalves: Neithea regularis (Schlotheim) [1] Durania spp. [several]; Veneridae [2]. Gastropods: ?Rhabdoconcha sp. [I]; ?Cerithiidae [1]; ‘Ampullina’ aff. splendida (Deshayes) [3]; Strom- bidae [1]; Cypraeidae [1]. Bivalves: ?Granocardium sp. [1]. Gastropods: ?Umboniinae gen. nov. [1]; ?Cimolithium sp. [2]; Strombidae, gen. nov. [1]. Coenholectypus inflatus (Cotteau & Gauthier) [1]; ‘Globator’ bleicheri (Thomas & Gauthier) [8]; Vologesia rawdahensis Ali [1]; Zuffardia morgani (Cotteau & Gauthier) [5]; Faujasia eccentripora Lees [19]; Arnaudaster cylindriformis sp. nov. [1]; Petalobrissus rawdahensis sp. nov. [73]; Petalobrissus cf. setifensis (Cotteau) [23]; Petalobrissus linguiformis (Peron & Gauthier) [5]; ?Stigmato- pygus pulchellus sp. nov. [1]; Pygurostoma morgani Cotteau & Gauthier [2]; Hemipneustes compressus Noetling [1]; Hemipneustes persicus (Cotteau & Gauthier) [1]. Bivalves: ?}Granocardium sp. [2]; Isognomon sp. [1]; ?Lucinidae indet. [1]; Dictyoptychus mor- gani (Douvillé) [1]; Veneridae, gen. indet. [1]. Gastropods:) ?Umboniinae, gen. nov. [9]; ?Rhabdoconcha sp. [1]; ?Cerithiidae [5]; Campanile sp. [4]; ‘Ampullina aff. splendida’ (Deshayes) [3); Tylostoma incerta (Forbes) [2]; ?Naticidae [4]; Strombidae, gen. nov. [5]; Cypraeidae, sp. [1]; Fasciolariidae or Buccinidae [1]; Canalicella sp. [1]; Volutidae [3]; Acteonella sp. [1]. Corals:| Neocaeniopsis reussi (Edwards & Haime) [1]; Astrogyra edwardsi (Reuss) [1]; cerioid colonies [2]; Cunnolites sp. [17]; Diploctentum| | sp. [1]; Aspidastraea sp. [8]; massive meandrine colonies [5]. Faujasia eccentripora Lees [2]; Petalobrissus linguiformis (Peron & Gauthier) [1]; Petalobrissus sp. [1]; Hemipneustes compressus Noetling [1]; Hemipneustes persicus (Cotteau & Gauthier) [1] Gastropods: ?Angaria sp. [1]; ?Umboniinae gen. nov. [2]: ‘Ampullina aff. splendida’ (Deshayes) [3]; Strombidae, gen. nov. [5]. [1]; ‘Globator’ bleicheri (Thomas & Gauthier) [1]; Conulus douvil lei (Cotteau & Gauthier) [2]; Faujasia eccentripora Lees [22] Zuffardia morgani (Cotteau & Gauthier) [5]; Pygurostoma mor gani Cotteau & Gauthier [7]; Arnaudaster cylindriformis sp. NOV [1]; Petalobrissus rawdahensis sp. nov. [45]; Petalobrissus cf setifensis (Cotteau) [44]; Petalobrissus linguiformis (Peron Gauthier) [4]; ?Stigmatopygus pulchellus sp. nov. [7]; Hemip n LATE CRETACEOUS CARBONATE PLATFORM FAUNAS neustes compressus Noetling [16]; Hemipneustes persicus (Cotteau & Gauthier) [22]. Bivalves: Barbatia cf. morgani (Douvillé) [3, bivalved]; Pycnodonte vesicularis (Lamarck) [1]; heterodont indet. [1]. Gastropods: Campanile sp. [1]; ‘Ampullina aff. splendida’ (Deshayes) [1]; Naticidae [2]; Tylostoma incerta (Forbes) [1]; Strombidae, gen. nov. [1]; ?Fasciolariidae [1]; bullaeform opistho- branch [1]. Ammonite: Desmophyllites diphylloides (Forbes) [1]. Bed 21 (middle). Gastropods: ‘Ampullina aff. splendida’ (Deshayes) [4]; Strombidae, gen. nov. [3]; Strombidae, sp. [1]; ?Hercorhyn- chus sp. [1]; ?Buccinacea, sp. [1]; neogastropod, gen. A [2]; neogastropod, gen. B [1]. Bed 22. Echinoids: ‘Globator’ bleicheri (Thomas & Gauthier) [1]; Faujasia eccentripora Lees [3]; Petalobrissus rawdahensis sp. nov. [13]; Hemipneustes compressus Noetling [1]. Gastropods: ‘Ampullina aff. splendida’ (Deshayes) [2]; ?Buccinacea [1]. Bed 23 (base). Echinoids: Petalobrissus sp. [1]. Gastropods: Pyrazus sp. [1]; Naticidae [1]. Beds 23-25. Echinoids: Petalobrissus sp. [5]; Faujasia eccentripora Lees [1]. Bivalves: ? Biradiolites aff. baylei Toucas [2]. Bed 25. Echinoids: Noetlingaster paucituberculatus (Noetling) [4]; Faujasia eccentripora Lees [3]; Zuffardia morgani (Cotteau & Gauthier) [1]; Petalobrissus sp. [11]. Bivalve: ?Lucinidae indet. [1]. Bed 26. Echinoids: Coenholectypus inflatus (Cotteau & Gauthier) [1]; Arnaudaster cylindriformis sp. nov. [1]; Pygurostoma morgani Cotteau & Gauthier [2]; Petalobrissus rawdahensis sp. nov. [5]; Petalobrissus cf. setifensis (Cotteau) [3]; Hemipneustes compressus Noetling [2]; Hemipneustes sp. fragment [1]. Bivalve: Neithea sp. {1]. ' Bed 27. Echinoids: Hemipneustes compressus Noetling [2]. / Bed 28. Bivalves: Indet. small rudists. Loose: a little below beds 21/22 and probably derived from them. Echinoids: Hemipneustes arabicus Ali [1]. Loose: scree from level of bed 11 (derived from beds 13-15 mostly). | Echinoids: Coenholectypus inflatus (Cotteau & Gauthier) [1]; Conulus douvillei (Cotteau & Gauthier) [1]; Faujasia eccentripora Lees [14]; Zuffardia morgani (Cotteau & Gauthier) [3]; Petalobris- sus linguiformis (Peron & Gauthier) [6]; Petalobrissus sp. [2]. | Loose, scree. Echinoids: Glyphopneustes hattaensis Ali [2]; Echinoti- ara perebaskinei Lambert [2]; Coenholectypus inflatus (Cotteau & | Gauthier) [1]; ‘Globator’ bleicheri (Thomas & Gauthier) [4]; | Faujasia eccentripora Lees [32]; Zuffardia morgani (Cotteau & | Gauthier) [11]; Arnaudaster cylindriformis sp. nov. [1]; Pyguros- | toma morgani Cotteau & Gauthier [1]; Petalobrissus sp. [62]; | Nucleopygus magnus sp. nov. [3]; Hemipneustes persicus (Cotteau | & Gauthier) [4]; Hemiaster hattaensis Ali [1]. Bivalves: ‘Modiolus’ | aff. typicus Forbes [1, bivalved]; Spondylus sp. C [1]; Amphidonte pyrenaicum (Leymerie) [1]; Dictyoptychus morgani (Douvillé) [1]; Radiolites sp. [1]; ?Biradiolites aff. baylei Toucas [2]. Gastropods: ?Umboniinae, gen. nov. [3]; Pyrazus sp. [1]; Campanile sp. [1]; ‘Ampullina aff. splendida’ (Deshayes) [1]; ?Perrisoptera sp. [1]; Caricella sp. [2]; Acteonella sp. [1]; Acteonella cf. caucasica Zekeli | [1]. Corals: Cunnolites sp. [12]; Aspidastraea sp. [1]. oose, lower part of section (from beds 1-11). Echinoids: Salenia nutrix Peron & Gauthier [3]; Goniopygus arabicus sp. nov. [4]; Glyphopneustes hattaensis Ali [4]; Hattopsis sphericus Ali [1]; Noetlingaster paucituberculatus Noetling [1]; Phymechinus? per- plexus sp. nov. [4]; Echinotiara perebaskinei Lambert [15]; Coen- holectypus baluchistanensis Noetling [5]; ‘Globator’ bleicheri (Thomas & Gauthier) [9]; Faujasia eccentripora Lees [80]; Zuffar- dia morgani (Cotteau & Gauthier) [29]; Petalobrissus spp. [136]. Asteroid: marginal [1]. Brachiopod: terebatulid, gen. nov. [1]. ose, mid-section. Echinoids: Salenia sp. [1]; Zuffardia morgani (Cotteau & Gauthier) [6]; ?Stigmatopygus pulchellus sp. nov. [1]. Corals: Cunnolites sp. [2]; Aspidastraea sp. [1]. 117 Jebel Rawdah, Section 3a Bed 2. Echinoids: Goniopygus arabicus sp. nov. [2]. Bivalve: Durania sp. [1]. Gastropod: Acteonella sp. [1]. Bed 4. Echinoid: Echinotiara perebaskinei Lambert [1]. Bivalves: Pycnodonte vesicularis (Lamarck) [1]; Amphidonte pyrenaicum (Leymerie) [2, bivalved]. Bed 5. Bivalves: ‘Modiolus’ aff. typicus Forbes [1, bivalved]; ?*Myzi- lus’ nitens Forbes [1, bivalved]; Barbatia sp. B [3]; Neithea regularis (Schlotheim) [3]; Amphidonte pyrenaicum (Leymerie) [3]; Ageros- trea ungulata (Schlotheim) [9]; Eligmidae [5, bivalved]; Crassitel- lites sp. [5]. Bed 7 (near top). Echinoid: ‘Globator’ bleicheri (Thomas & Gauthier) [1]. Bivalves: Pycnodonte vesicularis (Lamarck) [1]; Agerostrea ungulata (Schlotheim) [1, bivalved]; ?Clavagella sp. [tubes]. Gas- tropods: Naticidae, indet. [2]. Loose, derived from lower beds. Bivalves: Plicatula hirsuta Coquand [1]; ?Pycnodonte uncinella (Leymerie) [2, bivalved]. Gastropod: Turritellidae [1]. Jebel Rawdah, Section 3b Bed 2 (lower part). Echinoids: Orthopsis miliaris (d’Archiac) [1]; Glyphopneustes hattaensis Ali [1]; Circopeltis emiratus sp. nov. [1]; Faujasia eccentripora Lees [3]. Coral: Cunnolites sp. [1]. Decapod crustacean: Callianassa (limb segment) [1]. Bed 2 (upper part). Echinoids: Salenia nutrix Peron & Gauthier [1]; Glyphopneustes hattaensis Ali [1]; ‘Globator’ bleicheri (Thomas & Gauthier) [11]; Petalobrissus rawdahensis sp. nov [19]; Petalobris- sus cf. setifensis (Cotteau) [4]; Nucleopygus magnus sp. nov. [3]. Coral: ?Diploctenium sp. [1]. Bed 3. Echinoid: Echinotiara perebaskinei Lambert [1]. Bivalve: Pycnodonte vesicularis (Lamarck) [1]. Bed 4. Corals: large meandrine colonies [2]; placoid colony [1]. Bed 5. Echinoids: Heterodiadema buhaysensis sp. nov. [1]; Glyphop- neustes hattaensis Ali [1]; Coenholectypus inflatus (Cotteau & Gauthier) [3]; ‘Globator’ bleicheri (Thomas & Gauthier) [1]; Hemipneustes sp. [1]; Hemiaster hattaensis Ali [1]. Bivalve: Pycn- odonte vesicularis (Lamarck) [1]. Bed 5/6. Echinoids: ‘Globator’ bleicheri (Thomas & Gauthier) [8]; Hemipneustes persicus Cotteau & Gauthier [1]; Mecaster victoris Lambert [47]. Bed 6. Echinoid: Circopeltis emiratus sp. nov. [1]. Bivalve: Eligmidae [1]. Gastropod: Tylostoma incerta (Forbes) [1]. Bed 7. Echinoids: Coenholectypus inflatus (Cotteau & Gauthier) [1]; Hemipneustes persicus Cotteau & Gauthier [1]. Bivalves: Barbatia sp. B [1, bivalved)]; ‘Modiolus’ aff. typicus Forbes [2, bivalved]. Bed 8. Echinoids: Orthopsis miliaris (d’Archiac) [1]; Actinophyma spectabile Cotteau & Gauthier [1]; Coenholectypus sp. [1]; Vologe- sia rawdahensis Ali [1]; Faujasia eccentripora Lees [1]; Mecaster sp. [6]. Coral: Cunnolites sp. [1]. Decapod crustacean: pincer [1]. Bed 9. Echinoids: Orthopsis miliaris (d’ Archiac) [7]; Noetlingaster sp. [1, fragment]; Actinophyma spectabile Cotteau & Gauthier [1]; ‘Globator’ bleicheri (Thomas & Gauthier) [1]; Faujasia eccentri- pora Lees [3]; Hemipneustes compressus Noetling [2]; Hemiaster sp. cf. H. hattaensis Ali [1]; Mecaster victoris Lambert [1]; Pro- raster geayi Cottreau [2]. Bivalves: Cucullaea sp. A [4]; Scabrot- rigonia sp. [8, 1 bivalved]; ?Tancrediidae, sp. nov. [1]. Gastropods: Acteonella sp. cf. borneensis Nuttall & Leong [1]; Acteonella sp. [2]. Corals: Cunnolites sp. [8]; Diploctenium sp. [1]; thamnasteroid colony [1]. Bed 11. Echinoids: Coenholectypus inflatus (Cotteau & Gauthier) [1]; ‘Globator’ bleicheri (Thomas & Gauthier) [1]; Arnaudaster cylin- driformis sp. nov. [1]. Bivalves: Barbatia sp. B [2]; ‘Modiolus’ aff. typicus Forbes [1, bivalved]; ? Pycnodonte uncinella (Leymerie) [3, bivalved]; Amphidonte pyrenaicum (Leymerie) [2, 1 bivalved]; 118 Agerostrea ungulata (Schlotheim) [21]; Scabrotrigonia sp. [7, 1 bivalved]. Gastropod: Caricella sp. [1]. Loose. Echinoids: Prionocidaris morgani (Gauthier) [1]; Orthopsis miliaris (d’Archiac) [3]; Coenholectypus inflatus (Cotteau & Gauthier) [6]; ‘Globator’ bleicheri (Thomas & Gauthier) [3]; Conulus douvillei (Cotteau & Gauthier) [4]; Hemipneustes com- pressus Noetling [1]. Loose near top of section. Echinoids: Arnaudaster cylindriformis sp. nov. [1]; ?Linthia sudanensis (Bather) [1]. Jebel Rawdah, Section 4 Bed 1. Bivalves: Dictyoptychus morgani (Douvillé) [1]; Praeradiolites cf. subtoucasi Toucas [4]; Pseudosabinia aff. klinghardti Boehm [3]; Pseudosabinia sp. [1]. Bed 1/2. Echinoids: Phymosoma hexoaporum Lambert [1]; ‘Globator’ bleicheri (Thomas & Gauthier) [6]. Bivalves: Scabrotrigonia sp. [2]; Glabrobournonia arabica Morris & Skelton, sp. nov. [21]. Bed 2 (mostly near top). Echinoids: Echinotiara perebaskinei Lambert [1]; Coenholectypus sp. indet. [1]; ‘Globator’ bleicheri (Thomas & Gauthier) [3]; Faujasia eccentripora Lees [1]; Zuffardia morgani (Cotteau & Gauthier) [2]; Petalobrissus cf. setifensis (Peron & Gauthier) [1]; Hemiaster hattaensis Ali [1]. Bivalves: Neithea regularis (Schlotheim) [1]; Pycnodonte vesicularis (Lamarck) [1]; ?Pycnodonte uncinella (Leymerie) [3, 1 bivalved]; Amphidonte pyrenaicum (Leymerie) [1]; Scabrotrigonia sp. [1]. Gastropod: Acteonella sp. cf. borneensis Nuttall & Leong [1]. Bed 4. Echinoids: Glyphopneustes hattaensis Ali [1]; Circopeltis emiratus sp. nov. [1]; ‘Globator’ bleicheri (Thomas & Gauthier) [4]; Faujasia eccentripora Lees [2]; Petalobrissus sp. [1]. Corals: Cunnolites sp. |2|; Aspidastraea sp. [1]. Bed 5. Echinoid: ‘Globator’ bleicheri (Thomas & Gauthier) [1]. Bed 8. Echinoids: Orthopsis miliaris (d’Archiac) [1]; Echinotiara perebaskinei Lambert [1]; ‘Globator’ bleicheri (Thomas & Gauthier) [3]; Conulus douvillei (Cotteau & Gauthier) [1]; Petalo- brissus cf. setifensis (Cotteau) [3]. Bed 9. Bivalves: Amphidonte pyrenaicum (Leymerie) [1]; Scabrot- rigonia sp. [common]. Bed 10. Echinoids: Orthopsis miliaris (d’ Archiac) [6]; Salenia nutrix Peron & Gauthier [1]; ‘Globator’ bleicheri (Thomas & Gauthier) [10]. Bivalve: Dictyoptychus sp. [1]. Corals: cerioid colony [1]; placoid colony [1]. Bed 12. Echinoids: Coenholectypus sp. [1]; ‘Globator’ bleicheri (Tho- mas & Gauthier) [5]; Nucleopygus magnus sp. nov. [1]. Sponge: chaetetid [1]. Bed 13. Echinoids: Noetlingaster emiratescus Ali [2]; Coenholectypus inflatus (Cotteau & Gauthier) [1]; ‘Globator’ bleicheri (Thomas & Gauthier) [3]; Conulus douvillei (Cotteau & Gauthier) [1]; Hemi- aster hattaensis Ali [1]. Bed 15. Echinoids: Orthopsis miliaris (d’Archiac) [1]; ‘Globator’ beleicheri (Thomas & Gauthier) [1]. Coral: Actinacis sp. [1]. Bed 18. Echinoid: Coenholectypus sp. [1]. Bed 19. Echinoids: Coenholectypus inflatus (Cotteau & Gauthier) [1]; ‘Globator’ bleicheri (Thomas & Gauthier) [1]; Pygurostoma mor- gani Cotteau & Gauthier [1]. Bed 22/23. Echinoids: Coenholectypus inflatus (Cotteau & Gauthier) [1]; Faujasia eccentripora Lees {1]. 30 cm below top of measured section. Echinoids: Coenholectypus inflatus (Cotteau & Gauthier) [1]. Bivalve: Neithea regularis (Schlotheim) [1]. Loose, a little below the top of the section. Echinoids: Hemipneustes arabicus Ali [1]. Loose at level of bed 3. Echinoid: Petalobrissus linguiformis (Peron & Gauthier) [1]. £.oose. Echinoids: Orthopsis miliaris (d’ Archiac) [1]; Conulus douvil- lei (Cotteau & Gauthier) [1]; Petalobrissus sp. [1]. Bivalves: Amphidonte pyrenaicum (Leymerie) [2]. A.B. SMITH, N.J. MORRIS, W.J. KENNEDY AND A.S. GALE Jebel Faiyah, Section la Bed 6. Bivalves: Hippurites aff. lapeirousei Goldfuss [1 colony]. Bed 8. Bivalves: Agerostrea ungulata (Schlotheim) [2]; ?Pycnodonte uncinella (Leymerie) [18]; ?Amphidonte cf. pyrenaicum (Leyme- rie) [1]. Jebel Faiyah, Section 1b Bed 2 (lower part). Echinoids: Hattopsis paucituberculatus sp. nov. [6]. Corals: Polytremacis sp. [1]; Astraraea sp. [1]; Hydno- phoraraea sp. [1]. Sponge: chaetetid [1]. Bed 2 (upper part). Echinoids: cidarid [1]; Orthopsis miliaris (d’Archiac) [1]; Salenia nutrix Peron & Gauthier [2]; Glyphop- neustes hattaensis Ali [1]; Hattopsis paucituberculatus sp. nov. [1]; ‘Globator’ bleicheri (Thomas & Gauthier) [1]; Nucleopygus mag- nus sp. nov. [1]; Hemiaster sp. cf. H. hattaensis Ali [1]. Bivalves: Hippurites cf. lapeirousei Goldfuss [1 colony]; Hippurites aff. cornucopiae Defrance [1]; Dictyoptychus morgani (Douvillé) [2]; Biradiolites aff. baylei Toucas [1]; Durania cf. gaensis (Dacqué) [1]. Decapod crustacean: pincers [2]. Bed 4. Bivalves: Hippurites aff. lapeirousei Goldfuss [2]. Corals: Polytremacis sp. [1]; Hydnophoraraea sp. [1]; Paraplacocoenia orbignyana (Reuss) [1]; Moltkia isis Sreenstrup [1]. Bed 5. Echinoids: Glyphopneustes hattaensis Ali [2]. Bed 6. Echinoid: cidarid spine [1]. Coral: Moltkia isis Sreenstrup [1]. Bed 7. Echinoids: cidarid spine [1]; Mimiosalenia quinquetuberculata sp. nov. [21]; Hattopsis sphericus Ali [1]; ‘Globator’ bleicheri (Thomas & Gauthier) [36]. Bed 8. Coral: Cunnolites sp. [1]. Loose (derived from beds 2-5). Echinoids: Hattopsis paucituberculatus sp. nov. [5]. Bivalves: Spondylus sp. C [2]. Nautiloid: Deltoidonau- tilus salisfilius sp. nov. {1]. Corals: Polytremacis sp. [5]; Cunnolites sp. [1]; Paraplacocoenia orbignyana (Reuss) [1]; Hydnophoraraea sp. [5]; turbinate solitary [1]; cerioid colonies (6 genera) [14]; flabellate solitary [1]; phaceloid colony [1]. Branching algae [1]. Sponge: chaetetid [1]. Decapod crustacean: crab carapace [1]. Jebel Faiyah — southern nose of Jebel (bed numbers as in section la) Bed 8. Echinoids: Salenia sp. [1]; Glyphopneustes hattaensis Ali [8];| Hattopsis sphericus Ali [8]; ‘Globator’ bleicheri (Thomas & Gauthier) [2]; Pygurostoma morgani Cotteau & Gauthier [1]. Corals: Cunnolites sp. [2]. Loose. Echinoids: Hattopsis sphericus Ali [1]. Bivalves: ?Barbatia sp. A. cf. B. morgani (Douvillé) [1, bivalved]; Neithea regularis (Schlotheim) [7]; ?Pycnodonte uncinella (Leymerie) [18]; ?Pla- gioptychus sp. [1]; Glabrobournonia arabica Morris & Skelton, sp. nov. [1]; Hippurites 2cornucopiae Defrance [11]; Dictyoptychus morgani (Douvillé) [3]. Gastropod: Tylostoma incerta (Forbes) [1]. Corals: Moltkia isis Streenstrup [1]; Cunnolites sp. [3]; Hydno- phoraraea sp. [2]; Paraplacocoenia orbignyana (Reuss) [1]; cerioid colonies (3 genera) [4]; phlacoid colony [1]; thamnasteroid colony [1]. Jebel Faiyah, Section 2 Basal 1 m shell bed. Bivalves: Glabrobournonia arabica Morris & Skelton, sp. nov. [3]. Nautiloids: Cimonia cf. sowerbyan (d’Orbigny) [1]; Deltoidonautilus salisfilius sp. nov. [2]. Jebel Bu Milh, Section 1 Bed 1. Bivalves: Pseudosabinia aff. klinghardti (Boehm) [4]; Durant cf. gaensis (Dacqué). Bed 3. Bivalves: ‘Modiolus’ aff. typicus Forbes [1, bivalved]; Lyrio LATE CRETACEOUS CARBONATE PLATFORM FAUNAS chlamys ternatus (Minster) [2]; Neithea regularis (Schlotheim) [1]; ?Plagioptychus sp. [4, lids]; Dictyoptychus morgani (Douvillé) [1, juvenile]; Radiolites sp. [3]; Eodictyoptychus aff. arumaensis Skel- ton & El-Asa’ad [2]. Gastropods: ?‘Angaria’/Liotia sp. [1, frag- ment]; ?trochid [1]; Discotectus sp. [26]; Strigosella sp. cf. striolata (Stolizcka) [2]; Euchelus ornatus Stolizcka [3]; ?Umbonium greyi Lees [1]; ‘Turritella’ sp. [1]; ‘Turritella’ sp. 1 ?= Nairiella multi- striata (Reuss) [4]; Vermetus sp. [2]; Campanile curtum Douvillé [1]; Campanile persicum Douvillé [2]; Campanile aff. breve Lees [1, fragment]; Campanile morgani Douvillé [1]; Campanile sp. [6, internal moulds]; Campanile cf. ganesha Noetling [4]; Cimolithium sp. nov. [2]; Pyrazus sp. [1]; Paryphostoma morgani Douvillé [1]; Pugnellus sp. [2]; Ampullina aff. splendida (Deshayes) [30+]; ‘Euspira’ lirata J. de C. Sowerby [22]; ‘Natica’ pagoda Forbes [2]; Tylostoma incerta (Forbes) [5]; Confusiscala sp. [1]; ‘Cypraea’ kayeii Forbes [2]; Hipponyx sp. [1]; ‘Fulguraria’ multistriata Stolic- zka [1]; fasciolariid [1]; ?Rapanidae gen. indet. [1]; ‘Murex’ sp. [1]; ‘Trophon’ oldhamianum Stolizcka [1]; Volutoderma elongata Stolizcka [1]; Volutolithes latisepta Stoliczka [2]; Caricella pyrifor- mis Forbes [2]; Caricella sp. [2]; ‘Voluta’ sp. or spp. [3]; Trochac- taeon sp. [1]; Acteonella cf. caucasica Zekeli [30+]; Acteonella caucasica Zekeli subsp. grossouvrei Cossmann [10]; Acteonella laevis laevis (J. de C. Sowerby) [4]; Acteonella elongata Kollmann [4]. Corals: Aspidastraea sp. [1]; Hydnophoraraea sp. [2]. Jebel Bu Milh, Section 2 Bed 4. Gastropods: Acteonella cf. caucasica Zekeli [74]; Acteonella elongata Kollmann [1]; Acteonella laevis (J. de C. Sowerby) [3]; Acteonella cf. laevis zekelii Kollmann [1]. Bivalves: Lapeirousia sp. [2]. Beds 7/8. Echinoids: Petalobrissus cf. setifensis (Cotteau) [1]. eae Bivalves: Barbatia sp. B [12, 4 bivalved]; Cucullaea sp. A [2, bivalved]; Glycymeris sp. [1]; ‘Platyceramus’ sp. [1]; Lyriochlamys ternatus (Munster) [1]; Neithea regularis (Schlotheim) [1]; Spondy- lus sp. [4, bivalved]; Plicatula hirsuta Coquand [5]; Plagiostoma sp. [2]; Crassatellites sp. [5]; Eodictyoptychus aff. arumaensis Skelton & El-Asa’ad [1]; Biradiolites aff. baylei Toucas [1]. Gastropods: Calliomphalus sp. [2]; ?Calliomphalus or Helicanthus sp. [1]; ?Angaria’ sp. [2]; Cyclostrematid/Liotid gen. nov. [1]; cf. ‘Turbo’ punctatus Zekeli [1, fragment]; Discotectus sp. 1 [common]; Disco- tectus sp. 2 [common]; Strigosella sp. cf. striolata (Stolizcka) [common]; Nerita spp. [occurs]; ‘Lissocheilus’ persicus (Douvillé) [4]; ‘Turritella’ sp. 1 ?= Nairiella multistriata (Reuss) [7]; ‘Turri- tella’ sp. cf. morgani Douvillé [1]; ‘Turritella’ sp. 3 (2= Roemeriella nerinea Akopyan (non Roémer)) [2]; Campanile ?robustum Dou- 119 villé [1]; Campanile curtum Douvillé [26]; Campanile ?persicum Douvillé [2]; Campanile aff. breve (Lees) [5]; Campanile morgani Douvillé [1]; Cimolithium sp. nov. [72]; Semivertagus cf. arcotense Stolizcka [1]; cf. Pyrazella (Plicopyrazus) [1]; Pyrazus pyramidatus Douvillé [1]; Exechocirsus sp. 1 [2]; Exechocirsus sp. 2 [1]; ?Exechocirsus sp. [2]; ?Semivertagus sp. cf. arcotense (Stolizka) [7]; Hantkenia louristana Douvillé [5]; Strombidae gen. nov. giganteus Noetling [6]; Strombidae gen. nov. digitatus Noetling [2]; ‘Helicaulax’ sp. [4]; Ampullina aff. splendida (Deshayes) [100]; ‘Euspira lirata’ (J. de C. Sowerby) [1]; ?Tylostoma incerta (Forbes) [12]; ‘Confusiscala’ sp. cf. turbinata Forbes [5]; ‘Calyptraea’ elevata Forbes [1]; Thlacodes lamellosus Stolizcka [1]; Lathyrus sp. [1]; Lathyrus sp. cf. ‘Ornopsis’ digressa (Wade) [1]; ?Bellifusus sp. [1]; ?Pseudoliva sp. [2]; ‘Voluta’ citharina Forbes [1]; Volutoderma sp. [2]; Caricella pyriformis Forbes [20+]; ‘Voluta’ sp. [11]; ‘Voluta’ sp. [21]; Voluta cameleo Forbes [1]; Volutoderma sp. [1]; Lyria crassicostata Dujardin [2]; ?Naronia eximia Stolizcka [1]; Acteonella caucasica Zekeli grossouvrei Cossmann [46]; Acteonella caucasica caucasica Zekeli [31]; Acteonella caucasica Zekeli subsp. nov. [1]; Acteonella laevis laevis (J. de C. Sowerby) [3]; ?Acteonella laevis J. de C. Sowerby zekelii Kollman [1]; ?Acteonella elongata Kollman [4]; Neocylindrites minutus (Stolizcka) [2]. Ammonite: Nostoceras (Nostoceras) major Kennedy & Cobban [1]. Nautiloids: Deltoidonautilus salisfilius sp. nov. [9]. Corals: Cunnolites sp. [6]; cerioid colonies (2 genera) [3]. Sponges [7]. Bed 10. Bivalves: Vaccinites oppeli (Douvillé) [abundant, in situ]. Qarn Murrah Rudist level near top of section. Bivalves: Osculigera cf. vautrinioides Vogel [32]; Vaccinites vesiculosus (Woodward) [13];?Vaccinites loftusi (Woodward) [3]; Glabrobournonia arabica Morris & Skel- ton, sp. noy. [16]; Pseudosabinia aff. klinghardti (Boehm) [1]; Pseudopolyconites aff. parvus Milovanovic [1]; Colveraia variabilis Klinghardt [1]. Corals: Cunnolites sp. [2]; cerioid colonies (2 genera) [3]. Qarn Mulayh Lower part. Bivalves: Pironaea cf. polystylus Pirona [3]; Durania form A [2]; Durania sp. [1]. 7-7 m below the top of the section. Bivalyes: Torreites sanchezi (Douvillé) milovanovici Grubic [3]. Unspecified level. Bivalyes: Colveraia variablilis Klinghardt [1]. Oren RR A pCPalas bu) Gr etn rey ie Wea ie ate vrata > (ds oad As ad = 4 Fi LM my ae) bo. waa : SC) fart?) Sette et) Sea Vl age pile wit AN) \-. Sasi? . aon tive | (ih Vesa r A Ta) a wae skool iolaccbistwaree wee aleealne ceria eemtietee tae maniac coca aslede seeee oaige 130 SVStomatic GESCHIPLIOMS, HM... eemecemeeece seins «ote ciciins «sis atsebteelos waeaiste soelotiaaiel actemjaiacre da seiemeoee meee CoeeeGl eee es cideistess ceed. 130 IRELETENCES | cposcsta ist oa das teteee MOSSE Soe ee Ne «Sele ec wc eee Doiswienas ois» wa oe cetea Ot crate OM Pinee Sele ee eee teeta eink Stages ocean ge 239 Synopsis. Forty-five echinoid species, 14 of them new, are described from the Maastrichtian Simsima Formation exposed along the western margins of the North Oman Mountains. The stratigraphic distribution of over 2,500 individual fossil echinoids has been recorded and used to assess echinoid abundance quantitatively and identify recurrent assemblages of species. Although approximately equal numbers of regular and irregular echinoid species are known, irregular echinoids greatly outnumber regulars in abundance. Furthermore, most species are rare, with just five making up more than 67% of the total collection. Regular echinoids are divisible into four ecological groups, ranging from hard-ground dwellers living within active wave-surge environments, to herbivores living in protected subtidal soft-bottom environments. Five ecological groups of irregular echinoid are distinguished, largely on the basis of their inferred feeding strategies. Seven echinoid assemblages are distinguished and their palaeoecological setting interpreted on the basis of the autecology of included species, associated macrofauna and lithofacies evidence. One new genus of Goniopygidae, Mimiosalenia, is described and the following new species are erected: Prionocidaris? emiratus, Heterodiadema buhaysensis, Salenia microprocta, Goniopygus arabicus, Mimios- alenia quinquetuberculata, Codiopsis lehmannae, Hattopsis paucituberculatus, Circopeltis? emiratus, Phymechinus? perplexus, Coptodiscus magniproctus, Petalobrissus rawdahensis, Nucleopygus magnus, Stigmatopygus? pulchellus, Arnaudaster cylindriformis. INTRODUCTION _ Late Cretaceous echinoid faunas, though well-documented in _ both Europe and America, remain relatively little studied | elsewhere in the world. Those of the Middle East are particularly poorly known. Much of our knowledge of Tethyan late Cretaceous echinoid faunas comes from the | major monographic works of the last century such as those dealing with Algeria (Cotteau et al. 1881), Iran (Cotteau & Gauthier 1895; Gauthier 1902) and Baluchistan (Noetling 1897), all of which are now in great need of revision and updating. Additional major late Cretaceous faunas were | described during the early part of this century from Libya _(Checchia-Rispoli 1930, 1931a,b, 1932a,b, 1933) and from ) Madagascar (Cottreau 1908; Lambert 1933). _ Until recently the late Cretaceous faunas of the Arabian 'Peninsula remained virtually unknown. Duncan (1865) ‘described a few Cenomanian echinoids from south eastern Oman, while Lees (1928) described two new late Cretaceous ©The Natural History Museum, 1995 | echinoid species from north western Oman. Clegg (1933) described a large number of echinoids from the Persian Gulf, but only two of these were Cretaceous, and neither appears to be late Cretaceous in age. The first indication of the rich late Cretaceous echinoid fauna of the Arabian Peninsula came with the publication of Kier’s (1972) monograph on the Mesozoic and Tertiary echinoids around Riyadh, Saudi Arabia. In this work he described 11 species from the Campanian-Maastrichtian Aruma Formation, six of which were new to science and the remainder representing species already described by Cotteau & Gauthier (1895) and Gauthier (1902) from southern Iran. Subsequently, Ali (1989, 1992a,b) documented 20 echinoids, including seven new species, from the Simsima and Qahlah Formations of the western Oman Mountains. Smith (in Skelton et al. 1990) gave some preliminary records of the echinoid fauna from these beds and Roman et al. (1989) gave a preliminary report on the Cretaceous echinoids of the Dhofar region, Oman, including four Campanian/ Maastrichtian species. A.B. SMITH Cassiduloids 122 i 1) 83 oO B50 ©) OO oD) GS 7) Be S) WEA ) ‘ YW 2 YY E 3 YY), Fa 8 “ JWCCCCCCCCCCCCCIMMMMMwMwqqee) A erat WMM_VTJ#C@@J@™ #0. ao 28s YW WVVVVVVZ?l.-; tS seDig GJJVJ,0,0,0J].,J,.,V;S)G~GGJ]TJMqwqee” (”<89) SO he CALL YP 30 8 as Le co G) Z 3 MG I) WW SNAG | a 3 MW GQ IW et SERRE SACS SSS SSS % : cS Species (see Table 1 for key) Number of specimens ~ is) //" Regular echinoid species 0" CY J=@™— PPP P/»— PAZ] ]WWZ]—@-—MP|X] ]W@C]CZ7TZ7=ZPVZ7VV™ZJZ3—-; SS Sao SS) 6s eS FOeeCe eee ecm Seace is 6S) 2 = 98 (Seo ees SF oh eae SIJENPIAIPUI JO SIBQUINN faa SJENPIAIpU! JO SIBquUINN Species (see Table 1 for key) Emirates-Oman border region (based on ca. 2,500 individuals). Species 1—45 are listed in Table 1. Shading distinguishes the four major taxonomic groups of echinoids. A, species arranged systematically. B, species arranged in order of specimen abundance. Pie diagrams Fig. 1 Relative abundance of species collected from the Maastrichtian Qahlah and Simsima Formations along the United Arab represent the number of individuals and number of species for each major taxonomic group. LATE CAMPANIAN-MAASTRICHTIAN ECHINOIDS 123 Table 1. Echinoid species collected from the Qalah and Simsima Formations exposed in the Jebels along the Oman-United Arab Emirates border region. Class Echinoidea Leske, 1778 Subclass Cidaroidea Claus, 1880 Order Cidaroida Claus, 1880 Family Rhabdocidaridae Lambert, 1900 1. Gen. et sp. indet. Family Cidaridae Gray, 1825 Genus Prionocidaris Agassiz, 1863 Ds Prionocidaris morgani (Gauthier, 1902) 3. Prionocidaris? emiratus sp. nov. Subclass Euechinoidea Bronn, 1860 ?Cohort Diadematacea Duncan, 1889 ?Order Diadematoida Duncan, 1889 Family Heterodiadematidae Smith & Wright, 1993 Genus Heterodiadema Cotteau, 1864 4. Heterodiadema buhaysensis sp. nov. Cohort Echinacea Claus, 1876 Plesion (Order) Orthopsida Mortensen, 1942 Family Orthopsidae Duncan, 1889 Genus Orthopsis Cotteau, 1864 5. Orthopsis miliaris (d’ Archiac, 1835) Order Calycina Gregory, 1900 Family Saleniidae Agassiz, 1838 Genus Salenia Gray, 1835 Salenia nutrix Peron & Gauthier, 1881 Salenia microprocta sp. nov. Order Arbacioida Gregory, 1900 Family Goniopygidae Smith & Wright, 1993 Genus Goniopygus Agassiz, 1838 8. Goniopygus arabicus sp. nov. Genus Mimiosalenia gen. nov. 9. Mimiosalenia quinquetuberculata sp. nov. Family Glyphopneustidae Smith & Wright, 1993 Genus Glyphopneustes Pomel, 1883 10. Glyphopneustes hattaensis Ali, 1992 Family Arbactidae Gray, 1835 Genus Codiopsis Agassiz, 1840 11. Codiopsis lehmannae sp. nov. Genus Hattopsis Ali, 1992 12. Hattopsis sphericus Ali, 1992 | 3. Hattopsis paucituberculatus sp. nov. Genus Noetlingaster Vredenberg, 1911 («14 Noetlingaster paucituberculatus (Noetling, 1897) | iS. Noetlingaster emiratescus Ali, 1989 16. Noetlingaster? sp. Order Phymosomatoida Mortensen, 1904 Family Phymosomatidae Pomel, 1883 Genus Phymosoma Haime, 1853 if. Phymosoma hexoaporum Lambert, 1927 | Genus Actinophyma Cotteau & Gauthier, 1895 | 18. Actinophyma spectabile Cotteau & Gauthier, 1895 Genus Plistophyma Peron & Gauthier, 1881 19: Plistophyma asiaticum Cotteau & Gauthier, 1895 Family Stomechinidae Pomel, 1883 Genus Circopeltis Pomel, 1883 f 20. Circopeltis? emiratus sp. nov. ; Genus Phymechinus Desor, 1856 | 21. Phymechinus? perplexus sp. nov. Genus Echinotiara Pomel, 1883 22. 23. 24. 25. 26. 27. 28. 29. 30. 31. 32. 33. 34. 36. 35. 37. 45. Echinotiara perebaskinei Lambert, 1929 Cohort Irregularia Latreille, 1825 Order Holectypoida Duncan, 1889 Family Holectypidae Lambert, 1899 Genus Coenholectypus Pomel, 1883 Coenholectypus inflatus (Cotteau & Gauthier, 1895) Coenholectypus cf. baluchistanensis (Noetling, 1897) Genus Coptodiscus Cotteau & Gauthier, 1895 Coptodiscus magniproctus sp. nov. Family Conulidae Lambert, 1911 Genus ‘Globator’ Agassiz, 1840 ‘Globator’ bleicheri (Gauthier, 1889) Genus Conulus Leske, 1778 Conulus douvillei (Cotteau & Gauthier, 1895) Order Cassiduloida Claus, 1880 Family Clypeolampadidae Kier, 1962 Genus Vologesia Cotteau & Gauthier, 1895 Vologesia rawdahensis Ali, 1989 Family Faujasiidae Lambert, 1905 Genus Faujasia d’Orbigny, 1856 Faujasia eccentripora Lees, 1928 Genus Zuffardia Checchia-Rispoli, 1917 Zuffardia morgani (Cotteau & Gauthier, 1895) Unnamed family Genus Pygurostoma Cotteau & Gauthier, 1895 Pygurostoma morgani Cotteau & Gauthier, 1895 Family Cassidulidae Agassiz & Desor, 1847 Genus Petalobrissus Lambert, 1916 Petalobrissus rawdahensis sp. nov. Petalobrissus cf. setifensis (Cotteau, 1866) Petalobrissus linguiformis (Peron & Gauthier, 1881) Genus Stigmatopygus d’Orbigny, 1856 Stigmatopygus? pulchellus sp. nov. Genus Nucleopygus Agassiz, 1840 Nucleopygus magnus sp. nov. Family Echinolampadidae Gray, 1851 Genus Arnaudaster Lambert, 1918 Arnaudaster cylindriformis sp. nov. Order Holasteroida Durham & Melville, 1957 Family Holasteridae Pictet, 1857 Genus Hemipneustes Agassiz, 1836 Hemipneustes compressus Noetling, 1897 Hemipneustes persicus Cotteau & Gauthier, 1895 Hemipneustes arabicus Ali, 1989 Order Spatangoida Claus, 1876 Family Hemiasteridae Clark, 1917 Genus Hemiaster Agassiz, in Agassiz & Desor, 1847 Hemiaster hattaensis Ali, 1989 Hemiaster paronai Checchia-Rispoli, 1921 Genus Mecaster Pomel, 1883 Mecaster victoris (Lambert, 1932) Family Schizasteridae Lambert, 1905 Genus Linthia Desor, 1853 ?Linthia sudanensis (Bather, 1904) Genus Proraster Lambert, 1895 Proraster geayi Cottreau, 1908 124 Our field collecting in this region has more than doubled the known fauna of late Cretaceous echinoids, making it one of the most diverse assemblages of this age anywhere in the world. Much of the material is well preserved, allowing plating details to be recorded in many species and genera for the first time. Forty four species are recognized (Table 1) and sufficient material has been collected (over 2,500 specimens) to allow a detailed biometric study of most species. Further- more, bed by bed collecting has enabled me to recognize recurrent assemblages and to place these into their palaeoen- vironmental setting, something that has never previously been attempted. RELATIVE ABUNDANCES OF SPECIES A total of 2585 specimens were collected and identified to species level, allowing a quantitative assessment of species abundance. This is not entirely accurate in comparison with a previous study of echinoid abundance in the Cenomanian (Smith 1988) where every specimen seen was collected, because the three or four most abundant species are surely undercollected. A few species (e.g. Petalobrissus rawdahensis at Jebel Rawdah, section 2, or ‘Globator’ bleicheri at Jebel Buhays) were so abundant that only a proportion of the observed specimens were eventually collected. There is therefore a bias towards the rarer species. Nevertheless, a number of general conclusions can be drawn from the distri- butional and abundance data that have been collected. (a) There are approximately equal numbers of regular and irregular echinoid species (22 regular, 23 irregular) (Fig. 1B). A similar ratio of regular to irregular species is characteristic of the present day biota (Kier 1977) and for extensive collections from the Cenomanian of Great Britain and France (Smith 1988, Neraudeau & Moreau 1989). Amongst the irregular echinoids, cassiduloids (10 species) are more diverse than holectypoids (5 species), spatangoids (5 species) or holasteroids (3 species). (b) Irregular echinoids greatly exceed regular echinoids in numbers of specimens (Fig. 1B). In total 499 specimens of regular echinoid were collected (19%), and this is likely to overestimate their true relative abundance, since the five most common echinoids were all irregular species and are likely to have been significantly undercollected. This com- pares well with previous studies, where Smith (1988) found between 10-20% of the total number of echinoid specimens collected (1800 individuals) from the Cenomanian of Wilm- ington were regular species, and with Neraudeau & Moreau (1989) who reported 22.8% of 5133 individuals collected from the Cenomanian of North Aquitaine were regular species. Therefore regular echinoids achieve a similar species diver- sity to irregular echinoids, but occur in much lower absolute diversity. (c) Most species are rare. In terms of numbers of individu- als, just five species make up more than 67% of the total collection (Fig. 1A). Four of these five species are cassidu- loid, the fifth (8% of the total) is the holectypoid *‘Globator’. The most abundant species are cassiduloids and two of the three species of Petalobrissus together form more than 36% of the total number of individuals collected. In reality these species are even more dominant, since they were certainly A.B. SMITH undercollected in comparison to rare species, possibly by as much as a factor of two. In comparison, the most abundant regular echinoid species (Echinotiara perebaskinei and Glyphocyphus hattaensis) rep- resent only 3.2% of the total collection each, and the the most common spatangoid (Mecaster victori) and holasteroid (Hemipneustes persicus) represent a mere 1.6% and 1.4% respectively of the total collection. ECHINOID PALAEOECOLOGY Echinoids are adapted to live in a wide range of habitats and the relationship between skeletal structure, habitat and mode of life is now relatively well understood. Consequently they are an excellent group to use as palaeoenvironmental indica- tors. Virtually nothing is known about the late Cretaceous palaeoecology of echinoids in tropical carbonate shelf envi- ronments, and the Oman Mountains collection described here provides the first such opportunity to assess echinoid distribution and faunal associations quantitatively. Before discussing the assemblages that can be recognized and their palaeoenvironmental setting, it is necessary to look at the detailed functional morphology of the different taxa. Each has a preferred habit and life-style that is partially reflected and can be deduced from the skeletal morphology (e.g. Smith 1984). 1. Regular echinoids. All regular echinoids live epifaunally, but can be differentiated into a number of ecological groups based on their skeletal morphology. (a) Hard-ground dwellers living within the zone of active wave surge (0-5 m depth). This includes Codiopsis, Phy- mechinus, Circopeltis and Echinotiara. These forms are char- acterized by their flat oral surface and enlarged phyllodes composed of P3/P4-type pore-pairs (Smith 1978). They also typically have dense or modified aboral pore-pairs indicative of specialized respiratory tube-feet. The broad, flat base and numerous, large oral tube-feet are features characteristic of present-day echinoids living on rocky surfaces within the zone of active wave surge. Large and strong oral tube-feet are necessary to provide adhesion in a turbulent environment. Modern analogues would be Arbacia, Anthocidaris, Helioci- daris or Stomopneustes, all of which are coastal species living within the first few metres on rock or other hard ground surfaces. They are rock grazers, feeding on filamentous or fleshy algae which they obtain by rasping hard substrata. (b) Hard-ground dwellers living subtidally within the top few metres of water but subject to limited wave surge only. Here I include only Goniopygus. Goniopygus has moderately strong phyllodes, a depressed profile though without a broad flat oral surface, and well-developed aboral respiratory tube- feet (to judge from its pore-pairs). The aboral respiratory tube-feet suggest it had a relatively high metabolic rate and therefore lived in the shallow, warmest waters. It had strong oral tube-feet for adhesion, but these were not as highly developed as those of species in group (a) and it seems unlikely that Goniopygus could have lived in fully exposed habitats. (c) Shallow water (ca. 2-10 m) forms living in more protected environments, within wave base, but not subject to strong currents or wave surge. These echinoids might typi- LATE CAMPANIAN-MAASTRICHTIAN ECHINOIDS > Jebel Huwayyah, sections1/2: 50-4 Loftusia facies 404 N=57 Numbers of individuals 20 Faujasia Species 100) Jebel Huwayyah, section 1: 50-4 beds 14,15 40-4 N=62 Numbers of individuals 20 Species Globator bleicheri Fig. 2 Echinoid species abundances within specific units at Jebel Huwayyah. A, fauna from beds 10-12, section 1 and beds 1-6, section 2 (Loftusia-rich facies). B, fauna from beds 14 and 15, section 1. Species are listed numerically along the x-axis as in Table 1. N = total number of specimens collected. cally be expected to have lived on or close to firm bottoms, i.e. either rocky or reefal knolls or stabilized sedimentary ‘substrata. Taxa included here are Orthopsis, Phymosoma, | Plistophyma, Actinophyma, Mimiosalenia, and Glyphop- /neustes. These echinoids all have a depressed profile and ‘slight phyllodes, and often have increased densities of aboral _pore-pairs, possibly specialized for gaseous exchange. Their |spines are moderate to long. Modern equivalents, such as Lytechinus or Temnopleurus live in and around hardground substrata in reasonably well-protected habitats. They are predominantly grazers, feeding on encrusting or boring algae ‘or plants and removing upright algae down to the substratum. Those with no phyllode development and invaginated peris- tomes are likely to have been grazers, whereas those with oral phyllodes and a flat peristome were probably raspers. Heterodiadema is a large, motile diadematoid-like echinoid with long spines whose closest modern counterparts are forms such as Centrostephanus or Diadema, found living in algal turf (dense stands of filamentous algae) or in amongst thick- ets in the reef flat environment (Birkeland 1989). The cidarids Rhabdocidaris and Prionocidaris probably delong to this category. They are clearly shallow-water forms yecause of their well-developed respiratory tube-feet (conju- a pore-pairs), but are globular in profile and lack phyl- odes. They would have been confined to the most protected f the shallow-water habitats, along with Heterodiadema. ‘idarids and diadematoids are rather generalist feeders, referentially grazing on animals and plants, but also able to ake up bottom material (Birkeland 1989). (d) Protected subtidal (10 m plus) soft-bottom substrata below active wave base. Noetlingaster, Hattopsis and Salenia are all globular forms lacking aboral specialised respiratory tube-feet. Noetlingaster closely resembles the present-day Echinus or Tripneustes in morphology and presumably lived in much the same way, found largely on sandy substrata of lagoons or below active wave-base moving over stable sedi- mentary bottoms or living within algal stands. These forms are predominantly herbivorous browsers, cropping exposed algae and taking loose bottom material. The extreme globu- lar shape of Hattopsis is matched by that seen in some present day temnopleurids such as Microcyphus and, like those echinoids, it may have lived within algal turf, attached to and enveloped by fronds of filamentous algae. Irregular echinoids. These too can be divided into a number of discrete ecological groups based on observed skeletal characteristics. (e) Infaunal medium-fine sand-grade burrowers which are selective deposit feeders using penicillate tube-feet around the mouth to pick up food particles from sediment of the burrow floor. Here I include Hemiaster and Mecaster. Both have globular tests with no real frontal sulcus. Pore differen- tiation shows that funnel-building tube-feet would have been present in ambulacrum III adapically, and subanally in the posterior ambulacra also. These heart urchins clearly lived infaunally within relatively poorly permeable, rather fine- grained sediment. An apical funnel is required by infaunal spatangoids living in finer-grained sediments, as is the aboral fasciole (which is essential for maintaining a water-filled space surrounding the test). The lack of specialization of the 126 100 Jebel Rawdah, section 1: beds 3-5 N=170 Hattopsis sphericus Numbers of individuals 20 A.B. SMITH , Globator bleicheri , Petalobrissus Species Fig. 3 Echinoid species abundances within beds 3-5 at Jebel Rawdah, section 1. Species are listed numerically along the x-axis as in Table 1. N = total number of specimens collected. Unshaded box = fragments only. frontal groove implies that all food particles were derived from within the sediment, not from the sediment/water interface. (f) Infaunal medium-fine sand-grade burrowers, selective deposit feeders harvesting organic material from the sediment-water interface. Only Proraster falls into this group. It has a deep, highly specialized frontal groove which is characteristic of those modern taxa feeding by means of a mucus-string. It is very similar to the extant Schizaster, and presumably lived in very much the same way. The pore-pairs in ambulacrum III are specialized and indicate the presence of highly developed funnel-building tube-feet. There is little doubt that Proraster lived infaunally, feeding on detritus from the water/sediment interface that cascaded down the funnel. The implication (though untested as far as I am aware) is that Proraster is adapted for life in more organic-poor sediments than either Hemiaster or Mecaster since it preferentially harvests surface detritus. The well-developed petals show that Proraster was still a relatively shallow-water form. (g) Shallow infaunal or semi-infaunal ploughers in stable, unconsolidated bottoms. Bulk deposit feeders harvesting sediment at or close to the water interface. Only Hemip- neustes falls into this catagory. Hemipneustes has a well- developed anterior groove with specialized grill spines and tube-feet. Sediment would have entered the frontal groove adapically and passed to the mouth via a mucous string running down ambulacrum III. The lack of protection for petal tube-feet, and the asymmetry of the petals implies that the surrounding sediment was highly permeable and that the urchins lived only relatively shallowly burried. (h) Mobile high permeability (low fines) unconsolidated medium-coarse sands: infaunal bulk sediment swallowers. This category includes Petalobrissus, Nucleopygus, Pyguros- toma and Stigmatopygus. These urchins most closely resemble the present day Apatopygus, which is a bulk sedi- ment swallower. They have small phyllodes around the peristome, with moderately well developed bourrelets (spe- cialized regions of dense spines used for manipulating sand- grade particles into the peristome. Their large periproct and anal sulcus suggests they had to cope with copious faecal discharge. They are small, depressed forms streamlined for moving through loose sediment and have well-developed petals. Vologesia and Arnaudaster have similar phyllodes but | much larger animals, more closely resembling Echinolampas. Echinolampas is found today in subtidal (5-20 m) depths living infaunally in shell-sands (e.g. Thum & Allen 1975). Petalobrissus is likely to be adapted for more mobile sedi- ments, i.e. for shallow subtidal shoals. (i) Mobile, permeable unconsolidated sands: selective, infaunal deposit feeders. Here I place Zuffardia and Fauyasia. Both have small peristomes, surrounded by moderately well- developed phyllodes, and with slight bourrelet development. They both have rather small, posteriorly placed periprocts, and from this it would appear that they were rather more selective deposit feeders than Petalobrissus and other related taxa. They have large petals and clearly lived infaunally, because of their rounded shape. However, they could only have done so within highly permeable sands, finer sediments being insufficiently porous to allow sufficient water flow past the well-developed petals. (j) Infaunal selective particle feeders living within coarse, LATE CAMPANIAN-MAASTRICHTIAN ECHINOIDS A Mo Jebel Rawdah, section 2: = Ad on beds 3-10 5 2 90 N= 395 Dj i % o Yes in c 8 2 so a o 6 =) ies) < 3 Sh a S 70 oo = 5 Ww Q eo) £ > — Io} ie rs E 40 2 30 20 10 Species B a 700 2B Jebel Rawdah, section 2: 2§ 600-] beds 11, 12 Rs oO 2 Bs 5004 N = 790 9 ss Q 400 = g 5 3.9 GS YG S 25 300 be ss 200 > oy 2 ao 100 90 2 & = io} iy S 70 i 2 3 S iS i= a 6 5 g S re 40 =| 2 0 oa) Species Jebel Rawdah, section 2: 200— beds 14-26 Petalobrissus ‘awdahensis 1804 N = 508 160 140 oO 120 3 Q s 100 3 .s) 90 o Ny wo 7) a o 3% a = 3 7 o z Ss 2 5 3 > 70 re) iS SS so} 3 5 Ee [= Q ‘= mB oS re 2 = By no S 8 2 Se alhitiuast yo 0 a aS) o 2S 3 “ : eet ee ~-0 a i 2 a wo ie) = gs 8 38 5 & = 90 € Qa is = & 3 Y = 8 ro) a 2» st 2 Species Fig. 4 Echinoid species abundances within units at Jebel Rawdah, section 2. A, fauna from beds 3-10; B, fauna from beds 11 and 12; C, fauna from beds 14~26. Species are listed numerically along the x-axis as in Table 1. N = total number of specimens collected. 1 fT 128 A.B. SMITH Table 2 Inferred palaeoenvironmental settings for the late Cretaceous succession of the Oman-United Arab Emirates border region. The common echinoid taxa found in each habitat are listed, together with indications of their probable life style. Habitat A nearshore hardground dwellers living in zone of active wave surge. (i) Hardground grazers: Echinotiara perebaskinei, Phymechinus? perplexus, Codiopsis lehmannae. Habitat B nearshore hardground dwellers living in more protected environments (perireefal). (i) Hardground grazers: Goniopygus arabicus. Habitat C subtidal protected environments within normal wave base; mixed cobble and sand substrata with nearby level-bottom reefal thickets. (i) Epifaunal grazers: Glyphopneustes hattaensis, Mimiosalenia quinquetuberculata, Goniopygus arabicus, Phymosoma hexoaporum. (ii) Epifaunal browsers: Circopeltis? emiratus, Heterodiadema buhaysensis, Orthopsis miliaris, Hattopsis paucituberculatus, Plistophyma | asiaticum. (iii) Epifaunal generalists: Prionocidaris morgani, Prionocidaris? emiratus, Salenia nutrix, Salenia microprocta. (iv) Infaunal grazers/detritivores: ‘Globator’ bleicheri, Conulus douvillei, Coenholectypus cf. baluchistanensis. (v) Infaunal bulk sediment swallowers: Nucleopygus magnus, Pygurostoma morgant. } (vi) Infaunal selective deposit feeders: Vologesia rawdahensis, Arnaudaster cylindriformis, Hemiaster hattaensis. Habitat D Shore-face sand flats at or within normal wave base. (i) Bulk sediment swallowers: Petalobrissus rawdahensis, P. ct. setifensis. (ii) Selective deposit feeders: Zuffardia morgani, Faujasia eccentripora. (iii) Infaunal generalist detritivores: ‘Globator’ bleicheri, Coenholectypus cf. baluchistanensis. Habitat E Broad shallow, open shelf subtidal sand flats, within normal wave base (2-10 m) but distant from the platform margin and thus relatively protected. i (i) Epifaunal grazers: Noetlingaster paucituberculatus, Hattopsis sphericus. (ii) Semi-infaunal selective deposit feeders: Hemipneustes spp. (iii) Infaunal selective deposit feeders: Mecaster victoris, Hemiaster paronai, Stigmatopygus? pulchellus, Faujasia eccentripora. (iv) Infaunal bulk sediment swallowers: Petalobrissus rawdahensis, P. cf. setifensis, Pygurostoma morgani. Habitat F Deeper-water (20 m plus) platform basinal muddy sands, below normal wave base. (i) Epifaunal grazers: possibly Actinophyma spectabile. (ii) Infaunal selective deposit feeders, harvesting detritus from the sediment/water interface: Proraster geayi. permeable sands behind fringing reefs in 0-10 m water depth. stabilized bottoms in shallow water conditions protected from Conulus, ‘Globator’ and Coenholectypus belong here, though the influence of strong wave action. The presence of ‘Globa- the highly inflated C. inflatus may have reverted to a prima- tor implies that the sediment was unconsolidated, permeable rily epifaunal mode of life, below normal wave base. Globa- and rather coarse, while horizons with an abundance of | tor closely resembles the present day Echinoneus whose Hattopsis might suggest the presence of nearby algal stands. ecology was described by Rose (1976). Echinoneus lives The abundance and diversity of regular echinoids (most of beneath coral debris in coarse shell-sands passing suitable which are algivore grazers) implies a mixture of environments | grains into its peristome by means of its tube-feet. were close by, ranging from rocky (?patch-reef, as seen at Inferred depths and habitats for typical Omani echinoids Jebel Faiyah) to stabilised sedimentary bottoms with algal | are tabulated in table 2. stands. The palaeoenvironment is therefore most likely to represent a shallow (5-10 m) backreef or leaward environ- ment that supports a high algal diversity. ECHINOID ASSEMBLAGES (2). Echinotiara/Faujasia/Phymechinus assemblage. An assemblage which includes a number of rarer regular echi- Because careful count of the number of specimens collected noids absent from most other assemblages (e.g. Codiopsis, for each taxon was kept it is possible to identify recurring Phymechinus). Faujasia is the only common element found at assemblages. The following seven assemblages are differenti- other levels, but no other irregular echinoid occurs in any | ated here. abundance (Fig. 4A). This assemblage is best developed at Jebel Rawdah section (1). Conulid/arbacioid assemblage. An abundance of Conu- 2, in the basal calcarenitic beds with associated coral/rudist lus and ‘Globator’ which is usually accompanied by a diverse debris. mixture of regular echinoids, notably by the arbacioids Interpretation: this assemblage is dominated by hard- Goniopygus, Glyphopneustes, Mimiosalenia and Hattopsis. ground, shallow-water (0-5 m) epifaunal regular echinoids Other taxa making up a smaller component of the fauna (Echinotiara, Phymotaxis, Codiopsis) adapted for life in include Phymosoma, Nucleopygus and Pygurostoma (Figs strong wave-surge environments. Clearly none are preserved 2B, 3, 5B, 6). in situ, but have been transported into immediately adjacent This assemblage is characteristic of the lower beds of the basins of sediment accumulation, along with coral and stro- Simsima Formation at Jebel Buhays, Jebel Thanais, Jebel matoporoid debris and hippuritid rudist debris indicative of Faiyah, and Jebel Huwayyah. It is typically developed with reef habitat. Transportation has not been far, otherwise much the income of relatively coarse, carbonate bioclastic sands in more disintegration of tests might be expected. The occur- the succession. rence of large blocks of rolled coral with stromatoporoid and _Interpretation: the conulid/arbaciid assemblage has a very rudist debris at Jebel Rawdah section 3, implies that the high diversity of regular echinoids that would have preferred deposits accumulated close to actual reefal framework. Fauja- LATE CAMPANIAN-MAASTRICHTIAN ECHINOIDS , Mecaster victoris Petalobrissus rawdahensis Globator " bleicheri Species A A *°] Jebel Rawdah, section 3: beds 1-4 _E@ g 50 N = 88 g Jebel Rawdah, section 4: beds 1-7 ao N=25 & w 30 ° 2 5 20 E 5 10 z ee B | Jebel Rawdah, section 3: beds S+ @ sof N= 53 zg Jebel Rawdah, section 4: beds 8+ §S > 1 N=49 £ w 30 ° Ww 5 20 E 5 10 7a 60 : ; | i Jebel Faiyah, section 1: beds 8-10 ! —_ w = S 59 N=77 3 | 2 40 s aS) o | £ a 3 — fo} a) n es @ 20 28 | E 10 es | z Ss 5 10 15 20 Table 1. N = total number of specimens collected. fa may be the only in situ echinoid for the depositional avironment, living within the surrounding sand-fields in an nmediately subtidal environment, although none is pre- ‘rved with associated spines. Goniopygus is found in asso- ation, and it probably lived amongst the tallus derived from ie reef crest rather than on the crest itself. ) Petalobrissus assemblage. At certain levels of coarse, consolidated orbitoline limestones Petalobrissus occurs in ige abundance, accompanied by much smaller numbers of her infaunal echinoids (Zuffardia, Faujasia, Coenholecty- js and ‘Globator’). Rare Noetlingaster are the only regular hinoids encountered (Fig. 4B). Globator bleicheri . Species )Fig.5 Echinoid species abundances within units at Jebel Rawdah, sections 3 and 4. A, fauna from lower beds; B, fauna from higher beds. Species are listed numerically along the x-axis as in Table 1. N = total number of specimens collected. Globator bleicheri Species ‘ig.6 Echinoid species abundances within units at Jebel Faiyah, section 1, beds 8-10. Species are listed numerically along the x-axis as in This assemblage is seen only in the lower part of the section (beds 11-13) at Jebel Rawdah, section 2. Interpretation: this assemblage is composed entirely of small, bulk sediment swallowers and is suggestive of shallow subtidal unconsolidated and well-washed sand flats free of algae, presumably subject to constant disturbance. The beds can thus be envisaged as shallow (0-10 m) orbitoline sand flats developed behind a fringing reef. The abundance of individuals suggests that this was a region of relatively high productivity. (4) Hemipneustes/Noetlingaster assemblage. The abundance of large infaunal selective deposit feeders, notably Hemip- 130 neustes and Pygurostoma, characterises this assemblage. Usu- ally Noetlingaster-rich levels alternate with holasterid-rich levels, so it may not be entirely correct to treat these as one assemblage, though they occur intermixed within the same lithofacies. All the species are large in comparison to those elsewhere. This assemblage typifies the upper beds of section 2, Jebel Rawdah (Fig. 4C). Interpretation: the Hemipneustes/Noetlingaster assemblage is composed of selective deposit feeders specialized for life within coarse, permeable sands, together with occasional specific bands of the large generalist algivore Noetlingaster. The dominance of large selective deposit feeders implies a relatively nutrient-starved environment where large-scale harvesting of the sediment/water interface is needed to exploit the food supply. Associated lithofacies evidence sug- gests that they were living within shallow-water (2-10 m) flat-bottom shoals, probably immediately infratidal. Noetlin- gaster, like Echinus in today’s seas, may have been a general- ist detritivore/algivorous browser. The environment of deposition is therefore envisaged as relatively quiet sedimen- tary level bottom shoals that lay not particularly deep but distant from the high productivity reef area or from more exposed platform margin conditions. (5) Faujasia assemblage. Occasional, almost monospecific assemblages of Faujasia are found in calcified fine sands at Jebel Huwayyah near the base. In the associated Loftusia- rich levels there are also occasional Hemiaster and Proraster. These may be nearshore infratidal sands in relatively pro- tected environments (Fig. 2A). (6) Mecaster assemblage (Fig. 5A). Another virtually monospecific assemblage confined to a single level at Jebel Rawdah section 3. This time the only associated echinoids are the rare Hemipneustes. (7) Proraster assemblage. Only Proraster is found in the highest, muddiest limestones of Jebel Huwayyah. The rela- tively fine-grade sediments and the absence of infaunal cassiduloids or holasteroids suggests that these beds may represent shelf-basinal deposits more than ca. 20 m deep. Only highly specialist infaunal forms such as Proraster could apparently cope with life in these relatively nutrient-poor mud-rich deposits. PALAEOECOLOGICAL SYNTHESIS The faunal succession records a relatively rapid deepening from beach boulder beds and sands through nearshore, high-energy, subtidal conditions with off-beach coral-rudist reefal patches to somewhat deeper-water conditions at or immediately beneath wave base. Maximum echinoid diversity is found in the shallow sand fields strewn with shell and coral debris surrounding coral thickets that formed at about wave- base around the shores of the uplifted ophiolite. Subsequent shallowing over the platform led to the creation of broad, shallow shoals supporting a low-diversity infaunal echinoid | assemblage, dominated by holasteroids and cassiduloids. The only epifaunal echinoid here is Noetlingaster. The deepest water facies are probably those seen at the top of the section at Jebel Huwayyah, where only the infaunal spatangoid Proraster occurs. SYSTEMATIC DESCRIPTIONS Class ECHINOIDEA Leske, 1778 Subclass CIDAROIDEA Claus, 1880 Order CIDAROIDA Claus, 1880 Family RHABDOCIDARIDAE Lambert, 1900 Gen. et sp. indet. Pl. 1, fig. 1; Figeey MATERIAL. A single specimen, comprising three interambu- | lacral plates, BMNH EE3438. OCCURRENCE. From the scree at Jebel Buhays, section 1; derived from the lowest 2-3 metres of the Simsima Forma- tion. DESCRIPTION. Three interambulacral plates, rather badly weathered, from the ambital region of the test. Plates are, 18-7 mm wide by 8 mm tall and appear to have tesselate sutures. Each plate carries a single large tubercle with a circular, confluent areole that occupies the full height of the plate. The boss is surmounted by a broad, strongly crenulate Lithofacies 4 Well-sorted orbitoid- rich packstones oe | ! | Hemipneustes spp. Coenholectypus inflatus Tithotacies) 3b Pygurostoma Stigmatopygus 3 Poorly-sorted orbitoid- Noetlingaster y rich packstones with thodolites and Dictyoptychus Petalobrissus spp. Zuflardia Hattopsis Hemiaster Lithofacies 3c | Muddy calcarenites: very poorly sorted. Loftusia and Cunnolites prominent. 1 I Bedrock | | Shore-face I Echinotiara | clastic sands Codiopsis | | Phymechinus? Faujasia Lithofacies 3a Poorly-sorted bioclastic calcarenites Goniopygus Glyphopneustes Salenia Orthopsis Phymosoma Nucleopygus Globator Coenholectypus baluchistanensis | } Fig. 7 Schematic reconstruction of the sea-floor in Maastrichtian times showing the range of environments represented in the study area. See urchins are illustrated in their probable mode of life and the inferred environmental ranges for key taxa are shown. LATE CAMPANIAN-MAASTRICHTIAN ECHINOIDS Fig. 8 Camera lucida drawing of an interambulacral plate of Rhabdocidaridae gen. et sp. indet., BMNH EE3438. Scale bar = 5mm. platform while the mamelon itself is rather small (1-2 mm diameter) and has a large central perforation. The primary tubercle lies subcentrally on the plate with a broad adradial and interradial platform on either side. The broad zones outside the areole of the primary tubercle have a few scat- tered secondary tubercles preserved, but are otherwise too abraded to retain any evidence of fine tuberculation that may have been present. It is clear from the distribution of the tubercles that are preserved that these regions were covered in a rather heterogenous and open array of various-sized tubercles. REMARKS. The broad plates, confluent areoles, perforate crenulate tubercles and heterogeneous secondary tubercula- tion show this to be a member of the Rhabdocidaridae. It is impossible to place this specimen in any genus on the basis of such incomplete material. However, it most probably repre- sents a species of Rhabdocidaris itself, judging from the size of the specimen and the coarseness of the secondary tubercu- lation. Family CIDARIDAE Gray, 1825 Tribe CIDARINI Gray, 1825 Subtribe PHYLLACANTHINA Smith & Wright, 1989 Genus PRIONOCIDARIS Agassiz, 1863 Prionocidaris morgani (Gauthier, 1902) Pl. 1, figs 2-4; Fig. 9B Rhabdocidaris (Leiocidaris) morgani Gauthier: 145, pl. 20, figs 3-6. 1989 Cidaris cf. scabra Gauthier; Ali: 398, fig. 2.1. _ Types. The syntypes are the two specimens described and figured by Gauthier. One is a more or less complete test 35 _mm in diameter, the other an interambulacral segment. The whereabouts of this material is unknown: the specimens are 1902 131 not in the Morgan collection in the Museum National d'Histoire Naturelle, Paris. MATERIAL STUDIED. Two specimens: BMNH EE3433, from the scree derived from the lowest beds of the Simsima Formation at Jebel Buhays, section 1; BMNH EE34335, loose in the scree at section 3b, Jebel Rawdah. A third fragment, EE3434, probably belonging to this species, was also found loose in the scree at Jebel Buhays. OCCURRENCE. The syntypes come from the late Cretaceous (‘Senonien superieur’) of Louristan, Iran (Gauthier was unable to give a precise locality). In the United Arab Emirates the species comes from the Maastrichtian of Jebel Buhays and Jebel Rawdah. DIAGNOSIS. A species of Prionocidaris with broad conjugate pore-pairs occupying more than half of the ambulacral plate width, and a perradial zone with a single primary tubercle and a row of tubercle-like granules, up to four abreast, on each plate. Interambulacra with well differentiated scrobicular circles and aligned extrascrobicular tuberculation. DESCRIPTION. The following description is based on BMNH EE3433, the larger and better preserved of the two. The specimen comprises parts of two interambulacra and ambu- lacra. Interambulacral width at the ambitus is 23 mm, sug- gesting a test diameter of approximately 50 mm in life. Test height is 32 mm. The ambulacra are 6 mm in width and only slightly sinuous. The pore-pairs are very wide and conjugate (Pl. 1, fig. 4; Fig. 9B) and successive pore-pairs are closely spaced. The poriferous zone occupies approximately half of the plate width. There is a single primary tubercle with a mamelon immediately adjacent to each pore-pair. The remainder of the perradial zone is occupied by a row of large granules (non-mamelonate), some three or four abreast and organised into discrete vertical rows. Interambulacral plates are relatively broad, an ambital plate being 13-1 mm by 9-4 mm in height. All plates, except the most adapical in each column, have a single large primary tubercle. This has a sunken areole 7-2 mm in diameter and a mamelon 2.2 mm in diameter (PI. 1, figs 2, 4). Mamelons are perforate and the surrounding platform is non-crenulate. The primary tubercles lie offset towards the adradial suture leav- ing a broad interradial zone of miliary tuberculation. There is a clearly differentiated ring of scrobicular tubercles surround- ing each areole. Extrascrobicular tuberculation is well devel- oped with approximately 2 miliary granules abreast towards the adradial suture, eight abreast towards the interradial suture, and either a single row or no row developed adapical and adoral to the scrobicular circles. These extra-scrobicular tubercles are non-mamelonate and decrease in size towards Fig.9 Camera lucida drawings of ambital ambulacral plating in cidarid species. A, Stereocidaris persica (Cotteau & Gauthier), Museum @ Histoire Naturelle, Morgan Collection; B, Prionocidaris morgani (Gauthier), BMNH EE3433; C, Prionocidaris? emiratus sp. nov., | | | BMNH EE3431. Scale bar = 1 mm. o>) bo A.B. SMITH PLATE 1 Fig. 1 Rhabdocidarid gen. et sp. indet. BMNH EE3438; three interambulacral plates, x 3. Jebel Buhays, section 1; loose in the scree derived from the lowest 3 m of the Simsima Formation. Figs 2,4 Prionocidaris morgani (Gauthier). BMNH EE3433. 2, adapical portion of interambulacrum and adjacent ambulacral zones, X 2:5; 4, interambulacrum detail, x 4-5. Jebel Buhays, section 1; loose in the scree derived from the lowest 3 m of the Simsima Formation. Figs 3, 5,6 Prionocidaris? emiratus sp. nov. BMNH EE3431, holotype. 3, ambulacral zone detail, x 10; 5, lateral, x 2; 6, oral, x 2. Jebel Huwayyah, section 2, beds 3-6. Fig. 7 Cidarid spine, possibly belonging to Prionocidaris morgani (Gauthier). BMNH EE3434, x 3. Jebel Buhays, section 1; loose in the scree derived from the lowest 3 m of the Simsima Formation. LATE CAMPANIAN-MAASTRICHTIAN ECHINOIDS the interradius. There are neural grooves developed in the extra-scrobicular regions. The smaller specimen (BMNH EE3435) is more complete, but less well preserved. It has seven interambulacral plates in a column at an estimated test diameter of about 35 mm. Adapical interambulacral plates retain fully formed tubercles. Two cidarid spines were also collected from the scree at Jebel Buhays and probably belong to this species. The best preserved is BMNH EE3434 (PI. 1, fig. 7). It is 19 mm long by 2-6 mm wide. It has a short neck and a perforate, non-crenulate base. The shaft is widest midlength and tapers towards the tip, but then expands slightly at the very tip to end in a blunt crown. The shaft is covered in rows of thorns which coalesce to form ribs towards the tip. REMARKS. This species differs from the most common late Cretaceous cidarid described from Iran, ‘Cidaris’ persica Cotteau & Gauthier (1895), in having much finer and denser extra-scrobicular tuberculation and wider and more strongly conjugate pore-pairs at a comparable size (compare Figs 9A, B). ‘C’. persica belongs to the genus Stereocidaris and has a characteristic strong interporal ridge developed between pore-pairs. Cidaris scabra Gauthier (1902) was established for a 30 mm diameter individual of uncertain provenance within Louristan. It differs primarily in having two ambulac- ral tubercles to each plate but has very similar strongly conjugate pore-pairs. Ali (1989) described a specimen of P. morgani from Jebel Rawdah under the name Cidaris cf. scabra Gauthier. Spines comparable in morphology to the one described here have previously been described under the name Cidaris aftabensis Gauthier (1902). They can only tentatively be assigned to this species until specimens are found that are attached to a test. -Prionocidaris? emiratus sp.nov. Pi. figs 3, 5,6; Fig. XC TyPE. Holotype and only known specimen, BMNH EE3431. ) OCCURRENCE. Found loose in the Loftusia levels a little \ below Beds 6/7, section 2, Jebel Huwayyah. ‘DIAGNOs!s. A cidarid with conjugate ambulacral pore-pairs and a single primary ambulacral tubercle to each plate, adjacent to the pore-pair. The perradial zone of each ambu- lacrum is filled with small, dense miliary tuberculation, three or four to a plate and arranged in two rows. DESCRIPTION. Test 28-5 mm in diameter by 18-9 mm in height. Ambulacra 3-6 mm in width at the ambitus, slightly sinuous. Pore-pairs strongly conjugate, with individual pores af each pair ca. 0-3 mm diameter and separated by an |nterporal distance of about 0-7 mm (PI. 1, fig. 3; Fig. 9C). ?erradial zone elevated and more or less flat; comparatively ‘hort and tall. A large primary tubercle occupies about half of he tuberculate region on each plate (PI. 1, fig. 5; Fig. 9C). These tubercles are almost contiguous vertically. The central art of the perradial zone is occupied by very much smaller tanules, some three or four to a plate, irregularly scattered. | There are six interambulacral plates in a column, of which 1e most adapical in each zone has only a rudimentary dbercle. Ambital plates are relatively tall and narrow, 7-0 im wide and 5-8 mm high (Pl. 1, fig. 5). The primary ubercle lies centrally on the plate and has a sunken areole 4 1m in diameter (on ambital plates). The mamelon (1-1 mm 133 in diameter) is perforate and non-crenulate. The surrounding scrobicular circles are non-contiguous. Extra-scrobicular tuberculation is very fine and dense. There are approximately three to four granules abreast on the interradial margin, three abreast adapically and two abreast both adradially and adorally. REMARKS. The strongly conjugate pore-pairs indicate that this is a member of the Phyllacanthina, and in many respects it is close to P. morgani in tuberculation and appearance. The interambulacra are narrower than usual for Prionocidaris, and the interradial zones less well developed. Therefore, it is only tentatively assigned to this genus. P? emiratus can be easily differentiated from P. morgani by its ambulacral tuber- culation (compare Figs 9B and 9C). None of the cidarid species described by Cotteau & Gauthier (1895) or Gauthier (1902) have this form of tuberculation. Indet. cidarid plates In addition to the specimens described above, fragmentary material of cidaroids has been collected from the basal beds at Jebel Faiyah, section 1, and the Loftusia levels (beds 1-6) at Jebel Huwayyah. None of this material is adequate to determine to generic level. ?Order DIADEMATOIDA Duncan, 1889 Family HETERODIADEMATIDAE Smith & Wright, 1993 Genus HETERODIADEMA Cotteau, 1864 Heterodiadema buhaysensis sp. nov. Pl. 2, figs 1-3; Figs 10, 11 Types. Holotype BMNH EE3442-5, EES019. OTHER MATERIAL. One other specimen, BMNH EE3446. BMNH EE3441; paratypes, OCCURRENCE. Five specimens come from the lowest beds of the Simsima Formation and were collected in the scree derived from the lowest few metres of that formation at Jebel Buhays, section 1. One was collected from the lowest 1-5 m of Simsima Formation at Jebel Thanais. One specimen was collected from bed 5 at Jebel Rawdah, section 3b. DIAGNOsIS. Apical disc large, pentagonal, caducous; prob- ably monocyclic to judge from the outline. Ambulacra trigeminate, relatively wide with diadematoid-style plate compounding. Phyllodes absent. Primary tubercles perforate, crenulate; reducing in size sharply adapically and tending to become imperforate. Broad zones of granulation along the perradius, adradius and interradius. Peristome invaginated with reasonably deep and sharply defined buccal notches with tags. DESCRIPTION. Tests range in diameter from 18-36 mm. The test is circular in outline and depressed in profile, with a height 42-48% of the diameter (mean = 45%; Fig. 10). The ambitus lies at about mid-height and the sides are uniformly rounded (PI. 2, fig. 3). The apical disc and peristome are slightly invaginated in the largest specimen. Coronal plates are all firmly sutured together. The apical disc is caducous and has been lost from all specimens. The apical disc opening is pentagonal in outline with the angles projecting reasonably strongly into the inter- radii (slightly more into the posterior interradius, which isalso 134 14, = Ambulacrum e @ Interambulacrum 5 Exi2 ~ ” 3 = 210 l= tJ = =o ~ = = >>) 4 10 20 30 40 Test diameter (mm) 18 a E16 < ~ 14 @ £ ~ a 6 12 B ke 10 8 10 20 30 40 Test diameter (mm) 60 g Ambulacral pore-pairs Py er @ \Interambulacral plates Number in a column 10 20 30 40 Test diameter (mm) Fig. 10 Biometric data for Heterodiadema buhaysensis sp. nov. Kig. 11 Camera lucida drawing of three ambital ambulacral plates of Heterodiadema buhaysensis sp. nov., BMNH EE3444. Scale bar = 1 mm. A.B. SMITH more rounded). It is a little longer than broad with a length that is 36-43% of the test length. Ambulacra are broad and nearly parallel-sided at the ambitus but taper gradually both adapically and adorally. At the ambitus they have a width that is 23-25% of the test length. Pore-pairs are uniserial throughout, without any widening whatsoever at the peristomial margin. Plates are compound, with diadematoid-style triads in which the central element is the largest and upper and lower elements both taper towards the perradial and adradial suture (Fig. 11). Each triad bears a primary tubercle and there is a well- developed perradial band of miliary tuberculation that expands adapically. At the ambitus in the largest specimen perradial tubercles are about three abreast. Oral and ambital tubercles are perforate and crenulate, but tubercles reduce sharply in size above the ambitus, where they may become imperforate and non-crenulate (some remain perforate). This reduction in size occurs above about the seventh compound plate. Larger tubercles have circular, non-confluent areoles. At the ambitus pore-pairs are rather widely spaced and separated by a single row of miliary tubercles. There are no obvious sphaeridial pits around the peristome. Interambulacra are about 1-3-1-5 times as wide as the ambulacra. There are 15 plates in a column at 36 mm test diameter. All plates have a single large primary tubercle that is perforate and crenulate and is surrounded by a circular areole. The mamelon is notable for its small size in compari- son to the size of the areole. Areoles are non-confluent, being separated for the most part by a single band of miliaries. Tubercles decrease in size sharply towards the apex and may become imperforate. There are broad zones of uniform miliary tuberculation both adradially and interradially: about six abreast adradially and three or four abreast interradially at the ambitus in the largest specimen. Scrobicular tubercles are not differentiated. The peristome is relatively large, 28% of test diameter in a 36 mm diameter individual. It is strongly indented by the buccal notches. There are broad, smooth tongue-like regions running adradially from each buccal notch and extending up to the fourth interambulacral plate. REMARKS. This species is most closely related to the mid- Cretaceous (Cenomanian) Heterodiadema lybica (Agassiz & Desor). Both have a similar form of ambulacral compounding and a very similar tuberculation style. Granular zones are broad and well developed and in both there is a sharp reduction in the size of primary tubercles above the ambitus, with those adapically tending to become imperforate and non-crenulate. Finally, both have the distinctive tubercle-free tag to the buccal notches and uniserial pore-pairs throughout. H. buhaysensis differs from H. lybica in having fewer reduced tubercles, the primary tubercles becoming smaller consider- ably closer to the apex in H. buhaysensis in individuals of comparable size. It also differs in lacking the extreme prolon- gation of the apical disc into the posterior interambulacrum seen in medium to large specimens of H. lybica. Heterodiadema was placed in its own family by Smith & Wright (1993). Where this family fits into the higher tax- onomy of echinoids, however, is much less certain. They may be members of the aulodont group Diadematoida. This is suggested by the prominent buccal tags and the very delicate mamelons on the primary tubercles, which indicate that the species had small, fragile spines. Alternatively they may be stirodonts, early members of the Phymosomatoida (as LATE CAMPANIAN-MAASTRICHTIAN ECHINOIDS ) ATE 2 lis 1-3 Heterodiadema buhaysensis sp. nov. BMNH EE3441, holotype; 1, oral; 2, apical; 3, lateral; all x 3. Jebel Buhays, section 1; loose m the scree derived from the lowest 3 m of the Simsima Formation. Is4,5 Orthopsis miliaris (d’ Archiac). Topotype specimen of Orthopsis morgani Cotteau & Gauthier, from the Morgan Collection, Museum d’Histoire Naturelle, Paris, x 2. Senonian, Khianan, Iran. 136 Test height (mm) 30 40 50 Test diameter (mm) 20 @ Ambulacral pore-pairs ® \nterambulacral plates Number in column Test diameter (mm) Fig. 12 Biometric data for Orthopsis miliaris (d’ Archiac). defined by Smith & Wright, 1993). Diadematoids and phy- mosomatoids are fundamentally differentiated on the struc- ture of their spines and lantern apparatus, diadematoids having hollow spines and grooved teeth, phymosomatoids having solid spines and keeled teeth. In neither H. /ybica nor in our new species are the spines or lantern known, thus its higher taxonomic position must remain unresolved. Because of its delicate tuberculation, it is tentatively assigned here to the Diadematoida. Infraclass ACROECHINOIDEA Smith, 1984 Plesion (Order) ORTHOPSIDA Mortensen, 1942 Family ORTHOPSIDAE Duncan, 1889 Genus ORTHOPSIS Cotteau, 1864 Orthopsis miliaris (d’ Archiac, 1835) PI. 2, figs 4, 5; Pl. 3, figs 1-9; Figs 12-14 1835 Cidarites miliaris @ Archiac: 179, pl. 11, fig. 8. 1895 Orthopsis morgani Cotteau & Gauthier: 87, pl. 14, figs 6-9. 1933. Orthopsis sanfilippoi Checchia-Rispoli: 6, pl. 1, figs 5-15. 1985 Orthopsis miliaris (d’ Archiac); Geys: 143, pl. 5, figs 8-10 (see also for full prior synonymy). 1989 Orthopsis morgani Cotteau & Gauthier; Ali: 401, fig. A(T). 1991 Orthopsis miliaris (d Archiac); Smith & Bengtson: 30, pl. 8B-F, text-fig. 23. A.B. SMITH | 30 | ws Ambulacrum e @ Interambulacrum £ — $20 | oS | a ; E @ | s | @ £ 10 3 = | | 0 10 20 30 40 50 Test diameter (mm) 20 @ Peristome = ey @ Apical disc £ £ ~~ : | £10 | Ss = | | ) | 20 30 40 50 Test diameter (mm) MATERIAL STUDIED. Forty seven specimens of which the | following 11 were used in the biometric analysis: BMNH EE3720-21, E3723, EE3725-26, EE37/28, EE373iy EE3733, EE3738, EE3740, EE3749 and EES018. OCCURRENCE. In the western Oman mountains this species | occurs at the following localities and horizons (numbers in brackets are number of specimens): Jebel Buhays, section 1: loose in scree derived from lowest | few metres of the Simsima Formation (13) Jebel Thanais: lowest 2 m of Simsima Formation (1). | Jebel Huwayyah, section 2: Loftusia levels (beds 3-5; 1| fragment). Jebel Rawdah, section 1: beds 3 and 4, and loose (4). Jebel Rawdah, section 3b: bed 2 (1); bed 3, 1m above base (1), bed 8 (2); bed 9 (8); bed 10 (1); loose (3). Jebel Rawdah section 4: bed 12 (5);top of bed 15 (1); loose, | lower half of sandy beds (1). Jebel Faiyah, section 1a: top of bed 2 (1). DESCRIPTION. Tests range from 20 mm to 48 mm in diameter and are circular in outline and bun-shaped in profile. Test height is 39-52% of test diameter (Fig. 12) and in profile the ambitus lies about one third the height above the base. The apical disc is dicyclic, though occasionally one of the posterior oculars may just be exsert. The apical disc occupies 17-24% of the test diameter (mean = 21%, SD = 1-:9%, N= 10; Fig. 12). Genital plates are broad and crescentic in outline, except for the madreporite, which is larger and more pentagonal in outline (Pl. 3, fig. 7; Fig. 13A). Madrepores occupy almost the entire surface of the madreporite plate and LATE CAMPANIAN-MAASTRICHTIAN ECHINOIDS there are small scattered tubercles amongst the openings. Gonopores are present even in the 20 mm diameter indi- vidual. Ocular plates are small and pentagonal. All plates have small secondary tubercles, those on the genital plates tending to form a circle around the periproct. The periproct is irregularly oval in outline and occupies 10-14% of the test diameter. Ambulacra are 17-21% of the test diameter in width at the ambitus (mean = 20%, SD = 1:3%, N = 11). Plating is trigeminate throughout and pore-pairs are arranged either Fig. 13. Camera lucida drawings of plating in Orthopsis miliaris (d’Archiac). A, apical disc plating, BMNH EE3723; B, ambital ambulacral (left) and interambulacral (right) plates, BMNH EE3723; C, adoral plating, ambulacrum to left, interambulacrum to right, BMNH EE3733. Scale bars: A, B = 1 mm; C = 2 mm. A Fig. 14 Camera lucida drawings of lantern elements in Orthopsis miliaris (d’ Archiac): A-D, BMNH EE3735; E, BMNH EES5020. A, pyramid with rotula and tooth in position; B, same in lateral view; C, fragment of compass element; D, adapical view of pyramid, with proximal end of tooth (broken) towards the top; E, cross-section of a single tooth. Scale bars = 1 mm. 137 uniserially or in very weak arcs of three (PI. 3, fig. 6). All ambulacral elements are narrow and elongate and reach the perradius (Fig. 13B). A primary tubercle (perforate and non-crenulate) straddles two of the three elements in each compound plate (PI. 3, fig. 6). The third element carries two small secondary tubercles and an intermediate row of miliary granules. There are secondary and miliary tubercles down the perradius also. Adorally only the first five or so pore-pairs are offset to form a weak phyllode (Fig. 13C). There are 55 pore-pairs in a column at 20 mm test diameter, rising to around 90 at 46-48 mm diameter (Fig. 12). Sphaeridial pits are very shallow and are present on the two or three most adoral compound plates, immediately adjacent (perradial) to the pore-pair on the element in each triad that does not support a primary tubercle. Interambulacra are 38-44% of the test diameter in width at the ambitus (mean = 41%, SD = 2:0%, N = 11) (Fig. 12). Ambital plates are short and wide and slightly curved. At the ambitus most specimens have a single primary tubercle, centrally positioned, plus two smaller secondary tubercles one on either side (PI. 3, figs 3, 8; Fig. 13B). However, in a fragment of a large specimen (ca. 65 mm diameter, BMNH EE3717), there is a fourth large tubercle on the interradial margin. The remainder of the plate carries scattered tertiary tubercles and granules. All tubercles are perforate and non- crenulate. Adorally there is no primordial plate. There are 14 plates in a column at 20 mm test diameter, rising to 18 at 42-48 mm test diameter (Fig. 12). The peristome is 36-40% of the test diameter in diameter (mean = 38%, SD = 1:7%, N = 7). It is slightly invaginated. There are strong buccal notches that extend into the second interambulacral plate (PI. 3, fig. 9; Fig. 13C). The perignathic girdle is seen in BMNH EE3733. The auricles are well developed. They meet and are fused above the perradius to form a continuous arch. Lantern elements are preserved within the tests of several specimens, but are best seen in BMNH EE3735. Hemipyra- mids are tall and have a relatively shallow foramen magnum, only some 33% of their height (Figs 14A, B). There is a clear muscle attachment flange on the outer edge of each. The upper surface of the lantern is also pitted. There are well- developed processus superalveolares that extend to the tooth. The lateral (inter-pyramidal) face is denticulate along its inner edge and has the usual horizontal series of ridges for muscle attachment. The epiphyses are not clearly seen in any specimen but appear relatively short. They are definitely not extended above the foramen magnum nor are they fused together as is the case in camarodont lanterns. There is a single compass element that is siender and has a small head (Fig. 14C). The tooth is strongly keeled and is best seen in cross section in BMNH EE5S020 (Fig. 14E). Spines are seen associated with lantern elements in BMNH EE3735. They are relatively short and needle-like, with a small base and no cortex. The spines are solid, not hollow. REMARKS. The lantern of Orthopsis was described from late Cretaceous specimens of O. sanfilippoi Checchia-Rispoli (here treated as a synonym) by Serra (1934), who gave only a sketchy illustration of the apparatus, but pointed out its keeled teeth. Serra, and later Mortensen (1943: 11) described the latern as camarodont. However, this is clearly not the case since the epiphyses are not fused together to form a brace for the tooth. Instead both the hemipyramid and the A.B. SMITH PLATE 3 Figs 1-9 Orthopsis miliaris (d’Archiac). 1-3, BMNH EE3740; 1, oral; 2, apical; 3, lateral; all x 2. Jebel Rawdah, section 3, bed 9. 4, 5, 7, BMNH EE3725; 4, apical, x 2; 5, lateral, x 2; 7, apical disc, x 4. Jebel Buhays, section 1; loose in the scree derived from the lowest 3 m of the Simsima Formation. 6, 8, 9, BMNH EE3733; 6, ambital ambulacrum, x 6; 8, ambital interambulacrum, x 6; 9, peristomial region, x 6. Jebel Rawdah, section 1, bed 3. LATE CAMPANIAN-MAASTRICHTIAN ECHINOIDS epiphysis extend adaxially towards the tooth as in diadema- toid and stirodont lanterns (Jensen 1981, figs 31, 32). The epiphyses are well-separated and although they may actually reach the tooth and support it, they remain far apart. The tooth is undoubtedly keeled and similar to those of stirodonts and camarodonts. The phylogenetic position of orthopsids can now be clari- fied. The structure of their epiphyses and hemipyramids is primitive for euechinoids as a whole, thus there are no grounds for treating orthopsids as camarodonts. The keeled teeth and solid spines place orthopsids as acroechinoids and more derived than either diadematoids or echinothurioids. However, the simple acrosaleniid-style of ambulacral plate compounding and the perforate, non-crenulate nature of the primary tubercles differentiate it from all other acroechi- noids. They are best considered as an early plesion of the Acroechinoidea. Orthopsis morgani Cotteau & Gauthier (1895), from the late Senonian of southern Iran (PI. 2, figs 4, 5) is treated as a synonym. Order CALYCINA Gregory, 1900 Family SALENIIDAE Agassiz, 1838 Genus SALENIA Gray, 1835 Salenia nutrix Peron & Gauthier, 1881 PI. 4, figs 1-13; Pl. 17, figs 4, 5; Figs 15, 16A—-E, G, H 1881 Salenia nutrix Peron & Gauthier, in Cotteau, Peron & Gauthier: 167, pl. 18, figs 4-10. 1895 Salenia cossiaea Cotteau & Gauthier: 83, pl. 13, figs 13-19. 1902 Salenia cossiaea Cotteau & Gauthier; Gauthier: 149, pl. 18, fig. 12. 1928 Salenia cossiaea Cotteau & Gauthier; Lees: 659. 1932 Salenia lamberti Checchia-Rispoli: 6, pl. 2, figs 1-5. 1969 Salenia geometrica Agassiz; Devries: 167, pl. 1, fig. 1, pl. 4, figs 1-3. 1989 Salenia cossiaea Cotteau & Gauthier; Ali: 398, fig. 2.6. MATERIAL. 34 BMNH ~ EE3622-EE3654, EE3656. OCCURRENCE. In the western Oman Mountian sections stud- _ ied this species was found as follows: Jebel Aqabah; lowest few metres of the Simsima Formation (2). Jebel Buhays, section 1; loose in scree derived from lowest few metres of the Simsima Formation (7). Jebel Buhays, section 3: loose in scree (2). _ Jebel Faiyah, section 1: bed 8 (2). _ Jebel Rawdah, section 1; beds 3 and 4 (4); loose in scree (1). Jebel Rawdah, section 2; beds 6-8 (10); bed 11 (2); bed 13 (2); loose in scree (3). Jebel Rawdah section 3; bed 2b (1); bed 10 (1). Jebel Rawdah section 4; bed 10 (1). specimens, This species also occurs in the Upper Campanian of Nafun, 3-5 km west of Surayr, near Dumq, Oman (Skelton ef al. ' 1990). It was originally described from the ‘Etage dordonien _inferieur, couches a Heterolampas maresi’ (Late Campanian or early Maastrichtian) of Algeria. Later Gauthier (1902) described the same species from the Campanian of Tunisia. Cotteau & Gauthier described a very similar form from the 139 Senonian of Persia (Iran) under the name Salenia cossiaea (Pl. 4, figs 4-6). Finally, Devries (1967) described this species from Turkey under the name Salenia geometrica Agassiz. DIAGNOsIS. Rather inflated species of Salenia with seven to eight interambulacral plates at test diameters of 23-28 mm. Apical disc relatively small, circular and flat with periproct and suranal equal in size. Ocular I insert or exsert. Numerous small pits developed along all apical disc sutures in larger specimens, but largest along genital/ocular plate boundaries. Interradial granular zone well developed in specimens greater than 23 mm test diameter. DESCRIPTION. Tests range in diameter from 7 to 26 mm. Test height in specimens larger than about 10 mm is 60-76% of the diameter (mean 65%, N = 19; Fig. 15). The test is thus rather inflated in profile and in some specimens almost subglobular (Pl. 4, fig. 3). The apical disc is rather flat and only rises very slightly towards the apex. Its diameter is 42-55% of the test diameter (mean = 46%, N = 19). It is subcircular in outline. The suranal plate is relatively large, on average about 25% of the apical disc diameter. It is similar in size to the genital plates. The periproct is approximately the same size as the suranal plate, or very slightly larger, being on average 27% of the apical disc diameter. In some specimens ocular 1 is strongly exsert and forms the posterior wall of the periproct, but in other specimens ocular 1 is insert and separated from the periproct (Figs 16A-E). Approximately half of the specimens have the ocular plate insert. There is a slight elevation towards the periproct edge, but no true rim is developed. All plates are smooth and unornamented. The sutures are usually incised and may have a series of small pits along their length (Pl. 4, figs 1, 4, 7, 11). The ocular/genital plate boundaries always have pits that are more prominent than the rest. Gonopores are present on genital plates from approximately 10 mm diameter. Ambulacra are narrow and only very slightly sinuous towards the apex (PI. 4, fig. 13). They expand adorally to form a short phyllode. Plating is strictly bigeminate through- out. There are 45 pore-pairs in a column at 13 mm test diameter, rising to 78 at 26 mm test diameter (Fig. 15). Each compound plate has a primary tubercle that forms a contigu- ous row adjacent to the pore-pairs (Figs 16G, H). The perradial zone is narrow, but there is a single secondary tubercle on each compound plate and a single zig-zag row of miliaries also. On the oral surface up to 12 pore-pairs become crowded into a short phyllode in the largest specimens. There are approximately 15 or 16 ambulacral plates opposite an ambital interambulacral plate in individuals of 20-26 mm test diameter. Interambulacra remain relatively broad throughout. There are six interambulacral plates in a column from 10-16 mm test diameter, rising to eight by about 25 mm diameter (Fig. 15). Plates at the ambitus are slightly wider than tall and the primary tubercle lies towards the adradial margin, leaving a relatively broad interradial zone in specimens larger than about 22 mm diameter. Primary tubercles are imperforate and crenulate and are surrounded by five or six secondary tubercles (non-contiguous) (Pl. 4, fig. 13). The interradial zone has miliary tubercles from about 15 mm test diameter upwards and in larger specimens this forms a broad and distinctive band that runs almost to the peristome edge. The peristome is on average 48% of the test diameter 140 A.B. SMITH PLATE 4 Figs 1-13 Salenia nutrix Peron & Gauthier. 1-3, 13, BMNH EE3646; 1, apical, x 2; 2, oral, x 2; 3, lateral, x 2; 13, ambital detail, x 5. Jebel Buhays, section 1; loose in the scree derived from the lowest 3 m of the Simsima Formation. 4-6, B18723a, Museum d’Histoire Naturelle, Paris; topotype material from the Morgan Collection of Salenia cossiae Cotteau & Gauthier; 4, apical; 5, lateral; 6, oral; all x 3. Senonian, Kala é Melek, Iran. 7-9, BMNH EE3651; 7, apical; 8, oral; 9, lateral; all x 2. Jebel Buhays, lowest 2 m of the Simsima Formation. 10-12, BMNH EE3652; 10, lateral; 11, apical; 12, oral; all x 2. Jebel Buhays, lowest 2 m of the Simsima Formation. LATE CAMPANIAN-MAASTRICHTIAN ECHINOIDS 141 30 12 = = 10 ~ £20 Us z pied © 5 . Ege © £ = a a] ~ 2 10 ame cs FE = ab S 2 - 4 ae : 2 0 10 20 30 O 10 20 30 Test diameter (mm) Test diameter (mm) 4 15 5 ® Apical disc > r ® Periproct a - 3 a® BS eS ~ BS a £ o = a E + a o = i Ea = (® 2 & a B © a] — = 5 2 3 ) i~ 2 i ® a 2 4 6 8 10 12 14 «O 10 20 30 Test diameter (mm) Test diameter (mm) 80 3 ® |Interambulacral plates ® Ambulacral pore-pairs c E 2 r) 7) = _ @ 2 E = z 10) 10 20 30 Test diameter (mm) Fig. 15 Biometric data for Salenia nutrix Peron & Gauthier (squares) and S. microprocta sp. nov. (crosses). - across. It is hardly invaginated and has feeble buccal notches (Pl. 4, figs 2, 6, 8, 12). REMARKS. This species somewhat resembles S. geometrica - Agassiz, but differs from that species in having a relatively smaller apical disc and a larger periproctal opening. It also has much less pronounced pits developed at triple suture | junctions on the apical disc. Although there are clearly two | morphological forms, those with ocular 1 insert and those ) with ocular 1 exsert, the apical discs are in all other respects identical and the two forms co-occur in the various sections. I therefore treat the two forms as variants of the same species. ka form described by Checchia-Rispoli (1932a) as S. lam- berti from the Maastrichtian of Libya has all the characteris- tics of S. nutrix, except that ocular 1 is reportedly always insert. It too is treated as part of this same species complex. S. nutrix resembles S. loveni (Cotteau) in having a rather flat and smooth apical disc and in having expanded phyllodes adorally. However, it is very different both in the relative size of the apical disc (which in S. loveni occupies most of the upper surface) and in the coarseness of the tuberculation. S. maxima Arnaud and S. belgica Lambert are also similar in having fine sutural pitting but also have relatively much larger apical discs than is seen in S. nutrix. Fig. 16 Camera lucida drawings of plating in Salenia. A-E, apical discs of Salenia nutrix Peron & Gauthier; A, BMNH EE3646; B, BMNH EE3654; C, BMNH EE3647; D, BMNH EE3651; E, BMNH EE3627. F, apical disc of Salenia microprocta sp. nov. BMNH EE3657. G, H ambital ambulacral plating of Salenia nutrix Peron & Gauthier; G, BMNH EE3647; H, BMNH EE3638. I, ambital ambulacral plating of Salenia microprocta sp. nov., BMNH EE3657. Scale bars; A-D = 5 mm; E, F = 1 mm; G-I = 1mm. PIS, es 1S skies 15; 16F, I Salenia microprocta sp. nov Type. Holotype and only known specimen, BMNH EE3657. OCCURRENCE. Jebel Huwayyah, section 2: bed 1. DIAGNOsIS. A small, flattish Salenia with a relatively large, flat, pentagonal apical disc and a small oval periproct. Ambulacral tubercles are not contiguous and at the ambitus may be separated by granules. DESCRIPTION. Test 11-4 mm in diameter and 5-1 mm in height (45% of test diameter). Depressed in profile with a broad flat base and top. The ambitus is uniformly rounded. The apical disc is flat and pentagonal in outline, with the angles radial. The disc is 55% of the test diameter in length. The genital plates are approximately as broad as long and are all similar in size (Fig. 16F). Ocular plates protrude slightly. The suranal plate is relatively large, whereas the periproct is small (Pl. 5, fig. 1), being only 17% of the apical disc diameter along the plane of bilateral symmetry. There is no lip to the periproct. Sutural pits are present at all plate triple junctions and also mid-way between these junctions. Small gonopores are present. The ambulacra are slightly sinuous adapically, becoming straight adorally and expanding towards the peristome. There are 33 pore-pairs in a column and 16 primary tubercles. All plates are compound and bigeminate. At the ambitus the ambulacral width is 1-5 mm with the perradial tuberculate zone making up 1-0 mm of this width. Primary tubercles are relatively small and are not contiguous with their neighbours. There is a single row of miliary tubercles perradially and, at the ambitus, there is also a single row of miliaries between successive primary tubercles (Pl. 5, fig. 3; Fig. 161). Small phyllodes are developed adorally. A.B. SMITH Interambulacra are composed of five plates in each col- | umn. Each has a single primary tubercle surrounded by six widely-spaced scrobicular tubercles. There is a single row of miliary tubercles down the interradius. The peristome is very slightly invaginated and is 50% of the test diameter in diameter. There are feeble buccal notches. REMARKS. No other Salenia species has ambulacra with primary tubercles separated by rows of granules. S. micro- procta is also easily distinguished from small specimens of S. nutrix by the very small size of its periproct and the relatively large area occupied by the apical disc. Order ARBACIOIDA Gregory, 1900 Family GONIOPYGIDAE Smith & Wright, 1993 Genus GONIOPYGUS Agassiz, 1838 Goniopygus arabicus sp. nov. Pl. 6, figs 3-10; Pl. 7, figs 1, 3,5, 6; Figs 17, 18A, B; DEG 1972 Goniopygus superbus Cotteau & Gauthier; Kier: 68, pl. 42, figs 1-3. 1989 Goniopygus superbus Cotteau & Gauthier; Ali: 401, in. 2 (2), Types. The holotype is BMNH EE4012, paratypes are BMNH EE3983-84, EE39896, EE3992, EE3997, EE4005, EE4007, EE4015, EE4017 and EE4019. MATERIAL STUDIED. Forty-four specimens of which the type series was used in the biometric analysis. OCCURRENCE. In the western Oman Mountains this species is found at the following levels: Jebel Buhays, section 1: loose in scree derived from the lowest few metres of the Simsima Formation (29): bed 12 (1). Jebel Buhays, section 2: loose in scree derived from the basal few metres of the Simsima Formation (4). Jebel Buhays, section 3: basal 1 m of the Simsima Formation (4). Jebel Thanais: lowest 1 m of the Simsima Formation (2). Jebel Faiyah, section 1: bed 6 (1 fragment). Jebel Rawdah, section 2: bed 4 (1): beds 6-8 (10); bed 10 (1); bed 11 (1). Jebel Rawdah, section 3a, bed 2 (2). Elsewhere the species has been reported from the late Campanian of the Rihyad district of Saudi Arabia (Kier 1972). DIAGNOSIS. A species of Goniopygus with relatively narrow ambulacra with a single small secondary tubercle on each compound plate, a trigonal periproct with, predominantly, three perianal tubercles and apical disc plating that is smooth and unornamented. Gonopores lie on the genital plates. DESCRIPTION. Tests range in size from 25 to 41 mm in diameter and are circular in outline. Test height is 53-62% of test diameter (mean = 57%, SD = 3-0%, N = 9: Fig. 17). Both the base and top are flat in profile and the ambitus lies at approximately mid-height (PI. 6, figs 4, 5, 7). The apical disc plates are elevated above the corona. The apical disc is flat, large and prominent. It occupies 35-43% of the test diameter (mean = 40%, SD = 3-3%, N= 9). Genital plates are pointed distally and the gonopore opens LATE CAMPANIAN-MAASTRICHTIAN ECHINOIDS ) PLATE 5 | Figs 1-3 Salenia microprocta sp. nov. BMNH EE3657, holotype; 1, apical; 2, oral; 3, lateral; all x 6. Jebel Huwayyah, section 2, bed 1. ‘Figs 4-10, 12 Mimiosalenia quinquetuberculata gen. et sp. nov. Jebel Faiyah, section 1, bed 7. 4, 12, BMNH EE3981, holotype; 4, apical, | 4; 12, lateral, ambulacrum detail, x 8. 5, 6, BMNH EE3982, paratype, apical; 5, x 3; 6, x 6. 7-9, BMNH EE3978, paratype; 7, lateral; 8, apical; 9, oral; all x 3. 10, BMNH EE3980, paratype; apical, x 3. |Fig. 11 Glyphopneustes hattaensis Ali. BMNH EE4027, detail of ambital region, lateral view, x 8; Jebel Thanais, lowest 2 m of the Simsima | . Formation. 144 A.B. SMITH PLATE 6 Figs 1,2 Goniopygus superbus Cotteau & Gauthier. L12680, Lambert Collection, Geology Department, Université de Paris VI, Paris;1, | apical; 2, lateral; both x 2. Senonian, Derré-i-Chahr, Iran. Figs 3-10 Goniopygus arabicus sp. nov. 3, 4, 10 BMNH EE4005, paratype; 3, apical; 4, lateral; both x 2; 10, ambital detail, x 5. Jebel Buhays, section 1; loose in the scree derived from the lowest 3 m of the Simsima Formation. 5, 8, BMNH EE4017, paratype; 5, lateral; 8, : apical; both x 2. Jebel Rawdah, section 2, bed 6. 6, 7, BMNH EE4012, holotype; 6, apical, x 1-6; 7, lateral, x 2. Jebel Rawdah, section 3, | bed 2. 9, BMNH EE3983; apical disc, variety with five perianal tubercles, x 5. Jebel Buhays, section 1; loose in the scree derived from the | lowest 3 m of the Simsima Formation. | LATE CAMPANIAN-MAASTRICHTIAN ECHINOIDS 24 22 E E 20 — ‘3 S 18 © fe + 16 © Ee 14 12 30 40 50 Test diameter (mm) 20 @ Interambulacrum oe gs Ambulacrum - e E ~ on 3 = 2s 10 = © ~ © cS ~ 2 = 0 Test diameter (mm) 20 @ Apical disc @ Periproct -_ £ E — - © z « fa) Test diameter (mm) Fig. 17 Biometric data for Goniopygus arabicus sp. nov. beyond the tip of the apical disc platform, though still within the genital plate (Figs 18A, B). Ocular plates are relatively large and are insert. All plates are flat and smooth, without Ormamentation. The periproct is oval and lies slightly poste- rior of centre. It is approximately 9-12% of the test diameter in width along the anterior-posterior axis. In the great Majority of specimens the opening is trigonal and there are three perianal tubercles on genital plates (Pl. 7, fig. 1). However, there is a single specimen ( BMNH EE3983) that | has five perianal tubercles (PI. 6, fig. 9) and one that has four, thus the number of tubercles is not invariant. | The ambulacra are 13-14% of the test diameter in width at the ambitus. All plates are compound and trigeminate, with a demiplate and two full elements (Fig. 18D). The two major elements carry a single primary tubercle and the upper also Number of primary tubercles in a column 145 Peristome diameter (mm) 30 40 50 Test diameter (mm) 30 @ Interambulacrum ®@ Ambulacrum N oO = oO 30 40 50 Test diameter (mm) N le) - © fo) Number of pore-pairs in a column a ° wn fo) 30 40 50 Test diameter (mm) has a perradial secondary tubercle in addition (Fig. 18E). The perradial tuberculate zone thus is composed of an outer series of primary tubercles alternating with a distinct inner series of secondary tubercles (Pl. 7, fig. 3). All tubercles are imperfo- rate and non-crenulate. Below the ambitus pore-pairs are small and oval and become crowded close to the peristome to form a relatively well-developed phyllode (PI. 7, fig. 5). At the ambitus pore-pairs become markedly more elongate and the individual pores more widely separated. Individual pores in a pore-pair are distinctly conjugate in larger individuals. There are no sphaeridial pits. There are 56 pore-pairs and 19 primary tubercles in a column at 25 mm test diameter, rising to 77 pore-pairs and 25 primary tubercles at 41 mm test diameter (Fig. 17). Interambulacra are broad and each plate carries a large 146 A.B. SMITH PLATE 7 Figs 1,3, 5,6 Goniopygus arabicus sp. nov. 1, 2, BMNH EE4012, holotype; apical disc, x 5. Jebel Rawdah, section 3, bed 2. 3, 6, BMNH EE4015, paratype; 3, detail of ambulacrum at ambitus, x 5; 6, detail of interambulacrum at ambitus, x 3. Jebel Thanais, lowest 1 m of the Simsima Formation. 5, BMNH EE3997, adoral detail, x 3-5. Jebel Buhays, section 1; loose in the scree derived from the lowest 3 m of the Simsima Formation. Figs 2,4 Goniopygus superbus Cotteau & Gauthier. L12680, Lambert Collection, Geology Department, Université de Paris VI, Paris; 2, | apical disc, x 4; 4, detail of ambulacrum at ambitus, x 6. Derré-i-Chahr, Iran. LATE CAMPANIAN-MAASTRICHTIAN ECHINOIDS 147 Fig. 18 Camera lucida drawings of Goniopygus species. A, B apical disc plating of Goniopygus arabicus sp. nov.; A, BMNH EE4017; B, BMNH EE4005S. C, F, apical disc plating and ambital ambulacral tuberculation of Goniopygus superbus Cotteau & Gauthier, L12680, Geology Department, Universite de Paris VI; Senonian, Derre-i-Chahr, Iran. D, E, ambital ambulacral plating, Goniopygus arabicus sp. nov.; D, BMNH EE4017; E, BMNH EE4005. G, G. arabicus sp. nov., adoral plating, ambulacrum to left, BMNH EE3986. Scale bars = 1 mm. primary tubercle. At the ambitus interambulacral width is 42-46% of the test diameter. There are 8 plates in a column at 25 mm diameter, rising to 11 at about 40 mm test diameter (Fig. 17). Primary tubercles are stout, imperforate and non- crenulate at the ambitus, but reduce sharply in size adapically so that the top three or four tubercles are very small. At the -ambitus they are surrounded by about 6 widely-spaced scro- bicular tubercles (Pl. 7, fig. 6). Down the interradius there is a single column of secondary tubercles on each plate. Adorally both columns of plates reach the peristomial margin _and there is no primordial plate (Fig. 18G). The peristome is slightly invaginated and is 37-44% of the test diameter in diameter (mean = 42%, SD = 2:6%, N = 7). Buccal notches are relatively small and poorly differentiated, with only a weak rim. REMARKS. This species was described under the name Goniopygus superbus Cotteau & Gauthier by Kier (1972) and Ali (1989). It differs from that species in several important respects. Firstly, the apical disc of G. superbus has a larger periproctal opening and has either five, or rarely four peria- nal tubercles (PI. 6, figs 1, 2; Pl. 7, fig. 2; Fig. 18C). These perianal tubercles face upwards and the adjacent portions of the genital plates are raised in the form of a very characteris- tic stellate rim. The radial portions of this rim project upwards as blunt denticles. Furthermore, the apical disc plates of G. superbus are covered in fine granular ornament. The gonopores in all specimens studied open not in the genital plates, but within the interambulacral plates some one or two plates distant from the apical disc (Fig. 18C). Finally, he ambulacral tuberculation affords an easily distinguishable character: in G. superbus the ambulacra are wide and the inner series of secondary tubercles almost as large as the primary tubercles, whereas in G. arabicus the ambulacra are narrow and the secondary tubercles very much smaller (com- pare Pl. 7, fig. 3 and Fig. 18E, with PI. 7, fig. 4 and Fig. 18F). Finally, in G. superbus there are zones of small miliary granules separating successive tubercles which are totally absent in G. arabicus. Genus MIMIOSALENIA gen. nov. TYPE SPECIES. Mimiosalenia quinquetuberculata sp. nov. DIAGNOsIS. A goniopygid with a perianal tubercle on each genital plate and pits along apical disc sutures. Ambulacra bigeminate except adorally where occasional simple plates are intercalated. OCCURRENCE. Known only from the late Cretaceous (Maas- trichtian) of Jebel Faiyah, section 1, western Oman Moun- tains. REMARKS. This genus is closely related to Goniopygus on account of its distinctive apical disc plating. The five gonop- ores lie immediately beyond the genital plates and open in the interambulacra. The genital plates of the apical disc have perianal tubercles identical to those of Goniopygus and the stellate ridge surrounding them is very reminiscent of that seen in G. superbus Cotteau & Gauthier. However, there are well-developed sutural pits both at triple junctions and mid- length along the plate sutures on all apical disc plates, which 148 ~_ E E cd i= 2 © is ~ n © = 6 8 10 12 14 Test diameter (mm) 2.5 C= E E = 72{0) © ~ © : = 1.5 ~ 0 ° i= a = 10 © a 0.5 3 4 5 6 7 8 Apical disc diameter (mm) 8 @ B Peristome ) ‘ @ Apical disc i. E E o - o = fs (S) 6 8 10 Uz 14 Test diameter (mm) Fig. 19 Biometric data for Mimiosalenia quinquetuberculata sp. nov. are often seen in saleniids but are never seen in Goniopygus. Another major difference between Goniopygus and Mimios- alenia is that the ambulacra of Goniopygus are trigeminate or occasionally quadrigeminate, whereas those of Mimiosalenia are strictly bigeminate, except close to the peristome where plating tends to become unigeminate. Again, bigeminate plating is typical of saleniids. All ambulacral plates in Mimi- osalenia reach the perradius and are approximately equal in size, whereas in Goniopygus there is a demiplate and the other two plates in each triad are unequal in size. Despite the similarities to Salenia, Mimiosalenia lacks a suranal plate and has the highly characteristic apical disc structure of a goniopygid. For this reason I believe it to be a derived goniopygid that has developed sutural pitting and 16 16 Width at ambitus (mm) Number in column A.B. SMITH ® Ambulacrum 6 @ Interambulacrum oe 6 8 10 12 14 16 Test diameter (mm) 40 B® Pore-pairs a i @ Primary ambulacral tubercles 2" : 10 Test diameter (mm) 12 14 16 bigeminate plating through the loss of the demiplate in each| triad. Mimiosalenia quiquetuberculata sp. nov P1. 5, figs 4-10, 12; Figs 19, 20)7 Types. Holotype, BMNH EE3981; paratypes, BMNH| EE3971, EE3974, EE3977-78, EE3980, EE3982, EES014-17. MATERIAL STUDIED. Biometric data is taken from the type series. In addition there are six other specimens. OCCURRENCE. The species is known only from bed 7, section 1b, Jebel Faiyah, western Oman Mountains. LATE CAMPANIAN-MAASTRICHTIAN ECHINOIDS DIAGNOsIS. As for generic diagnosis: apical disc with five perianal tubercles. Gonopores lie just outside the genital plates in the interambulacral areas. DESCRIPTION. Tests range from 7-2 to 15-5 mm in diameter and are circular in outline. Test height is 42-57% of test diameter with the flatter tests possibly representing slightly crushed specimens (Fig. 19). In profile the apical disc is very slightly conical and the sides uniformly rounded so that the ambitus lies at about mid-height. The apical disc is relatively large and raised above the coronal plates (PI. 5, fig. 7). It is 48-57% of the test diameter across (Fig. 19) with the periproct more or less centrally positioned. Ocular plates are relatively large and square- ended. The five genital plates border the periproct and are bluntly pointed distally (Figs 20A, B). All plates are smooth and unornamented. Each genital plate has a single perianal tubercle adjacent to the periproct. The genital plates are raised to form a stellate rim surrounding these perianal tubercles (PI. 5, figs 4, 6; Figs 20A, B). All five genital plates are approximately the same size. Gonopores open beyond the genital plates and are found immediately adjacent in the interambulacral zones (Fig. 20B). They are only present in the larger specimens, ca. 14 mm diameter. The periproct is subcircular in outline and measures 9-15% of the test diam- eter in width (Fig. 19). Ambulacra are relatively broad and slightly concave perra- dially. They are 11-17% of the test diameter in width at the ambitus. From the apex to below the ambitus plating is strictly bigeminate (Pl. 5, fig. 12; Fig. 20C), with each element reaching the perradius (Fig. 20D). Each pair of elements bears a large primary tubercle. Close to the peris- tome there are occasional simple elements interspersed, each with a large primary tubercle. There are 31-32 pore-pairs and ‘16-17 primary tubercles in a column at 11-5-13 mm test ‘diameter, rising to 37 pore-pairs and 19 primary tubercles at ‘15-5 mm test diameter (Fig. 19). The perradial zone of tuberculation is very broad and contains a mixture of second- ary tubercles and granules, two or three abreast (PI. 5, fig. 12; Fig. 20C). Adorally the pore-pairs are slightly more widely separated and there is no phyllode development whatsoever. Ambulacra hardly taper either adorally or adapically. There \are no sphaeridial pits. Interambulacra are 38-44% of the test diameter in width at the ambitus. There are seven plates in a column at 6-2 mm ‘est diameter, rising to eight or nine at 15 mm test diameter. Bach plate carries a large primary tubercle which is non- vrenulate and imperforate. The most adapical two are gener- illy very much smaller than the remainder. The primary ubercles have six scrobicular tubercles, three on each side, hat are more or less contiguous (PI. 5, fig. 12). The interra- lial zone is broad and slightly concave. It is occupied by a eries of miliary tubercles, two to each plate (four abreast). ‘here is no primordial plate adorally and both columns reach € peristomial border. The peristome is circular, slightly invaginated and occupies 640% of the test diameter. Buccal notches are small and idistinct (PI. 5, fig. 9). Spines, lantern and perignathic girdle all unknown. | |EMARKS. The biserial nature of the ambulacra and the laracteristic apical disc structure make this species easy to stinguish from any other described here. 149 Fig. 20 Camera lucida drawings of plating in Mimiosalenia quinquetuberculata sp. nov. A, apical disc, BMNH EE3981; B, apical disc, BMNH EE3982; C, ambulacrum, from apical disc (top) to peristomial margin (bottom), BMNH EE3981; D, ambital ambulacral plating, BMNH EES5014~-17. Scale bars = 1 mm. Family GLYPHOPNEUSTIDAE Smith & Wright, 1993 Genus GLYPHOPNEUSTES Pomel, 1883 Glyphopneustes hattaensis Ali, 1992 PI. 5, fig. 11; Pl. 8, figs 1-12; Figs 21, 22 1992a Glyphopneustes hattaensis Ali: 68, fig. 3. Types. Holotype, the figured specimen, housed in the Geo- logical Museum, University of Al Ain, United Arab Emir- ates. MATERIAL STUDIED. 82 specimens, of which biometric data was taken from the following 30: BMNH EE3909, EE3913, EE3915, EE3919-20, EE3923-24, EE3926, EE3930-31, EE3934, EE3936-38, EE3940, EE3942-43, EE3945-47, EE3949, EE3953-5S4, EE3958, EE3960, EE3967-70. OCCURRENCE. The type material all comes from Jebel Raw- dah. This species was collected at the following levels: Jebel Huwayyah, section 2: bed 1 (1). Jebel Faiyah, section 1: bed 2 (1); bed 5 at base (1); bed 8 (9); loose approximately 10 m above base of the Simsima Formation (1). 150 Jebel Buhays section 1; loose, derived from lowest few metres of the Simsima Formation (50). Jebel Thanais: lowest 2 m of the Simsima Formation (6). Jebel Rawdah, section 1; bed 4 (1). Jebel Rawdah, section 2; beds 9/10 (2); bed 11 (5); bed 13 (1); loose (6). Jebel Rawdah, section 3; bed 2 (2); bed 5 (1). Jebel Rawdah, section 4; bed 4 (1). DESCRIPTION. Tests range in diameter from 10 to 30 mm, with the great majority around 18-24 mm diameter (Fig. 21). The test is circular in outline and depressed in profile, with a rounded ambitus, although some specimens are slightly more conical. Test height is 39-57% of the diameter (mean = 50%, 20 —_ wn Test height (mm) ° 30 Test diameter (mm) 10 20 swt 8 - os Peristome diameter (mm) 10 20 30 Test diameter (mm) 14 @ Interambulacrum AY s Ambulacrum 12 ee € -* yy 10 £ «3 a ° ome o%, 3 E « ~ i= ~ 1 = Test diameter (mm) Fig. 21 Biometric data for Glyphopneustes hattaensis Ali. 40 A.B. SMITH SD = 4:0%, N = 30; Fig. 21). The ambitus lies at about | mid-height. The apical disc is highly sculpted and occupies 25-42% of the disc diameter (mean = 32%, SD = 1-2%, N = 30). It is proportionally larger in small individuals (Fig. 21). The periproct is large and central, occupying 30-50% of the apical disc diameter (mean = 40%, SD = 4-9%, N = 28). It is oval | in outline with smoothly rounded edges (PI. 8, figs 1, 2). The apical disc is dicyclic and all five genital plates are approxi- | mately equal-sized (Fig. 22B). There is a rim surrounding the periproct which bears a large central tubercle and two lateral tubercles. The gonopores open at the outer edge of the genital plates. The madreporite has a horse-shoe-shaped zone of madrepores that open around its margin (Fig. 22B). | 25 gs Ambulacrum (compound) E @ Interambulacrum = ° 1S) @ = n © ~ w a Sa i) he 4 E =] z 70 (2) lo} wn jo) + °o Number of pore-pairs in a column 30 10 20 30 40 Test diameter (mm) 10 w Apical disc @ Periproct r 8 a a = = € & os z E S f*,* a a © r -_ ae oe s Se a 9 Ad ONE Ie e e 2F eo y ° 10 20 30 40 Test diameter (mm) LATE CAMPANIAN-MAASTRICHTIAN ECHINOIDS LATE 8 jigs 1-12 Glyphopneustes hattaensis Ali. 1, BMNH EE3930; apical disc, x 6. Jebel Buhays, section 1; loose in the scree derived from the lowest 3 m of the Simsima Formation. 2, 8, BMNH EE3943; 2, apical disc, x 5; 8, apical, x 2. Jebel Buhays, section 1; loose in the scree | derived from the lowest 3 m of the Simsima Formation. 3, 6, 9, BMNH EE3915; 3, ambital detail of ambulacrum, x 6. 6, ambital detail of _interambulacrum, 4; 9, peristomial detail, x 6. Jebel Buhays, section 1; loose in the scree derived from the lowest 3 m of the Simsima Formation. 7, BMNH EE3958; apical, x 2. Jebel Faiyah, section 1, 2 m above the base of the Simsima Formation. 10, 11, BMNH EE4027; } 10, oral, x 2; 11, lateral, x 2. Jebel Thanais, lowest 2 m of the Simsima Formation. 4, 5, 12, BMNH EE3945; 4, apical; 5, oral; 12, lateral; all x 3. Jebel Buhays, section 1, loose in the scree. | 152 Ocular plates are heart-shaped. The sutures are all deeply incised and there are typically three small tubercles along the ocular/genital plate boundary within these depressed regions. Ambulacra are 13-17% of the test diameter in width at the ambitus (Fig. 21). All plates are trigeminate and there is a single demiplate in each triad (Figs 22A, C). The two full elements bear a single large primary tubercle that is imperfo- rate and non-crenulate. The upper element is smaller than the lower. Towards the peristome the demiplate has a shallow pit immediately perradial to the pore-pair, which marks the site of a sphaeridium (PI. 8, fig. 3; Fig. 22D). There are four or five of these in each column. Pore-pairs are uniserially arranged and not noticeably enlarged ambitally and adapi- cally. There is no pore crowding whatsoever close to the peristome. There are around 33 pore pairs and 11 primary tubercles in a column at 11 mm test diameter, rising to about 60 pore-pairs and 21 primary tubercles (Fig. 21). There is a single row of scattered miliary tubercles down the perradius. Interambulacra are 39-46% of the test diameter in width at the ambitus (mean = 44%, SD = 1-8%, N = 30). There are 9 plates in a column at 10 mm test diameter, rising to 12 at 30 mm test diameter (Fig. 21). Ambital plates are much wider than tall and each bears a single large primary tubercle that is non-crenulate and imperforate (Pl. 8, fig. 6). The most adapical two or three plates have significantly smaller tubercles than the rest. The primary tubercles have three scrobicular tubercles on either side, but have confluent areoles within each column. Down the interradius there are two or three irregular rows of scattered secondary and miliary tubercles, forming a relatively broad granular zone (PI. 8, fig. 6). Adorally both columns of plates reach the peristome and there is no primordial plate. The peristome is rather small and not at all invaginated. It is 33-43% of the test diameter across (mean = 38%, SD = 2:9%, N = 27), proportionally smaller in larger individuals (Fig. 21). Buccal notches are relatively shallow. REMARKS. Ali (1992a) gave a detailed description of this species but based on only six specimens. The large number of well-preserved specimens now to hand allows a detailed biometric description of this species for the first time. Ali specifically stated that sphaeridial pits were lacking in this species, yet in well-preserved specimens such pits can be seen. The difference between this species and the Cenoma- nian G. problematicus rest almost entirely on the apical disc ornamentation and the absence of sutural pits on the interam- bulacral plates of G. hattaensis. Although Fell & Pawson (1966) placed Glyphopneustes in the family Hyposaleniidae, and were followed by Smith & Wright (1990), it is now evident from the material available that Glyphopneustes is an arbacioid, and has been placed in its own family Glyphop- neustidae by Smith & Wright (1993). It has the characteristic perianal tubercles and apical disc structure of that family, and also has a similar style of ambulacral plate compounding. The sphaeridial pits are shallow and clearly convergent with those in Hyposalenia. A.B. SMITH Fig. 22 Camera lucida drawings of plating in Glyphopneustes hattaensis Ali. A, Ambulacral plating, from ocular plate (top) to peristome margin (bottom), BMNH EE3926; B, apical disc, BMNH EE3912; C, ambital ambulacral plating, BMNH EE3915; D, adoral ambulacral plating, peristomial border at bottom, BMNH EE3915. Scale bars = 1 mm. Family ARBACIIDAE Gray, 1835 Genus CODIOPSIS Agassiz, 1840 Codiopsis lehmannae sp. nov PI. 9, figs 1-2; Pl. 12, figs 1-3; Fig. 23 DERIVATION OF NAME. After Mrs C. Lehmann, the finder of the holotype. Types. Holotype, BMNH EE5033; paratypes (both incom- plete test fragments), BMNH EE3439, EE3440. OCCURRENCE. One specimen comes from Bed 15, section 1, Jebel Buhays, a second comes from bed 10 (top), section 2, Jebel Rawdah. The third specimen was found loose in the basal scree at Jebel Rawdah, section 2, and is almost certainly derived from the lowest few metres of the Simsima Formation (beds 3-10). DESCRIPTION. The holotype is 18-3 mm in diameter and 8-2 mm in height (45% of the diameter). The two other speci- mens are larger, but incomplete, and by estimation would have been around 35—40 mm in test diameter. The base 1s flat, the upper surface domal, and the ambitus is very sharp and at the base. The apical disc is preserved in BMNH EE3440 and EE5033. It is dicyclic and firmly fixed to the corona (PI. 12, fig. 1; Fig. 23C). The periproct is oval and 1-7 mm in diameter in the 18-3 mm diameter individual (9-3%). LATE CAMPANIAN-MAASTRICHTIAN ECHINOIDS Fig. 23 Camera lucida drawings of plating in Codiopsis lehamannae sp. nov. A, ambulacral plating (one column only) from supra-ambital region (top) to peristomial margin (bottom), BMNH E3439; B, aboral ambulacral plating, a little above the ambitus, BMNH EE34339; C, apical disc (incomplete) showing parts of two genital plates and two ocular plates, BMNH EE3440. Scale bars = 1 mm. Ambulacra are narrow and parallel-sided above the ambi- 'tus, where they are trigeminate. Plate compounding is arbaciid in style with two demiplates (Fig. 23B). There is a single calcitic pustule on each triad (PI. 9, figs 1, 2). Pores are small and widely-spaced, possibly conjugate (though preser- vation is too poor to be certain). Below the ambitus the | pore-pairs reduce in width markedly (Fig. 23A) and the -ambulacra widen into extensive phyllodes. The pore-pairs adorally are crowded and circular in outline with well- _ developed periporal muscle attachment areas. Compounding becomes polygeminate in a complex fashion (Fig. 23A) and there is a single large tubercle on each compound plate. Interambulacral plates are geniculate at the ambitus and ) adoral portions bear a single primary tubercle. This arrange- /ment creates a row of slightly downward-facing tubercles ) along the ambitus (PI. 9, figs 1, 2). Adradial portions of these /plates have fine secondary tuberculation. Adradial to the | primary tubercles and continuing adapically along the adra- )dial margin are very large calcite pustules. These reduce in ‘size considerably above the ambitus and appear to continue adapically at least most of the way to the apical disc. The |remainder of the adapical plates may have a pitted epistroma \(traces are seen in BMNH EE34339), but the test is not well preserved. ‘REMARKS. There is no doubt as to the generic placement of ‘these specimens, on account of their distinctive pustules and acid style ambulacral compounding. Their sharp ambitus, flat base, extensive phyllodes and ambital ring of interambu- lacral tubercles are distinctive and are features unknown in other species. It differs from Codiopsis brunei Lambert, from the Maastrichtian of Maastricht, in being very much larger 153 and in lacking well-developed aboral pustules. It comes closer to C. disculus Peron & Gauthier (and its synonyms C. stephensoni Cooke and C. fontei Vidal) from the late Campanian/Maastrichtian of Algeria, southern Spain, Sene- gal, Brazil and southern U.S.A., but differs from that species in having a more depressed profile, sharper ambitus and more distinct ambital ring of large tubercles. In C. lehmannae the primary interambulacral tubercles form a peripheral row, whereas in C. disculus the tubercles form a V-shaped arrangement extending adorally. Finally, in C. Jehmannae the peristome appears highly scalloped. Genus HATTOPSIS Ali, 1992 Hattopsis sphericus Ali, 1992 Pl. 9, figs 3-8; Pl. 10, figs 1, 2, 4; Fig. 24, 25B-D, 26B, 27A 1992b Hattopsis sphericus Ali: 694, fig. 3. Types. Holotype 910401 in the Museum of the Geology Department, University of Al Ain, United Arab Emirates. Paratype 910402. MATERIAL STUDIED. 73 specimens, of which the biometric data was taken from the following: BMNH EE3658, EE3663, EE3690, EE3692-95, EE3698, EE3702, EE3705, EE3707-20. OCCURRENCE. The species was described from Jebel Raw- dah (?section 1) by Ali (1992). Material collected in situ is as follows: Jebel Faiyah, section 1: bed 7 (1); bed 8 (pycnodont level) (8). Jebel Rawdah, section 1: bed 3 (22); bed 4, mostly towards top and base of bed 5 (44). Jebel Rawda, section 2: bed 11 (2); loose in scree at base of section (1). DIaGNosis. A spherical arbacioid with a reticulate epistro- mal ornamentation. Every third ambulacral pore-pair reduced in size. Two interambulacral tubercles on each plate. Peristomial rim elevated as a lip interradially. DESCRIPTION. Tests range from 12-0 to 20-8 mm in diameter (Fig. 24) and are circular to rounded pentagonal in outline. Test height is 60-83% of the diameter (mean = 74%, SD = 6:5%, N = 16), and tests are globular in profile with a small base and apex (PI. 10, fig. 2). The apical disc is dicyclic and occupies 23-33% of the test diameter (mean = 27%, SD = 2:4%, N = 14). The periproct is oval in outline and 9-12% of the test diameter in diameter. Ocular plates project slightly beyond the ring of genital plates (Pl. 10, fig. 2; Figs 25B, C). Genital plates are large and flat except around the periproctal margin where they are raised to form a rim. All genital plates are similar in size. The madreporite pores extend over most of genital plate 2. Well-preserved specimens show a reticulate pattern of ridges and pits (Pl. 10, fig. 2). Ambulacra are 20-24% of the test diameter in width at the ambitus (Fig. 24). They are compound throughout with trigeminate plating. Close to the peristome both upper and lower elements are demiplates, but elsewhere it is only the upper element that is a demiplate (Fig. 26B). This demiplate has a pore-pair that is very much smaller than those on the other two elements. Each triad has a single large primary A.B. SMITH PLATE 9 Figs 1,2 Codiopsis lehmannae sp. nov. BMNH EE34339, paratype; 1, oral; 2, lateral; both x 3. Jebel Buhays, section 1, bed 15. Figs 3-8 Hattopsis sphericus Ali. 3-5, BMNH EE3710; 3, apical; 4, oral; 5, lateral; all x 3. Jebel Rawdah, section 2, bed 11. 6-8, BMNH EE3712; 6, apical; 7, oral; 8, lateral; all x 3. Jebel Rawdah, section 2, bed 11. Figs 9-11 Hattopsis paucituberculatus sp. nov. BANH EE3683, holotype; 9, apical; 10, oral; 11, lateral; all x 3. Jebel Buhays, section 1; loose in the scree derived from the lowest 3 m of the Simsima Formation. LATE CAMPANIAN-MAASTRICHTIAN ECHINOIDS 155 PLATE 10 )Figs1,2,4 Hattopsis sphericus Ali. BANH EE3660; 1, adoral detail showing sphaeridial pits at perradius and peristomial lip, 6; 2, apical disc, X 6; 4, detail of ambital interambulacrum, x 6. Jebel Faiyah, section 1, bed 11. \Fig.5 Noetlingaster paucituberculatus (Noetling). BMNH EE3680, juvenile, apical view, x 4 (see also PI. 11, Figs 4, 5). Jebel Buhays, section 1; loose in the scree derived from the lowest 3 m of the Simsima Formation. ‘Figs 3,6 Hattopsis paucituberculatus sp. nov. 3, BMNH EE3672, paratype, adapical detail, x 7. Jebel Faiyah, section 1, bed 2. 6, BMNH _ EE3682, paratype; ambital detail, interambulacrum to left, x 6. Jebel Buhays, section 1; loose in the scree derived from the lowest 3 m of | the Simsima Formation. 156 tubercle. There is a small secondary tubercle on the perradial margin of the lower element but other miliaries and second- ary tubercles are absent. The perradius is ornamented with a reticulate pattern of ridges and shallow pits (PI. 10, fig. 4). Close to the peristome there are up to four sphaeridial pits arranged uniserially down the perradius (PI. 10, fig. 1). There are 33 pore-pairs and 10 primary tubercles in an ambulacral column at 12 mm test diameter, rising to 45-50 pore-pairs and 14-16 primary tubercles at 20-21 mm diameter (Fig. 24). No phyllodes nor any hint of pore-pair crushing is seen towards the peristome, and the pore-pairs themselves become much smaller adorally (Pl. 9, figs 4, 7; Fig. 26B). Interambulacra are 36-40% of the test diameter in width at the ambitus. Plates are wider than tall and carry two small primary tubercles placed towards the adradial margin (PI. 10, fig. 4; Fig. 27A). The tubercles in each pair are contiguous V8 rs H. sphericus @ H. paucituberculatus = a a + a @ B C4 _ a =» 4 ~~, wo % s Test height (mm) o oN 10 12 14 16 18 20 Test diameter (mm) 60r gH sphericus E @ H. paucituberculatus = 6 50 £ : 8.40 © i & -— ° 30 8 E =I =20 10 12 14 16 18 20 Test diameter (mm) 16 8 H. sphericus @ H. paucituberculatus Number of Interambulacral plates 14 16 18 Test diameter (mm) 20 Fig. 24 A.B. SMITH but are well separated from pairs of tubercles on other plates. There are no secondary or miliary tubercles, the remainder of the plate being covered in the same reticulate ornament of ridges and pits. At the peristome edge the interambulacra are thickened to form a distinct lip (Pl. 10, fig. 1). There is a single T-shaped primordial interambulacral plate forming the border to the peristome (Fig. 25D). There are 11 interambu- lacral plates in a column at 12 mm diameter, rising to 14 or 15 at 20-21 mm test diamater (Fig. 24). The peristome is not at all invaginated and is 38-48% of the test diameter across (mean = 43%, SD = 3-1%, N = 13). Buccal notches are very slight and the raised interambulacral rim forms the most prominent feature (PI. 10, fig. 1). REMARKS. Ali (1992b) gave a detailed description of this species but new features reported here for the first time | g H. sphericus @ H. paucituberculatus Apical disc diameter (mm) 16 18 Test diameter (mm) 12 14 20 22 9p eH. sphericus @ H. paucituberculatus 20 12 14 16 18 22 Test diameter (mm) 22 Biometric data for Hattopsis sphericus Ali and Hattopsis paucituberculatus sp. nov. LATE CAMPANIAN-MAASTRICHTIAN ECHINOIDS include the perradial uniserial row of sphaeridial pits, the large T-shaped primordial interambulacral plate and the extensive development of reticulate ornamentation. The pri- mordial plate, the umiserial and perradially positioned sphaeridial pits and the style of ambulacral compounding clearly place Hattopsis in the Arbaciidae. The reduction of one pore-pair in each triad and the presence of primary interambulacral tubercles above the ambitus separate Hattop- sis from Codiopsis, which it resembles. Hattopsis comes most close in structure and appearance to juveniles of Noetlin- gaster, but differs from them in having fewer interambulacral tubercles at comparable sizes, and in being more globular in profile. Hattopsis paucituberculatus sp. nov. Pl. 9, figs 9-11; PI. 10, figs 3, 6; Pl. 11, fig. 9; Figs 24, 25A, 26A, 27B Types. Holotype EE3683, paratypes, EE3682, EE3678, EE3688, EE3684-85. MATERIAL STUDIED. There are 22 specimens in addition to _ the types. Biometric data is based on the following speci- _ mens: BMNH EE3672, EE3675, EE3678-79, EE3682-85, _ _EE3688. © OCCURRENCE. This species was found at the following locali- ‘ ties and horizons: _ Jebel Faiyah, section 1: bed 2 (12). | Jebel Buhays section 1: in scree derived from lowest beds of the Simsima Formation (11); base of bed 12 (1). Jebel Buhays, section 2: loose in scree, derived from lowest 3 m of the Simsima Formation (1). | Jebel Buhays, section 3: basal beds of the Simsima Formation (2). | Jebel Aqabah: bed 1 (1). | Jebel Rawdah, section 2, bed 8 (1); loose in scree derived from lowest part of section (1). _ DIAGNOsIs. A species of Hattopsis with only a single inter- | ambulacral tubercle on each interambulacral plate at all sizes. DESCRIPTION. Tests are 6-2 to 18-2 mm in diameter and circular to rounded pentagonal in outline (Pl. 9, figs 9-11). | Test height is 58-83% of test diameter (mean = 71%, SD = -6-7%, N = 9) and in profile the test appears globular. The apical disc is dicyclic, like that of H. sphericus (Fig. 25A). It is 28-37% (mean = 31%) of test diameter in diameter and there is an elevated rim around the periproct margin (Pl. 10, fig. 3). Gonopores are generally rather larger and more oval than those of H. sphericus. . Ambulacra are 20-25% of the test diameter in width at the -ambitus and the ambulacral compounding is as in H. spheri- cus (Fig. 26A). There are 27 pore-pairs and 8 tubercles in a column at 10-7 mm test diameter, rising to 44 pore-pairs and 14 tubercles at 18:2 mm test diameter (Fig. 24). Aborally from the ambitus, the pore-pair on the upper demiplate in each triad is greatly reduced in size (Pl. 10, fig. 6; Fig. 26A). Adorally all pore-pairs become small and they remain unise- \rial to the peristome edge. | Interambulacra are 36-39% of the test diameter in width. Each plate carries a small primary tubercle, situated towards \the adradial margin. (PI. 10, fig. 6; Fig. 27B). The remainder ‘of the plate is covered in fine reticulate ridges and pits. There ‘are no secondary tubercles developed, even in the largest specimens. The peristome is 43-47% of the test diameter in diameter Fig. 25 Camera lucida drawings of plating in Hattopsis. A-C, apical discs; A, H. paucituberculatus sp. nov, BMNH EE3672; B, H. sphericus Ali, BMNH EE3692; C, H. sphericus Ali, BMNH EE3681. D, H. sphericus, BMNH EE3693, adoral interambulacral plating, peristomial edge at base. Scale bar = 1 mm. Fig. 26 Camera lucida drawings of ambulacral plating in Hattopsis. A, H. paucituberculatus sp. nov., complete ambulacrum from apical disc (top) to peristomial margin (bottom); BMNH EE3678; B, H. sphericus Ali, ambulacrum from close to apical disc (top) to peristomial margin (bottom); BMNH EE3659. Scale bars = 1 mm. and has the usual interambulacral lip and shallow buccal notches. REMARKS. This species resembles H. sphericus in all details except that it has only a single interambulacral tubercle on each plate, rather than the two found on all specimens of H. sphericus greater than 12 mm. Although I have included A.B. SMITH PLATE 11 Figs 1-3 Noetlingaster emiratescus Ali. BMNH EE3285; 1, oral; 2, apical; 3, lateral; all x 1. Jebel Rawdah, section 4, bed 12. Figs 4-7 Noetlingaster paucituberculatus (Noetling). 4, 5, BMNH EE3680 (juvenile); 4, lateral; 5, oral; both x 4. Jebel Buhays, section 1; loose in the scree derived from the lowest 3 m of the Simsima Formation. 6, BMNH EE3286 (juvenile), lateral view, x 4. Jebel Rawdah, section 2, bed 11. 7, BMNH EE3282, detail of ambital region, adoral towards top and interambulacrum to left, x 4. Fig.8 ?Noetlingaster sp. BMNH EE36839, lateral, x 5. Jebel Huwayyah, section 2, beds 2-7. Fig. 9 Hattopsis paucttuberculatus sp. nov. BMNH EE3682, lateral view, x 3. Jebel Buhays, section 1; loose in the scree derived from the lowest 3 m of the Simsima Formation. LATE CAMPANIAN-MAASTRICHTIAN ECHINOIDS some specimens smaller than 12 mm in this species, they could possibly turn out to be juveniles of H. sphericus. However, the majority of specimens are larger than 12 mm and the distribution of the two species is also not the same. Whereas H. sphericus is found predominantly at Jebel Raw- dah, section 1, beds 34 and also in beds higher up the succession (bed 11) in section 2, as well as at the pycnodont level (bed 8) at Jebel Faiyah, H. paucituberculatus consis- tently occurs further down in the succession (basal beds of Jebel Buhays and Jebel Faiyah, and bed 8 in Jebel Rawdah, section 2). The two species are thus stratigraphically discrete. Genus NOETLINGASTER Vredenburg, 1911 TYPE SPECIES. Protechinus paucituberculatus Noetling, 1897 by original designation. REMARKS. Noetlingaster has previously been classified in the Stomechinidae on account of its imperforate tuberculation and dicyclic apical disc (e.g. Fell & Pawson 1966). However, it has a single large primordial interambulacral plate at the adoral end of each interambulacrum and a tuberculation style very similar to that of Hattopsis. Primordial interambulacral plates this well-developed are known only in the Arbaciidae. Finally, juvenile forms of Noetlingaster are extremely similar in appearance to Hattopsis, differing only in having a more depressed test profile and more interambulacral tubercles at corresponding sizes. Given the marked similarity between Hattopsis and Noetlingaster in apical disc, tuberculation, interradial epistroma and pore-pair development, there seem strong grounds for placing Noetlingaster as an arbaciid. There are nine named species of Noetlingaster, all from the late Cretaceous. The type species N. paucituberculatus (Noet- Fig. 27 Camera lucida drawings of ambital plating in Hattopsis and Noetlingaster? sp.: interambulacral plate on the left, ambulacral plate on the right. A, Hattopsis sphericus Ali, BMNH EE3693; B, Hattopsis paucituberculatus sp. nov., BMNH EE3683; C, Noetlingaster? sp., BMNH EE3689. 159 ling, 1897) comes from horizon 4 (?Maastrichtian) in the Des Valley, Mari Hills, West Pakistan. Other species are: N. emiratescus Ali (1989), described from Jebel Rawdah, west- ern Oman mountains; N. sanfilippoi Checchia-Rispoli (1930), N. millosevichi Checchia-Rispoli (1930) and N. lamberti Checchia-Rispoli (1930), all from the Maastrichtian of Gebel Misid, Tripolitania, Lybia; N. globulus Devries (1967) and N. hemisphericus Devries (1967), both from the Maastrichtian of Kahta, Turkey; N. monteili Gauthier (1901) from the ‘Senon- ian’ south of Bilma, Algeria; and N. boulei Lambert (1906) from the Maastrichtian of Marohite, Madagascar. Devries (1967) reviewed previous species and discussed their diagno- sis. He laid particular emphasis on the arrangement of interambulacral tubercles, recognizing two groups: those with a single row of interambulacral tubercles on each plate, and those with a double row. He pointed out that the actual number of tubercles in each row varied ontogenetically and also according to position on the test. All these species are closely related because of their highly distinctive ambulacral plate compounding and pore-pair arrangement (see below). However, all have been erected on the basis of very few specimens, often simply the holotype. Thus the intraspecific variability has never been properly documented. In addition, the relatively thin test means that specimens are rarely well preserved. The large collection of specimens from Jebel Rawdah allows ontogenetic variability to be assessed in this species complex for the first time. There are significant differences in size between the species that have been erected, and these may account for some of the morphological variation described. N. boulei is described from the smallest specimen, only 49 mm in diameter. N. globulus, N. hemisphericus and N. emiratescus are all based on specimens ranging from 65-83 mm. The remainder are described from large individuals between 95 and 120 mm in diameter (Fig. 28). As many of the characters previously used for species discrimination can be shown to vary with size in the Omani populations, it is important that similar-sized individuals are compared when differentiating species. Shape differences were used by Devries (1967) to separate some species. He identified species as either ‘subconical’ or ‘rounded convex’. However, there seems to be considerable variation in this feature within the sample described here, and thus the character has little worth. It is also difficult to use in practice since specimens are often crushed or distorted during preservation. Ambulacral differences have also been used for diagnosing species. N. boulei for example has just one primary tubercle on ambulacral plates, and this is found on alternate plates only. However, as Lambert’s (1906) photographs show, ambulacral tuberculation is not consistent. Adoral compound plates all have a single primary tubercle, whereas, adapically, tuberculation is more irregular with occasional plates lacking primary tubercles. Considering the small size of N. boulei one would not expect more than a single tubercle to be devel- oped, by comparison with the Omani sample. Furthermore, tuberculation is very irregular in the Omani population, especially adapically from the ambitus where it is relatively common to find primary tubercles missing from ocassional plates. Such irregular ambulacral tuberculation characterises all species and from about 60 mm test diameter upwards all species have plates bearing two or three tubercles irregularly arranged. Note that although Gauthier (1901) describes N. monteili as having eight rows of ambulacral tubercles, his figures show only four irregular columns and the ambulacral 160 Number of primary tubercles in row 10 2 [a G oe —- 5 ee se 2 see e Wee 2 B ao @ A.B. SMITH e N paucituberculatus 0 N emiratescus OH =double row of tubercles present 20 25 30 Plate width (mm) Fig. 28 Biometric data from Noetlingaster species. Type specimens for species are plotted as follows: B = N. boulei Lambert; E = N. emiratescus Ali (two syntypes); G = N. globulus Devries; H = N. hemisphaericus Devries; L = N. lamberti Checchia-Rispoli; Mi = N. millosevichi Checchia-Rispoli; Mo = N. monteili Gauthier. structure does not differ from that seen in equivalent-sized Omani specimens. Most emphasis has been placed on interambulacral tuber- culation for differentiating between species. Devries (1967) recognized two species groups: those with just a single row of tubercles to each interambulacral plate and those with a double row. He also distinguished between those with a naked interradial zone of fine granulation and those with primary tubercles extending more or less to the interradius. Unfortunately, the Omani specimens also show considerable variation in these features. The majority simply have a single row of primary tubercles extending more or less to the perradius. From the ambitus adapically there may or may not be a naked zone free of primary tubercles and when present this zone may or may not extend below the ambitus. In many specimens it is difficult to observe tuberculation interradially because of weathering, and the same is probably true for the specimens on which other species are based. Single rows of tubercles are characteristic of N. paucituberculatus, N. mon- teili, N. boulei and N. globulus, whereas N. sanfilippoi, N. millosevichi, N. hemisphaericus and N. emiratescus all have two or three additional tubercles forming a second row adradially, and N. lamberti has a full double row of tubercles developed on all plates. All tuberculation styles, except that seen in N. lamberti, are encountered in the Omani sample. As can be seen from Fig. 28, there is a strong correlation between the number of tubercles on a plate and the plate width (which is proportional to test diameter). Furthermore, it is primarily the larger specimens that have a second row of tubercles developed adradially. However, it is not completely size-dependent since the 69 mm diameter specimen EE3279 has a well developed secondary row of tubercles whereas others of that size do not. In conclusion, the variation observed within the 20 reason- ably well-preserved specimens from the western Oman mountains is almost as great as that observed between the nine described species, each based on one or a few specimens only. The presence or absence of an interradial naked zone and the extent of this zone when present, the number of interambulacral tubercles in a row in proportion to the plate width and the development of a second row of primary tubercles to a plate are all variable. Similar variation of tuberculation has been found in both Turkey (Devries 1967) and Libya (Checchia-Rispoli 1930), but was used to distin- guish ‘species’. By contrast the analysis of variation amongst Omani speci- mens suggests that tuberculation pattern may not be so rigidly developed. It does, however, support some subdivision of the genus. When tuberculation style is plotted against test diam- eter, two distinct growth trajectories emerge for Omani specimens (Fig. 28). In some specimens there are numerous densely-spaced tubercles with secondary tubercle rows present, even at 60 mm test diameter. Other specimens have more widely-spaced primary tubercles and only develop a secondary row of adradial tubercles on ambital plates at a very large size. Although these may simply represent end members of a continuous spectrum, the small sample does not indicate any significant overlap. Those with many tubercles also always lack a granular interradial zone, whereas those with relatively fewer tubercles typically have a naked zone. However, the development of this naked zone is quite variable, with some specimens showing a broad zone extending almost to the peristome, and others with the zone restricted to a narrow wedge-shaped area adapically. Mea- surements taken from the types of all nine species are plotted for comparison. The type species N. paucituberculatus falls into the growth series of forms with fewer tubercles, as do the types of N. millosevichi, N. boulei, N. monteili, N. sanfilip- poi, and N. lamberti. The more densely tuberculate form corresponds to N. emiratescus and may also include N. globulus and N. hemisphaericus, although both plot in an intermediate position. (This may be due to inaccuracies in the magnifications for the camera lucida drawings of interambu- lacral plates given approximately by Devries (1967, pl. 2)). LATE CAMPANIAN-MAASTRICHTIAN ECHINOIDS Plotting those with a secondary row of tubercles to inter- ambulacral plates shows that the secondary row appears at about 15 mm plate width in N. globulus/N. emiratescus, whereas it only develops at around 22 mm plate width in the N. paucituberculatus group. The one anomolous specimen is BMNH EE3282, which has numerous small interambulacral tubercles, but a wide interradial granular zone. It falls in between the two growth series in Fig. 28. In conclusion, only two species of Noetlingaster are recog- nized here: (1) Forms to ca. 125 mm diameter with granular interradial zone developed adapically or throughout and a single row of relatively widely spaced interambulacral tubercles. Adradially, a second shorter row of tubercles is developed in larger specimens. These forms occur at Jebel Rawdah, section 2 from bed 11 to 14 and corre- spond to the form first described as N. paucituberculatus (Noetling). (2) Forms to ca. 125 mm with a second row of interambulac- ral tubercles present adradially from about 60 mm test diameter, and no naked zone. At comparable sizes there are more interambulacral tubercles in a row than in the first species. This form occurs at Jebel Rawdah section 1, but at higher levels in the section than N. paucituber- culatus. It has been described from the Oman Moun- tains under the name N. emiratescus Ali. _Noetlingaster paucituberculatus (Noetling, 1897) Pl. 10, fig. 5; Pl. 11, figs 4-7; Figs 28, 29, 30A—D, F, H, 31 11897 Protechinus paucituberculatus Noetling, 1897: 16, pl. 2, fig. 3, pl. 3, fig.1. 11898 Noetlingia paucituberculata Noetling; Lambert: 126. 1901 Noetlingia Monteili Gauthier: 191, pl. 3, figs 1-3. / 1906 Noetlingia Boulei Lambert: 11, pl. 2, fig. 7. (1911 Noetlingaster paucituberculata Noetling; Vredenburg: 46. Noetlingaster Millosevichi Checchia-Rispoli: 14, pl. 2, figs 1, 2, pl. 4, figs 3, 4. 1930 Noetlingaster Sanfilippoi Checchia-Rispoli: 6, pl. 1, | fig. 1, pl. 3, fig. 2, pl. 4, fig. 2. 1930 Noetlingaster Lamberti Checchia-Rispoli: 20, pl. 1, fig. 2, pl. 3, fig. 1, pl. 4, fig. 1. Types. The type is the 95 mm diameter specimen described by Noetling (1897) from the late Cretaceous of the Mari Hills, West Pakistan. MATERIAL STUDIED. 19 relatively complete tests and 2 test fragments, including juveniles. The following ten specimens were used in the biometric analysis: BMNH EE3267, 3269, 3271-75, 3280, 3282 and 3286. ‘OCCURRENCE. In the western Oman mountain area N. pau- ‘cituberculatus was found at the following levels: Jebel Rawdah section 1, base of bed 4 (juvenile). Jebel Rawdah, section 2: bed 11 (3, plus fragments); bed 14 (5, plus fragments); bed 19 (2); bed 25 (4); bed 26 (2). Jebel Buhays, section 1: loose in scree derived from lowest | beds of the Simsima Formation (3, including one complete 13 mm diameter juvenile). Outside the eastern Arabian peninsula this species is 161 recorded from western Pakistan, Algeria, Libya and Mada- gascar. DESCRIPTION. Tests range in diameter from 15 to 126 mm. They are more or less circular in outline, but very slightly depressed both interradially and adradially. Test height is 54-79% of test diameter (mean = 66%, SD = 7:2%, N = 10) and juveniles tend to have more depressed tests than adults. In profile the test is subconical, with a broad base, narrow apex and low ambitus (PI. 11, figs 4, 6). The ambitus lies at about one quarter of the test height above the base. The apical disc is relatively small, occupying only 13-16% of the test diameter in medium to large individuals (mean = 14%, SD = 1:2%, N = 6). It is proportionally larger in small individuals, occupying 19% of the test diameter in the 15 mm diameter specimen (Fig. 29). Disc plating is dicyclic (Figs 30A, B). The madreporite is very much larger than other genital plates and is densely covered in small pores. Each genital plate has a large oval gonopore which may be sur- rounded by a slight rim. The ocular plates are small and pentagonal, each with a small ocular pore. The periproctal opening is large and oval. Ambulacra are straight and taper adapically. At the ambi- tus their width is 16-20% of the test diameter. Ambulacral plates are trigeminate throughout with a highly distinctive style of. compounding (Figs 30C, D, F, H). The lowest element is large and occupies the full width. There is a smaller demi-plate above, which always carries a large pore- pair adradially and an even smaller, fully occluded element above that (Figs 30D, F). This arrangement is found along the entire length of the ambulacrum in all medium to large individuals, except at the very apex, where the occluded plate may reach the adradial suture. In smaller individuals both of the smaller elements are demiplates (Fig. 30H), while in BMNH EE3286, a juvenile of only 15 mm test diameter (Fig. 30F), the upper element extends to the perradial suture. There are no phyllodes nor any pore crushing towards the peristome. Each element carries a pore-pair, but only on the large primary element are these well-developed throughout (Pl. 11, fig. 7). The pore-pairs on the small occluded upper element are always reduced to rudimentary structures and in places may simply be represented by a single pore. The middle element usually has well-developed pore-pairs, but in larger individuals around the ambitus these may also be very much reduced in size (Fig. 30F). Pore-pair differentiation is hardly developed in the 15 mm diameter juvenile. Each compound plate usually carries a single primary tubercle close to the pore-zone. However, tuberculation is irregular and occasionally two tubercles occur to a plate. In the largest individuals most ambulacral plates carry two adradial primary tubercles, slightly offset on alternate plates. The perradial band is devoid of large tubercles above the ambitus, but this zone is relatively narrow, typically only some 15% of the interambulacral width. There are approximately 112 pore- pairs in a column at a test diameter of 32 mm, rising to around 180 at 80 mm test diameter (Fig. 29). Interambulacral width at the ambitus is 41-47% of the test diameter. Plates are wide and low, and are slightly taller than ambulacral plates. There are 36 interambulacral plates in a column at 82 mm test diameter. The primary tubercle at the centre of the plate is set on a slight keel which runs down the midline of each column and is particularly prominent adapi- cally. There are multiple small tubercles on each plate, three in the smallest individual (PI. 11, fig. 4), rising to 11 in the A.B. SMITH 162 PLATE 12 | Figs 1-3 Codiopsis lehmannae sp. nov. BMNH EES033, holotype; 1, apical; 2, oral; 3, lateral; all x 4. Jebel Buhays, section 1; loose in the scree derived from the lowest 3 m of the Simsima Formation. Figs 4-7 Actinophyma spectabile Cotteau & Gauthier. BMNH EE3601; 4, apical, x 3; 5, oral, x 3; 6, lateral, x 3; 7, detail of ambital plating, x 6. Jebel Buhays, section 1; loose in the scree derived from the lowest 3 m of the Simsima Formation. LATE CAMPANIAN-MAASTRICHTIAN ECHINOIDS + a [es] fo) jo) (>) Test height (mm) N jo) 0) 20 40 60 80 100 120 140 Test diameter (mm) 140 Number of pores in column a 30 40 50 60 70 80 90 Test diameter (mm) 14, © Number of primary tubercles in row @ lamb. plate height (mm) 0 10 20 30 Interambulacral plate width at ambitus (mm) @ Ambulacrum A @ Interambulacrum BSS fo} Width at ambitus (mm) Ww io) 20 10 (0) 10) 20 40 60 80 100 120 140 Test diameter (mm) 30 @ Peristome ® Apical disc = E it ® -~ o E & 6 0 20 40 60 80 100 120 Test diameter (mm) Fig. 29 Biometric data for Noetlingaster paucituberculatus (Noetling) and N. emiratescus Ali. _ largest. These are arranged in a single, slightly arcuate row (Pl. 11, fig. 7), but the larger individuals may have two or three additional tubercles forming an upper row near the adradial suture. All tubercles are imperforate and non- crenulate and have relatively small mamelons and extensive areoles. Sutures between all plates are finely denticulate. The peristome is very slightly sunken and is strongly _ indented by wide buccal notches. Peristome diameter is 20-31% of test diameter (mean = 27%, SD = 4:0%, N = 7). REMARKS. This species is distinguished from N. emiratescus _ by its interradial granular zone, which may be small and | deveioped only adapically, or may be broad and extend _ adorally. Within a single population from bed 14, section 2, | Jebel Rawdah, the development of this granular zone was highly variable, with some having only a narrow adapical wedge of granules, others having a broad band extending adorally. It also differs from N. emiratescus in having fewer interambulacral tubercles at comparable sizes (Fig. 28). Smaller individuals (diameters less than ca. 100 mm) have single rows of tubercles on interambulacral plates, but a second row is present in larger specimens. The species was first described by Noetling (1897) on the basis of a single specimen from West Pakistan. Gauthier (1901) later described a smaller, incomplete specimen of this species under the name N. monteili from the eastern Sahara region of Algeria. Other species that are probably best treated as synonymous are the three species described by Checchia—Rispoli (1930) from the Maastrichtian of Libya. 164 Fig. 30 Camera lucida drawings of Noetlingaster. A-D, F, H, N. paucituberculatus (Noetling): A, B, apical disc plating: A, BMNH EE3269; B, BMNH EE3275; C, adapical plating in a juvenile, BMNH EE5049; D, Ambital plating, BMNH EE3266; F, ambital plating, BMNH EE3286; H, adapical plating, BMNH EE3283. E, G, N. emiratescus Ali, BMNH EE3285; E, adapical plating; G, ambital plating. Scale bars = 1 mm. Fig. 31 Camera lucida drawings of plating in a juvenile Noetlingaster paucituberculatus (Noetling), BMNH EE3680. A, Apical disc; B, ambital plating, ambulacrum to the left, interambulcaral plate on right; C, adoral interambulacral plating, showing the single primordial interambulacral plate. Scale bar = 1 mm. There is a general increase in size up the section at Jebel Rawdah, with small to medium-sized individuals found in the A.B. SMITH lower beds and only large individuals towards the top in the deeper water facies. Noetlingaster emiratescus Ali, 1989 P\. 11, figs 1-3; Figs 28, 29; 30E, G 1989 Noetlingaster emiratescus Ali: 398, Fig. 2 (3-5) Types. The types of N. emiratescus are three specimens and six fragments in the collections of the Geology Department, United Arab Emirates University, Al Ain. MATERIAL STUDIED. Three specimens, BMNH EE3279, EE3284-85. OCCURRENCE. All specimens, including the type material, come from Jebel Rawdah. The specimens reported here were collected from bed 4, Jebel Rawdah section 1 (1) and bed 13, Jebel Rawdah section 4 (2). DIAGNOsIS. A Noetlingaster with smaller and denser inter- ambulacral tubercles than other species, and completely lacking a naked interradial zone. DESCRIPTION. Tests range from 68-5 to about 105 mm in diameter. In outline the interradii and adradii are slightly depressed. Test height is 58-71% of test diameter and in profile the test is inflated and subglobular. The ambitus lies about one third of test height above the base. The apical disc occupies 11% of the test diameter and has the same arrangement of plates as N. paucituberculatus. Ambulacral structure is also more or less identical, with the pore-pair on the occluded plate rudimentary almost to the apex (Figs 30E, G). The middle pore-pair is also rudimentary from the ambitus adorally. Ambulacral tubercles are irregu- larly arranged and occupy the full width of the ambulacra, one or two to a compound plate. Interambulacral plates are slightly V-shaped, with a single well-developed row of tubercles along the lower edge and a second, irregular row of occasional tubercles above. There are 19 tubercles in a row at a test diameter of 71 mm, rising to 13 at 105 mm test diameter (Fig. 28). Tubercles occupy the full width of plates throughout and there is no granular interradial zone as is found in N. paucituberculatus. The peristome is small and somewhat invaginated, occupy- ing 26% of the test diameter. REMARKS. Ali (1989) described this species on the basis of the material collected from Jebel Rawdah. Its small, numer- ous interambulacral tubercles and lack of any interradial zone of granulation serve to separate it from the co-occurring species, N. paucituberculatus. Noetlingaster? sp. (juvenile?) Pl. 11, fig. 8; Fig. 27C MATERIAL. One specimen, BMNH EE3689. OCCURRENCE. From the Loftusia beds (beds 2-5), Jebel Huwayyah, section 2. DIAGNOsIS. A species of arbaciid with a single primary interambulacral tubercle on ambital and adoral interambulac- ral plates, but smooth adapically. Ambulacral tubercles developed to apex. Broad ambital and adapical interradial zone ornamented with vertical riblets. DESCRIPTION. The test is rounded pentagonal in outline and 13-2 mm in diameter. It has a broad flat base and domed ————— Eee TTS EE — ee a LATE CAMPANIAN-MAASTRICHTIAN ECHINOIDS upper surface. Test height is 57% of test diameter and the ambitus lies at around 40% of test height. The apical disc is 3-8 mm in diameter (29% of test diameter) and is dicyclic. It is not well preserved, but the genital plates are raised as a lip around the periproct margin and the periproct itself is 11% of the test diameter and oval in outline. Ocular plates protrude beyond the genital plates slightly. Ambulacra are 20% of the test diameter in width at the ambitus. All plates are compound and trigeminate, but details of plate compounding are not clear. There is clearly a single element and a double element in each triad, but it is impossible to tell whether one of the double elements is a demiplate, as I suspect. All pore-pairs are equally well- developed. There is no phyllode development. There are 33 30 E E 20 — ~ <£ 2 © £ * 10 © Ee (0) 30 40 50 Test diameter (mm) 20 8 Ambulacrum ’ @ Interambulacrum ~~ E E — n 3 = £10 @ ~ a <= - 3 = (0) 30 40 Test diameter (mm) 50 Number of Interamb. plates in column 30 40 Test diameter (mm) 50 Fig. 32 Biometric data for Phymosoma hexoaporum Lambertt. 165 pore-pairs and 10 primary tubercles in a column. Interambulacra are 39% of the test diameter in width at the ambitus. Each plate has a large adradial tubercle that extends adapically to within three or so plates of the apex. Adorally these tubercles lie subcentrally, but towards the apex they become positioned more and more closely towards the adra- dial suture. On ambital and adapical plates there is also a very much smaller mamelonate secondary tubercle lying at the centre of each plate within the zone of riblet ornamentation. The entire interradial zone from the ambitus upwards, has a well-developed ornamentation of vertically orientated riblets. The peristome is 5-2 mm in diameter (40% of the test diameter) and is not invaginated. There are no raised interra- dial lips to the peristome margin. Buccal notches are extremely feeble. -_ + o ® Total ¢@ Number arranged biserially Number of pore-pairs in column 30 40 Test diameter (mm) 60 50 60 ® Peristome @ Apical disc 18 a = (>) Diameter (mm) = N = fo) 30 50 Test diameter (mm) 60 40 60 60 166 REMARKS. This species is easily distinguished from Hattopsis sphericus on the basis of its interambulacral ornamentation of vertical riblets, and by its single row of primary interambulac- ral tubercles. It also lacks the reduced diameter pore-pair in each triad that is so characteristic of H. sphericus and H. paucituberculatus. It differs from H. paucituberculatus in its lower profile, less enlarged adapical pore-pairs and in its riblet ornamentation as opposed to the pitted ornamentation seen in H. paucituberculatus and H. sphericus. The vertical riblet ornamentation of Noetlingaster? sp. is characteristic of Codiopsis species such as C. doma, but Noetlingaster? sp. differs from Codiopsis in having no sharp reduction in the size of tubercles at the ambitus. It could possibly be the juvenile of a very much larger Codiopsis species, such as C. stephen- soni Cooke, but it has open gonopores, which suggests that it is a genuinely small species. It differs from species such as Codiopsis bruni Lambert & Thiéry (Maastrichtian of the Netherlands) and C. disculus Peron & Gauthier (late Campa- nian of Algeria, early Maastrichtian of Spain) in having a smooth, pustule-free upper surface. This specimen most likely represents a small Noetlingaster, one in which only the primary interambulacral tubercles have formed. However, neither of the named species of Noetlingaster shows the distinct vertical ribbing that characterises this specimen. Order PHYMOSOMATOIDA Mortensen, 1904 Family PHYMOSOMATIDAE Pomel, 1883 Genus PHYMOSOMA Haime, in d’Archiac & Haime, 1853 Pl. 13, figs 1-3; Figs 32, 33 1908 Phymosoma (Cyphosoma) Archiaci Cott. var: Cot- fieeaNs il, fol, 3), ws, Il 1927 Phymosoma hexoaporum Lambert: 35, pl. 2, figs 25-27. 1933 Phymosoma Paronai Checchia-Rispohi: 11, pl. 2, figs 4-7. Phymosoma hexoaporum Lambert, 1927 MATERIAL. Nineteen specimens of which the biometric data was drawn from the following: BMNH EE3607-08, EE3610-12, EE3614, EE3616, EE3618, EE3941. OCCURRENCE. This species is found almost exclusively in the lowest beds of the Simsima Formation at Jebel Buhays, section 1 (16), and the immediately adjacent Jebel Thanais (3). In addition one specimen was found loose in the scree in Jebel Rawdah, section 3, and another in Bed 2 at Jebel Rawdah, section 4. Cottreau (1908) recorded an identical specimen from the late Cretaceous (?Maastrichtian) of Maro- hita, Eastern Madagascar. DIAGNOsIs. A Phymosoma with a single large primary inter- ambulacral tubercle on all plates, and one or two small adradial tubercles. Plates are composed of six or seven elements at the ambitus and pore-pairs are biserial from about the ambitus adapically. Interradial zones of small granules are very well developed. DESCRIPTION. Tests range from 23 to 50 mm in diameter and are circular in outline. The test is depressed in profile (Pl. 13, fig. 3), with a height that is 34-46% of the test diameter (mean = 42%, SD = 4-4%, N = 8: Fig. 32). The ambitus lies slightly below mid-height. A.B. SMITH The apical disc is caducous and is missing from all speci- mens. The outline of the apical disc is pentagonal with angles pointing interradially and projecting slightly further into the posterior interambulacrum (PI. 13, fig. 2). Ambulacra taper slightly both adapically and adorally. They measure 22-26% of the test diameter in width at the ambitus. All plates are polygeminate (Fig. 33). Above the ambitus pore-pairs are biserial and plates are composed of six to eight elements. Plate compounding is in the phymosomatid style. At the ambitus the pore-pairs are in arcs of six or seven in specimens more than 25 mm diameter, while subambitally they are quinquegeminate. Immediately adjacent to the peristome there are a couple of quadrigeminate plates. There are no sphaeridial pits adorally. Each compound plate carries a single large imperforate, crenulate tubercle, as large as the adjacent interambulacral primary tubercles. This occupies most of the plate. However, perradially there is a narrow band of small secondary and miliary tubercles (Pl. 13, fig. 3). Primary tubercles more or less reach the apex. There are 57-58 pore-pairs in a column at 23-24 mm test diameter, rising to 138 at 50°mm test diameter. Biserial pores appear immediately above the ambitus in most specimens and com- prise 50-54% of the total number of pore-pairs in a column. In the very largest specimen, biserial pore-pairs extend to the subambital region. Interambulacra are 32-38% of the test diameter in width at the ambitus. There are 10 plates in a column at 23 mm test diameter, rising to 18 at 50 mm test diameter (Fig. 32). Each | plate carries a single large primary tubercle, centrally posi- tioned. However, areoles are not contiguous, but are sepa- | rated by a narrow band of miliary granules (PI. 13, fig. 3). | Tubercles are imperforate and crenulate and decrease in size gradually both adapically and adorally. There are one or two small secondary tubercles to each plate situated close to the | _ adradial suture. In a few specimens these tubercles enlarge | above the ambitus to about half the size of the primary | tubercles, so as to form a secondary row. However, this is | inconsistently developed. The interradius is broad and cov- ered in scattered small secondary and miliary tubercles (Pl. | 13, fig. 3). In some specimens (e.g. BMNH EE3614) this | tuberculate band is relatively dense, whereas in other speci- | mens (e.g. BMNH EE3618) tuberculation is more scattered. | ~ Both columns of plates reach the peristomial border. ii The peristome is very slightly invaginated and occupies | 40-47% of the test diameter. Buccal notches are small, but | clearly incised (PI. 13, fig. 1). Lantern elements are seen scattered inside the test in| — BMNH EE3941. Hemipyramids are largely concealed by sediment, but one keeled tooth is seen in cross-section. | REMARKS. The variation encountered in the secondary} tuberculation of this species is more marked than expected | and is matched by a difference in the degree of tubercle] — crenulation. In the specimens with the denser miliary tuber-|— culation interradially, there are usually well-developed sec- ondary tubercles along the adradial margin in the region) immediately above the ambitus, and primary tubercles are} only weakly crenulate. In the more usual variety, the interra-} dial zone has more scattered tuberculation, there is no enlarged secondary tuberculation and the primary tubercles have a well-developed and broad crenulate platform. These two forms may eventually prove to represent distinct species, but as they are both found at the same locality and the secondary tuberculation differences are not entirely consis- LATE CAMPANIAN-MAASTRICHTIAN ECHINOIDS 167 PLATE 13 Figs 1-3 Phymosoma hexoaporum Lambert. BMNH EE3614; 1, oral; 2, apical; 3, lateral; all x 3. Jebel Buhays, section 1; loose in the scree derived from the lowest 3 m of the Simsima Formation. Figs 4,5 Actinophyma spectabile Cotteau & Gauthier. B18731, Morgan Collection, Museum d’Histoire Naturelle, Paris; 4, lateral; 5, apical; both x 1-5. Senonian of Iran, no locality details. | | 168 Fig. 33 Camera lucida drawing of ambulacral plating, from apex (top) to peristomial margin (bottom), in Phymosoma hexoaporum Lambert, BMNH EE3617. Scale bar = 1 mm. Fig. 34 Camera lucida drawings of plating in Actinophyma spectabile Cotteau & Gauthier. A, adapical ambulacral plate from a large individual, BMNH EE3599; B, supra-ambital ambulacral plating in a juvenile, BMNH EE3601; C, adoral ambulacral plating in a juvenile, BMNH EE3601. Scale bar = 1mm. tent, the two forms are simply treated as varieties of the same species here. The species described here appears identical to the speci- men described from the Maastrichtian of Madagascar by A.B. SMITH Cottreau (1908) under the name Phymosoma archiaci Cot- teau var. It differs from true P. archiaci in having a broader, more granular interradial zone, and in having less well- | developed secondary tubercles on interambulacral plates. | The species P. hexoaporum was established by Lambert i (1927) for specimens from the Maastrichtian of Sopeira, — province of Aragon, Spain. P. hexoaporum differed from all | other species of Phymosoma described previously in having | compound plates composed of six elements at the ambitus | rather than five. Later, Checchia-Rispoli (1933) described the same species from the Maastrichtian of Libya under the name | P. paronai Checchia-Rispoli. This too has six or seven l) elements in ambital ambulacral compound plates. Genus ACTINOPHYMA Cotteau & Gauthier, 1895 Actinophyma spectabile Cotteau & Gauthier, 1895 PI. 12, figs 4-7; Pl. 13, figs 4, 5; Fig. 34) 1895 Actinophyma spectabile Cotteau & Gauthier: 98, pl. 25, figs 6-10. 1895 Cyphosoma persicum Cotteau & Gauthier: 93, pl. 25, | figs 3-4. 1902 Actinophyma spectabile Cotteau & Gauthier; Gauthier: 151, pl. 20, figs 7-10. 1935 Actinophyma_ spectabile Cotteau & Gauthier; ~ Mortensen: 489, fig. 287. | Types. The holotype is the specimen described by Cotteau & Gauthier from the upper Senonian of Endjir-kouh, Aftab district, Iran. i MATERIAL STUDIED. Six specimens, BMNH EE3598-3603,| — three of which are test fragments only. The following descrip-| - tion is based on the two more or less complete specimens} © BMNH EE3601 (a juvenile) and BMNH EE3603 (an adult).| ~ Topotype material of A. spectabile in the Natural History| © Museum and in the Morgan Collection, Museum d’Histoire Naturelle, Paris, has been studied for comparison. | OCCURRENCE. Five of the specimens come from scree collec- tions at Jebel Buhays, section 1 and are derived from the lowest few metres of the Simsima Formation. Two other specimens come from Jebel Rawdah section 3b, one from bed 8, the other from bed 9. The species range is ‘Upper Senonian’ of southern Iran and) ~ Maastrichtian of the Oman Mountains. DIAGNOsIS. A species of Actinophyma with strong radiating) © ridges on the boss of primary tubercles. Ambulacra polyploid adapically with up to 18 pore-pairs to an ambulacral plate arranged irregularly. DESCRIPTION. Tests range in diameter from 21-2 mm to approximately 90 mm and are circular in outline. Test height is 33 mm in the larger specimen and 9-4 mm in the smalle (ca. 35-45% test diameter). In profile the ambitus is more 0 less at mid-height and is smoothly rounded below and slightly more conical above. The apical disc is small; 5-8 mm diameter in the 2] m individual (21% of test diameter) and probably only about 15 mm in the larger individual (16% of test diameter). Apical) _ disc plates are lost from all specimens and were evidently not securely sutured to the corona. The opening is pentagonal in outline (Pl. 12, fig. 4; Pl. 13, fig. 5). LATE CAMPANIAN-MAASTRICHTIAN ECHINOIDS ' Ambulacra are almost as broad as the interambulacra, ' being 25-3 mm wide at the ambitus in BMNH EE3603, as _ compared to the interambulacral width of 28-7 mm. Pore- | pairs are in arcs of six on the oral surface, becoming biserial - subambitally in the larger specimen and continuing so to the apex (Fig. 34A). Adapical plates have up to 18 pore-pairs, and show phymosomatid-style compounding (Fig. 34A). In the small individual adapical pore-pairs are only just begining to become biserial and all plates have just six or seven _ elements (PI. 12, fig. 7; Fig. 34B). There are 10 or so plates in » acolumn at 21 mm diameter and 18 or 19 at 90 mm diameter. ‘Each plate carries a single large primary tubercle that occu- _ pies most of the plate. However, in the larger individual there is a narrow adradial band of heterogeneous small secondary and miliary tuberculation. Ambulacral tubercles are not ' contiguous, but are separated by a band of small secondary tubercles. Areoles are strongly sculptured by radial grooves ' that extend from the base towards the platform of the boss. ; The adoral plating in the small individual (BMNH EE3601) is noteworthy, since it comprises simple plating arranged in triads (Fig. 34C). However, standard phymosomatoid-style + compounding develops subambitally, and this appears to be simply a juvenile feature. There are 11 interambulacral plates in a column at 21 mm | test diameter, rising to about 18 at 90 mm test diameter. Each ) plate carries a large primary tubercle with a small imperforate ‘mamelon and a crenulate platform. Areoles at the ambitus ‘and adorally are confluent and are oval in outline. The areole ) bears strong radial grooves which extend up the sides of the ‘boss (PI. 12, figs 5, 6). These are most pronounced in the smaller individual. Tubercles decrease in size above the ‘ambitus. The primary tubercles are centrally positioned on \the plate and there are relatively broad adradial and interra- \dial bands of secondary tuberculation. On plates around the jambitus there is an enlarged secondary tubercle to the adradial side of the primary tubercle, but otherwise second- ary tuberculation is small and heterogeneous. The interradial ‘zone is depressed and tubercle-free towards the apex. The peristome is relatively small and invaginated. In the 21 ‘mm individual it is 7-1 mm in diameter (33% of test diam- eter), and is proportionally smaller in the larger individual. Buccal notches are very shallow (Pl. 12, fig. 5). ‘REMARKS. When first erected (Cotteau & Gauthier, 1895), ‘this species was based on a small individual 29 mm in diameter, which is virtually identical to BMNH EE3601 in morphology. In the same publication Cotteau & Gauthier (1895: 91) erected another species, Cyphosoma persicum on the basis of a larger, but fragmentary specimen from Derre-i- Chahr. Subsequently, with the collection of more material, Gauthier (1902) recognized that A. spectabile and C. persi- ‘\cum were simply different growth stages of the same species and synonymized the two, selecting A. spectabile as the valid ame. The characteristic radial striation, which is so strongly evident in juveniles, makes this an easily recognizable spe- cies. Only one other comparable species has been described, Actinophyma cf. A. spectabile Kier (1972: 68), from the ‘Campanian of Saudi Arabia. This differs from the Iranian and Omani species in having deep pits developed at the 2orners of interambulacral plates on interradial sutures. 169 Test height (mm) a N [o-) wn 10 20 30 Test diameter (mm) ® Ambulacrum @ Interambulacrum e Width at ambitus (mm) Test diameter (mm) @ Peristome s 10 ®@ Apical disc E = © ~ © E ol (=) 20 30 Test diameter (mm) Fig. 35 Biometric data for Plistophyma asiaticum Cotteau & Gauthier. Genus PLISTOPHYMA Peron & Gauthier, in Cotteau, Peron & Gauthier, 1881. Plistophyma asiaticum Cotteau & Gauthier, 1895PI. 14, figs 1-7; Figs 35, 36 1895 Plistophyma asiaticum Cotteau & Gauthier: 105, pl. 16, figs 11-14. Types. The holotype is the single specimen described and figured by Cotteau & Gauthier. It is not in the Morgan collection in the Museum d’Histoire Naturelle, Paris. MATERIAL STUDIED. Six specimens, BMNH EE3572-76, EE4932. Only BMNH EE3573, which is incomplete, was 170 omitted from the biometric analysis given below. OCCURRENCE. In the western Oman Mountains this species was found at the following levels and localities: Jebel Buhays, section 1: loose in scree, derived from lowest few metres of the Simsima Formation (4). Jebel Thanais: From the lowest metre of Simsima Formation (2). Jebel Rawdah, section 2, bed 4 (1). The species was described from the Senonian of Derre-i- Chahr, southern Iran and there are no other records. DIAGNOsIS. A Plistophyma in which the ambital interambu- lacral plates are very much smaller and narrower than either adoral or adapical plates. DESCRIPTION. Tests range in diameter from 12 to 25-5 mm and are rounded pentagonal in outline with the angles interradial. Tests are depressed and rounded in profile, with the ambitus at or very slightly below mid-height (PI. 14, figs 3, 4). Test height is 33-38% of test diameter (mean = 36%). The apical disc is large and pentagonal in outline with the angles interradial (Pl. 14, fig. 2). It is 47-50% of the test diameter in diameter and is not invaginated. No specimen retains any apical disc plates but, to judge from the size and shape of the opening, disc plating was presumably monocy- clic. Ambulacra are 13-16% of the test diameter in width at the ambitus. They are narrowest ambitally and expand slightly adapically (Fig. 36A). Plates both at the ambitus and adorally are trigeminate with all three elements reaching the perradial suture (Fig. 36C). The central element is the largest. Each plate carries a single imperforate, non-crenulate tubercle which straddles all three elements. Adorally, pore-pairs become crowded so as to form small phyllodes (Pl. 14, fig. 6). Immediately above the ambitus pore-pairs become biserially arranged and primary tubercles diminish in size and do not reach the apex (PI. 14, fig. 5). There are 48 pore-pairs in a column (of which the most adapical 18-20 are biserially arranged). There are 12 or 13 primary tubercles. Secondary tubercles lie immediately adjacent to each pore-pair and may also occur along the perradius on occasional elements. Interambulacra are 39-44% of the test diameter in width at the ambitus. Each plate is very wide and short and bears a row of small, equal-sized tubercles. These are imperforate and appear non-crenulate, but on close inspection of well- preserved material (e.g. BMNH EE3573) there are faint traces of crenulation to larger tubercles. At the ambitus the interambulacral plates become very much narrower and the size of the tubercles more or less halves (PI. 14, figs 3, 4; Fig. 36B). All plates are arranged to form a pronounced downward-pointing V. Both columns of plates reach the peristomial border. The peristome is 42-52% of the test diameter in diameter and is hardly invaginated. There are very feeble buccal notches. The perignathic girdle consists of small auricles which do not meet above the perradius. REMARKS. The species is distinguished from the type species P. africanum Peron & Gauthier (Cotteau et al. 1881) by its somewhat sharper decrease in interambulacral plate size at the ambitus. However, the two species are very similar indeed in other features and the Algerian and Omani-Iranian species may eventually turn out to be conspecific. The residual crenulation on well-preserved tubercles and the large A.B. SMITH Fig. 36 Camera lucida drawings of plating in Plistophyma asiaticum Cotteau & Gauthier, BMNH EE3575. A, ambulacrum, from apex (top) to peristomial margin (bottom); B, interambulacrum from apex (top) to peristomial margin (bottom); C, detail af ambital ambulacral plating. Scale bars = 1 mm. Fig. 37 Camera lucida drawings of plating in Circopeltis? emiratus sp. nov. A, apical disc, BMNH EE3584; B ambital ambulacral plating, BMNH EE3582; C, ambital plating, ambulacrum to right, interambulacral plate to left, BMNH EE3584. Scale bar = 1 mm. LATE CAMPANIAN-MAASTRICHTIAN ECHINOIDS PLATE 14 Figs 1-7 Plistophyma asiaticum Cotteau & Gauthier. 1-3, 6, BMNH EE4932; 1, oral, x 5; 2, apical, x 5; 3, lateral, x 5; 6, detail of peristomial region, x 8. Jebel Thanais, lowest 1 m of the Simsima Formation. 4, 7, BMNH EE3575; 4, lateral; 7, oral; both x 3. Jebel | Thanais, basal 1 m of the Simsima Formation. 5, BMNH EE3572; apical detail, x 8. Jebel Buhays, section 1; loose in the scree derived from the lowest 3 m of the Simsima Formatio 172 pentagonal caducous apical disc suggests that Plistophyma belongs within the phymosomatoids. Family STOMECHINIDAE Pomel, 1883 Genus CIRCOPELTIS Pomel, 1883 Pi isetiess—3 elle fos 2hig1 37 Circopeltis? emiratus sp. nov. Types. The holotype is BMNH EE3584 and paratypes are BMNH EE3582, EE3583, EE3585 and EE3596. OTHER MATERIAL. There is one other fragment tentatively attributed to this species, BMNH EE3586. OCCURRENCE. In the western Oman mountains this species is found at the following levels: Jebel Buhays section 1: loose in scree (derived from lower beds of the Simsima Formation) (1). Jebel Buhays section 3: immediately overlying prominent red marly sand level (ca. 3 m above the base of the Simsima Formation) (2). Jebel Rawdah section 3: bed 6 (1). Section 3b basal 1 m (1). Jebel Rawdah, section 4: bed 4 (1). It is thus known from the Maastrichtian of the western Oman Mountains. DIAGNOSIS. Test low,. domal; apical disc hemicyclic with plates firmly bound to the corona. Ambulacra straight, plating compound in the phymosomatoid style with five or six elements to a compound plate; pore-pairs arranged in arcs from apex to peristome. There is a single large primary tubercle to each plate with broad adradial and interradial zones of secondary tuberculation. DESCRIPTION. Tests range in diameter from 20-1 mm to 47-2 mm and in height from 11-8 to 23-5 mm (mean test height is 53% of test diameter). The test is circular in outline and flattened ovate in profile, with the ambitus at approximately 40% of test height above the base. The apical disc is hemicyclic, with the three anterior oculars insert and the posterior two exsert (Fig. 37A). The apical disc is 19-24% of the test diameter in diameter (mean = 22%) and the periproct occupies approximately 55% of the apical disc diameter. The periproct is D-shaped in outline, with the slightly flattened edge abutting the madreporite. Genital plate 2 is the largest and is tumid and covered in dense madrepores. Other genital plates and all ocular plates have a scattering of small miliary tubercles. The ambulacra are 22-24% of the test diameter in width at the ambitus. They are straight and taper gradually adapically. All plates are compound in the phymosomatid style (Fig. 37B) and most are composed of five, or occasionally six, elements. The three most adoral plates however are com- posed of just four elements. Pore-pairs are arranged in arcs and phyllodes are developed around the peristomial edge. Each compound plate has a single large primary tubercle (imperforate and non-crenulate) that overlaps all five (or six) elements that make up the compound plate (Fig. 37C; Pl. 17, fig. 1). There is a much smaller secondary tubercle lying perradially, plus one or two scattered miliary granules. The perradial zone is relatively broad. There are 14 compound plates and 73 pore-pairs at 20 mm test diameter, rising to 18 A.B. SMITH compound plates and 88 pore-pairs at 37 mm test diameter. There are no sphaeridial pits. Interambulacra are broad and carry two rows of primary tubercles, set close to the adradial margin. There are 13 plates in a column at 20 mm test diameter, rising to 17 at 37 mm test diameter. Areoles are almost contiguous adorally but are separated adapically. The mamelon is large and imperforate and there is a narrow ledge that may retain faint traces of crenulation in well-preserved individuals. Adradi- ally there is a narrow band of small secondary tubercles and miliary granules. Interradially the plates are slightly | depressed and there are small and irregularly scattered sec- | ondary tubercles throughout (PI. 17, fig. 2). Both columns reach the peristomial margin. The peristome is about 42% of the test diameter across. It is slightly invaginated. Buccal notches are moderately well- developed and have a thickened lip. Lantern and spines are unknown. REMARKS. This species has a phymosomatid style of ambu- lacral compounding. However, the apical disc structure dif- ferentiates it from the great majority of phymosomatids, since these almost all have monocyclic apical discs that are typically caducous. Only Glyptocidaris has a comparable apical disc. Furthermore, this species has tuberculation that is virtually non-crenulate, whereas other genera, including Glyptoci- | daris, show stronger crenulation. The species is here tenta- | tively assigned to the genus Circopeltis. Circopeltis has polygeminate plate compounding and a hemicyclic apical disc. Circopeltis also has non-crenulate tuberculation.. How- ever, its ambulacral compounding style is unreported and it is not yet known whether it is phymosomatoid. It differs from Phymechinus? perplexus in its finer tubercu- lation, with more extensive scattered secondary tubercula- tion. More importantly it has ambulacral pore-pairs in simple | © arcs throughout. In Phymechinus pore-pairs become irregu- | larly biserial adapically. Furthermore, Circopeltis? has a | larger peristome and much less well developed phyllodes than | does Phymechinus? perplexus from the same levels. How- ever, there is little doubt that the two forms are rather closely | | related. Genus PHYMECHINUS Desor, 1856 Phymechinus? perplexus sp. nov PI. 15, figs 4-10; Figs | 38, 308 Types. Holotype EE3579, paratypes BMNH EE3581, | JB E25 oil, HSs5 98), BSI). MATERIAL STUDIED. 14 specimens, of which biometric data | - was derived from the following: BMNH EE3578-79, EE3581, EE3989, EE3991-94, EE3619. OCCURRENCE. All specimens come from Jebel Rawdah as follows: Section 1: bed 4 (2). Section 2: bed 6 (2); bed 8 (4); bed 11 (1); loose in scree (4). Section 4: bed 8/9 (1). DIAGNOsIS. Apical disc small, caducous. Ambulacra polygeminate with seven or eight elements to a compound plate. Plate compounding phymosomatid-style. Pore-pairs arcuate or irrgularly multiple above. One primary imperfo- rate, crenulate tubercle on each ambulacral and interambu- LATE CAMPANIAN-MAASTRICHTIAN ECHINOIDS 173 PLATE 15 figs 1-3 Circopeltis? emiratus sp. nov. BMNH EE3584, holotype; 1, apical; 2, oral; 3, lateral; all x 2. Jebel Buhays, section 3, ca. 3m | above the base of the Simsima Formation. jigs 4-10 Phymechinus? perplexus sp. nov. 4-6, BMNH EE3591, paratype; 4, oral; 5, apical; 6, lateral; all x 2. Jebel Rawdah, section 2, | bed 8. 7, BMNH EE3579, holotype; lateral, x 1. Jebel Rawdah, section 2, bed 8. 8-10, BMNH EE3581, paratype; 8, lateral; 9, apical; 10, | oral; all x 2. Jebel Rawdah, section 2, loose in scree derived from beds 3-10. 174 40 Ww o N °o Test height (mm) fo) Width at ambitus (mm) 50 20 30 40 Test diameter (mm) = [2) fo) Number of pore-pairs in column 20 30 Test diameter (mm) Number of interamb. plates in column 20 30 Test diameter (mm) Fig. 38 Biometric data for Phymechinus? perplexus sp. nov. lacral plate. Mamelons very large and crenulation feebly developed. Lower surface flat, peristome small with feeble buccal notches. Phyllodes extremely well-developed. DESCRIPTION. Tests are 11 to 60 mm in diameter and 5-8 to 31-2 mm in height (test height 50-60% of diameter; mean = 54%, SD = 3-2%, N = 9). They are circular in outline and bun-shaped in profile, with a broad base and depressed conical upper surface (Pl. 15, fig. 8). The ambitus lies at approximately 30% of test height above the base. The apical disc is small and circular, only 16-28% of the test diameter across (PI. 15, fig. 9). It is proportionally smaller in larger individuals (Fig. 38). No specimen retains any apical disc plating, but to judge from the small size of the 60 Diameter (mm) 40 40 A.B. SMITH 30 8 Ambulacrum @ Interambulacrum Test diameter (mm) ®@ Peristome ® Apical disc 50 20 30 40 60 Test diameter (mm) opening, plating is almost certain to have been dicyclic or hemicyclic. Ambulacra are only slightly narrower than interambulacra at the ambitus and measure 23-28% of the test diameter in width (mean = 25:5%, SD = 1:8%, N = 9). They are more or less straight, tapering slightly adorally and more signifi- cantly adapically. All plates are polygeminate with seven (rarely six or eight) pore-pairs to an ambital plate. Plate compounding is in the phymosomatid style (Figs 39A, B). Pore-pairs are strongly arced at the ambitus, but tend to become irregularly biserial or pleuriserial adapically and adorally (Figs 38C, D). Close to the apex, pore-pairs once again become uniserially arranged. Adapically, small second- ary tubercles occur scattered within the pore zone. However, LATE CAMPANIAN-MAASTRICHTIAN ECHINOIDS there are a few specimens (e.g. EE3581) in which the pore-pairs are in uniserial arcs right to the apex. Each plate carries a single large primary tubercle that occupies most of the plate (Pl. 15, fig. 7; Figs 38A, B). This has a relatively well-developed imperforate mamelon and is distinctly crenu- late, though the surrounding platform is not very broad. Areoles are circular and separated by a single ring of small secondary and miliary tubercles. There are 53 pore-pairs in a column at 11 mm test diameter, rising to about 100 pore-pairs at 34 mm test diameter (Fig. 38). Tubercles are largest at the ambitus and decrease in size gradually both adapically and adorally. Adorally there are very strong phyllodes composed of circular pore-pairs with well-developed periporal muscle attachment areas (PI. 15, fig. 10). These phyllodes are so large as to make up more than half of the ambulacral width. Interambulacra are relatively narrow, being only 30-37% of the test diameter in width at the ambitus. There are 10 plates in a column at 11 mm test diameter, rising to 16 at 34 mm test diameter. Each plate carries a large primary tubercle which is imperforate and weakly crenulate. In smaller speci- mens (i.e. less than 35 mm) these are the only large tubercles present, although a distinct small secondary tubercle is present both adradially and interradially. In specimens around 58 mm test diameter the adradial secondary tubercle reaches approximately half the diameter of the primary tubercle at the ambitus and adorally, and there are additional smaller secondary tubercles close to the adradial suture. Primary and secondary tubercles extend to the apex and _peristome. The primary tubercles are non-confluent, being separated by a single row of miliaries. The peristome is 30-55% of the test diameter in width (proportionally smaller in larger individuals) and is hardly invaginated, the entire lower surface being flat. Buccal ' notches are small and poorly developed. Spines, lantern and _ pedicellariae are all unknown. | REMARKS. It is with some slight hesitation that I refer the _ new species P? perplexus to this genus. This species has stout 'tubercles with large mamelons and virtually no platform. ‘Nevertheless fine crenulation is developed around the mamelon, though it is often difficult to see unless preserva- tion is near perfect. The ambulacral plate compounding is ‘Fig. 39 Camera lucida drawings of plating in Phymechinus? perplexus sp. nov., BMNH EE3581. A, ambital ambulacral plate; B, adapical ambulacral plate; C, adapical pore-pair arrangement; D, adoral pore-pair arrangement. Scale bar = 1 mm. iD phymosomatid in style, with adapical pore-pairs either strongly arcuate or, more often, actually multiple in an irregular way. The apical disc is small and unlikely to be monocyclic, but plating is not preserved. The type species of Phymechinus, P. mirabilis (Agassiz) comes from the Middle Oxfordian and has a similar overall shape, small apical disc and stout tuberculation. Well-preserved specimens (e.g. Hess 1975, pl. 37, fig. 5) apparently show feeble crenulation. Unfortunately, none of the specimens I have examined show the style of ambulacral compounding. P? perplexus differs from the type species in having a proportionally smaller peristome and better developed phyllodes. It also has much less well-developed buccal notches. In tuberculation style it is very similar to Schluter’s species Phymechinus cretaceus from the B. mucronata Zone, Upper Campanian of Ciply, Bel- gium, but this species has a much larger peristome and much less well-developed phyllodes. This species is easily distin- guished from Phymosoma cf. paroni Checchia-Rispoli by its very much smaller apical disc, more subconical profile and compound amulacral plates that incorporate more than five elements. It is also easily distinguished from Actinophyma. Although Actinophyma has a similar arrangement of pore- pairs adapically, forming rather irregular multiple columns with intersperced tubercles, it is very different adorally. The peristome in Actinophyma is invaginated and the pore-pairs remain uniserial and rather widely spaced across the entire oral surface. In P? perplexus the pore-pairs form a very strong phyllode and the oral area is broad and flat. It is distinguished from Circopeltis emiratus by its very much coarser tuberculation, smaller peristome, more polygeminate ambulacral compounding and cadoucous apical disc. Furthermore, C. emiratus never developed biserial pore-pairs adapically. Genus ECHINOTIARA Pomel, 1883 Echinotiara perebaskinei Lambert, 1930 Pl. 16, figs 1-6; Pl. 17, figs 3, 6, 7; Figs 40, 41 1930 Echinotiara perebaskinei Lambert, in Lambert & Pere- baskine: 472, pl. 38, figs 1-5. Types. The two specimens figured and described by Lambert and presumably in the Lambert Collection, Université de Paris VI, France. MATERIAL STUDIED. 82 specimens of which only the follow- ing were used in the biometric analysis: BMNH EE3756-57, EE3761-63, EE3767-69, EE3772, EE3774, EE3782, EE3785-88. OCCURRENCE. Along the western margins of the Oman mountains this species is confined to the lowest arenaceous levels of the Simsima Formation at Jebel Rawdah. It occurs as follows: Jebel Rawdah, section 2: bed 4 (6); bed 6 (9); bed 8 (49); bed 11 (3); loose in lower scree (17). Jebel Rawdah, section 3b: bed 3 (1). Jebel Rawdah, section 4: bed 2 (1); bed 8/9 (1). The species was originally described from the ‘Calcaires inferieur a Libycoceras, Maastrichtien’ at Oued Tarinkat, Tchi-Dermine and Oued Tinamassine in tke district of Gao, 176 Test height (mm) Test diameter (mm) & Amulacral pore-pairs @ Interambulacral plates 40 Number in column N o 10 20 30 Test diameter (mm) 40 Fig. 40 Biometric data for Echinotiara perebaskinei Lambert. Niger (Lambert & Perebaskine 1930). Amard ef al. (1981: 124) reported an Echinotiara cf. perebaskinei Lambert from the Upper Maastrichtian of Tademait, Algeria, but as no figures or description are given it is not possible to confirm this record. DiAGnosis. An Echinotiara with a relatively small caducous apical disc, ambulacral pore-pairs in arcs of three, with very strong phyllodes developed adorally. Interambulacral plates with a single large primary tubercle and well-developed adradial and interradial secondary tubercles that diminish in size adapically. DESCRIPTION. Tests range from 9 to 32 mm in diameter and are more or less circular in outline. Test height is 40-52% of test diameter (mean = 46%, SD = 3:7%, N = 14). In profile the test is low conical (Pl. 16, fig. 3), with the ambitus approximately 40% above the base. The apical disc is always missing and plating can only have been loosely fixed to the corona. The apical disc outline is irregularly circular (Pl. 16, fig. 1) and is 21-29% of the test diameter in length (mean = 24%, SD = 2:1%, N = 13). To judge from the size, it must have been dicyclic or hemicyclic. Ambulacra are trigeminate throughout with pore-pairs arranged in distinct arcs of three (Fig. 41A). Ambulacra are relatively broad, being approximatley 22% of the test diam- eter in width at the ambitus. They taper gradually adapically, but primary tubercles continue to the apex (Pl. 17, fig. 3). Adorally the ambulacra remain broad and there are large, well-developed phyllodes, which in specimens 25 mm in diameter include at least 22 pore-pairs in each column (PI. 16, Diameter (mm) Number in column ® Peristome @ Apical disc Test diameter (mm) ® Amulacral pore-pairs ¢@ |Interambulacral plates 60 BS lo) 20 20 Test diameter (mm) 30 40 fig. 5). Each compound plate carries a single large imperfo- rate and non-crenulate tubercle that overlaps all three ele- | ments (Fig. 41). Plate compounding is diadematoid in style, with all three elements reaching the perradius and the middle | plate pinched towards the centre of the plate (Fig. 41C). The lowest element in each compound plate carries a small | secondary tubercle adradially and perradially. There are 32 | pore-pairs in a column at 9 mm test diameter, rising to 71 at | 32 mm test diameter (Fig. 40). At the ambitus the interambulacra are 35-38% of the test | diameter in width. Both columns reach the peristomial bor- | der, although the interambulacra taper considerably towards | the peristome. Plates at the ambitus are longer than wide and are slightly curved. Each plate carries a single large imperfo- | rate and apparently non-crenulate tubercle at its centre (PI. 17, fig. 7). The mamelon is slightly undercut but there is little | surrounding platform developed. On the adradial side there are one or two smaller secondary tubercles, while on the interradial edge there is a single secondary tubercle (Fig. | 41C). These primary and secondary tubercles are relatively | coarse and occupy most of the available space. There are, | however, miliary granules and small tertiary tubercles along | the adapical margin and in spaces adjacent to the primary tubercle. There are 10 plates in a column at 9 mm test diameter, rising to 16 at 32 mm test diameter. Primary tubercles continue to the apex and there is no median naked zone. The peristome is circular and measures 35-50% of the test diameter in diameter (it is proportionally larger in smaller individuals (Pl. 16, figs 4, 5). It is only slightly sunken and tp LATE CAMPANIAN-MAASTRICHTIAN ECHINOIDS 177 |PLATE 16 (Figs 1-6 = Echinotiara perebaskinei Lambert. Jebel Rawdah, section 2, beds 6-8. 1, 2, BMNH EE3756; 1, apical, 2, oral; both x 3. 3, 4, BMNH EE3768; 3, lateral; 4, oral; both x 3. 5, 6, BMNH EE3788; 5, oral; 6, lateral; both x 3. 178 Fig. 41 Camera lucida drawings of plating in Echinotiara perebaskinei Lambert. A, BMNH EE3769, ambulacrum from apex (top) to peristome margin (bottom); B, BMNH EE3763, ambulacrum from apex (top) to peristome margin (bottom); C, ambital ambulacral and interambulacral plate, BMNH EE3763. Scale bar = 1 mm. buccal notches are small but distinct. The perignathic girdle structure is seen in BMNH EE3783 and EE3788. It consists of two long peg-like auricles that do not meet above the perradius. Spines and lantern are unknown. REMARKS. This. species lacks crenulate tuberculation, although preservation is usually inadequate to be certain for most specimens. It could easily be mistaken for Orthopsis miliaris on account of its very similar tuberculation and test shape. However, Orthopsis has perforate tuberculation and its ambulacral pore-pairs are strictly uniserial, not arranged in arcs of three as in Echinotiara. Furthermore, Echinotiara has well-developed phyllodes that are never seen in Orthopsis. The type material described and illustrated by Lambert differs in apparently having slightly less well developed phyllodes adorally at comparable sizes, but for the present the two populations are treated as conspecific. A.B. SMITH Cohort IRREGULARIA Latreille, 1825 Order HOLECTYPOIDA Duncan, 1889 Family HOLECTYPIDAE Lambert, 1899 Genus COENHOLECTYPUS Pomel, 1883 Coenholectypus inflatus (Cotteau & Gauthier, 1895) Pl. 18, figs 7-11; Figs 42, 43A, C Holectypus inflatus Cotteau & Gauthier: 73, pl. 12, figs 14. 21989 Holectypus (Caenholectypus) inflatus Cotteau & Gauthier; Ali: 401, fig. 4 (1). Types. The specimen described and illustrated by Cotteau & Gauthier, from the late Cretaceous of Aftab, southern Iran. 1895 MATERIAL STUDIED. Thirty specimens, of which 16 were | used in the biometric analysis (BMNH E82644, EE3399, EE3401-04, EE3406-07, EE3409, EE3411-15, EE3417, EE3429). OCCURRENCE. There are two morphologies found along the western foothills of the Oman Mountains: Depressed variety: this occurs at Jebel Huwayyah, from an uncertain horizon. It is also found at Jebel Buhays (sections 1 and 2) in the lowest limestones. It is more common at Jebel Rawdah: in section 3 it occurs reasonably abundantly in bed 6 (6) and is found at a comparable level in section 4 (beds 19 and 21—22 and at top of measured section) (10). It also occurs frequently at and immediately above the level of the first major red parting in section 3 (bed 5). More inflated, rounded forms are recorded from Jebel Huwayyah (unknown horizon); and from Jebel Rawdah, Section 2, bed 6, section 3, bed 10, section 4, bed 13. DESCRIPTION. Tests are circular in outline and range from around 13 mm up to 56 mm in diameter. In profile their height varies from low conical to almost subglobular. Test height is 47-77% of test length (Fig. 42). The ambitus is well rounded and lies a little below midheight (Pl. 18, fig. 11). The apical disc is small and compact with five gonopores. Genital plates are small and pentagonal and separated from one another by ocular plates that are almost as large (Fig. 43C). Madrepores occupy the entire central region. Ambulacra are simple and uniserial, except adapically in larger specimens (40 mm diameter plus), where pore-pairs become slightly offset creating an incipient biserial arrange- ment. There is no pore crowding towards the peristome (Fig. 43A) and all plating is simple. Only close to the peristome does ambulacral plating become differentiated into triads with every third plate becoming enlarged. All pore-pairs are nonconjugate. There are approximately 100 pore-pairs in a column at 18 mm test length, rising to 185 at 56 mm test length (Fig. 42). Interambulacra are standard in their structure. The periproct is oval in outline and lies close to the peristome and well separated from the ambitus (PI. 18, fig. 7; Fig. 43A). It opens between interambulacral plates 2a,b and 6b,7a or 7a,b. Its width is 55-81% of its length (mean = 65%, SD = 7%, N = 17). The distance separating the periproct and peristome is small, only 20-50% of the periproct length; whereas the distance separating the periproct from the ambitus is much greater, being 110-260% of the periproct length (greatest in largest individuals). The peristome is circular with feeble buccal notches. It is 15-35% of test length in diameter (mean = 23%, SD = t LATE CAMPANIAN-MAASTRICHTIAN ECHINOIDS 179 ‘PLATE 17 Figs 1,2 Circopeltis? emiratus sp. nov. BMNH EE3584, holotype, details of ambital tuberculation; 1, ambulacrum, x 1325 | interambulacrum, x 5. Figs 3, 6,7 Echinotiara perebaskinei Lambert. 3, 6, BMNH EE3774; 3, apical, x 6; 6, ambital detail, x 5. Jebel Rawdah, section 2, bed 4. | 7, BMNH EE3785, lateral detail, x 6. Jebel Rawdah, section 2, beds 6-8. Figs 4,5 Salenia nutrix Peron & Gauthier. 4, BMNH EE3627, apical, x 5. Jebel Rawdah, section 1, bed 3. 5, BMNH EE3634, lateral, x 5. Jebel Rawdah, section 2, loose in scree at level of bed 11. 180 Fig. 42 4-6%, 40 é @ C. cf baluchistanensis @ C. inflatus e ao) £ & £ 220 © £ r) Ww © F 10 20 30 40 50 60 Test diameter (mm) C. cf. baluchistanensis C. inflatus 40 w fo} N (o} Number of interamb. plates in column rS) 20 30 40 Test diameter (mm) 50 = lo} a C. cf baluchistanensis C. inflatus o fee) Periproct length (mm) o 2 10 20 30 40 50 60 Test diameter (mm) 10r w C cf. baluchistanensis @ C. inflatus Periproct width (mm) Periproct length (mm) Biometric data for Coenholectypus inflatus (Cotteau & Gauthier) and C. cf. baluchistanensis (Noetling). N = 16), being relatively larger in small individuals. REMARKS. This species is easily recognised by its rounded profile, and its small periproct that lies close to the peristome and well separated from the ambitus. Although Cotteau & Gauthier’s (1895) description is sketchy, it is readily recognis- Distance between peristome and periproct (mm) Distance from periproct to ambitus (mm) able from the illustrations given. Ali (1989) described two) species of Coenholectypus from the western Oman mountains| ¢ a Number of pore-pairs in column A.B. SMITH 200 BS C. cf baluchistanensis @ C. inflatus ey 180 160 140 120 100 30 40 60 Test diameter (mm) 10) ® C. cf. baluchistanensis © C. inflatus 2 4 6 8 Test length (mm) B C. cf baluchistanensis © C. inflatus ° Peristome length (mm) 20 30 40 Test diameter (mm) SO 60 8 C. cf. baluchistanensis © C. inflatus Test length (mm) . inflatus and C. larteti (Cotteau). Unfortunately the latter is Cenomanian species (Smith et al. 1990). Because Ali gave! — no description and only illustrated the aboral surfaces of his LATE CAMPANIAN-MAASTRICHTIAN ECHINOIDS 181 Pie yee og SG 56 | PLATE 18 )Figs 1-6 Coenholectypus cf. baluchistanensis (Noetling). 1-3, BMNH EE3389; 1, apical; 2, oral; 3, lateral; all x 2. Jebel Buhays, section 1; loose in the scree derived from the lowest 3 m of the Simsima Formation. 4-6, BMNH EE 3395; 4, apical; 5, oral; 6, lateral; all « 2. Jebel Rawdah, section 2, bed 11. Figs 7-11 Coenholectypus inflatus Cotteau & Gauthier. 7, BMNH EE4330; oral, x 1. Jebel Buhays, section 1; loose in the scree derived | from the lowest 3 m of the Simsima Formation. 8, 9, BMNH EE3412; 8, lateral; 9, oral; both x 2. Jebel Buhays, section 2, lowest part of | the Simsima Formation. 10, 11, BMNH EE3424; 10, apical; 11, lateral; both x 1-5. Jebel Rawdah, section 3b, loose, from higher part of the section. 182 two forms, it is impossible to tell to which species either belongs. His assignment of one to C. inflatus is therefore accepted tentatively. C. inflatus differs from C. cf. baluchistanensis (Noetling), the other species described here, on several counts. Firstly, it has finer tuberculation on its oral surface at comparable sizes. Secondly, its peristome is proportionally smaller at compa- rable sizes, and thirdly, its periproct is smaller and more removed from the ambitus at all sizes (Fig. 42). There is a stratigraphical variation in profile. The taller forms, exactly comparable in form with the Iranian type, are found lower down in the sections in the coarse calcarenites: in bed 2 at Jebel Rawdah section 4, In bed 6 at section 2, and in bed 13 at section 4. The more depressed forms are found higher up in the more orbitoline-rich limestones and may represent deeper water morphological varieties. Coenholectypus cf. baluchistanensis (Noetling, 1897) Pl. 18, figs 1-6; Figs 42, 43B, D cf. 1897 Holectypus baluchistanensis Noetling: 18, pl. 3, fig. 3. MATERIAL. Thirteen specimens (BMNH EE3386-98). OCCURRENCE. In the western Oman mountains this species was found at the following localities and horizons: Jebel Buhays, section 1: loose in scree (3) and bed 8 (1). Jebel Thanais, lowest 2 m of limestone section (1). Jebel Rawdah, section 1: 20 cm below the top of bed 5 (1). Jebel Rawdah, section 2: bed 11 (9). A Fig. 43 Camera lucida drawings of plating in Coenholectypus. A, C, C. inflatus (Cotteau & Gauthier), BMNH EE3412: A, oral plating; C, A.B. SMITH It was originally described from the Maastrichtian of Bal- uchistan. DESCRIPTION. Tests are circular in outline and range from 18-37 mm in diameter. They are moderately inflated in profile (Pl. 18, fig. 3), with a test height 55-62% of test length. The ambitus is rounded and lies a little below mid-height. The apical disc has five gonopores. The four genital plates are small and pentagonal and separated from each other by ocular plates that are similar in size (Fig. 43D). The madreporite plate is large and often tumid. Madrepores are well developed over the central area. Ambulacra are uniserial throughout, with simple noncon- jugate pores. There is no hint of incipient biseriality adapi- cally, nor any pore crowding adorally. All plates are simple, | with triad development only appearing towards the peris- tome. The interambulacra are standard in structure. The periproct is relatively large and opens between interambulac- ral plates 2a,b and 7a,b. It is oval in outline, pointed at both ends. Its width is 60-76% of its length (mean = 70%, SD = 5:7%, N = 7). It opens close to the peristome (PI. 18, fig. 2; Fig. 43B), separated by 20-45% of the periproct length from the peristome (mean = 32%, SD = 8-4%, N = 7). It is separated from the ambitus by only a small distance in smaller individuals, some 28% of periproct length at 17 mm test length, but this increases in larger individuals to reach 65% of periproct length at a test length of 35-37 mm. The peristome is circular in outline with deep and well marked buccal notches. It is 10-22-5% of test length in apical disc. B, D, C. cf. baluchistanensis (Noetling), BMNH EE3389: B, oral plating; D, apical disc. Scale bars: A, B = 5 mm; C, D = 1 mm. | LATE CAMPANIAN-MAASTRICHTIAN ECHINOIDS diameter (being relatively larger in small individuals). REMARKS. This species is easily distinguished from the other Coenholectypus that occurs here, C. inflatus, by its larger peristome, larger periproct that extends much closer to the ambitus, and by its coarser adoral tuberculation. It also has deeper buccal notches. C. baluchistanensis Noetling, from the Maastrichtian of Baluchistan, was established on the basis of two specimens 31 and 38 mm in diameter. Like the species described here, this has a relatively large peristome occupying much of the lower surface. Unlike the arabian specimens, the periproct of the Baluchistan species extends closer to the ambitus in the illustrated specimen. However, Noetling states that all speci- mens are crushed to some degree, and the illustrated speci- men is damaged posteriorly. Therefore I suspect that the periproct is in reality slightly more distant from the ambitus than is actually shown. Until new material of C. baluchistan- ensis is available, the identification of Omani material as this species must remain tentative. C. subcrassus Peron & Gauthier, from the early Maastrich- tian of Algeria and Tunisia, differs from C. baluchistanensis _ in having a much larger periproct that occupies virtually the » entire oral surface of the posterior interambulacrum. It is illustrated as having a broad, flat, adambital margin, unlike { the arabian species. However, C. subcrassus and C. baluchis- ' tanensis are sister taxa. Genus COPTODISCUS Cotteau & Gauthier, 1895 | Coptodiscus magniproctus sp. nov. PI. 19, figs 4-7; Figs 44, 45 BMNH | | Types. Holotype, BMNH_ EE3716, _ EE3715. There are no other specimens. paratype, OCCURRENCE. Both specimens come from the base of the “silty Loftusia beds (bed 1), at Jebel Huwayyah, section 2. | DIAGNOsIs. A Coptodiscus with a relatively large periproct occupying most of the oral surface of the posterior interam- bulacrum and opening between interambulacral plates 2 and 7. Aboral interambulacral ornament comprising a series of fine sutural pits and a set of pits along the midline of the each | plate. DESCRIPTION. Both specimens are small, but have open ‘gonopores and are thus mature individuals. The smaller ‘specimen is 10 mm in diameter, the larger 16 mm. Both are ‘circular and low conical in profile, with the ambitus posi- \tioned relatively low down. Test height is about 40% of test ‘diameter. The apical disc is small and compact (Fig. 45A). There are five genital plates, each perforated by a gonopore. The ocular plates are almost as large as the genital plates, but are exsert and only oculars III and II abut the madreporite plate. The Zenital plates are contiguous around the posterior and lateral \margins of the madreporite. | Ambulacra are uniserial and simple throughout, pores secoming slightly more widely spaced towards the peristome. There are three and a half ambulacral plates to an ambital mterambulacral plate. The peristome is relatively large, 25% of test diameter in liameter. It is not much invaginated. The periproct is rela- ‘ively large, being 3-6 mm in length (22% of test diameter) by @C. magniproctus Periproct width (mm) “10 20 30 Test diameter (mm) 4 2 @C. magniproctus <= ~ Le,) r= oe ~ ° 3 _ 2a © a 2 10 20 30 Test diameter (mm) Fig. 44 Biometric data for species of Coptodiscus. Data from the type series of Coptodiscus noemiae Cotteau & Gauthier. The holotype of C. magniproctus sp. nov. is also plotted. A B O fe a8 COS Fig. 45 Camera lucida drawings of Coptodiscus magniproctus sp. nov., BMNH EE3716. A, Apical disc; B, ambital interambulacral plate, adradial margin to right. Scale bar = 1 mm. 2:3 mm. It is pointed at both ends and lies close to both the peristome and the ambitus. Only 1-3 mm separates the periproct from the peristome and 1-2 mm separates the periproct from the ambitus. The periproct opens at interam- bulacral plate 2 and extends to interambulacral plate 7 (Fig. 44). Tuberculation is standard with three or four primary tubercles on an ambital interambulacral plate. The surface of the test is ornamented by rows of fine pits (PI. 19, fig. 4; Fig. 45B). These are arranged along the horizontal sutures, but are also developed along the midline of the plate in between the primary tubercles. REMARKS. Amongst holectypoids, ornamentation of the test, as seen in this species, is found only in the genus Coptodiscus. Only one late Cretaceous species of Coptodis- cus has ever been described, C. nomiae Cotteau & Gauthier (1895) (Pl. 19, figs 1-3; see Kier 1972 for a detailed descrip- tion). C. nomiae, which comes from the ‘Senonian’ of south- 184 A.B. SMITH | PLATE 19 Figs 1-3 Coptodiscus noemiae Cotteau & Gauthier. Syntype, from the Morgan Collection, Museum d’Histoire Naturelle, Paris; 1, apical, x 5; 2, apical, x 2; 3, oral, x 2; Senonian, Khianan, Iran. } Figs 4-7 Coptodiscus magniproctus sp. nov. Jebel Huwayyah, section 2, bed 1. 4, 6, 7, BMNH EE3716, holotype; 4, detail of adapical ornamentation, x 10; 6, oral, X 3; 7, apical, x 3. 5, BMNH EE3715, paratype, apical surface, x 5. Figs 8-11 Conulus douvillei Cotteau & Gauthier. BMNH EE4306; 8, apical; 9, oral; 10, lateral; 11, posterior; all x 2. Jebel Thanais, lowest 2 m of the Simsima Formation. } LATE CAMPANIAN-MAASTRICHTIAN ECHINOIDS 185 40 8 G. bleicheri @ C. douvillei = x E E 30 E E * y a [>] = 3 20 2 5 E ” © is E 10 2 LJ i= © a (0) (0) 10 20 30. 40 50 2 3 4 5 6 U/ 8 AAG Test diameter (mm) 5a Peristome major axis (mm) 8 G. Dleicheri 4 S G. bleicheri © C. dowvillei on 50 © C. dowillei 40 5 -_-_ 5 E ° ~ 30 : g a AS) = 2 20 3 3 NM = é 10 10) 10 20 30 40 50 Test diameter (mm) Test length (mm) 14 8 G. bleicheri 12 @ C. douvillei a Height to base of periproct (mm) Test diameter (mm) Fig. 46 Biometric data for ‘Globator’ bleicheri (Thomas & Gauthier) and Conulus douvillei (Cotteau & Gauthier). ern Iran and the late Campanian of Saudi Arabia and Oman, Family CONULIDAE Lambert, 1911 differs from our species in several important details. Firstly, Genus ‘GLOBATOR’ Agassiz, 1840 its periproct is very much smaller at comparable sizes (Fig. |44) and it lies well separated from the peristome, with at least three and usually four interambulacral plates in each column Separating the two openings, as opposed to the two in C. magniproctus. The periproct is also considerably more rounded in C. nomiae. A second immediately apparent difference is in the style of ornamentation developed abo- REMARKS. The genus Globator was erected by Agassiz (1840) for small, ovoid conulids with large periprocts opening above the ambitus. The type species is Globator nucleus Agassiz, but this is based on a juvenile Conulus rotundus Goldfuss and thus Globator falls into synonymy with Conu- lus. However, there is a distinct clade of ovoid conulids with 2 . 3 ; large, supra-ambital periprocts that can be recognized. This rally. In C. nomiae there is a single laterally extensive sutural one is ie a. Pare! (Smith & Wright, in prep.) pit on either side of the primary tubercle, also, intraplate pits ee | : : 5 and for the moment we retain its members under the name are not as well developed, whereas in C. magniprocta there is ‘Globator’ 4 well-developed row of sutural pits (compare PI. 19, figs 1, 4), } 186 ‘Globator’ bleicheri (Gauthier, 1889) PI. 20, figs 1-10; Figs 46, 47A, B, F-I, 48A 1889 Pyrina bleicheri Gauthier: 51, pl. 3, figs 15-18. 1895 Pyrina orientalis Cotteau & Gauthier: 68, pl. 11, figs 1-8. 1897 Pyrina zumoffeni de Loriol: 158, pl. 7, fig. 1. 1967 Pyrina ovulum Agassiz; Devries: 177, pl. 5, figs 19-21. 1987 Pseudopyrina bleicheri (Thomas & Gauthier); Zhagbib-Turki: 167. 1989 Globator orientalis (Cotteau & Gauthier); Ali: 403, fig. 5 (4-5). MATERIAL STUDIED. This is a common species in the lower beds of the Simsima Formation and a large number of specimens were available for study. There are 208 specimens in the collections made. Of these the following were mea- sured: BMNH EE4107, EE4116-19, EE4121, EE4129, EE4130, EE4132-34, EE4139-41, EE4144-45, EE4154-57, EE4163, EE4172, EE4174, EE4186, EE4208, EE4217, EE4232, EE4249, EE4251, EE4253-54, EE4257. OCCURRENCE. The species was first described from the late Cretaceous (late Campanian) of Jebel Atra, Tunisia. It has also been described from the Upper Senonian of Derre-i- Chahr and Endjir-kouh, southern Iran (Cotteau & Gauthier 1895) and the late Cretaceous of Palestine (de Loriol 1897) and Turkey (Devries 1967). In the western margins of the central Oman Mountains this species is found at the following localities and levels: Jebel Buhays, section 1: loose in scree (96). Jebel Buhays, section 2: loose in scree (3). Jebel Buhays, section 3: lowest 2 metres (1). Jebel Thanais: lowest couple of metres of section (5). Jebel Agabah: basal shell bed (2). Jebel Faiyah, section 1: arbaciid level ca. 4 m above base (12). Jebel Huwayyah, section 1: beds 14 and 15 (54). Fig. 47 Camera lucida drawings of plating in ‘Globator’ and Conulus. A, B, F-I, ‘G.’ bleicheri (Thomas & Gauthier). A, B, BMNH EE4187; A, adapical ambulacral plating; B, adoral ambulacral plating, peristomial margin at base; F, BMNH EE4186, apical disc; G, BMNH EE4148, apical disc; H, BMNH EE4154, apical disc; I BMNH EE4151, peristomial plating. C-E, Conulus douvillei (Cotteau & Gauthier), apical disc plating: C, BMNH EE4204; D, BMNH EE4277; E, BMNH EE4211. Scale bars = 1 mm. A.B. SMITH Jebel Rawdah, section 1: bed 6 (1); top of bed 4 (15); bed 3 (49); loose (2). Jebel Rawdah, section 2: bed 6 (1); bed 11 (10); bed 13 (1); bed 14 (7); bed 16 (3); bed 19 (8); bed 20 (1); bed 21 (1) | loose in scree just above bed 12 (2); loose in scree, mostly | near base (14). Jebel Rawdah, section 3: bed 1 (11); bed 5 (9); bed 9 (1); bed 11 (1); loose in scree (3). Jebel Rawdah, section 4: bed 2 (9); bed 4 (4); bed 5 (1); bed 8 (3); bed 10 (10); bed 14 (3); bed 20 (1). DIAGNOsIS. An oval, rather depressed ‘Globator’ with a | large, strongly ellipsoidal peristome. The periproct lies high | on the posterior surface and is visible from above but not | | from below. Pore-pairs in weak arcs only towards the peris- | tome, not forming multiple rows. Genital plates 3 and 4] separated by ocular plate IV in adults. DESCRIPTION. Tests range from 9 to 35 mm in length and are | oval in outline and profile. Test width is 82-91% of test length (mean = 86%, SD = 2:6%, N = 32; Fig. 46) with the | widest point on the test coincidental with the posterior portion of the anterior ambulacra. Test height is 61-80% of | test length (mean = 70%, SD = 5:3%, N = 32) and the tallest | point on the test is subcentral. Tests in profile have a |} relatively broad, flat apex and base and a rounded ambitus | — (Pl. 20, figs 3, 8). The apical disc is more or less central and is tetrabasal (Fig. | 47F-H). Genital plate 2 is considerably larger than the other | _ four genital plates and is covered in madrepores. Genital Fi plate 3 is the smallest and in the great majority of specimens . is separated from genital plate 4 by ocular plate 4, which | — abuts genital plate 2. Genital plates 3 and 4 are found in tl contact only in small individuals. The posterior pair of genital | ) plates are in contact posterior to genital plate 2. Ocular plates | | are pentagonal in outline and project. ti Ambulacra are uniserial and pore-pairs are undifferenti- | | ated. Above the ambitus they are very strictly uniserial (Fig. | LATE CAMPANIAN-MAASTRICHTIAN ECHINOIDS 187 sei) OY 2 3 ae S209 SSo 0 £95 ot LATE 20 igs 1-10 ‘Globator’ bleicheri (Peron & Gauthier). 1-4, Topotype specimen of Pyrina orientalis Cotteau & Gauthier, from the Morgan Collection, Museum d’Histoire Naturelle, Paris; 1, oral; 2, apical; 3, lateral; 4, posterior; all x 2. Senonian, Poucht-e-Kouk, Iran. 5, 6, 10, BMNH EE4251; 5, apical; 6, oral; 10, posterior; all x 2. Jebel Buhays, section 1; loose in the scree derived from the lowest 3 m of the | Simsima Formation. 7-9, BMNH EE4208; 7, posterior; 8, lateral; 9, apical; all x 2. Jebel Buhays, section 1; loose in the scree derived from ' the lowest 3 m of the Simsima Formation. igs 11-16 Conulus douvillei (Cotteau & Gauthier). 11, 12, topotype specimen from the Morgan collection, Museum d’Histoire Naturelle, | Paris; 11, oral; 12, lateral; both x 2. Senonian, Khianan, Iran. 13-16, BMNH EE4308; 13, oral; 14, apical; 15, posterior; 16, lateral; all | 2. Jebel Thanais, lowest 2 m of the Simsima Formation. 188 Fig. 48 Camera lucida drawings of adoral pore arrangement. A, ‘Globator’ bleicheri (Thomas & Gauthier), BMNH EE4187; B, Conulus douvillei (Cotteau & Gauthier), BMNH EE4309. Scale bar = 1 mm. 47A), but towards the peristome they become weakly arcuate and reduce in pore-diameter size (Figs 47B, 48A). There are about 88 pore-pairs in a column at 18 mm test length, rising to 127 at 32 mm test length (Fig. 46). Plates are compound in the pyrinoid style, with a single small demiplate in each triad (Figs 47A, B). All plate sutures are denticulate. The periproct is large and tear-drop shaped, pointed adapi- cally (Pl. 20, figs 7, 10). It lies on the posterior surface relatively high on the test, so that it is visible when viewed from above, but not from beiow. Periproct height is 28-46% of the test height (mean = 36%, SD = 4:6%, N = 29) and periproct width is 56-80% of its height (mean = 65%, SD = 5:8% , N = 27). The distance from the base of the periproct to the base of the test is 38-62% of the test height (mean = 49%, SD = 5-6%, N = 31). The peristome is oblique and broadly fusiform in outline, with the long axis running from interambulacrum 3 to ambu- lacrum I (PI. 20, fig. 6). There is hardly any invaginated lip A.B. SMITH | developed around the peristome, although the oral surface ! does curve inwards towards the periproct slightly. | Tuberculation is uniform throughout, with semi-regular and slightly sunken primary tubercles scattered over the) surface, surrounded by a very dense miliary granulation. | There is no internal butressing. REMARKS. This species is easily distinguished from the other — species of Conulidae, Conulus douvillei, that occurs here. C. | douvillei has a periproctal opening that lies close to the base}, of the test, whereas the periproct in ‘G.’ bleicheri lies high on! the posterior and is separated by a considerable distance from the base. This is not a size-related character since there is clear separation of the two species at all sizes (Fig. 46). A’ second difference concerns the development of phyllodes adorally. In ‘G.’ bleicheri the pore-pairs become slightly) arcuate adorally (Fig. 48A), but even in the largest specimens, they never become triserially arranged. Adoral pore-pairs in’ C. douvillei, by contrast, are arranged triserially across much of the oral surface (Fig. 48B). Finally, in the apical disc’ plating of ‘G.’ bleicheri genital plate 2 almost always reaches to ocular IV separating genital plates 3 and 4. In C. douvillei genital plate 2 does not reach ocular plate IV and genital plates 3 and 4 maintain firm contact. : This species has previously been recorded from the Oman) mountain region by Ali (1989) and Smith (in Skelton et al. 1990) under the name Globator orientalis (Cotteau & Gauthier). ‘G.’ orientalis (P1. 20, figs 1-4) was described from) the late Cretaceous of southern Iran by Cotteau & Gauthier (1895). However, it appears virtually indistinguishable in form to ‘Globator’ bleicheri Thomas & Gauthier, from the) late Campanian of Tunisia (Zhagbib-Turki 1987). The only slight difference between these two forms is that “G.’ orienta- lis may have a slightly smaller peristome. For the present,| however, the two species are synonymized. Genus CONULUS Leske, 1778 | Conulus douvillei (Cotteau & Gauthier, 1895) PI. 19, figs 8-11; Pl. 20, figs 11-16; Figs 46, 47C-E, 48B 1895 Echinoconus douvillei Cotteau & Gauthier: 70, pl. 11, figs 9-13. 1932 Pyrina mortenseni Checchia-Rispoli: 21, pl. 2, figs 1-3, le 3h, whys I 2. 21967 Conulus douvillei (Cotteau & Gauthier); Devries: 184, pl. 5, figs 22-25. 1972 Globator mortenseni (Checchia-Rispoli); Kier: 70, figs 35, 36, pl. 44, figs 1-7. 1989 Globator mortenseni (Checchia-Rispoli); Ali: 403, Fig. 4 (6-7). TypPES. The syntypes are the five specimens whose dimen- sions are cited by Cotteau & Gauthier (1895: 70). They may be represented amongst material in either the Cotteau Col-| lection (Lyon) or the Morgan Collection (Museum d’Histoire Naturelle, Paris), but none have been definitely identified. MATERIAL STUDIED. This species is less common than “Glo- bator’ bleicheri, but is never the less well represented in the collections. There are 39 specimens, of which the following were measured: BMNH EE4211, EE4250, EE4277, EE4279-80, EE4283, EE4285, EE4287, EE4291, EE4898-99, EE4301, EE4304-06, EE4308, EE4310. LATE CAMPANIAN-MAASTRICHTIAN ECHINOIDS OCCURRENCE. This species is found in the western Oman Mountains at the following levels: Jebel Huwayyah, section 1: beds 3-5 (1). Jebel Buhays, section 1: loose, derived from lowest few metres of section (19). Jebel Thanais: lowest few metres of the Simsima Formation (11). Jebel Rawdah, section 2: bed 14 (1); bed 15 (2); bed 21 (3); loose, mid-section (1). Jebel Rawdah, section 3: loose (4). Jebel Rawdah, section 4: beds 8/9 (1); bed 13 (1); loose in scree (1). _ Outside Oman, the species is known from Libya, Saudi Arabia, southern Iran and Assam, India. DIAGNOsIS. A species of Conulus with a rounded to strongly fusiform peristome which is not sunken. Periproct situated relatively low on the posterior surface, not visible from above. Pore-pairs adorally arranged triserially. Apical disc with genital plates 3 and 4 always in contact. Test profile subconical. DESCRIPTION. Tests range from 17 to 45 mm in length and are ovoid to rounded pentagonal in outline. Test width is | 82-96% of test length (mean = 88%, SD = 3:2%, N = 18) _ and the widest point coincides with the posterior part of the _ antero-lateral ambulacra. Test height is 64-81% of test length | (mean = 72%, SD = 4.9%, N = 20) and in profile the test has a broad, flat base and is rounded subconical above (PI. 20, figs 11-16). The apical disc lies centrally and is tetrabasal. Genital plate | 2abuts the other three genital plates but never reaches ocular ' plate 4 to separate genital plates 3 and 4 (Figs 47C-E). Genital plates 4 and 1 are always in contact behind the ' madreporite. There appears to be some degree of differentia- tion in gonopore size, suggestive of sexual dimorphism. Ambulacra are straight and compound in the pyrinoid | style. Above the ambitus pore-pairs are strictly uniserial, but / below the ambitus they become offset into three discrete columns and these continue to the peristome edge (PI. 19, fig. / 9; Pl. 20, fig. 11; Fig. 48B). The periproct lies on the posterior border, close to the base (Pl. 20, fig. 15). It is tear-drop shaped, being pointed adapically. Its height is 28-40% of the test height (mean = 35%, SD = 3:2%, N = 20) and its width is 55-77% of its height (mean = 65%, SD = 6:2%, N = 19). The base of the periproct lies 3-17% of the test height above the base (mean = 11%, SD = 3-6%, N = 20). The periproct is just visible from beneath, but is not seen from above (PI. 19, figs 8, 9). The peristome is oval to fusiform in outline and is not \invaginated, although the oral surface may be slightly ‘depressed towards the peristome. Its length is 14-23% of the “test length (mean = 20%, SD = 2:7%, N = 13). Its width is 56-81% of its length (mean = 70%, SD = 8-1%, N = 13). It is oblique, with its long axis orientated from interambulacrum 3 to ambulacrum I (PI. 19, fig. 9). if |REMARKS. The distinction between this species and ‘Globa- tor’ bleicheri is detailed above. This species comes closest to — Conulus giganteus Noetling (C. gigas Cotteau, C. ataxaensis Cotteau) but in this species complex the peristome is less ovoid and is distinctly more invaginated than C. gigas at least. Whether the Middle Eastern species turn out to be suffi- ciently distinct remains to be seen. 189 Order CASSIDULOIDA Claus, 1880 Family CLYPEOLAMPADIDAE Kier, 1962 DIAGNOsIs (following Kier, 1962). Cassiduloids with a domed test and flat base. Petals long and straight, apical disc tetrabasal, periproct inframarginal and transverse, bourrelets with three or more columns of pores and with buccal pores present. TYPE GENUS. Clypeolampas, type ovatus Lamarck, 1816. Goeoes (C- OTHER GENERA INCLUDED. Hungaresia Szorenyi, type spe- cies Hungaresia hungarica Szorenyi (junior synonym of Cly- peolampas ovum Grateloup); Vologesia Cotteau & Gauthier, type species V. tataosi Cotteau & Gauthier. REMARKS. Lambert (1919) claimed that the original figure and description of Clypeolampas ovatus Lamarck (1816: 22) was inadequate for certain identification, and thus used the name Clypeolampas leskei Goldfuss (1829) as the oldest available name. However, Kier (1962: 190) accepted C. ovatus Lamarck as a valid designation and sunk C. leskei Goldfuss in synonymy. There are eighteen nominal species assigned to the three genera listed above. The differentiation of the three genera is, however, unclear. Vologesia was established by Cotteau & Gauthier (1895: 65) for a late Cretaceous species from Aftab, Iran, V. tataosi Cotteau & Gauthier. This is based on a single small individual, subcircular in plan view, with poorly devel- oped bourrelets and the peristome positioned close to the anterior border. No other species were included in this genus until Lambert (1919) revised the then known members of Clypeolampas. He separated species into two groups; those with uniform aboral tuberculation, all tubercles being scro- biculate, and those forms which had a second kind of aboral tuberculation composed of nonscrobiculate pustules. The former he assigned to Vologesia, the latter to Clypeolampas. Lambert (1919) placed the following species in Vologesia: C. ovum Grateloup, C. acuta Desmoulins, C. conica Arnaud, C. toucasit Lambert, C. gossauviensis Lambert and a small undescribed Maastrichtian form. In Clypeolampas Lambert placed C. leskei Goldfuss [=C. ovatus Lamarck], C. perova- lis, Arnaud, C. orbicularis Arnaud, C. lestelei Cotteau, C. vishnu, Noetling, C. douvillei Lambert and C. mengaudi Lambert. In 1955, Szorenyi erected the genus Hungaresia for the new Santonian species H. hungarica Szorenyi of Hungary. This species appears to be identical in many important respects to the common Santonian species Clypeolampas ovum Grate- loup, and Kier (1962: 191) synonymized the two genera, making Hungaresia a junior synonym of Vologesia. Kier (1962, 1966) followed Lambert’s generic differentia- tion, distinguishing Clypeolampas from Vologesia by its non- scrobiculate pustules developed adapically, and by its better developed floscelle. However, Kier’s concept of Vologesia was based not on the type species but on Clypeolampas ovum Grateloup and is thus misleading. There are very few stable characters on which to subdivide the group, the following being amongst the most informative. (1) Aboral pustules developed. In some species there are characteristic pustules over the adapical surface that are slightly larger than the normal tuberculation and give the surface a rugose appearance. These are not tubercles for spine articulation, as they have no articular surface or sur- 190 rounding scrobicule. Instead they resemble the pustular cal- cite formed in species of Conulus or Echinoneus (Smith 1980). Their function is unknown but it may be to do with deterring parasitic and commensal settlement. Pustules are developed in a number of species, including the type C. ovatus Lamarck. (2) The position of the peristome seems highly stable and distinctive. In Vologesia tataosi and V. rawdahensis the peristome lies close to the anterior border, with the anterior edge lying between 20 and 25% of the test length from the anterior. In almost all other species the peristome is subcen- tral, lying between 30 and 40% of the test length from the anterior. Only one species, Clypeolampas toucasi Lambert is intermediate in this respect, with its peristome between 25 and 30% of test length from the anterior. (3) Elongation of the peristome. Only Vologesia rawdahen- sis Ali has such a laterally elongate peristome. In other species the peristome is suboval. (4) The degree to which the floscelle projects as prongs over the peristome is to some extent size dependent, with more prominent floscelles in larger individuals. However, at a similar size it is apparent that Vologesia tataosi, V. rawda- hensis and Clypeolampas toucasi show virtually no floscelle development, whereas C. ovatus and its synonyms and C. perovalis have very pronounced floscelle projection. In C. lestelei, C. ovum, and C. conicus as well as probably C. helios, the floscelle is slightly swollen but not projecting. (5) The periproct is usually unambiguously positioned on the flat oral surface, but in a few species such as C. helios and C. ovum, the periproct lies subambitally because of the strongly inflated test profile. (6) The apical disc is definitely tetrabasal in C. /estelei, C. ovum and Vologesia rawdahensis, but monobasal in C. ova- tus. (7) The arrangement of pores in the bourrelets, though to some extent size dependent, offers some differentiation. In C. ovatus there are many pores irregularly scattered between inner and outer columns in each half ambulacrum. By con- trast C, ovum and Hungaresia hungarica have only the inner and outer series of pores. If C. conicus is just a variety of C. ovum as is suspected, then some specimens may have a few pores forming a mid row. The same is true of Vologesia rawdahensis where individuals have either two or three rows. From this the following supraspecific taxonomy is pro- posed: Clypeolampas Lamarck: Large forms with moderate to well developed bourrelets, subcentral peristome and aboral calcite pustules. Species included; C. ovatus Lamarck (includes C. leskei Goldfuss, C. mengaudi Lambert, C. douvillei Lambert, C. orbicularis Arnaud), Upper Campanian-Maastrichtian of Spain, southern France, Tur- key; C. perovalis Arnaud, Lower and Middle Campanian of Gironde, France; ?C. lestelei Cotteau, ?Danian of Saint Cirac, Ariége, France. Vologesia Cotteau and Gauthier. Distinguished from Cly- peolampas by its anterior peristome without floscelle devel- opment. Type species V. fataosi Cotteau & Gauthier, Upper Senonian of Louristan, Iran; V. rawdahensis Ali, Maastrichtian of the Oman Mountains; V. toucasi (Lam- bert), Campanian of the Pyrenees, France. Hungaresia Szorenyi. Smaller, ovoid forms with subcentral, pentagonal peristome with swollen but not projecting flos- celles, no aboral pustules and subambital rather than fully adoral periproct. Type species, H. ovum (Grateloup) A.B. SMITH [includes H. hungarica Szorenyi], Upper Santonian of France, Pyrenees, Hungary. Other species included: H. helios (Noetling), ?Maastrichtian, Mari Hills, Baluchistan. Unplaced taxa: Clypeolampas conicus Arnaud (U. Santonian|_ to L. Campanian, SW France) and C. acuta Desmoulins (pores in bourrelets shown as forming two columns only). material of these species has not been seen and so they cannot be placed with any confidence from the published! descriptions and figures. C. vishnu Noetling: based on a single worn specimen, inad- equately known nomen dubium. C. gossaviensis Lambert: based on a single, poorly preserved specimen and indeterminate from description and figure} nomen dubium. Genus VOLOGESIA Cotteau & Gauthier, 1895 TYPE SPECIES. Vologesia tataosi Cotteau & Gauthier, 1895, i by original designation. OCCURRENCE. Late Cretaceous (‘Upper Senonian’) of Iran; Maastrichtian of the United Arab Emirates and Oman. DIAGNOSIS. Clypeolampadids with an anteriorly positioned! peristome lying 20-30% from the anterior border. Peristome wide, pentagonal with bourrelets hardly developed. Phyl- lodes with two or three rows of pores in each half ambu lacrum. Apical disc tetrabasal. - REMARKS. Vologesia is distinguished from Clypeolampas b its lack of aboral pustules and its poorly developed floscelles _ and bourrelets. It is distinguished from Hungaresia by its lac of floscelles, its more transverse peristome and its mor anterior peristome. Pl. 21, figs 1-5; Figs Vologesia rawdahensis Ali, 1989 : 49-5]. 1989 Vologesia rawdahensis Ali: 406, fig. 5 (13). TyPEs. Three specimens in the Geology Museum, United Arab Emirates University, Al Ain, United Arab Emirates. MATERIAL STUDIED. Seven specimens, of which fiv (BMNH EE3383-85, EE4326, EE4329) are well enoug preserved to be included in the biometric analysis. OCCURRENCE. This species is known only from the Simsim Formation of the western margins of the Oman Mountains. Specimens were found at the following localities and hori zons: Jebel Buhays, section 1: loose in the scree, derived from th lowest few metres of Simsima Formation (5). Jebel Buhays, section 2: loose in the scree, derived from th lowest 2 m of Simsima Formation (1). Jebel Thanais: lowest 2 m of Simsima Formation (1). Jebel Rawdah, section 2: bed 19 (1). Jebel Rawdah, section 3: bed 8 (1). DESCRIPTION. Tests are flat-based and rounded to subconi cal in profile (Pl. 21, figs 1-4), with a relatively sharp ambitu situated low down. In outline the test is ovoid with a rounde anterior and a distinctly more pointed posterior. Tests rang in length from 46 to 68 mm. Test width is 75-77% of tes length and test height 53-62% of test length (Fig. 49). The tallest part of the test is central or slightly anterior of centre. LATE CAMPANIAN-MAASTRICHTIAN ECHINOIDS 60 @ Test width (mm) @ Test height (mm) 50 60 70 Test length (mm) 30 @ to apical disc @ to peristome Distance from anterior (mm) N ° = v2 °o 60 70 Test diameter (mm) Length of petal (mm) 60 Test length (mm) Fig. 49 Biometric data for Vologesia rawdahensis Ali. The apical disc is tetrabasal (Fig. 51B), with the madreporite very large and occupying the centre. Other genital plates are much smaller and pushed far out into the adjacent interambulacra. The apical disc lies 37-39% of test length from the anterior border (Fig. 49). Petals are relatively long and straight or only very slightly powed. They are open distally and extend most of the listance towards the ambitus. The posterior petals are always blightly longer than the anterior three petals. They are pomposed of an inner circular pore and a highly elongate buter slit-like pore, joined by a furrow. There are 51 pores in 1 petal column in a 53 mm individual and 57 in a 68 mm individual. Pores below the petal are all single. The peristome is pentagonal in outline and very much vider than long (width is 1-7-2-0 times greater than length). 191 It is straight sided and only slightly invaginated (PI. 21, fig. 5; Fig. 50) and bourrelets are hardly developed. It lies 21-27% of the test length from the anterior border. Phyllodes are relatively short and are not depressed in the slightest. In some individuals there are only two columns of pores in each half ambulacrum, whereas in others there are three columns. The outer series is composed of about 12 pores, the inner series of 7 or 8 pores and the mid series, where present, of 4 or 5 pores (Fig. 51). The inner and middle series of pores are borne on occluded plates. Buccal pores are present. The periproct lies on the oral surface at the posterior. It is oval in outline, approximately twice as wide as long and approximately the same size as the peristome. Aboral tuberculation is fine and uniform, oral tubercula- tion slightly coarser and becoming less dense towards the midline. There is a broad tubercle-free band down the midline in the posterior interambulacrum running between the peristome and periproct. This is lightly pitted (PI. 21, fig. 5). REMARKS. This species was erected by Ali (1989) on the basis of three specimens from Jebel Rawdah, Oman. It differs from V. tataosi Cotteau & Gauthier, from a similar horizon in southern Iran, by being more elongate and pointed posteri- orly, and by having a smaller, more transversely elongate mouth (if the original figures of this species are true to life). The Iran species was, however, based on a single individual 27 mm in test length, and there is the possibility that with more material the two species may eventually prove to be synonymous. Family FAUJASIIDAE Lambert, 1905 Genus FAUJASIA d’Orbigny, 1856 TYPE SPECIES. Pygurus apicalis Desor, by subsequent desig- nation of Lambert & Thiery, 1921: 273. OTHER SPECIES INCLUDED. Only one other species, Faujasia eccentripora Lees. Two species previously ascribed to Fauja- sia were transfered to other genera by Kier (1962): F. faujasi (Desmoulins) to Eurypetaium and F. chelonium Cooke to Domechinus. DIAGNOSIS. Small ovoid cassiduloids with a monobasal api- cal disc in which the genital pores open through interambu- lacral plates. Petals broad, closed distally and strongly petaloid in form. Periproct small, circular and inframarginal. Peristome small, anterior with short, arcuate phyllodes in two columns and with buccal pores, and strongly projecting bourrelets. OCCURRENCE. Maastrichtian of Belgium, France, Oman and the United Arab Emirates. REMARKS. Kier (1962: 137), in discussing this genus, was uncertain whether F. eccentripora Lees truly belonged here, since its apical disc plating had never been described. As shown below, this species has an identical arrangement of gonopores opening within the adapical portion of the inter- ambulacra as characterizes F. apicalis Desor and thus is clearly closely related. The derived position of gonopores outside apical disc plating distinguishes these two species from all other cassiduloids. | | 192 A.B. SMITH | “f o oo "<= yp : ay i = Hips, Br. Z Z ; 1 Ba a ai ~. Ps ZZ, 5 2 MMi STATA ~ Pa i” we WWYE™ oat i \\\ SS x Uh Fig. 61 oral. Scale bar = 1 cm. three. The anterior petal is also typically narrower and more parallel-sided than the other petals. The two pores are subcircular and widely separated, united by a well developed groove. The petals converge slightly distally but do not close (Fig. 61). The interporal zone is about twice the width of a single pore zone. Pores below the petals are all single. The peristome is quinquestellate in outline with the promi- nent bourrelets projecting into the opening (PI. 23, fig. 7). The bourrelets are triangular and have long lateral bands of fine tuberculation. They are blunt-ended. The phyllodes are distinctly sunken and arcuate. They consist of many rows of pores (Fig. 63). There is a well defined outer series and inner series in each half ambulacrum, but the central zone consists of a broad band of unorganised pores, four or five abreast. The anterior ambulacrum, however, has notably fewer median pores than other ambulacra, with only one or two median pores abreast. There are 16-18 pores in the outer series and 7 or 8 in the inner series. There are many occluded plates in the phyllodes (Fig. 63). The peristome is 1-4 to 2-1 times as wide as it is long and is situated 33-43% of test length from the anterior border (mean = 39%, N = 13). The peristome is small and lies inframarginally, typically forming a small pointed rostrum. There is a distinct, well- formed invagination of the test on the adoral margin of the periproct but not adambitally. The periproct is oval and longer than wide (width = 45-70% of length). It opens between interambulacral plates 6 and 8 (Fig. 61). Tuberculation is fine and dense aborally, slightly less fine and less dense adorally. There is a very pronounced fusiform A.B. SMITH Camera lucida drawings of plating in Pygurostoma morgani Cotteau & Gauthier. A, BMNH EE3303, apical; B, BMNH EE3304, naked zone along the sternum between the peristome and periproct. This is slightly raised above the surrounding test and is covered in small pits. There is a similar, smaller naked zone developed anterior of the peristome in ambulacrum III. REMARKS. The species was described and figured by Cotteau & Gauthier (1895). In the same work they described a number of smaller species of Parapygus under the name Pseudocatopygus. Some of these may possibly turn out to be juveniles of P. morgani. This species is very different from the only other possible species assigned to this genus, P. pasionensis Cooke (1949), from the ?Campanian of Guatamala. P. pasionensis has much less well developed bourrelets and its periproct is more equant and lies completely flush and is not invaginated. As discussed above, it is not clear that P. pasionensis is conge- neric with P. morgani. Family CASSIDULIDAE Agassiz & Desor, 1847 Genus PETALOBRISSUS Lambert, 1916 Petalobrissus rawdahensis sp. nov. Pl. 24, figs 1-12; Figs 64, 65 Types. Holotype, BMNH EE3485; paratypes, BMNH EE3467-84, EE3486-87, EE4321-22. OTHER MATERIAL. Over 650 specimens were collected. { _ LATE CAMPANIAN-MAASTRICHTIAN ECHINOIDS Fig. 62 Camera lucida drawing of apical disc plating in Pygurostoma morgani Cotteau & Gauthier, BMNH EE3288. Scale bar = 1 mm. ‘Fig. 63 Camera lucida drawing of phyllode plating in Pygurostoma morgani Cotteau & Gauthier, BMNH EE3304. Scale bar = 5 mm. ‘OCCURRENCE. This species occurs abundantly at Jebel Raw- ‘dah, section 2 but is found virtually nowhere else. The only ‘other occurrence of this species is in the lowest bed (bed 1) at Jebel Rawdah, section 3b. At section 2 the species is found at ithe following levels: bed 4 (3); bed 8 (4); bed 11 (247); bed 12/13 (230); bed 14 (17); bed 15 (25); bed 19 (73); beds 20-21 (45); bed 22 (13); bed 26 (2). Additional material was collected loose from the scree at this locality. DiaGNosis. A Petalobrissus with a monobasal apical disc and a small quinquestellate peristome, anterior in position and surrounded by small pointed bourrelets. Phyllodes well- 203 developed, composed of outer and inner series of pores. Buccal pores present. Periproct posterior, longitudinal. Nar- row, smooth tubercle-free zone present both anterior and posterior to the peristome. In outline the posterior projects slightly and adorally there is a characteristic interradial keel in the posterior interambulacrum. DESCRIPTION. Tests are oval in outline, uniformly rounded at the anterior, but slightly pointed posteriorly (Pl. 24, figs 5, 6). Test length ranges from 10-3 to 21-2 mm. Test width is 88-95% of test length (mean = 91%, SD = 1:8%, N = 23) and the widest point is slightly posterior of midlength. In profile the test is depressed with a relatively flat upper surface, a uniformly rounded anterior and an obliquely truncated posterior (PI. 24, figs 4, 7, 10, 11). The tallest point coincides with the apical disc or is slightly posterior to it. The ambitus is rounded and relatively low. Test height is 51-62% of test length (mean = 56%, SD = 2-4%, N = 23; Fig. 64). The oral surface is flat anteriorly, but slopes away towards the posterior. There is a relatively sharp and well-defined posterior keel on the oral surface along the midline and two less sharply defined keels in the two postero-lateral interam- bulacra (PI. 24, figs 5, 8). The apical disc lies 37-46% of test length from the anterior border (mean = 43%, SD = 2:5%, N = 23). It is monocyclic, with four gonopores that open at around 10-13 mm test length. The gonopores are oval and there is no sign of any suture separating them from the central area of madrepores (Fig. 65C). Ocular plates are small and subtriangular in outline. Petals are bowed and converge distally, though remaining open (Fig. 65A). The anterior petal is the longest and the least bowed. It has about 20 pore-pairs in a column at test length of 10 mm, rising to 39 at 21 mm test length. The perradial portion of the ambulacrum is about 1-5 times the width of a single pore zone. Lateral and posterior petals are 60-86% of the anterior petal in length and are more strongly bowed. In all petals the two columns are of equal length. The posterior petals end well short of the periproct. All pores below the petals are single. The phyllodes are well developed and strongly bowed but are only slightly sunken (PI. 24, figs 1, 5, 12). The first ambulacral plates are elongate and bootshaped in outline, with small buccal pores (Fig. 65D). There is an outer series of closely packed pores and a shorter inner series of equally closely spaced pores (PI. 24, fig. 12; Fig. 65D). There are 10 or 11 pores in the outer series of lateral and posterior phyllodes, and 9 or 10 in ambulacrum III. There are 4 or 5 pores in the inner series in all ambulacra. Despite the large number of individuals avail- able, none show the detailed plating of the phyllode region adequately. The outer series of pores lie on a series of narrow plates that do not reach the perradius. Internally there is a second series of plates housing the inner series of pores. These may be oblique, but in no specimen is the plating in this region clear. The peristome is pentagonal, slightly broader than long in larger individuals, and situated 33-39% test length from the anterior border (mean = 36%, SD = 1:5%, N = 23). The peristome width is 7-12% of test length and its length is 85-100% of its width. The peristome is sunken with well developed vertical walls. The surrounding interambulacral areas are developed into short, projecting, knob-like bourre- lets, which do not, however, impinge on the peristome (PI. 24, fig. 12). 204 A.B. SMITH PLATE 24 Figs 1-12 Petalobrissus rawdahensis sp. nov. 1-4, 12, BMNH EE4321, holotype; 1, oral; 2, apical; 3, posterior; 4, lateral; all x 3; 12, detail | of peristomal region, x 6. Jebel Rawdah, section 2, loose in scree at level of bed 11. 5-7, BMNH EE3485, paratype; 5, oral; 6, apical;7, | lateral; all x 3. Jebel Rawdah, section 2, bed 11. 8-11, BMNH EE4322, paratype; 8, oral; 9, apical; 10, lateral; 11, posterior; all x 3. Jebel | Rawdah, section 2, loose in scree at level of bed 11. | LATE CAMPANIAN-MAASTRICHTIAN ECHINOIDS 30 © P. rawdahenss: to apical disc a E @ P. rawdahenss: to periproct £ @ P. cf. setifensis: to apical disc - © P. cf. setifensis: to periproct _ fe} @ 20 i © cd s oa rs ° © mR . Se 1@) oe [ao] oO © o La Pe ad = s ed = e a ac) 10 20 30 Test length (mm) 14 @ P. rawdahensis © P. cf. setifensis = = i) (-) co [o} N To peristome from anterior (mm) bd 20 30 Test length (mm) + fo) @ P. rawdahensis o @ P. cf. setifensis Ww (“al Ww [o) Number of pore-pairs in column N on 4 6 8 Length of petal in ambulacrum Ill © Number of pore-pairs in petal 205 © P. rawdahensis: test width @ P. rawdahensis: test height @ P. cf. setifensis: test width © P. cf. setifensis: test height 10 20 30 40 Test diameter (mm) es P. rawdahensis @ P. cf. setifensis Test length (mm) 40 @ P. rawdahensis @ P. cf. setifensis 3 4 5 6 7 8 Length of petal in ambulacrum Il Fig. 64 Biometric data for Petalobrissus rawdahensis sp. nov. and P. cf. setifensis (Cotteau). The periproct is clearly visible from above and opens 71-87% test length from the anterior border (mean = 82%, SD = 2:7%, N = 23). It is longitudinal, with a width that is 41-65% of its height. The opening is near vertical, with slightly invaginated walls forming a short, parallel-sided anal sulcus. There is a subanal rostrum (PI. 24, figs 2, 6, 9-11). Tuberculation is fine and uniform aborally, slightly coarser adorally. On the oral surface there is a narrow band free of tubercles down the midline of the posterior interambulacrum. This is finely granular and tapers towards the posterior. ‘REMARKS. This species is problematic to place on account of its monobasal apical disc. In general shape and plating it conforms closely to species of Petalobrissus, having very ‘similar phyllodes and bourrelets, a similar longitudinal | /Periproct set far back on the test, and only a remnant anal | sulcus. However, the type species of Petalobrissus has a tetrabasal apical disc according to Kier (1962), whereas P. rawdahensis has a monobasal disc. P. rawdahensis also bears a strong resemblance to small Hardouinia species, especially in the characteristic keeled oral surface, bowed petals and well developed bourrelets. However, in the type species of Hardouinia, H. mortonis, the phyllodes are more strongly arcuate with the inner series arranged distally as an integral part of the arc. Furthermore, the peristome opens subcen- trally. In the type species of Procassidulus, P. lapiscancri (Goldfuss) from the Maastrichtian of Maastricht, the phyl- lodes are less arcuate and the inner series is parallel to the outer series of pores. P. rawdahensis also differs from P. lapiscancri (Goldfuss) in being larger, more rounded in profile and oval in outline, and in having more pores in its 206 A.B. SMITH Fig. 65 Camera lucida drawings of plating in Petalobrissus rawdahensis sp. nov. A, apical surface, BMNH EE3485; B, oral surface, BMNH EE3487; C, apical disc, BMNH EE3476; D, One phyllode, peristome margin at base, BMNH EE3484. Scale bars = 1 mm. phyllodes at comparable sizes. Pending revision of this group, P. rawdahensis is placed in the genus Petalobrissus. Petalobrissus rawdahensis is easily distinguished from Peta- lobrissus cf. setifensis (Peron & Gauthier), which occurs in the same section, by its more angular outline, its posterior keel and slope on the oral surface, its smaller, less quinquelo- bate peristome and its more bowed and densely pored petals. Petalobrissus cf. setifensis (Cotteau, 1866) al, 25); figs 1-10; Figs 64, 66 cf. 1866 Echinobrissus setifensis Cotteau: 267, pl. 14, figs 13-15. cf. 1962 Petalobrissus setifensis (Cotteau); Kier: 125, pl. 16, figs 10-13. 1989 Petalobrissus inflatus Gauthier & Thomas; Ali: 405, fig. 5 (9). MATERIAL STUDIED. Thirty one specimens were used for the biometric analysis: BMNH EE3505-10, EE3512-28, EE3530, EE3532, EE3535, EE3537-41. An additional 269 specimens were collected. OCCURRENCE. Apart from five specimens collected from the scree at Jebel Buhays, section 1, and a single specimen from bed 9 at Jebel Bu Milh, all the material comes from sections at Jebel Rawdah. The distribution of specimens is as follows: Jebel Rawdah, section 1: bed 3 (18). Jebel Rawdah, section 2: bed 8 (9); bed 11 (124); beds 12/13 (4); bed 14 (15); bed 19 (23); bed 21 (44); bed 26 (3). Jebel Rawdah, section 3b: bed 2 (4). Jebel Rawdah, section 4: bed 4 (1); beds 8/9 (3). DESCRIPTION. Tests are oval in outline, uniformly rounded at the anterior and slightly flattened at the posterior (PI. 25, figs 1, 2). Test length ranges from 10 to 33 mm. Test width is 82-93% of test length (mean = 87%, SD = 2-:9%, N = 3}), with the widest point slightly posterior of midlength. Test height is 49-61% of test length (mean = 54%, SD = 3-1%,N = 31), with the highest point at or a little posterior of the apical disc. In profile the anterior is uniformly rounded, the posterior obliquely truncated (PI. 25, figs 4, 10). The ambitus is rounded and about one-quarter of the test height above the base. The oral surface is flat, with a very slight depression towards the peristome. The apical disc lies 37-48% test length from the anterior border (mean = 42%, SD = 2-4%, N = 31) being relatively more anterior in larger individuals (Fig. 64). It is monocyclic with no evidence of any sutures between the gonopores and central madreporite region in any specimen, even where ocular plate sutures are clear (Fig. 66D). However, the LATE CAMPANIAN-MAASTRICHTIAN ECHINOIDS 207 PLATE 25 Figs 1-10 Petalobrissus cf. setifensis (Cotteau). 1, 10, BMNH EE4341; 1, oral; 10, lateral; both x 3 Jebel Buhays, section 1; loose in the _ scree derived from the lowest 3 m of the Simsima Formation. 2-5, BMNH EE3519; 2, apical; 3, oral; 4, lateral; 5, posterior; all x 3. Jebel Rawdah, section 2, bed 11. 6-9, BMNH EE3536; 6, apical; 7, oral; 8, lateral; 9, posterior; all x 3. Jebel Rawdah, section 2, bed 19. Figs 11,12 Arnaudaster cylindriformis sp. nov. BMNH EE4324, depressed variety; 11, oral; 12, apical; both x 2 (see also Pl. 29, Figs 6, 9). Jebel Rawdah, section 2, in scree at level of bed 14. 208 A.B. SMITH Fig. 66 Camera lucida drawings of plating in Petalobrissus cf. setifensis (Cotteau). A, apical surface, BMNH EE3514; B, oral surface, BMNH EB3535; C, phyllode plating, BMNH EE3524; D, apical disc, BMNH EE3524; E, single phyllode, peristome margin to base, BMNH EE3535. Scale bars: A, B = 2 mm, C-E = 1 mm. madrepores do not usually extend up to the gonopores. Gonopore diameter varies markedly between individuals of the same test length, indicating sexual dimorphism in this species. Gonopores appear at around 10-12 mm test length. Petals are lanceolate, with almost straight lines of inner pores and only slightly bowed lines of outer pores in all petals (Pl. 25, fig. 2; Fig. 66A). All are distally open and both inner and outer pores are subcircular in outline. The anterior petal is the longest and has about 22 pore-pairs in a column at 15 mm test length, rising to 39 at 33 mm test length. Petals in ambulacra II and IV are 65-93% of the length of the anterior petal (mean = 80%, SD = 6:6%, N = 28), whereas the posterior petals are 73-100% of the length of the anterior petal (mean = 87%, SD = 6:6%, N = 28). The posterior LATE CAMPANIAN-MAASTRICHTIAN ECHINOIDS petals end well short of the periproct. All pores below the petals are single. The phyllodes are only slightly expanded and are flush with the surrounding test (Pl. 25, figs 1, 3). Irrespective of size there are 7 or 8 pores forming an outer series and 3 or 4 pores forming an inner series in the phyllodes (Figs 66C, E). The first ambulacral plates are boot-shaped and carry small buccal pores, but are not particularly elongate. The inner series of pores are found on small occluded plates which are separated from one another (i.e. they do not form a continuous inner series of plates). Each is separated from its neighbours by two of the outer plates. Sphaeridial pits are confined to the area between the most adoral of the inner series of pores and the buccal pores. There are just two or three in each column. The peristome is quinquelobate, and relatively large (Fig. 66C). Its length is 9-16% of the test length (mean = 11%, SD = 1-6%, N = 29) and 81-120% of its width. There is a distinct vertical-walled well to the peristome, covered in fine miliary tubercules. Bourrelets are not developed. However, the interambulacral margins to the peristomial well are elevated slightly, forming a lip-like rim (PI. 25, fig. 1). The periproct is longitudinal and clearly visible from above (Pl. 25, figs 2, 6). It lies 76-92% of test length from the anterior border (mean = 85%, SD = 3:5%, N = 31). It is about twice as tall as wide (mean width = 48% of height). There is a short parallel-sided anal sulcus that stops short of the ambitus. All tubercles are sunken, those adorally being noticeably larger than the aboral ones. There is a median naked zone on | the oral surface in both the anterior ambulacrum and the posterior interambulacrum (PI. 25, figs 1, 3, 7). The anterior zone is short and does not reach the ambitus, but the posterior zone is broader, approximately parallel-sided and reaches the posterior border. It is finely granular. REMARKS. This species could at first glance be confused with Petalobrissus rawdahensis, but is distinguished on several counts. It has a larger, more quinquelobate peristome, a flatter base without the posterior keel, it lacks bourrelets and has less inflated phyllodes with fewer pores. It has more parallel-sided petals with less densely packed pore-pairs. It _ resembles certain other species of Petalobrissus in its overall | shape and form, but is distinguished by its monobasal apical disc. In particular it most closely resembles P. cubensis (Weisbord), particularly in its floscelle structure. However, it | differs from P. cubensis in having a narrower posterior naked | zone on its oral surface, in having its apical disc less anterior and in having a proportionally longer anterior petal. It also comes close to certain species currently assigned to Rhynchopygus in test shape and floscelle structure, but differs in having a longitudinal rather than a transverse | peristome. It is closest to P. setifensis (Peron & Gauthier), from the Maastrichtian of Algeria, but differs in some minor details, having a flatter base and more quadrate outline with | less rounded ambitus, and in having fewer inner-series pores. __ This species was identified by Ali (1989) as Petalobrissus _inflatus (Gauthier & Thomas), but P. inflatus is a Cenoma- | nian species that differs in several respects, notably in having “much longer and more densely pored petals at equivalent ' sizes. These petals are strongly bowed in P. inflatus and the posterior pair extend back to a point level with the opening of the periproct. Cassidulus oldhamianus Stolizcka, from the Maastrichtian Arrialoor Group of Southern India may be comparable, and 209 appears similar in overall form. However, that species is inadequately described and figured and no examples of it are available for comparison. At present it is impossible to say what genus the species belongs to. Petalobrissus_ linguiformis (Peron & Gauthier, in Cotteau, Peron & Gauthier, 1881) Pl. 23, figs 5, 6; Pl. 26, figs 1-6; Figs 67, 68 Types. Syntypes are the two measured specimens in the Peron & Gauthier collections referred to in Cotteau ef al. (1881: 162). MATERIAL STUDIED. Ten specimens, BMNH EE3334-35, EE3337, EE3341-45, EE4319, EE5021, were measured for the biometric analysis. An additional 11 specimens were also collected. OCCURRENCE. This species was originally described from the Maastrichtian of Algeria, and has since been reported from Tunisia and Egypt. Apart from one specimen collected loose at Jebel Buhays, section 3, all specimens come from Jebel Rawdah. It is found at the following localities and horizons: Jebel Buhays, section 1: loose in the scree, derived from the lowest few metres of the Simsima Formation (1). Jebel Rawdah, section 2: bed 14 (4); bed 19 (5); bed 20 (1); bed 21 (4); loose from upper part of section (beds 14-21) ©) he Jebel Rawdah, section 4: loose in scree (1). DIAGNosiIs. An elongate Petalobrissus with a narrow, almost vertical periproct and short anal sulcus set very far towards the posterior on the upper surface. Phyllodes are very strongly developed, with prominent bourrelets and outer and inner series of pores forming well-defined lines. There are up to 20 pores in the outer series and 10 in the inner series. In addition there are well developed rows of sphaeridial pits down the perradius. DESCRIPTION. Tests are ovoid in outline, uniformly rounded at the anterior and slightly more pointed towards the poste- rior, with a slight, but distinct, anal cleft (PI. 26, figs 1, 2). Tests range in size from 13 to 50 mm in length. The widest point on the test lies about two-thirds the distance from the anterior border. Test width is 76-87% of test length (mean = 82%, SD = 46%, N = 10). In profile the test is depressed, with a height that is 43-48% of test length (mean = 46%, SD = 1-8%, N = 9). The upper surface is slightly domed with the tallest point slightly posterior to the apical disc. The ambitus is uniformly rounded and lies at about one-third test height. The posterior is more or less truncated (PI. 23, fig. 6). The apical disc lies 34-44% test length from the anterior border (mean = 40%, SD = 3-0%, N = 7). It is tetrabasal, with a large madreporite plate and three small genital plates projecting into the interambulacra (Fig. 68D). These genital plates are U-shaped because of the gonopore that opens along their outer margin. Genital plates lie separated from adjacent ocular plates. Petals are well developed (PI. 26, fig. 5: Fig. 68B). The anterior and posterior pair of petals are both rather straight and remain open distally, whereas the antero-lateral pair of petals are more bowed and converge distally (PI. 23, fig. 5). The perradial zone is about twice the width of a pore zone. The anterior petal is noticeably longer than other petals and extends more than three-quarters of the distance to the ambitus. Petals II and IV are 61-74% of the length of the 210 A.B. SMITH PLATE 26 Figs 1-6 Petalobrissus linguiformis (Peron & Gauthier). 1, 6, BMNH EE4319; 1, oral, x 3; 6, detail of peristome, x 6. Jebel Rawdah, section 2, beds 19/20. 2-5, BMNH EE3345; 2, oral; 3, lateral; 4, posterior; 5, apical; all x 2. Jebel Rawdah, section 2, loose, derived from beds 14—21. LATE CAMPANIAN-MAASTRICHTIAN ECHINOIDS 40 = Test width Py @ Test height a (mm) 20 30 Test length (mm) 40 Length of petals (mm) 50 Length (mm) Distance (mm) 211 ®B Peristome * Bourellet e 20 30 40 Test length (mm) 50 @ Anterior to peristome @ Anterior to apical disc = jo) (0) 30 40 Bye) 10 20 30 40 SO Test length (mm) Test length (mm) Fig. 67 Biometric data for Petalobrissus linguiformis (Peron & Gauthier). anterior petal, wheras petals I and V are 71-89% of the length of the anterior petal. The posterior petal extends less than two-thirds the distance from the apical disc to the periproct. All pores below the petals are single. The phyllodes are well developed and bow outwards strongly towards the peristome. They become slightly sunken towards the peris- tome, with the buccal pores situated in a shallow depression separated from the peristomial well (PI. 26, fig. 6). There are two series of pores, well separated from each other and with pores closely packed together in each row. The outer row comprises some 17 to 20 pores, the inner series 7 to 9 (Fig. 68C). The first ambulacral plates are squat and broad, with buccal pores situated near their adambital edge and well separated from the peristome. Pores in the outer series are bowed and every third plate is smaller and does not reach the inner series of plates. These pores may be slightly offset, marking the incipient development of a median series of pores. The inner series of pores are situated towards the outer edge of a double column of occluded plates. Each plate carries a sphaeridial pit on its perradial side, there being some 8-10 sphaeridial pits forming a well defined column on either side of the perradial suture. The peristome is subcircular to pentagonal and is approxi- mately as broad as it is wide. It is 69% of test length in | diameter and lies 37-42% of test length from the anterior | border. There are short, pointed bourrelets which project outwards but do not impinge on the peristome (PI. 26, fig. 6). The periproct lies close to the posterior border, and is clearly visible from above (Pl. 26, fig. 5). It is longitudinal, being 27-40% as wide as it is tall (Pl. 26, fig. 4). It is slightly V-shaped, narrowing adapically, and opens into a short anal sulcus with parallel-sided walls. The periproct is almost vertical in orientation. Tubercles are minute and densely packed aborally, but coarser and less dense adorally. There is a relatively broad naked zone in the posterior interambulacrum that runs from the peristome to the posterior border. This is 13-18% of the test width in width. It is covered in a reticulate pattern of pits. REMARKS. This species is readily differentiated from other cassiduloids described here by its highly developed phyllodes, its almost posterior periproct and its characteristic shape. The only species that come close are the southern Indian P. testudo (Forbes) and P. emys (Stoliczka), both from the Maastrichtian. Both have a similar test shape, with the posterior slit-like periproct, the anal notch and the projecting bourrelets. P. testudo has much less developed phyllodes with few pores in the inner series and only 8-10 pores in its outer series. P. emys is too poorly known to be compared closely as the original figures and description omit many important details such as phyllode structure. It may eventually prove to be synonymous with P. testudo. ANY i A.B. SMITH Fig. 68 Camera lucida drawings of plating in Petalobrissus linguiformis (Peron & Gauthier). A, oral surface, BMNH EE3344; B, apical surface, BMNH EE3345; C, oral phyllode, peristomial margin at base, BMNH EE3334; D, apical disc, BMNH EE3345. Scale bars: A, B = 5mm; C, D = 1 mm. Genus STIGMATOPYGUS d@ Orbigny, 1856 Stigmatopygus pulchellus? sp. nov. Pl. 27, figs 1-8; Figs 69, 70 Types. Holotype BMNH EE4314, paratypes, BMNH EE3324-25, EE3329-30, EE3332-33, EE4312-13: all nine specimens were used for the biometric analysis given below in the description. OTHER MATERIAL. In addition a further six specimens were collected. OCCURRENCE. The species has been found only at Jebel Rawdah, section 2, at the following horizons: bed 14 (6); bed 19 (1); bed 21 (7); loose in scree, derived from beds 14-21 (1). DESCRIPTION. Tests are elongate oval in outline, with a rounded anterior and a small posterior indentation (Pl. 28, figs 2-7). Tests range in length from 37 to 68 mm. Test width is 77-84% of test length (mean = 80%, SD = 1:9%, N = 9) and the widest point is slightly posterior of midlength. Test height is 41-56% of test length (mean = 47, SD = 4.9%, N = 9). The test has a depressed profile with the highest point coincidental with the apical disc and thus anterior of centre. There is a posterior notch marking the position of the periproct, and a small heel underneath the periproct in profile (Pl. 27, fig. 5). The apical disc lies anterior of centre, some 32-40% of test length from the anterior border (mean = 37%, SD = 3-4%, N = 7). It appears to be tetrabasal, although the madrepores extend almost up to the start of the gonopores (Fig. 70D). There is no evidence for sexual dimorphism amongst the specimens to hand. The petals are bowed, and are widest about one third of their distance from the apex (PI. 27, fig. 7). The anterior petal has columns of equal length and the inner series of pores are almost straight. It is 22-27% of test length in length. The interporal zone is about twice as wide as the pore zone. Petal III is significantly longer than the rest. The petals in ambu- lacra II and IV are bowed and converge distally but remain open. The pore columns in the posterior petals are signifi- cantly different in length, with the inner series some 8-10 pore-pairs shorter than the outer column in larger specimens. These petals are also the narrowest. All pores below the petals are single. Towards the peris- tome phyllodes are well-developed, and expand as an open LATE CAMPANIAN-MAASTRICHTIAN ECHINOIDS 213 |PLATE 27 'Figs 1-8 Stigmatopygus? pulchellus sp. nov. 14, 8, BMNH EE4314, holotype; i, lateral, x 1; 2, oral, x 1; 3, apical, x 1; 4, posterior, x 1; 8, detail of peristomial region, x 5. Jebel Rawdah, section 2, bed 19. 5-7, BMNH EE4313, paratype; 5, lateral; 6, posterior; 7, apical; all | 2. Jebel Rawdah, section 2, bed 14. / | | | 214 50 mia @ Test wdth @ Test height a 40 RY 0) 60 Test length (mm) 26 a E24 : E — ra) ) =] ws ) a @ ° ~ = 2 = © - c < 40 50 60 Test length (mm) 10 @ Bourrelets * Peristome ~~ E E ~~ <= ~ a i= © = Test length (mm) Fig. 69 Biometric data for Stigmatopygus pulchellus sp. nov. ‘V’ (Pl. 27, fig. 8). There are five or six pores in each series along the margin with an additional one, or rarely two inner pores at the distal end of the phyllodes (Fig. 70C). Usually there is only one occluded plate in each phyllode, all other plates extending from the adradial to the perradial suture. The first ambulacral plates are long and boot-shaped, with buccal pores that are much smaller than those composing the remainder of the phyllodes (Fig. 70C). Sphaeridial pits occur on each plate close to the midline, forming an alternating series, 4 or 5 per column. The phyllodes lie sunken relative to the surrounding test with buccal pores on the adoral walls of this depression, rather than in the peristomial well. The peristome is pentagonal and either equilateral or slightly longer than wide (PI. 27, fig. 8). Bourrelets are well developed, being straight-sided to weakly wedge-shaped in Anterior to peristome (mm) . SMITH N a 20 50 60 70 Test length (mm) 40 50 60 70 Test length (mm) outline and projecting upwards strongly. They do not, how- ever, project over the peristome. The periproct is only just visible from above, being situated more or less posteriorly (Pl. 27, figs 6, 7). It is triangular in shape with an upper pinched portion and a broader, more rounded lower portion. It is typically slightly taller than broad. There is a subanal platform that extends as an invaginated floor to the periproct. There is a slight aboral lip to the periproct. It lies 21-34% above the base of the test (mean = 27%, SD = 5-5%, N = 7). Tuberculation is fine and uniform aborally, slightly coarser orally. There is a very broad and well-developed naked zone running the length of the posterior interambulacrum. A smaller naked zone is also present anteriorly along the midline. These zones are covered in a very fine granulation LATE CAMPANIAN-MAASTRICHTIAN ECHINOIDS | =1mm. but appear smooth to the naked eye. ' REMARKS. This is a very distinct species, on account of its / asymmetric posterior petals and keyhole-shaped periproct. Small forms resemble Petalobrissus linguiformis (Peron & Gauthier), but are easily distinguished from that species by | their phyllode structure and periproct shape. P. linguiformis has a longitudinal periproct with a short parallel-sided anal sulcus, and also has much better developed phyllodes, with a separate series of inner occluded plates. The new species undoubtedly comes closest to the type species of Stigmatopygus, S. galeatus d’Orbigny from the late Cretaceous of Angouleme, France. Both have a very similar test shape and periproct position and shape. Unfortunately, this species is very poorly known, the original description and | figures being quite inadequate by today’s standards. In par- ticular its phyllode structure is unreported. Kier (1962) described the phyllode structure of another species S. lam- berti Bessairie, from the Campanian of Madagascar, but this \differs in being much wider and taller, with a much larger periproct. It also has better developed phyllodes, which are clearly bowed and comprise an outer series of some 12 pores i) — Nn | Fig. 70 Camera lucida drawings of plating in Stigmatopygus pulchellus sp. nov. A, oral surface, BMNH EE4312; B, apical surface, BMNH EE4314; C, phyllode plating, peristomial margin at base, BMNH EE4312; D, apical disc, BMNH EE3325. Scale bars: A, B = 5mm; C, D and an inner series of 4 or 5 pores. I am therefore not certain that S. /Jamberti is truly congeneric with the type species S. galeatus. Only re-study of the type (apparently lost), or topotype material will solve the problem. If the phyllode structure of S. galeatus is similar to that of S. pulchellus, then S. lamberti should be transferred to a new genus. On the other hand, if S. galeatus proves to have a phyllode structure similar to that of S. lamberti, then S. pulchellus should be made the type of a new genus. Consequently, S. pulchellus can only tentatively be placed in the genus Stigmatopygus. S. galeatus can be distinguished from S. pulchellus by the fact that its petals are illustrated as being of equal length. 216 40 ws Test wdth @ Test height (mm) 30 40 Test length (mm) 20 gs Amb. lll @ Amb. Il r O Amb. | L —) jo) fee} Petal length (mm) 30 40 Test length (mm) Peristome width (mm) 1 2 3 Peristome length (mm) Fig. 71 Biometric data for Nucleopygus magnus sp. nov. Genus NUCLEOPYGUS Agassiz, 1840 Nucleopygus magnus sp. nov. PI. 28, figs 1-7; Figs 71, 2 Types. Holotype BMNH EE4339, paratypes, BMNH EE3340, EE3356, EE3358, EE3363, EE3365, EE3367-68, EE4327, EE4335-38 OTHER MATERIAL. An additional 29 specimens were col- lected. Biometric data was taken from the type series only. OCCURRENCE. This species was found at the following locali- ties and horizons: Jebel Buhays, section 1: loose in scree, derived from the lowest few metres of the Simsima Formation (19). 50 Anterior to peristome (mm) A.B. SMITH 20 30 40 50 Test length (mm) 40 ss a Amb. lll » oO - @ Amb. Il & O Amb. | a £ 2 ss a © ro a i) h o 2 E =) z 20 30 40 50 Test length (mm) 40 ® to apical disc * tostart of anal sulcus Distance from anterior (mm) 20 30 Test length (mm) 40 50 Jebel Buhays, section 3: loose, derived from the lowest bed of the Simsima Formation (1). Jebel Thanais: lowest 2 m of the Simsima Formation (4). Jebel Rawdah, section 2: bed 14 (8); loose in scree (3). Jebel Rawdah, section 4: bed 12 (1). Jebel Faiyah, section 1b: bed 2 (1). DIAGNOsIs. A very large, elongate Nucleopygus with a deep median depression on the oral surface. The periproct opens relatively close to the posterior border and there is only a short anal sulcus. The posterior petals end at a level slightly anterior to the start of the anal sulcus. DESCRIPTION. Tests are subquadrate in outline, with a rounded anterior and a somewhat truncated posterior with a shallow anal notch (PI. 28, figs 1, 4, 5). Test length ranges LATE CAMPANIAN-MAASTRICHTIAN ECHINOIDS 217 & & a a é, Serve rere re * % * PLATE 28 Figs 1-7 Nucleopygus magnus sp. nov. 1-3, BMNH EE4327, paratype; 1, apical, x 2; 2, lateral, x 2; 3, detail of apical disc, x 4. Jebel Thanais, lowest 2 m of the Simsima Formation. 4-6, BMNH EE4339, holotype; 4, apical; 5, oral; 6, lateral; all x 1. Jebel Buhays, section 1; loose in the scree’derived from the lowest 3 m of the Simsima Formation. 7, BMNH EE3367; detail of peristomial region, x 4. Jebel | Buhays, section 2; loose in the scree derived from the lowest 3 m of the Simsima Formation. Figs 8-10 Nucleopygus iranicus (Cotteau & Gauthier). Specimen in the Morgan Collection, Museum d'Histoire Naturelle, Paris; 8, lateral; 9, apical; 10, oral; all x 2. Senonian, Poucht-e-Kouh, Iran. 218 A.B. SMITH Fig. 72 Camera lucida drawings of plating in Nucleopygus magnus sp. nov. A, apical surface, BMNH EE4339; B, oral surface, BMNH EE4337; C, apical disc, BMNH EE4339; D, phyllode plating, BMNH EE4337. Scale bars: A, B = 5 mm; C, D = 1 mm. from 18 mm to about 60 mm (estimated from a broken specimen). Test width is 76-82% of test length (mean = 79%, SD = 2%, N = 11), and the widest point on the test is about two thirds of the distance back from the anterior. The test has a low profile, with test height 42-51% of test length (mean = 39%, SD = 3%, N = 9). It is rounded towards the anterior but obliquely truncated towards the posterior (PI. 27, figs 2, 6). The ambitus is rounded, but relatively low. The oral surface has a marked median depression and the peristome is sunken (PI. 28, fig. 5). The apical disc lies 36-43% of the test length from the anterior border (mean = 39%, SD = 1%, N = 9). It is tetrabasal with large gonopores projecting into the interam- bulacra (Fig. 72C). Ocular plates are small and U-shaped. The small genital plates are separated from one another by ocular plates. Petals are open and subparallel to very slightly bowed (lateral and posterior pairs) (PI. 28, figs 1, 4; Fig. 722A). Pores are both approximately circular and are joined by a well- marked furrow (PI. 28, fig. 3). The interporal zone is approxi- mately 1-5 times as wide as a pore-zone. The anterior petal is the longest, but is only slightly longer than the posterior petals. It has 30 pore-pairs in a column at 30 mm test length, rising to 40 at 40 mm test length. The posterior petals end slightly in front of the anal sulcus. All pores are single below the petals. Phyllodes are hardly expanded adorally (PI. 28, fig. 7; Fig. 72D). There are buccal pores at the rim of the peristome, situated on relatively short and squat first ambulacral plates. The phyllodes have both an outer and an inner series of pores, although the two series are not well separated (Fig. 72D). There are six pores per column in the outer series in ambulacrum III with 7 or 8 in the other | ambulacra, and | or 2 pores in the inner series of ambulacrum III with 2 or 3 in other ambulacra. Every third ambulacral plate is occluded, but the occluded plates do not form a continuous inner series. The phyllodes are not depressed relative to the surrounding test. There are no bourrelets, but the walls to the peristome are covered with fine, dense tubercules. The posterior interambulacrum differs from other interambulacra in that it does not turn sharply in towards the peristome, but instead forms a sloping shelf into the opening. The peristome is pentagonal in outline, approximately 1-6 times as wide as long. It lies 38-42% of the test length from the anterior border (mean = 40%, SD = 1%, N = 10). The periproct is tall and narrow and has a width approxi- mately 40% of its height. It opens into a narrow, parallel- : | : LATE CAMPANIAN-MAASTRICHTIAN ECHINOIDS sided anal sulcus that slopes posteriorly. The periproct opens 73-85% of test length from the anterior border (mean = 78%, SD = 4%, N = 11). The upper surface is covered in fine, slightly sunken ~ tubercles. Adorally tubercles are coarser but are still sunken. _ There is a narrow, zig-zagged granular zone in the posterior -interambulacrum that follows the interradial suture and tapers out before reaching the pesterior border (PI. 27, fig. 7). It is never more than about 78% of the test width at its ' widest. REMARKS. Few other species of Nucleopygus come any- _ where near the size of N. magnus. Both N. iranicus (Cotteau } & Gauthier) (Pl. 28, figs 8-10), from the late Cretaceous of southern Iran, and N. pullatus Stolizcka, from the Maastrich- tian of southern India are much smaller species, reaching little more than 14 mm in test length. Furthermore, they are squatter. N. geayi (Cottreau), from the Maastrichtian of Madagascar, is even smaller, never being reported larger ' than 8 mm test length. Similarly, the European species N. . coravium and N. scrobiculatus are equally small forms. Only in the late Cretaceous of North Africa do any _ Nucleopygus approach the size of N. magnus. N. inaequalis (Peron & Gauthier) resembles N. magnus in shape but its posterior petals extend posteriorly beyond the start of the _ anal sulcus. N. meslei (Peron & Gauthier), from the Campa- | nian, is probably the closest. N. meslei reaches 30 mm in test ‘length, but differs from N. magnus in having a more anterior _ apical disc, a longer anal sulcus and more petaloid ambulacra. / It is also less markedly depressed along the midline on the _ oral surface. . Family ECHINOLAMPADIDAE Gray, 1851 Genus ARNAUDASTER Lambert, 1918 REMARKS. Until now there has only been one species assigned to this genus, A. gauthieri Lambert, from the _Cenomanian of France (Aquitaine). Kier (1962: 105) noted that Arnaudaster might best be considered a synonym of Parapygus, as it differs only in having a more cylindrical shape and more unequal poriferous zones in the same petal. | The discovery of a Maastrichtian species very close in form to _A. gauthieri supports the maintenance of Arnaudaster as a separate genus. Pseudocatopygus longior Cotteau & Gauthier, from the late Cretaceous of Iran, also has unequal columns in its petals and may belong to Arnaudaster. How- ever, Kier (1962) placed Pseudocatopygus as a synonym of _ Parapygus. _Arnaudaster cylindriformis sp. nov. PI. 25, figs 11, 12; Pl. 29, figs 1-9; Figs 73-75 Types. The holotype is BMNH EE4334, paratypes are |BMNH EE4324, EE4331-33, EE3378-81. OTHER MATERIAL. Three poorly preserved specimens, BMNH EE3374-76. BMNH EE3373 may also belong here, but only a fragment of the upper surface is preserved and this apparently shows equally developed columns of pore-pairs in -ambulacrum petal V. OCCURRENCE. This species was found at the following locali- ties along the western margin of the Oman Mountains: Jebel Buhays, section 1: top of bed 1 (1); loose in the scree, 219 40 ws Test width (mm) @ Test height (mm) Test length (mm) 18 ae B To peristome oO @ To apical disc = a Distance from anterior (mm) N 10 8 6 20 30 40 50 60 Test length (mm) 20 a Ambulacrum iil =e @ Ambulacrum Il JN £ A Ambulacrum! = 15 ve = = A 5 ° a A er ¢ - A a ° 010 ] r : € oe? = ° ® —_ 20 30 40 50 60 Test length (mm) Fig. 73 Biometric data for Arnaudaster cylindriformis sp. nov. derived from the lowest few metres of the Simsima Forma- tion (6). Jebel Rawdah, section 2: bed 14 (1); loose at the level of bed 19 (2); loose in scree, derived from beds 14~21 (2); bed 21 (1); bed 26 (1). Jebel Rawdah, section 3: bed 9 (1); loose near top of section (1). DIAGNOsIS. An Arnaudaster with a wide, pentagonal peris- tome with weak bourrelets and approximately four pores in the inner series of phyllodes in each half ambulacrum. DESCRIPTION. Tests range from 21 to approximately 50 mm in length, although many specimens are lacking the very 720 A.B. SMITH PLATE 29 | Figs 1-9 Arnaudaster cylindriformis sp. nov.14, BMNH EE4334, holotype; 1, oral, x 2; 2, apical, x 2; 3, detail of peristomial region, x 4, | 4, lateral, x 2. Jebel Buhays, section 1; loose in the scree derived from the lowest 3 m of the Simsima Formation. 5, 7, 8, BMNH EE4323;, 5, apical; 7, lateral; 8, oral; all x 2. Jebel Rawdah, section 2, bed 21. 6, 9, BMNH EE4324 (depressed variety); 6, posterior; 9, lateral; both | x 2 (see also Pl. 25, Figs 11, 12). Jebel Rawdah, section 2, bed 14. | LATE CAMPANIAN-MAASTRICHTIAN ECHINOIDS 221 | at ACN, PAARL ua = catty , My “Ny ly Uy My A ww My ny HAWN Fig. 74 Camera lucida drawing of plating in Arnaudaster cylindriformis sp. nov, A, oral surface, BMNH EE4324; B, apical surface, BMNH EE4333. Scale bar = 5 mm. Fig. 75 Camera lucida drawings of plating in Arnaudaster cylindriformis sp. nov. A, apical disc, BMNH EE3379; B, phyllode plating, BMNH EE4324. Scale bars = 1 mm. 222 posterior end of the test. Tests are ovoid in outline with a rounded anterior and a very weakly pointed posterior (PI. 25, figs 11, 12; Pl. 29, figs 1, 8). Test width is 71-78% of test length (mean = 74%, SD = 2:5%, N = 10), with the widest point on the test approximately two thirds of the distance from the anterior and well behind the level of the apical disc. Test height is 41-60% of the length (mean = 51%, SD = 6:1%, N = 10). In profile the upper surface is broad and almost flat, uniformly rounded at the anterior, but undercut at the posterior (PI. 29, figs 4, 7, 9). The oral surface is flat or very slightly sunken towards the peristome, while the ambitus is smoothly rounded. The apical disc lies well anterior of centre, some 31-41% of test length from the anterior border (mean = 37%, SD = 2:9%, N = 8). Plating is tetrabasal, although genital plate 2 is enormously enlarged in comparison to the other genital plates and occupies the centre of the disc (Fig. 74A). The entire madreporic plate is covered in dense, fine perforations. Other genital plates are very small and are dominated by the gonopores. There may be sexual dimorphism in the size of gonopore openings. Ocular plates are small, approximately the same size as the genital plates. Ambulacra are petaloid with the interporal zone slightly inflated. The anterior and two posterior petals are similar in size, but the lateral petals are shorter. Petals are broad, lanceolate with more or less parallel columns of pore-pairs at the distal tip (Pl. 29, fig. 2; Fig. 74B). They extend about 80% of the distance to the ambitus. Pore-pairs are conjugate. The anterior petal has columns of equal length, with 41 pore-pairs in a column at 32 mm test length, rising to 57 at 50 mm test length. The anterior column of pore-pairs in the lateral petals is shorter than the posterior column by about five pore-pairs (Fig. 74B). Distally they are slightly narrower, but remain broadly open. The posterior petals also have unequally developed columns of pore-pairs, with the posterior column shorter than the anterior column by seven or so pore-pairs. They extend approximately 70% of the distance to the ambitus and remain broadly open distally. Below the petals pores are single. Phyllodes are not strongly developed and phyllode plating is best seen in BMNH EE3379. Pores of the outer series become enlarged close to the peristome, but the rows of pores do not bow out (Fig. 75B). There are only eight or nine pores forming the outer series in one half ambulacrum. There are three or four pores forming an inner series in each half ambulacrum. Each inner series pore is found on an occluded plate (Fig. 75B). There are small but obvious buccal pores which are not separated from the other pores of the phyllode but which open right at the peristomial rim. The first interambulacral plates are short and wide and are slightly swollen to form a weak floscelle (Pl. 29, fig. 3; Fig. 75B). The peristome is pentagonal, approximately twice as wide as long, and with well-developed vertical walls. The periproct is small and transversely oval, with a width approximately 50-70% of its height. It lies at approximately mid-height on the posterior face, but is angled slightly downwards so that it is just visible from the oral surface and not from the adapical surface (PI. 29, figs 5, 8). Because the posterior of the test is slightly drawn out, the periproctal region is often damaged. The base of the periproct lies between about 30 and 40% of the test height above the base. Tubercles are small, sunken and densely crowded over the whole surface. A.B. SMITH REMARKS. This species comes very close to the type species | A. gauthieri in almost every respect. Both have a very similar, cylindrical test, anterior apical disc and strongly unequal | | same columns that are asymmetric). The only difference of | columns of pores in both lateral and posterior petals (the | significance appears to be in the development of phyllodes. In | A. gauthieri the inner series of pores is rather poorly devel- oped and not well separated from the outer series. In A. | | cylindriformis the inner series pores are somewhat more numerous and form a distinct series. However, this is a minor difference, and there is very little else to separate the two forms. The species also resembles Parapygus inflatus Cotteau & | Gauthier in its overall shape and appearance. However, Cotteau & Gauthier catagorically state (1895: 55) that F | columns of pore-pairs in the petals of the species are equally developed. Order HOLASTEROIDA Durham & Melville, 1957 Family HOLASTERIDAE Pictet, 1857 Genus HEMIPNEUSTES Agassiz, 1836 DIAGNOsIS. Heart-shaped holasteroids showing pronounced petal asymmetry, with the posterior column of pores well) developed and anterior column rudimentary. Periproct sub- | ambital and clearly visible in oral view because of the distinct | posterior notch. The madrepores extend over all genital) plates as well as the anterior three ocular plates. Plastron| © plates wedge-shaped, each just reaching the opposite adradial suture and thus biserially arranged. | OCCURRENCE. Late Campanian and Maastrichtian, Europe, North Africa and the Middle East. REMARKS. Hemipneustes is very close in appearance to Car-. diaster. Both have a cordiform test with a deep, well-defined anterior sulcus which has enlarged primary tubercles along its|_ border. Both genera also have a similar style of plastron) , plating, composed of a short, broad labral plate followed by! up to five wedge-shaped plates which more or less occupy the full width of the plastron (eg. Ernst 1972, text-fig. 20).\ Finally, both have rather similar petals, with the anterior - | ; ! ; column greatly reduced in comparison to the posterior col-_ umn. The primary difference between Cardiaster and Hemip-| neustes lies in the fact that Hemipneustes has little more than) rudimentary pore-pairs in the anterior columns of its petals, whereas Cardiaster has small but distinctly conjugate pore- pairs in these anterior columns. Furthermore, madrepores are more extensively developed in Hemipneustes, extending beyond genital plate 2 to cover genital plate 3 and ocular plates II, III and IV. Many species names have been erected in the past, mostly based on relatively few specimens and often on rather poorly preserved material. From Noetling (1897) onwards workers have relied on two simple biometric indicators to distinguish species of this genus; width/length and height/length ratios) (see for example Devries 1967; Aziz & Badve 1990). How-| ever, not only do these ratios vary ontogenetically and thus vary according to absolute test size, but they have also rarely been applied rigorously using large populations. There are three species found amongst the material col- lected from the Oman Mountains. These appear to have some stratigraphic value. The lowest forms found at Jebel Rawdah belong to H. persicus var. sardanyolae Vidal, and t v ~ LATE CAMPANIAN-MAASTRICHTIAN ECHINOIDS are found in the basal sandy facies. These have the broad shallow anterior sulcus characteristic of the H. delettrei com- plex, a group restricted to the late Campanian of Tunisia and Algeria (Zhagbib-Turki 1987). Above this, or possibly par- tially overlapping, H. arabicus is found. This species is similar in profile and in the positioning of its apical disc, but has a _ much narrower, deeper and more sharply defined anterior sulcus. The first elevated Hemipneustes forms, belonging to _ H. compressus, appear in bed 14, having more uniformly rounded profiles. However, by the time bed 21 is reached, H. - compressus has a much more quadrate profile with strongly _ pinched ridges to the frontal groove. Finally, by bed 26 some _ H. compressus appear that are very elevated and which have developed a strong slope towards the posterior similar to the late Maastrichtian H. striatoradiatus. In trying to identify the Oman/United Arab Emirates species, a number of other species have been examined and the | following are accepted as valid: Hemipneustes delettrei Peron & Gauthier (PI. 30, figs 9, 10), late Campanian of Algeria. Rounded in outline and vari- able in profile, from highly inflated (H. africanus form) to depressed (H. delettrei form). w/l = 0-89-0-95 (mean = 0-91): h/l = 0:58-0:79 (mean =0-70) [size range 73-98 mm]. Distinguished from all other species by the large, broad anterior ambulacral sulcus; about twice as broad as in any other species. Hemipneustes striatoradiatus (Leske) (Pl. 30, figs 7, 8; Fig. | 78C), Upper Maastrichtian of the Netherlands. Rounded in outline with a very narrow and shallow anteal sulcus. Variable in profile from flat-topped to distinctly raised anterior to the apical disc. Sides rather vertical giving quadrate cross-section. Apical disc anterior of centre. w/l = 0:87-0:97, mean = 0-91: h/l = 0-61-0-80, mean 0-69 [Size range 50-100 mm]. Distinguished from other species by (i) its narrow, shallow anteal sulcus, (ii) rounded outline and (iii) quadrate lateral cross-section. _ Hemipneustes pyrenaicus Hebert (Pl. 30, figs 5, 6) — closest to H. compressus in having a short narrow frontal groove with a strong vertical component, anterior apical disc, and quadrate profile. It differs in having a flat apical surface | and in lacking adapical pinched ridges to the anterior sulcus. | H. persicus Cotteau & Gauthier (PI. 31, figs 1-7; Pl. 32, figs 1-4; described below). | H. compressus Noetling (PI. 31, figs 8-11; described below). 'H. arabicus Ali (PI. 30, figs 14; described below). In addition the following species are treated as synonyms or rejected names, or are based on inadequate material: _H. minor Peron & Gauthier, Upper Senonian of Iran (a small H. persicus Peron & Gauthier). H. oculatus Cotteau, Maastrichtian of Ciply, Belgium. This appears to be a large variety of H. striatoradiatus, some- what crushed but with a deeper frontal groove as illustrated by Cotteau. The specimens referred to under this name by van der Ham (1989) are simply rather tall and posteriorly inclined H. striatoradiatus. ‘H. arnaudi Cotteau 1892, Upper Senonian of Dordogne. This is very like high forms of H. compressus in having an elevated keel to the ambulacrum. It has too wide an anterior sulcus to be a H. striatoradiatus. Treat as a probable synonym. EEE 223 H. indicus Aziz & Badve 1990; Maastrichtian of S India, is identical in profile to H. arnaudi but larger. Its deep frontal groove and anterior apical disc makes it part of the compressus group. H. sardanyolae Vidal 1921, Campanian of Spain, is treated here as a shallow grooved variety of H. persicus. H. nicklesi Vidal 1921, Campanian of Spain, is a Hemip- neustes sp. based on crushed and badly preserved speci- mens that are basically indeterminate. It is probably a synonym of H. persicus var. sardanyolae Vidal. Spatangoides martelli Checchia-Rispoli, Maastrichtian of Libya. Here synonymized with H. compressus Noetling. Spatangoides tripolitanus Checchia-Rispoli, Maastrichtian of Libya, is clearly a species of Opisopneustes. Its primary adapical interambulacral tubercles are well developed and it represents a valid species, differing from O. cossoni in having a vertically positioned periproct. Spatangoides aichinoi Checchia-Rispoli, Maastrichtian of Libya. Here synonymized with H. persicus. Hemipneustes batheri Lambert [=H. leymeriei Noetling]. This is probably a variety of H. persicus, having the posterior apical disc and oval outline with shallow and open anterior sulcus, but it is unusually tall. H. noetlingi Lambert was erected as a replacement name for H. pyrenaicus of Noetling, from the Maastrichtian of Baluchistan. However, like Devries (1967), I can find no significant difference between Noetling’s specimens and the type material of H. pyrenaicus and treat H. noetlingi as a junior synonym. Hemipneustes leymeriei Hebert has the very narrow and shallow groove and circular outline of a H. striatoradiatus. Topotype material from Gansec, southern France, con- firms this. Spatangoides tripolitanus Checchia-Rispoli is a species of Opisopneustes. Hemipneustes compressus Noetling, 1897 PINS iesties 8-11; Figs 76, 77, 78A 1897 Hemipneustes compressus Noetling: 34, pl. 7, figs 3, 4, pl. 8, figs 1, 2. 1931 Spatangoides Martellii Checchia-Rispoli: 7, pl. 1, figs 3), 1967 Hemipneustes compressus Noetling; Devries: 188, pl. 6, figs 31-33. TyPES. The syntypes are the four specimens illustrated by Noetling and presumably in the Geological Survey of India collections. MATERIAL STUDIED. Thirty specimens were collected. Bio- metric data was taken from the following 16 specimens: BMNH_ EE3744-45, EE3748, EE4070-71, EE4073-74, EE4076, EE4078-82, EES022-24. OCCURRENCE. Maastrichtian of Libya, Oman and Bal- uchistan. In the western Oman Mountains it occurs at the following levels: Jebel Rawdah, section 2: bed 14 (2); bed 15 (2); bed 19 (1); beds 20/21 (17); bed 22 (1); bed 26 (2 specimens plus fragments); bed 27 (2). Jebel Rawdah, section 3: beds 9/10 (1). Jebel Rawdah, section 4: loose, towards top of section (1). DIAGNOSIS. Test rather more elongate than in other species, typically flat-topped in side-view and arched in cross-section. 224 60 @ H. persicus th O H. arabicus ° 3 50 ¢ H. compressus s ~— ~ <£ 2s © = ~ e © = > fo) A.B. SMITH | 70 = H persicus #0 % rene ° | O H. arabicus * ~~ ’ @ H. compressus r E 60 y 2 | — = ' re) $ | a) | ® | © - | 30 40 50 60 70 80 90 40 50 60 70 80 90 Test length (mm) Test length (mm) | 16 50 =e = H. persicus a a H. persicus 0 E 0 H arabicus o . 0 H arabicus fo) als ¢ H compressus 4 ¢ H. compressus a s re} ao) €12 = £ 2 = = 210 i) é 19 © 8 = ra c 3 < = 6 20 30 40 50 Length of Amb. Ili (mm) Test length (mm) 7 @ H persicus 6 re) O H arabicus @ H compressus 5 w + Sulcus depth at ambitus (mm) N 10 Sulcus width at ambitus (mm) 12 14 Fig. 76 Biometric data for species of Hemipneustes. w/l = 80-87% of length, mean = 83%; height 54-67% of length, mean = 60% (size range 63-83 mm). Frontal groove narrow but moderately deep at the anterior; typically slightly keeled. Apical disc lies well towards the anterior (distance between anterior and apical disc = 34-44% of length: mean = 39%) and is pinched up. Distinguished from other species by (i) its anterior apical disc, (ii) the deep anterior groove which is relatively short and geniculate with a strong vertical component, (iii) being more subquadrate outline, typically inclined from the apical disc posteriorly. DESCRIPTION. Tests range in length from 63 to 83 mm in length and from 53 to 69 mm in width (width = 80-87% of length; mean = 83%, SD = 1-8%, N = 13). In outline the test is cordate with a sharp, deep anterior sulcus, and a 16 truncated and even slightly indented posterior (PI. 31, figs 8, 9). The widest point on the test lies approximately mid- | length. Test height is variable, ranging from 38 to 54 mm | (height = 54-66% of test length: mean = 60%, SD = 3-7%, | N = 13). In side view the upper surface varies from almost flat to strongly vaulted, with the tallest point on the test lying | immediately anterior of the apical disc (Pl. 31, fig. 11). The test slopes towards the posterior which is truncated. Anterior to the apical disc the test curves uniformly to become almost | , vertical. There is a sharp curve from the anterior to the more or less flat base (PI. 31, fig. 11). In anterior profile the test appears vaulted with rounded sides sloping up to the crest. The anterior sulcus becomes slightly pinched towards the | , base and its edges are sharply delimited (PI. 31, figs 10, 11). | ) 4 LATE CAMPANIAN-MAASTRICHTIAN ECHINOIDS 225 PLATE 30 Figs 1-4 Hemipneustes arabicus Ali. BMNH EE5027; 1, oral; 2, apical; 3, anterior; 4, lateral; all x 1. Jebel Rawdah, section 1, bed 4. Figs 5,6 Hemipneustes pyrenaicus Hebert. Specimen in the Museum d’Histoire Naturelle, Paris; Maastrichtian of Montbéraud, Haute | Garonne, France. 5, apical; 6, lateral; both x 1. | Figs 7,8 Hemipneustes striatoradiatus (Leske). BMNH 75822; 7, apical; 8, lateral; both x 1. Maastrichtian of Maastricht, The Netherlands. Figs 9,10 Hemipneustes delettrei africanus Peron & Gauthier. BMNH E3654; 9, apical; 10, lateral; both x 0-75. Upper Campanian of Ain | Joutu, Algeria. 226 ) Z i) i) i \ yp ; ‘ A.B. SMITH Fig. 77 Camera lucida drawings of plating in Hemipneustes compressus (Noetling), BMNH EE3745. A, apical surface; B, oral surface. Scale bar = 5 mm. Fig. 78 Camera lucida drawings of apical disc plating in species of Hemipneustes. A, H. compressus (Noetling), BMNH EE3745; B, H. arabicus, BMNH EES027; C, H. striatoradiatus (Leske), BMNH 38630; D, H. persicus Cotteau & Gauthier, BMNH EE3747. Scale bars = 1 mm. The apical disc lies at or slightly in front of mid-length (distance to the anterior border = 34-44% of test length: mean = 39%, SD = 3-1%, N = 13). It is elongate with 4 gonopores occupying most of the area of the genital plates (Fig. 78A). The anterior pair of genital plates are separated from the posterior pair by the lateral ocular plates. Madrepores extend across genital plates 2 and 3, as well as ocular plates II, III and IV, even extending into the wall of the gonopores in some instances. The two posterior ocular plates are slightly larger and abut each other. The anterior ambulacrum is non-petalloid. It is almost flush at the apex, becomes slightly depressed towards the anterior and then forms a deep vertical sulcus close to the anterior (PI. 31, figs 8, 10). This groove continues up to the peristome and is widest near the top of the anterior sulcus becoming narrower towards the base of the sulcus. Pore-pairs are oblique isopores, closely spaced in the upper half, but becoming smaller and more widely spaced towards the bot- tom of the sulcus. The margins of the sulcus are relatively sharp, especially towards the top of the anterior sulcus, where they may form small crests on either side. The width of the sulcus is 13-20% of the test width (mean = 16-3%, SD = 1-86%, N = 12) and at its deepest it is about 3-5 to 4-5 mm. The anterior petals are strongly asymmetric, with only the posterior column well-developed (Pl. 31, fig. 8; Fig. 77). These curve forward and, towards the tip, turn slightly down. The inner pore is circular, the outer one elongate, and the distance between the pairs is as wide as the outer slit-like pore. Pore-pairs are conjugate and are separated from one another by a single row of small tubercles. The posterior column tapers both adapically and adambitally. There are 45 pore-pairs in a column in a 65 mm individual, rising to 52 in an 81 mm individual (Fig. 76). The anterior column of pores remains rudimentary throughout. All are pore-pairs, but the pores always remain small and close together, never becom- ing conjugate. However, they do gradually increase in size towards the ambitus. The posterior petals are similar to the anterior petals in their pore-pair development and column asymmetry, with only the posterior column bearing large conjugate pore-pairs (Fig. 77). This column is flexed outwards and backwards and extends only about two thirds of the distance towards the ambitus. There are 32 pore-pairs in a column in a 65 mm individual, rising to 42 in an 81 mm individual. Close to the apex the two ambulacra are almost parallel, but further away they diverge at about 90°. Pores beneath the petals remain double, but are microscopic. Close to the mouth there are | | | LATE CAMPANIAN-MAASTRICHTIAN ECHINOIDS two to three larger peribuccal pores in ambulacrum III, four to five in ambulacra II and IV, and three to four in ambulacra Tand V. The interambulacra are biserial to the apex, although they become extremely narrow, especially posteriorly. On the oral surface there is a short, broad labral plate followed by a series of 5 alternating triangular-shaped plates which either just or almost reach the opposite suture (Fig. 77). Further towards the posterior the columns become more typically biserial. The peristome is oval, slightly more than twice as wide as long, and lies at the base of the anterior sulcus, some 15-18% of test length from the anterior (mean = 17%, SD = 1:2%,N = 11). The peristome faces forward and is visible in anterior view. The labral plate hardly indents the posterior of the peristome. The periproct is oval, typically about 70% as wide as tall, and is strongly overhung so that it is visible from the oral surface but not from the apical surface (Pl. 31, figs 8, 9). The test beneath the periproct is indented and there are two projections on either side. The periproct opens between interambulacral plates 5 and 10, the lower plates being strongly curved. The periproct lies low on the posterior surface and the distance to the base of the periproct from the lower surface is some 18-27% of the test height. Tuberculation is fine and uniform throughout, except along the iner (interambulacral) margin of the frontal sulcus where a double or triple row of noticeably larger tubercles is developed. There is no trace of a lateral fasciole to be seen. Oral tubercles are slightly larger than adapical ones, but there is no real difference in size between the tubercles of the plastron and those of the latero-ventral regions. The oral ambulacra appear to be tubercle-free. REMARKS. This species is relatively common at certain levels in Jebel Rawdah. It co-occurs with small varieties of H. persicus but can be distinguished from that species by its Narrower, more vertical and more sharply defined anterior sulcus, its more anterior apical disc, its lack of adapical primary tubercles, and its more quadrate profile. It differs from H. arabicus, whose size range is more-or-less coinci- (as (a9 Li /? l eee DOT dent, in having the apical disc positioned anterior of centre, and in having a more developed vertical component to the anterior sulcus. The stratigraphically lowest species tend to have a more rounded profile, while higher samples become progressively more peaked, with a stronger posterior inclination as the region immediately in front of the apical disc becomes taller. Thus in the beds immediately above bed 13, Jebel Rawdah, section 2, H. compressus is rather flat and the vertical component of the anterior sulcus is small. By bed 21 the tests are much more quadrate in outline, with a small but obvious peak in front of the apical disc, and by bed 26 some highly elevated tests are found. Hemipneustes persicus Cotteau & Gauthier, 1895 PI. 31, figs 1-7; Pl. 32, figs 14; Figs 76, 78D, 79 1895 Hemipneustes persicus Cotteau & Gauthier: 15, pl. 2, figs 1-6. 1895 Hemipneustes minor Cotteau & Gauthier: 17, pl. 2, figs 7-9. 1921 Hemipneustes sardanyolae Vidal: 11, pl. 2, fig. 2; pl. 3, fig. 2. TYPES. The syntypes are the specimens from Aftab and Derre-i-Chahr, Iran, described by Cotteau & Gauthier (1895: 15). Although the authors stated that they had many examples, their description appears to have been based on only one specimen 54 mm in length, which was illustrated. This is here designated lectotype. MATERIAL STUDIED. There are five large individuals of this species, BMNH EE4083, EE4084, EE4090, EE5025-—26, all rather poorly preserved. Small individuals that appear indis- tinguishable are relatively common and well-preserved at one horizon at Jebel Rawdah, section 2. Biometric data is based on the following eleven specimens: BMNH_ EE3742, EE3746—-47, EE4091, EE4097, EE4099, EE5028-32 in addi- tion to the five larger specimens cited above. OCCURRENCE. In the Oman Mountains region this species Lens 1 Fig. 79 Camera lucida drawing of plating in Hemipneustes persicus Cotteau & Gauthier. A, oral surface, BMNH EE3747; B, apical surface, BMNH EE3746. Scale bar = 5 mm. 228 occurs at Jebel Rawdah at the following localities and hori- zons: Section 1: bed 4 (2). Section 2: bed 19 (1); beds 20/21 (23); loose in scree, derived from beds 14-21 (4). Section 3: bed 5 (1); bed 7 (1). Section 4: loose a little below the top of the measured section (1). The type series comes from the ‘Senonienne’ (probably Maastrichtian) of Aftab and Derre-i-Chahr, southern Iran. DIAGNOSIS. Test ovate, depressed in side-view and in cross- section arched. Width 80-88% of length, mean = 85%: height 42-55% of length, mean = 49% [size range 44-89 mm test length]. Frontal groove broad and open, expanding anteriorly, comparatively shallow at the anterior; never keeled. Apical disc lies approximately mid-length (distance between anterior and apical disc 45-52% of test length; mean = 48%). Distinguished from other species by (i) its depressed, rounded profile, (1) broad, relatively shallow anterior groove (iii) central apical disc. DESCRIPTION. Tests range from 44 to 89 mm in length and from 38 to 68 mm in width (width = 80-88% of length; mean = 85%, SD = 2:1%, N = 4). In outline the test is oval with a prominent anterior sulcus and a truncated posterior (PI. 31, figs 1, 3, 4, 6). The widest point on the test is approximately mid-length. In profile the test is depressed, more or less uniformly curved both in front and behind, with the tallest point of the test anterior of centre (PI. 31, figs 2, 5). Test height is 42-53% of length (mean = 48%, SD = 4:2%, N = 5). In anterior profile the sides are uniformly rounded. The apical disc lies approximately mid-length, 43-52% of test length from the anterior border (mean = 48%, SD = 2-4%, N = 15). Plating appears to be similar in arrangement to that of H. compressus, with the anterior pair of genital plates separated from the posterior pair by ocular plates II and IV, which meet centrally (Fig. 78D). Madrepores extend over genital plates 2 and 3, and over ocular plates II, III and The pores of the anterior groove are small and closely spaced adapically, becoming slightly more widely separated towards the ambitus. The anterior sulcus is rather broad and shallow, becoming more or less parallel-sided towards the anterior border (Pl. 31, figs 1, 3, 6). The margins of the groove are gently rounded, never crested. The width of the sulcus is 17-22% of the test width (mean = 19%, SD = 2:0%, N = 13) and at its deepest it is only 2:0-3-0 mm in depth. The sulcus does not have a vertical component at the anterior, as is seen in H. compressus, but curves uniformly towards the ambitus. Petals are as in H. compressus, though the anterior pair are slightly more curved forwards than in that species (Fig. 79). The anterior column of pore-pairs in each petal is composed of small, rudimentary pore-pairs which become slightly larger towards the ambitus. The posterior column in each petal is composed of wide, conjugate pore-pairs, with the inner pore circular and the outer pore distinctly slit-like. There are 36 pore-pairs in the posterior column of ambulacrum II petal, and 28 in ambulacrum I petal in a 44 mm length individual, rising to 65 and ?55 pore-pairs respectively in an 80 mm individual. The anterior petals curve forwards to diverge at an angle of approximately 30° to the midline, before tapering A.B. SMITH and turning laterally slightly at their distal end. They extend almost to the ambitus in adapical view (Fig. 79). The poste- rior petals are slightly shorter, extending approximately two- thirds of the distance to the ambitus. They too are flexed, but for most of their length they diverge at approximately 50° to the midline. Pores below the petals are microscopic, but remain double. Peribuccal pores are well-developed, with four or five to a column in ambulacrum III, six or seven in ambulacra II and IV, and four or five in ambulacra I and V. The interambulacra become very narrow adapically (Fig. 79), but remain biserial. The plastron consists of a small, broad, labral plate followed by five wedge-shaped plates arranged uniserially (Fig. 79). The peristome is oval to crescentic, slightly more than twice as wide as long, and lies at the base of the anterior sulcus, some 17-21% of test length from the anterior (mean = 19%, SD = 1:8%, N = 5). The labral lip projects downwards slightly and is visible in anterior view. The labral plate indents the posterior of the peristome. The periproct is oval, 65-75% as wide as tall (mean = 71%), and more or less vertical on the posterior face. It is therefore not visible from either the apical or oral surfaces (Pl. 31, figs 1-4). There are no subanal protuberances devel- oped in small individuals, although small protruberances are found on larger individuals (Pl. 32, fig. 2). The periproct opens between interambulacral plates 5 and 10, the lower plates being strongly curved. The periproct lies low on the posterior surface and the distance to the base of the periproct from the lower surface is some 16-25% of the test height (mean = 22%). Aboral tuberculation consists of small primary tubercles scattered amongst dense miliaries. Larger primary tubercles are found along the inner (interambulacral) margins of the frontal sulcus, where they are approximately three abreast. They are also found close to the apical disc interambulacrally on many juveniles. These adapical tubercles are only slightly larger than other tubercles, but form a characteristic feature where preservation is good. There is no trace of a lateral fasciole. Oral tubercles are slightly larger than adapical ones, but there is no real difference in size between the tubercles of the plastron and those of the latero-ventral regions. The plastron tubercles are largest towards the adambulacral mar- gins and decrease in size towards the midline. Oral ambulacra are smooth and free of tubercles. REMARKS. I have examined topotype material of this species from Iran and feel secure that the Omani Mountain material is conspecific. However, there is one small difference; the Omani specimens in general have a slightly shallower anterior groove at the ambitus than do the Iranian specimens. This is not considered sufficient to merit separation, since all inter- mediates can be found. Similarly, Hemipneustes sardanyolae Vidal from the late Campanian of Sardanol, Spain, is slightly flatter and more oval in outline, but is otherwise very similar. It too is synonymized here. The species is readily distinguished from H. compressus by its very different, more depressed profile, shallower and very much broader frontal sulcus and central apical disc. It also has better developed phyllodes. It differs from H. arabicus in its very broad and shallow frontal sulcus, and from Opisop- neustes in the lack of aboral primary tubercles. There is little doubt that the closest species to H. persicus is H. delettrei (Peron & Gauthier) from the late Campanian of North Africa. Both have a very similar broad, shallow frontal ~the oe ann lem LATE CAMPANIAN-MAASTRICHTIAN ECHINOIDS 229 PLATE 31 Figs 1-7 Hemipneustes persicus Cotteau & Gauthier. 1, 2, BMNH EE4084; 1, apical; 2, lateral; both x 1. Jebel Rawdah, section 3, bed 5. 3-5, BMNH EE5032; 3, apical; 4, oral; 5, lateral; all x 1. Jebel Rawdah, section 2, bed 21. 6, BMNH EE3746, apical, x 1. Jebel Rawdah, section 2, bed 21. 7, BMNH EE3747, lateral, x 1. Jebel Rawdah, section 2, bed 21. Figs 8-11 Hemipneustes compressus Noetling. BMNH EE3745; 8, apical; 9, oral; 10, anterior; 11, lateral; all x 1. Jebel Rawdah, section 2, bed 21. 230 groove that expands continuously towards the anterior, a character seen in no other Hemipneustes. H. persicus differs from H. deletrettei primarily in being very much more depressed in profile and more elongate, with more strongly inflexed petals. However, the differences are not great. Hemipneustes arabicus Ali, 1989 _ PI. 30, figs 1-4; Figs 76, 78B, 80 1989 Hemipneustes arabicus Ali: 408, figs 6 (1-3), 7. 1989 Hemipneustes persicus Cotteau & Gauthier; Ali: 408, fig. 6 (4). Types. The syntype series are the seven specimens and three fragments mentioned by Ali (1989) as being housed in the Geology Department Museum, Al Ain University, United Arab Emirates. The lectotype, here designated, is his figured specimen (op. cit. fig. 6 (1-3)). MATERIAL STUDIED. Nine specimens: the following descrip- tion is based on the four best-preserved of these, BMNH EE4085, EE4087-88, EE5027. OCCURRENCE. This species was first described from Jebel Rawdah, Oman. Specimens were collected from the follow- ing horizons at Jebel Rawdah: Section 1: bed 4 (3). Section 2: bed 11 (4); bed 14 (1). Section 4: loose in scree, a little below beds 21/22 (1). DIAGNOSIS. Test ovate and rather elongate, depressed in side-view and in cross-section arched. Width 76-79% of length, mean = 77%; height 45-46% of length, mean = 45% (size range 76-89 mm). Frontal groove narrow, sharply defined with a small but prominent rim; deeply sunken at the anterior. Apical disc lies at or slightly behind mid-length (distance between anterior and apical disc = 49-55% of test length: mean = 52%). Distinguished from other species by (i) its depressed, elongate profile, (11) long, sharp and deep anterior groove (iii) central apical disc. DESCRIPTION. Tests range in length from 77 to 89 mm and in width from 60 to 69 mm (width = 76-79% of length, mean = 77%, SD = 1-:1%, N = 4). The widest point lies at or slightly in front of mid-length. In outline the test is oval with a sharp and narrow anterior sulcus and a rather broad posterior truncation (PI. 30, fig. 1). In side-view the test is depressed and gently rounded (PI. 30, fig. 4). Test height is 44-46% of length (mean = 45%, SD = 0-8%, N = 4). The tallest point on the test is at, or slightly in front of, the apex. In anterior view the test is uniformly rounded, but there are small crests on either side of the sulcus. The apical disc is clearly seen in BMNH EE5S5027 (Fig. 78B). Genital pores are large and separated, with ocular plates If and IV meeting centrally and separating genital plates 2 and 3 from genital plates 1 and 4. Madrepores cover genital plates 2 and 3 and ocular plates II, II] and IV. A couple of pores also appear on the margin of genital plate 1. It lies 49-55% of the test length from the anterior border (mean = 52%, SD = 2:5%, N = 4). Ambulacrum III is narrow and parallel-sided adapically (Pl. 30, fig. 2). It is slightly depressed from the apex until it approaches the anterior border, then it turns rather sharply downwards into a deep groove, 4-6 mm in depth. The sulcus remains narrow throughout, only 11-16% of the test width at the anterior (mean = 13%). Pore-pairs are densely packed A.B. SMITH rN a d, et et Fig. 80 Camera lucida drawing of adapical plating of Hemipneustes arabicus, BMNH EES027. Scale bar = 5 mm. along its length until it turns adorally, where they become much more spread out. The floor of the sulcus is covered in fine tuberculation which decreases in size away from the pore-pairs towards the perradius. The petals are similar in form to those of other species of Hemipneustes, the adapical pores in the anterior columns being particularly rudimentary. The anterior petals are flexed forward and diverge at an angle of about 15° to the frontal groove before curving laterally near their distal end (Fig. 80). The posterior petals diverge at about 110—120° to each other. Adorally there are well-developed phyllodes, with five or six pores in a column in the lateral ambulacra. The plastron consists of a small, broad labral plate and a series of four or five wedge-shaped plates that extend across the full width of the plastron (Fig. 80). Posterior plates are biserial. The peristome is oval, indented slightly by the labral plate, and predominantly forward-facing. It lies at the base of the anterior sulcus, some 16% of the test length from the anterior border. The periproct is oval and downward facing; clearly visible from below but hidden from above (PI. 30, figs 1, 2). The test is developed into two prominent bulges, one on either side of the periproct. The periproct lies between interambulacral plates 5 and 10. The base of the periproct lies between 19 and 26% of the test height above the base. Tuberculation is fine and uniform over the apical surface, except along the inner interambulacral margins-of the frontal groove, where larger primary tubercles are developed. Oral tuberculation is slightly coarser, with ambulacral zones free of any tuberculation. No lateral fasciole is seen. REMARKS. This species appears somewhat intermediate between the broad grooved H. persicus and the tall, narrow- grooved but upright H. compressus. It differs from H. persicus in the sharpness and narrowness of its anterior LATE CAMPANIAN-MAASTRICHTIAN ECHINOIDS 231 PLATE 32 Figs 1-4 Hemipneustes persicus Cotteau & Gauthier. Specimen in the Morgan Collection, Museum d’Histoire Naturelle, Paris; 1, apical; 2, oral; 3, lateral; 4, anterior; all x 2. Arkouraj, Iran. Figs 5-8 Hemiaster hattaensis Ali. 5, BMNH EE4060; apical, x 2. Jebel Thanais, basal 2 m of the Simsima Formation. 6-8, BMNH EE4059; 6, lateral; 7, apical; 8, oral; all x 1-5. Jebel Buhays, section 1; loose in the scree derived from the lowest 3 m of the Simsima Formation. 232 sulcus. In H. persicus the frontal groove is much broader and lacks the interambulacral keels. It is also much less deeply indented at the ambitus. However, like H. persicus, H. arabicus has its apical disc at or slightly behind mid-length, in contrast to H. pyrenaicus where the test is more quadrate and the apical disc lies anterior of centre. Like H. compressus, H. arabicus shows a sharp change from apical to anterior sectors of its anterior sulcus. Order SPATANGOIDA Claus, 1876 Family HEMIASTERIDAE Clark, 1917 Genus HEMIASTER Agassiz, in Agassiz & Desor, 1847 Pl. 32, figs 5—8; Figs 81C-E Hemiaster hattaensis Ali, 1989 21903 Hemiaster sp. Lambert: 87, pl. 3, figs 6-8. 1989 Hemiaster (Bolbaster) hattaensis Ali: 409, fig. 6 (5-8). Types. Eleven specimens referred to by Ali form the syntype series since no holotype was designated. These are in the Geology Department Museum, University of Al Ain, United Arab Emirates. MATERIAL STUDIED. Thirteen specimens, of which the fol- lowing six were used for the biometric study: BMNH EE4055, EE4057, EE4059-60, EE4064 and EE4320. OCCURRENCE. The species is known for certain only from the western Oman mountain. It has been found at the following localities: Jebel Buhays, section 1: loose in the scree, derived from the lowest few metres of the Simsima Formation (5). Jebel Thanais: lowest 2 metres of the Simsima Formation (1). Jebel Rawdah, section 1: bed 6 (3). Jebel Rawdah, section 2: loose in scree (1). Jebel Rawdah, section 3: bed 5 (1). Jebel Rawdah, section 4: bed 2 (1); bed 13 (1). We ‘ Ly Le Vy Fig. 81 A.B. SMITH DIAGNOsIS. An ovate Hemiaster with cruciform petals in which the posterior pair are about half the length of the anterior pair. Apical disc ethmophract, with the posterior genital plates L-shaped and separating the posterior ocular plates; lying posterior to midlength. Ambulacrum III long, narrow, depressed adapically but becoming flush with sur- rounding area towards the ambitus. DESCRIPTION. Tests oviform with a uniformly rounded ante- rior and a slightly pointed posterior (Pl. 32, fig. 5). Test length is 15-32 mm and test width 11-26 mm (width = 82-87% of length in larger individuals but only 75% in a juvenile 15 mm long). The widest point lies about mid-length. Test rather depressed in profile (Pl. 32, fig. 6), with a height 55-67% of length (mean = 64%, SD = 4-6%, N = 6). Lower surface gently convex with a slight keel to the plastron towards the peristome. Upper surface flat towards the poste- rior, sloping uniformly towards the anterior (PI. 32, fig. 6). The tallest point on the test lies just posterior to the apical disc. The apical disc is ethmophract (Fig. 81D) and lies 58-62% of the test length from the anterior border in adults, but further (73%) in the juvenile 15 mm in length. Anterior gonopores are set closer together than the posterior pair. Genital plates 2 and 3 are relatively small whereas genital plates 1 and 4 are longer and L-shaped and are broadly in contact (Fig. 81D). There are relatively few madrepores developed. Ocular plates are small, pentagonal and project. The anterior ambulacrum is sunken adapically but the sulcus is lost about two-thirds of the way towards the ambitus and there is no indentation at the ambitus (PI. 32, figs 5, 7). The adapical sulcus is parallel-sided and very narrow. There are 14 pore-pairs in a column in 17-18 mm individuals and these are strongly oblique with a prominent interporal parti- tion. The floor of the sulcus is covered in fine and dense granulation. Pores beyond the peripetalous fasciole are single and minute. The interambulacra on either side of the frontal sulcus are keeled, especially in the larger specimens. ———— Camera lucida drawings of plating in Hemiaster species. A, B, Hemiaster paronai Checchia-Rispoli, BMNH EE5034: A, apical surface (fasciole stippled); B, apical disc. C-E, Hemiaster hattaensis Ali: C, peristome and labral plate, BMNH EE4064; D, apical disc, BMNH EE4060; E, apical surface (fasciole stippled), BMNH EE4059. Scale bars: A, E = 5 mm; B, D = 1 mm; C = 2mm. cE LATE CAMPANIAN-MAASTRICHTIAN ECHINOIDS The petals are cruciform and sunken, the anterior pair being twice as long as the posterior pair (PI. 32, figs 5, 7; Fig. 81E). Pore-pairs are broad and conjugate and the columns close distally. The perradial zone is slightly narrower than the pore-pair zones on either side. There are 12-14 pore-pairs in a column in the posterior petals and 24-26 in the anterior petals in individuals 15-18 mm in length. The periproct is tear-drop shaped, pointed adapically, and lies high on the posterior face. It is about 3 mm wide and 4 mm in height in an individual 17 mm in length. The base of the periproct lies 55-58% of test height above the base. The peristome is D-shaped and is 1-6 to 1-8 times as broad as wide. It lies 24-29% of the test length from the anterior border in 17-18 mm length individuals. It is neither invagi- nated nor does it have a surrounding rim, as is seen in many Hemiaster species. The plastron is broad, with a straight, median suture (Fig. 81C). It is covered in dense orderly rows of tubercles. The labral plate is vase-shaped and relatively small. Surrounding the peristome there are four phyllode pores in each column of a lateral ambulacrum and three in the posterior ambulacrum. There are also two or three enlarged subanal pore-pairs. There is a well-developed peripetalous fasciole that is without sharp angles. The remainder of the upper surface has scattered primary tubercles and dense miliaries. REMARKS. Ali (1989) erected this species for 11 specimens from Jebel Rawdah. It resembles H. aquisigranensis Schluter in the form of its petals and frontal groove, but differs from that species in having a larger peristome and having a vertical or slightly outward-sloping posterior. In H. aquisigranensis the posterior is strongly retrenched, such that the periproct is visible from the oral surface rather than the aboral surface. H. hattopsis undoubtedly comes closest to H. punctatus _d@Orbigny, from the late Campanian of France. However, ~ this species differs in being very much smaller, more elevated and in having a well developed rim to the peristome. Lambert (1903) described and figured a badly preserved _ specimen that may be conspecific. Lambert’s specimen came from the late Cretaceous of Fanivelona, eastern Madagascar. _ It has a very similar overall shape, but unfortunately the apex of the test is damaged and only the anterior petal on one side is preserved. It is only tentatively assigned to this species. Hemiaster paronai Checchia-Rispoli, 1921 Pl. 33, figs 1-4; Figs 81A, B 1921 Hemiaster Paronai Checchia-Rispoli: 27, pl. 8, fig. 24, pl. 9, figs 14-18. | 1932 Hemiaster Paronai Checchia-Rispoli; Rispoli: 8, pl. 2, figs 1-8. 21967 Hemiaster punctatus d’Orbigny; Devries: 194, pl. 6, . figs 34-41. 21967 Hemiaster regulusanus d@’Orbigny; Devries: 194, pl. 6, figs 42-44. Checchia- MATERIAL. Seven specimens, only one of which is well- _ preserved, BMNH EE5034. ) OCCURRENCE. Specimens were found at the following locali- ties and horizons in the western Oman mountains: | Jebel Huwayyah, section 1: bed 9 (1); bed 11 (1). | Jebel Huwayyah, section 2: beds 3-5 (3). _ Jebel Faiyah, section 1: bed 4 (1). Jebel Rawdah, section 3: bed 9 (1). | | 233 Elsewhere the species has been recorded from the Maastrich- tian of northern Libya. DIAGNOsIs. Like H. hattaensis but more inflated and more circular in outline, with deeper petals and deeper and wider anterior sulcus adapically. DESCRIPTION. The best preserved specimen is 37-1 mm in length, 35-6 mm in width (96% of length: widest point approximately mid-length) and 28-8 mm in height (78% of length). In outline the test is more or less circular and in profile the upper and lower surfaces are broad and flat with the anterior uniformly rounded and the posterior sloping steeply outwards (PI. 33, figs 1-4). The apical disc is ethmophract with the posterior two genital plates rather stout and broadly in contact (Fig. 81B). The posterior ocular plates are separated. Gonopores are large and occupy most of the plate, but in some individuals (?males) may be relatively smaller. Genital plate 2 has a small central zone of madrepores. The apical disc lies 58% of the test length from the anterior border. The anterior ambulacrum lies in a frontal sulcus which is bordered by sharp interambulacral crests adapically but which shallows and is lost towards the ambitus (PI. 33, fig. 2). There are 22 pore-pairs in a column between the apex and the peripetalous fasciole and these are strongly oblique. The sulcus is rather narrow and slightly lanceolate in outline. The petals are cruciform with the anterior pair twice the length of the posterior pair (Fig. 81A). There are 40 pore- pairs in a column in the anterior pair and 21 in the posterior pair at 37 mm test length. Petals close distally and the perradial interporal zone is narrower than either of the bordering pore zones. The periproct is oval, slightly pointed adapically and lies high on the posterior side. It is just visible from above because of the outward slope of the posterior face. It is about 70% as wide as tall and is 18% of the test height in height. Its base lies 60% of test height above the base (PI. 33, fig. 4). The peristome is D-shaped, twice as wide as long and lies 28% of the test length from the anterior border. It is not rimmed. There are five phyllode pores in each column of lateral and posterior ambulacra. There are also five subanal pore-pairs in the two posterior interambulacra. REMARKS. This species closely resembles H. hattaensis in the shape of petals, apical disc structure and the form of the anterior ambulacrum. However, it differs consistently in shape, being both more rounded in outline and more inflated in profile. This is not simply an attribute of size, since the small specimens from Jebel Huwayyah (eg. BMNH EE4061, 22 mm in length) are very different in shape from similar- sized individuals from Jebel Buhays. In addition the pore- pairs in the frontal groove are more numerous and more densely packed. The specimens appear indistinguishable from _ those described from the Maastrichtian of Libya by Checchia- Rispoli (1921) under the name H. paronai. Very similar material was also described by Devries (1967) from the Maastrichtian of Turkey under the names Hemiaster puncta- tus d’Orbigny and H. regulusanus d’Orbigny. Unlike the specimens described here and by Devries (1967), H. puncta- tus has a large flush peristome lacking a rim. The species is also very similar to Hemiaster noemiae Cotteau & Gauthier, from the late Senonian of southern Iran (Pl. 33, figs 5-8). However, H. noemiae differs in having a shorter anterior 234 ; A.B. SMITH PLATE 33 Figs 1-4 Hemiaster paronai Checchia-Rispoli. BMNH EES034; 1, oral; 2, apical; 3, lateral; 4, posterior; all x 2. Jebel Rawdah, section 3, bed 9. Figs 5-8 Hemiaster noemiae Cotteau & Gauthier. B18727, Morgan Collection, Museum d’Histoire Naturelle, Paris; 5, oral;6, apical; 7, posterior; 8, lateral; all x 2. Senonian of Awasa, Iran. Figs 9-11 ?Linthia sudanensis (Bather). BMNH EE4054; 9, lateral; 10, apical; 11, oral; all x 1-5. Loose near top of section at Jebel Rawdah, section 3. LATE CAMPANIAN-MAASTRICHTIAN ECHINOIDS 2B5 PLATE 34 Figs 1-3, 8 Proraster geayi Cottreau. 1-3, specimen in Museum d’Histoire Naturelle, Paris; 1, apical; 2, oral; 3, lateral; all x Maastrichtian of Antanihody, Madagascar. 8, BMNH EE4067, apical, x 1. Jebel Huwayyah, section 1, bed 18. | Figs 4-7 Mecaster victoris (Lambert). Jebel’ Rawdah, section 3, bed 5. 4, BMNH EE4050; apical, x 2. 5, BMNH EE4049; oral, * 25 Grills BMNH EE404?2; 6, apical; 7, lateral; both x 2. i) 236 sulcus with many fewer pore-pairs, and in having a much smaller peristome at comparable sizes. Genus MECASTER Pomel, 1883 Mecaster victoris (Lambert, 1932) PI. 34, figs 4-7; Figs 82-84A 1932 Hemiaster Victoris Lambert: 127, pl. 4, figs 18, 19. 1967 Hemiaster sp. Devries: 194, pl. 6, figs 45—47. 1990 Periaster subsexangulatus Airaghi; Ali: 410, fig. 5 (8). MATERIAL STUDIED. Ten reasonably complete specimens were used in the biometric analysis, BMNH EE4035—36, EE4038-40, EE4042, EE4045—-46, EES035-36. Another 32 specimens were collected. OCCURRENCE. In the study area, this species was found at the following localities and horizons: Jebel Buhays, section 1: loose in the scree, derived from the lowest few metres of the Simsima Formation (1). Jebel Huwayyah, section 1: bed 14 (1). Jebel Huwayyah, section 2: beds 3-5 (3). Jebel Rawdah, section 3 (top of bed 5 (28): bed 8 (2); beds 9/10 (7). The species was originally described from the Campanian of El Kantara, Algeria. A poorly preserved specimen that also appears identical was described from the Maastrichtian of Cortinek, Turkey. DESCRIPTION. Specimens range in size from 24 to 33 mm in length. Mean test width is 98% of the length (range 93-100%, SD = 3:2%, N = 9) and test height 76% of the length (range 72-80% , SD = 3-:1%, N = 7). The test is weakly cordiform in outline, tapering posteriorly to a rounded point and with the widest point in front of the mid-line (Pl. 34, figs 4-6). In profile the test is inflated with a rounded posterior and anterior (Pl. 34, fig. 7). The tallest point is posterior of the apical disc. The apical disc lies 36-40% of the test length from the anterior border (mean = 38%, SD = 1-4%, N = 8). It is ethmolytic with the posterior oculars separated by the madreporite (Fig. 84A). The gonopores on each side of the mid-line lie close together. Although ambulacra are sunken, they are all rather shallow and relatively narrow (PI. 34, figs 4, 6; Fig. 833A). Pore-pairs in the anteal sulcus are small and situated along the adradial margins. The anterior petals are long, narrow and straight- sided. They extend most of the way to the ambitus and diverge from one another at an angle of 130°. There are 33 pore-pairs in a column at 25 mm test length, rising to 36 at 29 mm test length. The pore-pairs are narrow, with the outer pore in each pair slightly more elongate than the inner. The posterior petals extend about 75% of the distance to the ambitus and are equally narrow and parallel-sided. They are about 85-90% of the length of the anterior petals and have 28 pore-pairs in a column at a test diameter of 25 mm, rising to about 32 at 28 mm. The periproct is high on the posterior and is usually just visible from above. The base lies more or less half way up the posterior face (mean = 50%, SD = 4:5%, N = 4). The opening is vertically elongate, being about 1-6—1-8 times as tall as broad, and is pointed adapically. A.B. SMITH The peristome is D-shaped and lies about 20% of the test length back from the anterior border (range 18-5—21-5%, SD | = 1:1%, N = 6). The labrum projects slightly over the | opening. The labral plate is relatively long and narrow, almost parallel-sided (Fig. 83B). It meets only one of the | sternal plates. Tuberculation is fine adapically, slightly coarser adorally. | There is always a peripetalous fasciole, some 0-6 mm in width. Although preservation is usually too poor to allow its course to be traced fully, there appears to be little indentation of the peripetalous fasciole laterally and it runs close to the | ambitus. In one specimen (BMNH EE4050) there is a distinct | latero-anal fasciole also, but other specimens, equally well preserved in this region have only rudimentary traces of such a fasciole or no fasciole at all. It is clear that the latero-anal | fasciole is variably developed in this species. } REMARKS. Lambert (1932: 127) created this species on the basis of material from the late Campanian of El Kantara, | Tunisia. It has narrower, shallower petals than other Mecaster species and in profile is inflated with the tallest point lying well to the posterior. Lambert noted that there 5 traces of a latero-anal fasciolt in six out of twelve specimens, | while another three had a distinct latero-anal fasciole. Lam- bert separated those with a latero-anal fasciole and placed them in his ‘Periaster Victoris’, though recognising that they were completely intergradational with Hemiaster victoris. As Lambert quite correctly pointed out, the presence/absence of a latero-anal fasciole is highly variable and the various species classified under the genus Periaster are polyphyletic in origin. |— Ali (1989) figured a specimen of this species without . description under the name Periaster subsexangulatus Air- aghi. However, P. subsexangulatus has a much broader, |_ deeper anteal sulcus, is less rounded and less inflated and i most particularly, the petals are broader and less parallel- | sided and the posterior pair are distinctly shorter than the | anterior pair. The plastron structure is significant in that the labral plate is in contact with sternal plate 2b only (Fig. 83B). This is also the situation in /raniaster and other Somaliasteridae. /rant- aster has been placed in the Holasteroida on account of its plastron structure, yet it has a compact apical disc in which the posterior genital plates, but not the posterior ocular plates, are separated by the madreporite (Kier 1972, figs 41-42). The fact that a meridoplacous plastron can be devel- oped in at least one Mecaster species, and the similarity of apical disc plating strongly indicates that somaliasterids are derived from the Mecaster lineage. ge a eS ee Genus LINTHIA Desor, 1853 4 ?Linthia sudanensis (Bather, 1904) __ PI. 33, figs 9-11; Figs 84B, 85 21904 Hemiaster sudanensis Bather: 299, pl. 11, figs 6-13. Family SCHIZASTERIDAE Lambert, 1905 i MATERIAL. One specimen, BMNH EE4054. OCCURRENCE. The specimen was found loose about 8 m below the top of section 3 at Jebel Rawdah, western Oman. The section here is capped by a two metre thick conglomerate of reworked Simsima Limestone and thus the specimen could be of late Maastrichtian age. LATE CAMPANIAN-MAASTRICHTIAN ECHINOIDS 40 | = Test width | @ Test height 22 24 26 28 30 32 Test length (mm) gs Anterior to apical disc @ Anterior to peristome — Distance (mm) Test length (mm) | Fig. 82 Biometric data for Mecaster victoris (Lambert). EE4049. Scale bar = 2 mm. DESCRIPTION. The specimen is not well preserved but retains ‘sufficient features to be able to place it generically. The test is 32-5 mm in length, 30-5 mm in width (93% of length) and 24 mm in height (74% of length). It is heart-shaped in outline ‘with a squarely truncated posterior and a rather deep anteal ‘sulcus, approximately 2-5 mm deep (7% of test length). The anteal sulcus is relatively broad and deepens gradually away from the apex. In profile the test is rather flat above and below and has a rounded anterior and steep posterior. Length of petal (amb. Il) (mm) ye \ raga 237 22 24 26 28 30 32 34 Test length (mm) 5.0 a a 24 26 28 30 32 34 Test length (mm) see | Fig. 83 Camera lucida drawings of plating in Mecaster victoris (Lambert). A, apical surface, BMNH EE4050; B, oral surface, BMNH The apical disc is ethmolytic and gonopores are large, those on either side almost touching (Fig. 84B). It lies well anterior of the centre, the anterior gonopores being 38% of the test length from the anterior border. The posterior ocular plates are separated by the posteriorly elongated madreporite plate. The antero-lateral ocular plates project and hardly indent the adjacent genital plates. The anterior sulcus has small isopores that are not crowded together. The anterior petals are 13-5 mm in length and Fig. 84 Camera lucida drawing of apical discs. A, Mecaster victoris (Lambert), BMNH EE4050. B, ?Linthia sudanensis (Bather), BMNH EE4054. Scale bar = 1 mm. NN ee a Fig. 85 Camera lucida drawing of ?Linthia sudanensis (Bather), BMNH EE4054. A, apical; B, oral; C, lateral. Scale bar = 5 mm. widely divergent (Fig. 85). They extend for most of the distance to the ambitus when viewed from above. Inner and outer pores are equally slit-like and there are 33 pore-pairs in a column. The posterior petals are shorter, only 10 mm in length, and reach approximately half-way towards the ambi- tus. There are 24 pore-pairs in a column. The periproct is just visible from above. It is 6-5 mm in height and 4 mm in width, being pointed adapically. The base of the periproct lies just above mid-height, some 11-3 mm above the base of the test (52% of test height). The plastron is asymmetric, with the median suture dis- placed towards the right. The labral plate is short (11-5 of the test length) and trapezoidal in outline. The peristome is D-shaped although the labral plate does not project over the peristome much. The peristome is set rather far back from the anterior border, its anterior margin being 23% of the test length from the anterior. REMARKS. The specimen is unfortunately very badly pre- served and cannot be identified to species level with any certainty. However, from the general shape and petal form it cannot be distinguished from the common Palaeocene species Linthia sudanensis described by Bather (1904) from Sudan. A.B. SMITH Genus PRORASTER Lambert, 1895 Pl. 34, figs 1-3, 8 | Proraster geayi Cottreau, 1908 1908 Proraster Geayi Cottreau: 26, pl. 4, fig. 5. | Types. The syntypes of P. geayi are the two specimens described from Marohita, eastern Madagascar by Cottreau (1908) stated to be in the Museum d’Histoire Naturelle, Paris. MATERIAL STUDIED. Six specimens, BMNH EE4065-69, EE5037, none of which are particularly well-preserved. occurs as juveniles in beds 3—5 at Jebel Huwayyah, section 2 (2). Large individuals occur in bed 18 at the top of section 1, Jebel Huwayyah (7), and in bed 9 at Jebel Rawdah, section 1 | (2). It is also found in the Maastrichtian of Madagascar. | OCCURRENCE. In the western Oman Mountains this species | \ DIAGNOsIs. A Proraster with a very deep anteal sulcus that is closed or almost closed distally by convergence of the lateral — walls. DESCRIPTION. No test is well-preserved, the best specimens being BMNH EE4067 and EE4068. The former is a large individual 71 mm in length, 56 mm in width and about 27 mm in height, whereas the latter is a much smaller individual 27 mm in length, 24 mm in width and 15-6 mm in height. The test is oval in outline with its widest point slightly posterior of midline (Pl. 34, fig. 8). In profile the test is depressed and | 7 wedge-shaped, sloping gradually towards the anterior. The | posterior is almost vertical and the tallest point on the test lies | close to the posterior. The apical disc lies well towards the posterior, 66-69% of test length from the anterior border. Plating is not seen in any specimen. The anterior ambulacrum is extremely deep and at the) ~ ambitus is about 10% of the test length in depth. The walls are concave with the adjacent interambulacra curving over | | the groove. Towards the anterior the two sides almost touch | so as to roof over the sulcus. The groove thus appears pinched shut close to the anterior (PI. 34, figs 1, 3, 8). Petals are sunken. The anterior pair curve forward and run) - parallel with the anteal sulcus for about two thirds of its) — length. The posterior petals are very much shorter (about one third of the length of the anterior petals) and diverge strongly at about 120°. There is a well developed peripetalous fasciole developed at the base of the petals, which is presumably | continuous, but is not seen other than in small patches because of poor preservation. There is no evidence of a} | latero-anal fasciole. 3 The peristome is positioned far forward, lying at the base) » of the anteal sulcus, and faces anteriorly. The periproct is) ~ large and situated high on the posterior face. | REMARKS. Although the Omani specimens to hand are not well-preserved they show sufficient detail to be unambiguously |, assigned to this species. P. geayi was first erected for large| . individuals from the Maastrichtian of Madagascar by Lambert (1908). Lambert (1905) had previously erected the genus Pro- raster for Schizaster atavus (Arnaud) from the Campanian of Charante, France and two Iranian species Opissaster morgani (wrongly cited as O. Douvillei) and O. centrosus Cotteau &| ~ Gauthier (1895). The genus was erected to encompass Schizaster- like forms lacking a latero-anal fasciole. The species of Cotteau & Gauthier are unfortunately based LATE CAMPANIAN-MAASTRICHTIAN ECHINOIDS on small individuals and are thus not directly comparable with P. geayi. Differences between P. morgani and P. centrosus seem slight and restudy may prove them synony- mous. Because they are small, there is the possibility that they represent juveniles of P. geayi. However, this seems unlikely since they have much wider and less pinched anterior sulci than very slightly larger individuals of P. geayi found in the lower beds at Jebel Huwayyah. This pinching of the frontal groove, whereby the two sides converge and almost touch near the anterior border, becomes very much more pronounced in the larger individuals higher in the section. This character serves to distinguish P. geayi from both P. atavus, and an undescribed ?Campanian species from Nafun, Oman. Cotteau & Gauthier’s species also have a much less well developed anterior notch and their peristome is standard in orientation rather than being subvertical and directed towards the anterior. It is noteworthy that at Jebel Huwayyah only small indi- viduals are found in the lower levels, in the Loftusia facies, while large individuals are found near the top of the section in carbonate marls representing shelf basinal facies. This could be because the two samples represent different species with consistent morphological differences, or because the smaller individuals are juveniles of the larger, but inhabit a different biotope. Without considerably more and better preserved material it is impossible to determine which is correct. For the present I treat the two forms as juveniles and adults of the same species, partially because the small forms are much closer in appearance to the large forms than they are to, for example, P. morgani. REFERENCES Agassiz, A. 1863. 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(Geol.) 51(2):241-250 Issued 30 November 1995 Maastrichtian ammonites from the United Arab Emirates-Oman border region W.J. KENNEDY Department of Earth Sciences, University of Oxford, Parks Road, Oxford OX1 3PR. INTRODUCTION The ammonites described are from the late Cretaceous Qahlah and Simsima Formations of the United Arab Emirates-Oman border region and come from two collec- tions. Some of the specimens were collected by Dr. P.W. Skelton (Open University) in 1990 and are housed in Oxford University Museum (OUM prefixes). The remainder are housed in the Natural History Museum, London (BMNH prefixes) and were mostly collected by A.B. Smith and N.J. Morris during fieldwork in 1991 and 1992. A few specimens were collected by amateur enthusiasts and subsequently donated to the Natural History Museum. Wherever possible, ammonites have been tied down to specific levels within measured sections. Details of the locali- ties of these sections, and measured lithological logs are given in a preceding section (Smith ef al.). SYSTEMATIC DESCRIPTIONS Order AMMONOIDEA Zittel, 1884 Suborder AMMONITINA Hyatt, 1889 Superfamily DESMOCERATACEAE Zittel, 1895 Family DESMOCERATIDAE Zittel, 1895 Subfamily DESMOCERATINAE Zittel, 1895 Genus DESMOPHYLLITES Spath, 1929 (= Schliiteria de Grossouvre, 1894: 126 (non Fritsch in Fritsch & Kafka, 1887: 33); Schliitericeras Collignon, 1938: 92 (non Hyatt, 1903: 110)) TYPE SPECIES. Desmoceras larteti Seunes, 1892: 19, pl. 12 (3), fig. 2; pl. 13 (4), figs 2, 3, by subsequent designation by Spath 1921: 46, as type species of Schliiteria de Grossourvre, of ’ which Desmophyllites is the replacement name. _ Desmophyllites diphylloides (Forbes, 1846) Plate 1, figs il, 2 1846 Ammonites diphylloides Forbes: 105, pl. 8, fig. 8. 1992 Desmophyllites diphylloides (Forbes); Kennedy & | Henderson: 405, pl. 6, figs 1-9; pl. 16, figs 1-3, 7-8; pl. 17, figs 4~7; fig. 3F (with full synonymy). 19834 Desmophyllites diphylloides (Forbes); Kennedy & Cobban: 120, pl. 1, figs 1-8; text-fig. 5c. TyPEs. Lectotype, by the subsequent designation of Matsu- © The Natural History Museum, 1995 moto & Obata (1955: 122) is BMNH C22682, the original of Forbes 1846: pl. 8, fig. 8; paralectotypes are BMNH C22683-85, all from the Upper Maastrichtian Valudavur Formation of Pondicherry, southern India. DESCRIPTION. BMNH C9399? consists of 270° of body cham- ber and the nucleus of an individual with an estimated original diameter of 62 mm. Coiling is very involute, with a tiny, pitlike umbilicus, the umbilical wall subvertical and narrowly rounded. The whorl section is compressed, with whorl breadth to height ratio 0-71, the greatest breadth around mid-flank, the flanks feebly convex and subparallel, and the ventrolateral shoulders and venter evenly rounded. The surface of replaced shell and internal mould are smooth, but for a single constriction approximately 70° from the aperture. This is narrow, shallow and markedly prorsiradiate, straight on the inner flank, feebly convex on the outer flank and concave on the outermost flank. Sutures not seen. DISCUSSION. The types of Desmophyllites diphylloides are all rather small (Kennedy & Henderson 1992, pl. 6, figs 1-9; pl. 16, figs 1-3, 7, 8), but Henderson and McNamara (1985: 54, pl. 4, figs 1-4) described larger specimens comparable to the present individual from the Upper Maastrichtian of Western Australia. These show constrictions that are flexuous on the flanks, as with the present material. The other Maastrichtian Desmophyllites is D. larteti (Seunes, 1892) (p. 19, pl. 12 (3), fig. 2; pl. 13 (4), figs 2, 3; see Hancock & Kennedy 1993: 154, pl. 2, figs 1-3, 10, 11, 14; pl. 3, figs 1, 4, 5), which is a larger, much more compressed species with a narrower, arched venter and much more prominent and markedly fiexuous constrictions. OCCURRENCE. The Oman specimen is from bed 21, Jebel Rawdah, section 2. Elsewhere, the species ranges from Lower Santonian to Upper Maastrichtian. There are records from southern India, Western Australia, Japan; Alaska, British Columbia, California, and Arkansas in the U.S.A.; Argentina, Angola; Pondoland and Zululand (South Africa), Tunisia, and southern France (Corbieres). Family KOSSMATICERATIDAE Spath, 1922 Subfamily KOSSMATICERATINAE Spath, 1922 Genus BRAHMAITES Kossmat, 1897 TYPE SPECIES. Ammonites Brahma Forbes, 1846: 100, pl. 8, fig. 1, by original designation. PLATE 1 Figs 1,2 Desmophyllites diphylloides (Forbes, 1846). BMNH C93992, Simsima Formation, Jebel Rawdah, section 2, bed 21. Figs 3-7 Pachydiscus (Pachydiscus) neubergicus neubergicus (Hauer, 1858). 3, 4, OUM KY 1991, from an unknown level in the Simsima Formation, southern embayment of Jebel Rawdah. 5, 6, OUM KY 1990, from an unknown level in the Simsima Formation, southern | embayment of Jebel Rawdah. 7, OUM KY 1995, from the basal part of the Simsima Formation, Jebel Huwayyah, close to where the road | cuts the north-western part of the outcrop. All figures are < 1. , | | MAASTRICHTIAN AMMONITES Subgenus ANABRAHMAITES Yabe & Shimizu, 1924 (= Subbrahmaites Yabe & Shimizu, 1924: 75). TYPE SPECIES. Ammonites vishnu Forbes, 1846: 100, pl. 7, fig. 9, by original designation. Brahmaites (Anabrahmaites) vishnu (Forbes, 1846) Pl. 2, figs 9-14 1846 Ammonites vishnu Forbes: 100, pl. 7, fig. 9. 1992 Brahmaites (Anabrahmaites) vishnu (Forbes); Kennedy & Henderson: 418, pl. 6, figs 25, 26; pl. 9, figs 5—7, 17-20; pl. 10, fig. 5; pl. 17, figs 8, 10-11 (with full synonymy) Brahmaites (Anabrahmaites} vishnu (Forbes); Ken- nedy & Hancock: 582, pl. 1, figs 5, 6. TYPES. Lectotype, by the subsequent designation of Kennedy & Henderson (1992: 418), is BMNH C51026, the original of Forbes 1846, pl. 7, fig. 9; BMNH C51027 is a paralectotype. Both are from the Upper Maastrichtian Valu- davur Formation of Pondicherry, southern India. DESCRIPTION. BMNH C93892 (Pl. 2, figs 11-14) consists of 200° of adult phragmocone and body chamber with a maxi- mum preserved diameter of 83 mm, and part of the septate inner whorls, 48 mm diameter. Coiling is very evolute, the shallow umbilicus comprises 55% of diameter, with a broadly rounded umbilical wall and shoulder. The whorl section is depressed reniform, with greatest breadth just outside the umbilical shoulder, and the whorl breadth to height ratio is 1-1. The inner whorls (PI. 2, figs 13, 14) are badly preserved, but traces of delicate crowded ribs are present on one flank. Both flanks have well-developed, prorsiradiate, deep, narrow constrictions, 2 per half whorl, flanked by a strong adapical collar-rib with feeble umbilical bulla, and a much weaker adapertural rib, the ribs extending across the venter. The outer whorl of the specimen is worn, but shows similar constrictions and collar ribs, with traces of coarse prorsiradi- ate flank ribs between constrictions. BMNH C93891 is a much larger body chamber fragment (PI. 2, figs 9, 10), with maximum preserved whorl height 27-7 mm and whorl breadth to height ratio 1-0. There is a single strong constriction, flanked by a bullate adapical collar rib that persists over the venter, where it is strengthened into an incipient bulla (the specimen is worn). There is a much weaker adapertural collar-rib, and blunt non-bullate ribs, straight and prorsiradi- ate on the flanks, weakened and feebly convex across the venter, cover the remainder of the fragment. Sutures not seen. 1993 DISCUSSION. These fragments are referred to Anabrahmaites rather than Brahmaites sensu stricto on the basis of the absence of bullae on intermediate ribs and the development of an incipient siphonal bulla. The inner whorls of BMNH C93892 differ in no respects from those of the paralectotype of B. (A.) vishnu figured by Kennedy & Henderson (1992, pl. 10, fig. 5); the outer whorls of this specimen and BMNH / €93891 also find a match in the Pondicherry material (see also Stoliczka 1865, pl. 79, fig 5). OCCURRENCE. The Oman specimens are from bed 3 or 4, Jebel Rawdah, section 1. The types are from the Upper Maastrichtian of southern India; the holotype of Puzosia haugi Seunes, 1892, a synonym, is from the Upper Maastrich- tian fresvillensis Zone of southern France. The species also 243 occurs in the Maastrichtian of northern Spain and Armenia. Family PACHYDISCIDAE Spath, 1922 Genus PACHYDISCUS Zittel, 1884 Subgenus PACHYDISCUS Zittel, 1884 TYPE SPECIES. Ammonites neubergicus Hauer, 1858: 12, pl. 2, figs 1-3; pl. 3, figs 1, 2, by the subsequent designation of de Grossouvre 1894: 177. Pachydiscus (Pachydiscus) dossantoi (Maury, 1930) PI. 3: BIA fhigsrie 2 Parapachydiscus dossantoi Maury: 136, pl. 16, fig. 1; pl. 17, figs 1, 2. 1930 21944 Parapachydiscus sp. Olsson: 107, pl. 16, fig. 1. 1985 Pachydiscus (Pachydiscus) dossantoi (Maury, 1930); Zaborski: 20, figs 17, 18, 20. Types. Maury (1930: 136, pl. 16, fig. 1; pl. 17, figs 1, 2) based this species on a number of specimens, referring to an individual 190 mm in diameter as the type. All are from the Maastrichtian on the right bank of Rio Gramame, Fazenda do Congo, Parahyba do Norte, Brazil. DESCRIPTION. Large, septate to whorl heights of up to 110 mm. Evolute, with 50% of previous whorl covered; umbilicus broad, shallow, with flattened, outward-inclined umbilical wall and broadly rounded umbilical shoulder. Whorl section compressed, with whorl breadth to height ratio of 0-8 in the best-preserved specimen; greatest breadth low on broadly convex flanks; outer flanks convergent; ventrolateral shoul- ders and venter arched. Coarse, distant ribs arise at blunt umbilical bullae, are straight and prorsiradiate across the flank, alternating regularly with shorter ribs that arise on the outer flank. Ribs sweep forwards across the ventrolateral shoulders and are weakened and broadly convex across the venter. Ornament of this type extends to the end of the phragmocone in BMNH C93895-96 and onto the body cham- ber in BMNH C93894. Suture poorly preserved (Pl. 3); intricately and deeply subdivided, as is typical for the genus. DISCUSSION. Compressed whorl section plus persistence of coarse, alternately long and short ribs to a large size show these specimens to belong to P. (P.) dossantoi. Of other Maastrichtian species, P. (P.) neubergicus (Hauer, 1858) (see revision in Kennedy & Summesberger 1986), P. (P.) gollevil- lensis (d’Orbigny, 1850) (see revision in Kennedy 1986) and P. (P.) egertoni (Forbes, 1846) (see revision in Kennedy & Henderson 1992) are also compressed, but all are more delicately ribbed, are mature at smaller diameters, and have adult growth stages characterized by loss of outer flank and ventral ornament. P. (P.) jacquoti Seunes, 1890 (see revision in Kennedy 1986) has a depressed whorl section, distant ribs and effacement of outer flank and ventral ornament on the phragmocone. OCCURRENCE. Most of the Oman specimens come from the lower Loftusia-rich beds at Jebel Huwayyah. BMNH C93894 comes from bed 10 or 11, section 1, Jebel Huwayyah, while C93895 comes from bed 10. OUM K1998 (= Skelton 84/32:-2) is also from Jebel Huwayyah, section 1, from an unspecified level. One specimen, BMNH C93896, comes from a loose block derived from the basal bed of the Simsima Formation at Jebel Buhays, section 1. The type occurrence is of Maastrich- 244 tian age, but is difficult to place more precisely within the stage. Kennedy (1986: 44) regarded Pachydiscus sumneri Maury, 1930 (p. 155, pl. 13, figs 1, 2), Parapachydiscus poseidon Maury, 1930 (p. 155, pl. 15) and Canadoceras riogramense Maury, 1930 (p. 169, pl. 21, fig. 2), which are said to occur in the same unit as P. (P.) dossantoi, as possible synonyms of the Upper Maastrichtian Anapachydiscus fresvil- lensis (Seunes, 1890). The original figures are so poor, however, and the relative position of species within Maury’s ‘grey limestone’ is unknown, so that only the possibility of an Upper Maastrichtian date can be considered. P. (P.) dossan- toi from Nigeria (Zaborski 1983, 1985) comes from the Nkoporo Shale, and was regarded as ‘probably Lower Maas- trichtian’ (no definition of the Lower/Upper Maastrichtian boundary was given). It co-occurs with Gaudryceras beantaly- ense Collignon, 1956, Baculites sp. and Sphenodiscus lobatus costatus Zaborski, 1982. On this evidence it can be dated no more precisely than Maastrichtian. Pachydiscus (Pachydiscus) neubergicus neubergicus (Hauer, 1858) Plate 1, figs 3-7 1858 Ammonites neubergicus Hauer: 12 (pars), pl. 2, figs 1-3 (non pl. 3, figs 1, 2). Pachydiscus (Pachydiscus) neubergicus neubergicus (Hauer, 1858); Hancock & Kennedy: 158, pl. 3, figs 6, 7; pl. 9, figs 5-8; pl. 12, figs 7-9; pl. 13, figs S—7 (with synonymy). 1993 Types. Lectotype, by the subsequent designation of de Gros- souvre 1894: 209, is no 1858.01.6 in the collections of the Geologisches Bundesanstalt, Vienna; three paralectotypes bear the same number, and all are from the Lower Maastrich- tian of Neuberg, Steiermark, Austria. DESCRIPTION. The best-preserved specimen is OUM KY 1990 (PI. 1, figs 5, 6), an internal mould of a phragmocone 93 mm in diameter. Coiling is fairly involute, the umbilicus small, with a flattened, subvertical low wall and narrowly rounded umbilical shoulder. The whorl section is com- pressed, with a whorl breadth to height ratio of 0-74, the greatest breadth below mid-flank, inner flanks feebly convex, outer flanks flattened and convergent, ventrolateral shoul- ders broadly rounded, and the venter only feebly convex. There are an estimated 14 umbilical bullae per whorl. These give rise to single ribs, sometimes feebly concave on the innermost flank. They are prorsiradiate and weak across the flanks, where long and short intercalated ribs are inserted, so that there are many more coarse concave ribs at the ventro- lateral shoulder, although the number per whorl cannot be determined. OUM KY1994 is a worn but conspecific frag- ment of phragmocone with a maximum preserved whorl PLATE 2 W.J. KENNEDY height of 36 mm and whorl breadth to height ratio of approximately 0-89. OUM K1991-93 are fragments of body chamber (PI. 1, figs 3, 4), with a maximum preserved whorl height of 51-5 mm and whorl breadth to height ratio of 0-75. At the adapical end of the fragment strong bullate primary ribs at the umbilical shoulder correspond to 3-4 times as many ribs at the ventro- lateral shoulder. On the adapical part of the fragment the secondary ribs are lost, and the ornament is of distant primaries that efface across the flanks. OUM K1995 (PI. 1, fig. 7) is a very worn individual 150 mm in diameter, probably adult, with a phragmocone diameter of 110 mm and whorl breadth to height ratio of 0-78. There appear to be 14-16 primary ribs per whorl, and more numerous primary plus secondary ribs at the ventrolateral shoulder. Sutures not seen. DISCUSSION. Coiling, whorl proportions and ribbing show these specimens to belong to the Pachydiscus (P.) neuber- gicus neubergicus (Hauer, 1858) — P. (P.) gollevillensis (d’Orbigny, 1850) group. Topotypes of the former generally have 14-17 umbilical bullae and 58-60 ventral ribs per whorl (Kennedy & Summesberger 1986), the latter 9-11 umbilical bullae and approximately 80 ventral ribs (Kennedy, 1986), suggesting the present material belongs to the former. Nuclei of P. (P.) neubergicus neubergicus and P. (P.) neubergicus dissitus Henderson & McNamara, 1985, are identical, but the latter has a mature ornament with many ventral ribs (Hender- son & McNamara 1985, pl. 7, fig. 7), not seen in the present material (Pl. 1, figs 3, 4), which are thus referred to the nominate subspecies. OCCURRENCE. OUM KY1990, 1991-93 (1 specimen) and KY 1994 were collected in the southern embayment of Jebel Rawdah. Their position in the succession is unknown. KY1995 comes from the lower part of the Simsima formation in the north-western part of Jebel Huwayyah, close to where the road cuts the outcrop. Elsewhere the species first appears low in the Lower Maastrichtian, and is best known from the Lower Maastrichtian of Austria, Poland, Ukraine, Armenia, Russia, SW France, northern Spain, Nigeria, Brazil and Zululand (South Africa). It occurs in the lower Upper Maastrichtian of Denmark, and the Upper Maastrichtian of southern India. Indeterminate pachydiscid Discussion. BMNH C93987 is a fragmentary pachydiscid, still septate at a whorl height of 150 mm. It bears distant primary ribs, but is indeterminate even as to genus. OCCURRENCE. The specimen comes from the gritty calcare- ous bed (bed 9), of the Qahlah Formation, immediately Figs 1-3, 15, 16 Lewyites ambindense (Collignon, 1971). 13, BMNH C93890; from the Loftusia-rich beds, Qahlah Formation, Jebel Huwayyah, section 1, bed 10 or 11. 15, 16, OUM KY1996; from the basal part of the Simsima Formation, north-western Jebel Huwayyah, close to where the road cuts the outcrop. Figs 4-8 Nostoceras (Nostoceras) major Kennedy and Cobban, 1993. 4-6, BMNH C€93994, from beds 10 or 11, Qahlah Formation, Jebel Huwayyah, section 1; 7, 8, BMNH C93993, Basal Simsima Formation, bed 9, Jebel Bu Milh, section 2. Figs 9-14 Brahmaites (Anabrahmaites) vishnu (Forbes, 1846). 9, 10, BMNH C93891; 11-14, BMNH C93892. Both from the Simsima Formation, bed 3 or 4, Jebel Rawdah, section 1. Figs 17-19 Libycoceras? sp., BMNH C93887, from the conglomeratic basal bed of the Simsima Formation, Jebel Buhays, section 2. Fig. 20 Nostoceras (Nostoceras) sp., BMNH C93888, from bed 6, Jebel Huwayyah, section 2. All figures are x 1 | MAASTRICHTIAN AMMONITES 246 W.J. KENNEDY PLATE 3 Pachydiscus (Pachydiscus) dossantoi (Maury, 1930). BMNH C93896, from the basal bed of the Simsima Formation, Jebel Buhays, section 1; _ slightly reduced. MAASTRICHTIAN AMMONITES below the Loftusia-rich beds at Jebel Huwayyah, section 1. Superfamily ACANTHOCERATACEAE de Grossouvre, 1894 Family SPHENODISCIDAE Hyatt, 1900 (= Libycoceratidae Zaborski, 1982: 306) Genus LIBYCOCERAS Hyatt, 1900 TYPE SPECIES. Sphenodiscus ismaelis Zittel, 1895, p. 451, text-fig. 631, by original designation. Libycoceras ? sp. Plate 2, figs 17-19 DESCRIPTION. BMNH C93887 is a very corroded internal mould of a small adult with 240° of body chamber preserved and a maximum diameter of 80 mm. The phragmocone is oxycone, with a whorl breadth to height ratio of 0:4. The body chamber develops subparallel flanks, and the venter rounds progressively; the whorl breadth to height ratio is 0-39 at the aperture. No ornament is preserved on the heavily corroded surface, other than a faint trace of low ribs on the outer flank. The suture (Text-fig. 1) shows a well-developed adventitious lobe in the first lateral saddle, the ventral saddles are feebly indented, and the umbilical saddle is entire. DISCUSSION. This poorly preserved specimen is referred to Libycoceras? rather than Sphenodiscus on the basis of its sutural characteristics; it is specifically indeterminate. OCCURRENCE. BMNH (C93887 is from the basal conglomer- atic bed of the Simsima Formation at Jebel Buhays, section 2. Species of Libycoceras first occur in the Upper Campanian and may range into the Lower Maastrichtian (Zaborski 1982). Suborder ANCYLOCERATINA Wiedmann, 1966 Superfamily TURRILITACEAE Gill, 1871 Family DIPLOMOCERATIDAE Spath, 1926 (= Neocrioceratinae Spath, 1953) Subfamily DIPLOMOCERATINAE Spath, 1926 (= Scalaritinae Ward, 1976: 455) Genus GLYPTOXOCERAS Spath, 1925 (= Neohamites Brunnschweiler, 1966) | TYPE SPECIES. Hamites rugatus Forbes, 1846: 116, pl. 11, fig. 6, by original designation (Spath 1925: 30, as Hamites (Aniso- ceras) rugatus (Forbes) Kossmat). A 10mm ; Fig. 1 Idealised suture of Libycoceras? sp., based on BMNH C93887, a badly corroded specimen. 247 Glyptoxoceras sp. DESCRIPTION AND DISCUSSION. BMNH C93889 is a straight fragment with a whorl height of 6-5 mm, the whorl section is compressed oval, with strong even prorsiradiate flank ribs, transverse on the venter and weakened on the dorsum. It is specifically indeterminate, but ribbing and coiling suggest that it is a Glyptoxoceras, a genus recorded from Lower Santonian to Upper Maastrichtian. OCCURRENCE. BMNH C93889 is from the basal 60 cm shell bed immediately overlying unweathered ophiolite (bed 1) at Jebel Aqabah. Genus LEWYITES Matsumoto & Miyauchi, 1984 TYPE SPECIES. Idiohamites (?) oronensis Lewy, 1969: 127, pl. 3, figs 10, 11, by original designation. Lewyites ambindense (Collignon, 1971) Plate 2, figs 13, 5; 16 1971 Neancyloceras ambindense Collignon: 11, pl. 644, fig. 2380. TyPe. Holotype, by monotypy, is the original of Collignon 1971, pl. 644, fig. 2380, from the Maastrichtian of locality 504 of Collignon, Mont Ambinda-Mikoboko (Manera), Mada- gascar. DESCRIPTION. BMNH C93890 is a 45 mm long fragment of a straight shaft showing slight curvature at the adapical end. The fragment is wholly septate, with traces of recrystallized shell. The whorl section is compressed oval, with a whorl breadth to height ratio of 0-9 and maximum preserved whorl height of 20 mm. There are 11 ribs in a distance equal to the whorl height. They are narrower than the interspaces, sharp, transverse to feebly convex on the dorsum, sweeping for- wards and feebly convex on the dorsolateral margin, mark- edly prorsiradiate and strengthening across the flank, and transverse on venter. Alternate ribs bear small ventral clavi; occasionaly a second rib is feebly linked to a clavus. Much larger is OUM KY1996, a body chamber fragment with a maximum preserved whorl height of 29 mm. Ornament is as in the smaller specimen, but for the marked effacement of ribs on the dorsum of the internal mould. Sutures not seen. Discussion. I was unable to trace the holotype of Neancylo- ceras ambindense during a recent examination of the Colli- gnon Collection, housed in the Université de Bourgogne, Dijon. The ribs of the Oman material are more markedly prorsiradiate than in the holotype, with fewer non- tuberculate ribs, but they are otherwise similar. Reference to Lewyites is based on a comparison with topotypes of the type species in the Oxford University Museum (OUM KY2021—25), and large fragments from New Jersey figured by Cobban (1974, pl. 10, figs 22-35) and Kennedy & Cobban (1993b, figs 5.1-5.18, 5.22-5.26; 7.19, 7.20; 9.4, 9.7). OCCURRENCE. BMNH_ C93890 is from bed 10, Jebel Huwayyah, section 1. OUM K1996 (Skelton 84/16-2) comes from the basal part of the Simsima Formation of the north- western part of Jebel Huwayyah, close to where the section is cut by the road. The holotype is from the so-called Lower Maastrichtian Zone a Pachydiscus gollevillensis et P. neuber- gicus of Collignon, but associated ammonites from the type 248 PLATE 4 Pachydiscus (Pachydiscus) dossantoi (Maury, 1930). BMNH C93895 from the Qahlah Formation, bed 10, Jebel Huwayyah, section 1, x 0-75. | locality in the Collignon Collection indicate Upper Maastrich- tian only. Family NOSTOCERATIDAE Hyatt, 1894 (= Jouaniceratidae Wright, 1952: 218; Bostrychoceratinae Spath, 1953: 16; Emperoceratinae Spath, 1953: 17; Hyphantoceratinae Spath, 1953: 16) Genus NOSTOCERAS Hyatt, 1894 Subgenus NOSTOCERAS Hyatt, 1894 TYPE SPECIES. Nostoceras stantoni Hyatt, 1894: 569 (= Ammonites approximans Conrad, 1855: 266), by original designation. W.J. KENNEDY Nostoceras (Nostoceras) major Kennedy & Cobban, 1993 Plate 2, figs 4-8) 1993c Nostoceras (Nostoceras) major Kennedy & Cobban: | 6-1, fig. 4. | Type. Holotype in Texas Memorial Museum Collections no} 77981, from the Upper Maastrichtian Corsicana Formation | 1-2 km (0-75 miles) SE of New Sweden, Travis County, | Texas. DESCRIPTION. There are 2 fragments, BMNH C93993, with a whorl height of 13 mm, and BMNH C€93994, with a whorl | height of 39-5 mm. Both are derived from helices with the whorls in tight contact, there being a shallow concave | | MAASTRICHTIAN AMMONITES impressed zone on the upper whorl face; the outer and lower whorl faces are broadly convex, the inner whorl face is flattened. Ornament is effaced on the impressed zone of the upper whorl face, but wiry narrow ribs strengthen, sweep back and are concave across the juncture of upper and outer whorl faces and are markedly prorsiradiate and narrower than the interspaces on the outer whorl face, sweeping backwards and feebly concave across the juncture of outer and lower whorl faces, straight and prorsiradiate on the lower whorl face and convex and effaced on the inner. All ribs are single on the larger fragment, but occasionally join in pairs at the juncture of outer and lower whorl faces in the smaller fragment. Sutures not seen. DISCUSSION. The simple coiling and ornament of these frag- ments occurs in nostoceratids from the Turonian onwards. Given the Maastrichtian age of the specimens, reference to Nostoceras (Nostoceras) is indicated. The fragments differ in no significant respect from the holotype of N. (N.) major, other than their coiling direction, absence of flared ribs and constrictions, possibly reflecting no more than the short lengths preserved, there being only 2-3 constrictions and associated flared ribs per whorl in the type. OCCURRENCE. BMNH C€93993 comes from bed 9, Jebel Bu Milh, section 2: C93994 comes from bed 10 or 11, Jebel Huwayyah, section 1. The holotype is from the Upper Maastrichtian of Texas. Nostoceras (Nostoceras) sp. Plate 2, fig. 20 DESCRIPTION AND DISCUSSION. BMNH C93888 is an poorly preserved U-shaped body chamber with a maximum pre- served whorl height of 21 mm. Ornament is of coarse single ribs with traces of ventral tubercles on at least some ribs. The specimen is specifically indeterminate, but recalls the Nosto- ceras (N.) hyatti group, of the uppermost Campanian Lower Maastrichtian (Kennedy & Cobban 1993b). OCCURRENCE. The specimen comes from bed 6, Jebel Huwayyah, section 2. REFERENCES Brunnschweiler, R.O. 1966. Upper Cretaceous ammonites from the Carnavon Basin of Western Australia. 1. The heteromorph Lytoceratina. Bulletin of the Bureau of Mineral Resources, Geology and Geophysics, Australia, 58: 58 pp.. 8 pls. Collignon, M. 1938. Ammonites Campaniennes et Maestrichtiens de l’ouest et du sud de Madagascar. Annales géologiques du Service des Mines de Madagascar, 9: 55-118 (1-65), pls 1-9. 1956. Ammonites néocrétaces du Menabe (Madagascar) IV. Les Phyllo- ceratidae. V. Les Gaudryceratidae. VI. Les Tetragonitidae. Annales géologiques du Service des Mines de Madagascar, 23: 1-106, pls 1-11. — 1971. Atlas des fossiles caracteristiques de Madagascar (Ammonites) XVII (Maestrichtien). Service Géologique, Tananarive. iv + 44 pp., pls 640-658. Conrad, T.A. 1855. Descriptions of eighteen new Cretaceous and Tertiary fossils. Proceedings of the Philadelphia Academy of Natural Sciences, 7: 265-268. Forbes, E. 1846. Report on the Fossil Invertebrata from southern India, collected by Mr. Kaye and Mr. Cunliffe. Transactions of the Geological Society of London, (2) 7: 97-174, pls 7-19. Fritsch, A. & Kafka, J. 1887. Die Crustaceen der bodhmischen kreideforma- tionen. Selbstverlag, Prague. 53 pp. Gill, T. 1871. Arrangement of the Families of Mollusks. Smithsonian Miscella- neous Collections, 227: xvi + 49 pp. Grossouvre, A. de 1894. Recherches sur la craie supérieure, 2, Paléontologie. Les ammonites de la craie supérieure. Mémoires du Service de la Carte 249 géologique deétaillée de la France. 264 pp., 39 pls. (mis-dated 1893). Hancock, J.M. & Kennedy, W.J. 1993. The high Cretaceous ammonite fauna from Tercis, Landes, France. Bulletin de l'Institut des Sciences Naturelles de Belgique, 63: 149-209, 20 pls. Hauer, F. von 1858. Uber die Cephalopoden aus der Gosauschichten. Beitrdge zuir Palaontologie von Osterreich, 1: 7-14, pls 2-4. Henderson, R.A. & McNamara, K.J. 1985. Maastrichtian non-heteromorph ammonites from the Miria Formation, Western Australia. Palaeontology, 28: 35-88, pls 1-9. Hyatt, A. 1889. Genesis of the Arietidae. Smithsonian Contributions to Knowledge, 673, xi + 239 pp., 14 pls. 1894. Phylogeny of an Acquired Characteristic. Proceedings of the American Philosophical Society, 32: 349-647, pls 1-14. 1900. Cephalopoda, pp.502-604 in ZITTEL, K.A. VON 1896-1900, Textbook of Palaeontology, transl. EASTMAN, C.R. Macmillan, London and New York. — 1903. Pseudoceratites of the Cretaceous. United States Geological Survey Monograph, 44: 351 pp., 47 pls. Kennedy, W.J. 1986. The ammonite fauna of the Calcaire a Baculites (Upper Maastrichtian) of the Cotentin Peninsula (Manche, France). Palaeontology, 29: 25-83, 6 pls. — & Cobban, W.A. 1993a. Upper Campanian ammonites from the Ozan- Annona Formation boundary in southwestern Arkansas. Bulletin of the Geological Society of Denmark, 40: 115-148, 8 pls. & Cobban, W.A. 1993b. Ammonites from the Saratoga Chalk (Upper Cretaceous) Arkansas. Journal of Paleontology, 67: 404-434. — & Cobban, W.A. 1993c. Maastrichtian ammonites from the Corsicana Formation in northeast Texas. Geological Magazine, 130: 57-67. — & Hancock, J.M. 1993. Upper Maastrichtian ammonites from the Marnes de Nay between Gan and Rébénacq (Pyrénées-Atlantiques), France. Geo- bios, 26: 575-594, 7 pls. — & Henderson, R.A. 1992. Non-heteromorph ammonites from the Upper Maastrichtian of Pondicherry, South India. Palaeontology, 35: 381-442, 18 pls. ~ — & Summesberger, H. 1986. Lower Maastrichtian ammonites from Neu- berg, Steiermark, Austria. Beitrdge ziir Palaontologie von Osterreich, 12: 181-242, 16 pls. Kossmat, F. 1895-1898. Untersuchungen tiber die Sudindische Kreideforma- tion. Beitrdége ztir Paldontologie Osterreich-Ungarens und des Orients, 9 (1895): 97-203 (1-107), pls 15-25 (1-11); 11 (1897): 1-46 (108-153), pls 1-8 (12-19); 11 (1898): 89-152 (154-217), pls 14-19 (20-25). Lewy, Z. 1969. Late Campanian heteromorph ammonites from southern Israel. Israel Journal of Earth Sciences, 18: 109-135, pls 1-4. Matsumoto, T. & Miyauchi, T. 1984. Some Campanian ammonites from the Soya area. Palaeontographical Society of Japan Special Paper, 27: 33-93, pls 10-31. —— & Obata, I. 1955. Some Upper Cretaceous desmoceratids from Hokkaido and Saghalien. Memoirs of the Faculty of Science, Kyushu University, Series D, Geology, 5: 119-151, pls 24-30. Maury, C.J. 1930. O Cretaceo da Parahyba do Norte. Monografias Servico Geologico e Mineralogico do Brasil. 305 pp.; Album das estampas i—xxiii, 35 pls. Olsson, A.A. 1944. Contributions to the Paleontology of Northern Peru. VII. The Cretaceous of the Paita Region. Bulletin of American Paleontology, 28: 11-46, pls 1-17. Orbigny, A.d’. 1850. Prodrome de Paléontologie stratigraphique universelle des animaux mollusques et rayonnés. 2, Masson, Paris. 428 pp. Seunes, J. 1890. Contributions a l'étude des céphalopodes du Crétacé Supérieur de France. 1. Ammonites du Calcaire a Baculites du Cotentin. Mémoires de la Société Géologique de France. Paléontologie, 1, Mémoire 2: 1-7, pls 2-3 (1-2). — 1892. Contributions a l'étude des céphalopodes du Crétacé Supérieur de France. I. Ammonites du Calcaire 4 Baculites du Cotentin (Suite). II. Ammonites du Campanien de la région sous pyrénéenne. Départment de Landes. Mémoires de la Société Géologique de France; Paléontologie, 1, Mémoire 2: 8-22, pls 12-15 (3-6). Spath, L.F. 1921. On Upper Cretaceous Ammonoidea from Pondoland. Annals of the Durban Museum, 3: 39-56, pls 5-9. 1922. On the Senonian ammonite fauna of Pondoland. Transactions of the Royal Society of South Africa, 10: 113-147, pls 5-9. — 1925. On Senonian Ammonoidea from Jamaica. Geological Magazine, 62: 28-32, | pl. — 1926. On new ammonites from the English Chalk. Geological Magazine, 63: 77-83, table. — 1929. Corrections of cephalopod nomenclature. Naturalist, 871: 269-271. — 1953. The Upper Cretaceous Cephalopod fauna of Grahamland. Scientific Reports of the British Antarctic Survey, 3, 1-60, pls 1-13. Stoliczka, F. 1865. The fossil Cephalopoda of the Cretaceous rocks of southern 250 India. Palaeontologia Indica, Calcutta, Series 3, part 9: 143-154, pls 71-80. Ward, P.D. 1976. Upper Cretaceous Ammonites (Santonian-Campanian) from Orcas Island, Washington. Journal of Paleontology, 50: 454-461, pl.1. Wiedmann, J. 1966. Stammesgeschichte und System der posttriadischen Ammonoideen; ein Uberblick. Neues Jahrbuch ftir Geologie und Paldontolo- gie Abhandlungen, 125: 49-79, pls 1, 2; 127: 13-81, pls 3-6. Wright, C.W. 1952. A classification of the Cretaceous Ammonites. Journal of Paleontology, 26: 213-222. Yabe, H. & Shimizu, S. 1924. A new species of Brahmaites from the Upper Cretaceous of S. Saghalien, with some remarks on the genus Brahmaites. Japanese Journal of Geology and Geography, 76: 77-80, pl. 13. W.J. KENNEDY Zaborski, P.M.P. 1982. Campanian and Maastrichtian sphenodiscid ammonites from southern Nigeria. Bulletin of the British Museum (Natural History), Geology, 36: 303-332. —— 1983. Campano-Maastrichtian ammonites, correlation and palaeogeogra- phy in Nigeria. Journal of African Earth Sciences, 1: 59-63. —— 1985. Upper Cretaceous ammonites from the Calabar region, southeast Nigeria. Bulletin of the British Museum (Natural History), Geology, 39: 172. Zittel, K.A. yon 1884. Handbuch der Palaeontology. 1, Abt. 2; Lief 3, Cephalopoda, pp. 329-522. R. Oldenbourg, Munich & Leipzig. — 1895. Grundziige der Palaeontologie (Palaeozoologie). R. Oldenbourg, Munich & Leipzig, vii + 972 pp. Bull. nat. Hist. Mus. Lond. (Geol.) 51(2):251-255 Issued 30 November 1995 Maastrichtian nautiloids from the United Arab Emirates-Oman border region NOEL J. MORRIS Department of Palaeontology, The Natural History Museum, Cromwell Road, London SW7 5BD Synopsis. The two nautiloids Deltoidonautilus salisfilius sp. nov. and Cimomia aff. sowerbyana (d’Orbigny) are described from the Maastrichtian Simsima Formation of the United Arab Emirates and Oman. INTRODUCTION Cephalopods form only a tiny percentage of the macrofossils collected from the late Cretaceous of the Emirates-Oman border area. They consist only of ammonites and nautiloids and, in spite of the small numbers, it is apparent that the distribution of the two groups is rather different and may reflect their partial ecological separation. Two distinct species of nautiloids were collected from a number of the Maastrichtian localities on the Emirates-Oman borders. The possibility that they might have been sexual dimorphs is discounted because there are no features in common between the two. For instance, the change in shape on the final whorl of the smaller species does not occur on the larger species. The nautiloids contrast with those described by Noetling from the Late Cretaceous of the Mari Hills (Pakistan), where species of Cimomia and Deltoidonautilus do not seem to occur (Noetling 1897). There seems to be little change in the relatively low nautiloid diversity after the end of the Cretaceous. The apparent loss of Cymatoceras is matched by the increase in the Hercoglossidae, mostly in the form of the new suture-line pattern of Hercoglossa. This change in shell architecture could represent a wider range of depth tolerance within the superfamily. SYSTEMATIC DESCRIPTIONS Superfamily NAUTILOIDEA de Blainville, 1824 [Name elevated from family rank, Shimanskiy, 1957] Family HERCOGLOSSIDAE Spath, 1927 The distinction between some of the genera in this family is rather dubious: Cimomia and Deltoidonautilus in particular seem to overlap. Kummel (1964, K456) suggests that Cimo- mia is ‘a morphologically transitional form between Eutre- Phoceras and Hercoglossa’. Both Cimomia and | Deltoidonautilus, if indeed they are distinct, seem to fore- shadow Nautilus itself in their shell morphology. © The Natural History Museum, 1995 Genus DELTOIDONAUTILUS Spath, 1927 TYPE SPECIES. Nautilus sowerbyi J. de C. Sowerby, 1843 (see footnote by Kummel, 1964: K456-57), from the Lower Eocene, London Clay. SYNONYMY. Deltoidonautilus may yet prove to be a junior synonym of Angulithes de Montfort, 1808, because of the various intrepretations of the type species of Angulithes, Nautilus triangularis de Montfort, 1802. Kummel accepted the name in 1956 but rejected it as a nomen dubium in the Treatise (1964). It is not clear in de Montfort’s (1808: 8) description, whether the type specimen came from the Lower Kimmeridgian or from the Cenomanian. It is beyond the scope of the present work to establish the type material of this taxon, with the result that Kummel’s (1964: K456) later opinion that Angulithes should be treated as a nomen dubium is followed here. DESCRIPTION. Characteristically smooth, involute and often compressed with a sub-carinate venter in early to middle growth stages. Suture sinuous and projected forwards on the venter. Distinguished from Cimomia by the carinate venter. RANGE. Upper Cretaceous (Cenomanian) to Oligocene; cos- mopolitan. Deltoidonauitilus salisfilius sp. nov. Plate 1, figs 1-3 21861 Nautilus rota Blandford: pl. 25, fig. 2 only. 1928 Nautilus jordani Zittel; Lees: 663, pl. 44, fig. 6. HOLOTYPE. BM C.59590, from the ‘Main Gastropod Bed’, bed 6, at Jebel Bu Milh (JBM 2), NNW of Al Ain; base of Simsima Formation, Maastrichtian, probably Lower Maas- trichtian. OTHER MATERIAL. Thirteen paratypes from the same locality and horizon, BM C.59591—603, Morris and Smith Collection. Three additional paratypes apparently from the same locality and horizon, G.M.Lees collection (mentioned Lees, 1928), BM C.31034-36. Two specimens from the basal 2 meters of the Simsima Formation at Jebel Faiyah (JF2), U.A.E., BM C.59607-08. ?Lower Maastrichtian, Morris and Smith Collec- tion. One specimen from Jebel Faiyah (JF1b), loose, but probably from the basal Simsima Formation, ?Lower Maas- trichtian, Gale, Morris and Smith collection, BM C.59609. One specimen from Jebel Buhays (JB1b) also loose but ?basal Simsima Formation, Gale, Morris and Smith collec- tion, BM C.59610. All the specimens found apparently occur 252 just above the flooding surface at the base of the Simsima Formation, probably at a similar horizon to Libycoceras sp. and Nostoceras spp. DESCRIPTION. Shell rather smail, the two near complete specimens from Jebel bu Milh are both approximately 102 mm in diameter. Shell planospiral, smooth, involute, non- umbilicate on outer shell surface; internal mould with a narrow, shallow umbilicus. Compressed, discoidal, sub- oxycone with rounded to subcarinate venter on inner and middle whorls; outer whorl, ie. body-chamber, rounded and much less compressed, occupying approximately one third of the last whorl. Siphuncle well dorsal of centre. Suture with well-rounded but relatively narrow ventral saddle that bends forward towards the aperture in a way that is more prominent in the sub-oxycone stages than in the more rounded whorls where the few final sutures occur; broad, shallow, sub- symmetrical lateral lobe; relatively low, smallish lateral saddle close to the umbilical margin. Sutures show consider- able crowding towards the body-chamber in what seems to be a fully grown individual. Siphonal sinus moderate on the penultimate whorl, observed only on specimen BM C.31034. COMPARISON WITH OTHER SPECIES. Nautilus fleuriausianus d’Orbigny (1840: 82, pl. 15) has a rounded venter similar to the body-chamber of the present species, but does not have the earlier, more oxycone stage. It is apparently of Upper Cenomanian age from the ‘craie a Caprines’ at Ile Madame, Charente, France. It may prove to be a more rounded specimen of the widespread species named Nautilus triangu- laris (de Montfort; d’Orbigny, 1840: pl. 12; ?=Angulithes triangularis de Montfort 1808). Two nearly complete speci- mens from the Cenomanian of Sidmouth (BM C.931), and from the Upper Grey Chalk near Folkestone (BM C.8320), are both carinate and show no sign of change to a round ventered body-chamber at approximately 160 mm and 180 mm diameter respectively. Nautilus mermeti Coquand, 1862, from Algeria, may also prove to be a synonym of this Cenomanian species. Nautilus westphalicus Schliiter, 1876 from the Late Santo- nian or Lower Campanian Quadraten Kreide from Dulmen in Germany also has a carinate body-chamber. Nautilus galea Fritsch, 1872, from the Iserschichten in Bohemia, has a carinate venter on the body chamber, has much broader whorls and apparently has less sinuous sutures. There may be three separate taxa amongst Blanford’s (1861) illustrations of his species Nautilus rota: his pl. 24 fig. 3 and pl. 25, fig. 1 clearly belong to Cymatoceras, while the smooth internal mould of his pl. 25, fig.2 has the flexed suture typical of Deltoidonautilus and could well belong in the present species. Genus CIMOMIA Conrad, 1866 TYPE SPECIES. Nautilus burtini Galeotti, 1837, from the Eocene of Belgium (a very similar species to C. imperialis (J. Sowerby, 1812)). N.J. MORRIS Cimomia aff. sowerbyana (d’Orbigny, 1840) Plate 1, fig. 4; Plate 2 Nautilus sowerbyanus d’Orbigny: 83, pl. 16. Nautilus sowerbyanus d’Orbigny; d’Orbigny: 189. Nautilus jordani Wanner: 143, pl. 19, fig. 21. Cimomia jordani (Wanner); Kummel: 451. aff. 1840 aff. 1850 1902 1956 MATERIAL. A single fairly well-preserved specimen, BM C.59611, from the base of the Simsima Formation at Jebel Fayah (JF 2), associated with Deltoidonautilus salisfilius sp. nov.; the body chamber is missing. A single poorly preserved but virtually complete specimen from Jebel Buhays (JB 1b), also apparently from the base of the Simsima Formation, BM C.59612. Three doubtful specimens from Jebel Rawdah (JH 1), one of them in situ in the base of bed 5, BM C.59613, and the other two loose on the scree below but apparently from the same horizon, BM C.59614-15. In his original description, d’Orbigny stated that the local- ity of the type specimen of Nautilus sowerbyanus was unknown, but later, in the Prodrome (d’Orbigny, 1850: 189), he gave the locality as Montrichard, France. The specimen was in the Musée d’Histoire Naturelle, Paris, the locality details having been sent by M. Cordier, and it seems quite likely that Fischer, who organised d’Orbigny’s collection, must have sorted out the locality before publication of the Prodrome. Wanner’s holotype of Nautilus jordani came from the uppermost white chalk near Bab-el-Jasmund to the north of Dachel, western Egypt. DESCRIPTION. Shell medium-sized, planospiral, smooth, involute. External shell non-umbilicate, with the umbilicus of the internal mould plugged by a slight recurved extension of the lateral saddle, almost in the form of a small very shallow lobe. Shell inflated to spheroidal with flanks converging to the rounded but slightly flattened venter. Suture with a very low, gently rounded ventral saddle, a symmetrical, relatively shallow lateral lobe and a relatively prominent well-rounded lateral saddle. COMPARISON WITH OTHER SPECIES. ?Cimomia applanatus (Wanner, 1902: 143), another species named from western Egypt, could be another variant of this species but is consid- erably more evolute than any of the specimens considered here from the same area. The identification of Nautilus desertorum Quaas (1902: 299) is also uncertain, because it is not possible to judge from the original illustration whether the suture-line is sinuous like that of Cimomia or straighter like that of Eutrephoceras. Cimomia aff. sowerbyana is similar in general shape to Nautilus sublaevigatus d’Orbigny (1850: 189; nom. nov. for N. laevigatus d’Orbigny, 1840: pl. 17, non Reinecke, 1818). The first locality listed where d’Orbigny collected this species is Martrou near Rochefort. He also listed a number of other localities including Montrichard, Uchaux and several from Cotentin; these span a range from Turonian to Maastrichtian. The sutures of N. sublaevigatus seem much less sinuous than those of Cimomia sowerbyana and suggest that this species PLATE 1 Figs 1-3 Deltoidonautilus salisfilius sp. nov. Base of Simsima Formation Maastrichtian, ? Lower Maastrichtian, Jebel Bu Mil;ch, NNW of Al Ain. 1, holotype, BM C.59590; 2, paratype, BM C.59591; 3, paratype, BM C.59592, arrow shows poistion of the siphuncle; all x 1. Fig. 4 Cimomia aff. sowerbyana (d’Orbigny, 1840). Base of Simsima Formation, ?Lower Maastrichtian, Jebel Fayah, south of Dayid, ventral view; BM C.59611; x 1. | f ’ MAASTRICHTIAN NAUTILOIDS 253 J. MORRIS 254 rte “SL-0 x ‘apouNYdis ay Jo UONIsod ay SMOYS MOLE !MIIA [PINJIOde “QT :MIIA OPIS “BT +1 196S'O WE :PIAe Jo yinos ‘yeAvy oqo ‘uenYoINseRJ] (I9MOT{) “UONLULIOY BUISUIIS Jo aseg “(PET ‘AUsIGIO,p) DUDAqQuamos ‘ffo DnUOUND ¢ ALV 1d MAASTRICHTIAN NAUTILOIDS belongs to Eutrephoceras. Similarly N. vastus Kner, 1850, Cimomia cf. forbesi dArchiac & Haime, 1854 (a Danian species), and Eutrephoceras sphaericus (Forbes, 1846) (holo- type from Pondicherry, India, BM C.73524; not pre-occupied by Nautilites sphaericus Martin, 1809), may belong to Eutre- phoceras. A number of Late Cretaceous specimens in the BM(NH) collection from the United States also indicate that Nautilus dekayi Morton, 1833, does not have the sinuous suture-line of Cimomia and properly belongs in Eutrephoceras. REFERENCES Blainville, H. M. D. de 1824. Dictionaire des science naturelles; Mollusques, 32: 392 pp. Strasbourg and Paris. Blandford, H. F. 1861. The Fossil Cephalopods of the Cretaceous rocks of Southern India, (Belemnitidae - Nautilidae). Memoirs of the Geological Survey of India, Palaeontologia Indica, 1: 40pp., 25pls. Conrad, T. A. 1866. Observations on Recent and fossil shells, with proposed new genera and species. American Journal of Conchology, 2: 101-03. Coquand, M. H. 1862. Géologie et Paléontologie de la région sud de la Province de Constantine. Marseille. Forbes, E. 1846. Report on the fossil Invertebrata from southern India collected by Mr. Kaye and Mr. Cunliffe. Transactions of the Geological Society of London, (2) 7: 97-174, pls. 7-19. Fritsch, A. & Schloenbach, U. 1872. Cephalopoden der béhmischen Kreidefor- mation. 52 pp., 16 pls. Prague. Galeotti, M. H. 1837. Mémoir sur la constitution géognostique de la province de Brabant, en réponse a la question suivante: décrire la constitution géologique de la province de Brabant, déterminer avec soin les éspeces minérales et les fossiles que les divers terrains renferment et indiquer la synonymie des auteurs qui en ont déja traité. Mémoires couronneés de l'Académie Royale des Science et Belles Lettres, 12: 1-192, pls. 1-4. Brux- elles. 255 Kner, R. 1850. Versteinerungen des Kreidemergels von Lemberg und seiner Umgebung. Naturwissenschaftliche Abhandlungen (Haidinger), 3 (2): 1-42, pls 1-S. Kummel, B. 1956. Post-Triassic Nautiloid Genera. Bulletin of the Museum of Comparative Anatomy at Harvard College, 114 (7): 321-494, 28 pls. 1964. Nautiloida-Nautilida. Jn Moore, R.C. (editor), Treatise on Inverte- Brate Paleontology, K, Mollusca 3, 519 pp. The Geological Society of America and the University of Kansas Press. Lees, G. M. 1928. Geology and tectonics of Oman and parts of southeastern Arabia. Quarterly Journal of the Geological Society of London, 84: 585-670. Montfort, D. de 1808. Conchyliologie systématique et classification méthodique des coquilles; offrant leur figures, leur arrangement générique, leurs descrip- tions caractéristiques, leurs noms; ainsi que leur synonymie en plusieurs langues. 1: 409 pp., 100 pls. Paris. Morton, S. G. 1833. Supplement to the ‘Synopsis of the Organic Remains of the Ferruginous Sand Formation of the United States’, contained in Vols XVII and XVIII of this Journal. American Journal of Science, 23 (2): 288-94. Noetling, F. 1897. Fauna of Baluchistan, Vol. 3, part 1, Fauna of the Upper Cretaceous (Maastrichtian) Beds of the Mari Hill. Memoirs of the Geological Survey of India, Palaeontologia Indica, (16) 1 (3): 79 pp., 23 pls. Orbigny, A. d’ 1840. Paleontologie Francaise. Terraines Cretacées. Céphalopo- des. 1: 662 pp., 148 pls. 1850. Prodrome de Paléontologie stratigraphique universelle des animaux mollusques et rayonnes faisent suite au cours elementaire de Paléontologie. 3 vols. Masson, Paris. Quaas, A. 1902. Beitrag zur Kenntniss der Fauna der obersten weissen Kreidebildungen in der libyschen Wuste (Overwegischichten und Blatter- thone). Palaeontographica, 30 (2) (4): 153-355, pls 20-33. Schliiter, C. 1876. Cephalopoden der oberen Kreide. Palaeontographica, 24 (2): 123-263, pls. 36-55. Shimanskiy, V. N. 1957. Sistematica i Filogenia otryada Nautilida. Bulletin de la Societe des Naturalistes de Moscou, Geologie, 32 (4): 105-20 [In Russian]. Sowerby, J. & Sowerby, J. de C. 1812-1846. A Mineral Conchology of Great Britain. Vols 1-7, 650 pls. Wanner, J., 1902. Die Fauna der obersten weissen Kreide der libyschen Wiste. Palaeontographica, 30 (2) (3): 91-151, pls 13-19. Bull. nat. Hist. Mus. Lond. (Geol.) 51(2):257-265 Issued 30 November 1995 Maastrichtian Inoceramidae from the United Arab Emirates-Oman border region NOEL J. MORRIS Department of Palaeontology, The Natural History Museum, Cromwell Road, London SW7 5BD Synopsis. Descriptions of Inoceramids from Upper Cretaceous post-Semail Nappe emplacement deposits include Endocostea (Selenoceramus) semaili sp. nov. Most specimens are from the Simsima Formation in Jebel Rawdah, immediately below the massive limestone facies, and they indicate a mid-Maastrichtian age. INTRODUCTION Small collections of Inoceramidae from two localities on the western edge of the Omani Mountains, close to the border between the United Arab Emirates and Oman, are preserved largely as uncrushed internal moulds. The first is from the Simsima Formation in Jebel Rawdah just north of the Al Ain to Hatta Road, about 10 Km east of the Madam Roundabout, just east of the Omani Border Post, and is of mid- Maastrichtian age. The second is from the Loftusia-rich marls below the Simsima Formation in Jebel HuwaYyah (known as Fossil Valley), to the east and north-east of Al Ain, and is apparently of Lower Maastrichtian age. It is quite clear that the specimens from the higher horizon, i.e. in Jebel Rawdah, assist with the age determination of the strata, and they may have potential for international correlation. The Inoceramidae show that at least three distinct lineages existed in the eastern Arabian area in mid-Maastrichtian times. Each lineage seems to fall in some part of Whitfield’s (1877) genus, Endocostea, which is in much need of revision. Any serious revision is, however, beyond the scope of this paper. Subgeneric names are used tentatively until we have a better understanding of the phylogeny of the group. The apparent lack of specimens of Trochoceramus, which is widespread across Africa in late Campanian and Maastrich- tian times, may reflect the palaeoecology. The water may have been either too shallow or too warm, or the sediment too coarse. The specimens described here were collected by Drs Gale, Morris and Smith in January 1992, by Drs Nolan and Skelton during an earlier visit (Nolan et al. 1990), and by members of the Emirates Natural History Group. Class BIVALVIA Linnaeus 1758 Subclass PPERIOMORPHIA Beurlen, 1944 [nom. trans. Newell, 1965] Superfamily INOCERAMOIDEA Giebel, 1852 [?=Ambonychioidea Miller, 1877] | : ‘ ‘ si The Inoceramidae share with Ambonychia the thick calcitic _ inter-umbonal ligament area not present in adult Pterioidea. © The Natural History Museum, 1995 Family INOCERAMIDAE Giebel, 1852 DESCRIPTION. Variously shaped, concentrically lamellose or plicated or radially plicated Pteria-like bivalves which some- times grew to a large size. The outer calcareous shell layer consists of vertical calcite prisms which are modified to pyriform laths on the ligament area. The ligament, usually multivincular, is attached largely to this external layer; inner shell layer nacreous; usually lacking hinge teeth; usually equivalve or subequivalve, some species inaequivalved. A complex of muscle scars close to the umbones is consistant with byssal fixation. Apparently monomyarian with the pos- terior adductor often close to the posterior ventral margins but often obscure. It is very difficult to be certain whether an isolated muscle scar situated about halfway below the umbones and towards the anterior margin is a large pedal muscle scar or a small anterior adductor. The Inoceramidae became extinct by the beginning of the Tertiary. COMMENTS. The ligament area of these later Cretaceous species is seldom well-exposed and the pits characteristic of earlier species have only been observed on one of the species. It remains possible that lineages other than Tenuipteria (Dhondt, 1983b) have no pits for the differing ligament types. THE INTERNAL SHELL RIB. Many post-Coniacian Inocerami- dae have an internal shell rib, sillon or Hohlkehle. The nature of this internal rib has been discussed on a number of occasions (especially Seitz, 1967: 14-41). It occurs commonly in a number of Inoceramidae from at least as early as the Santonian. It is here considered to be an architectural charac- ter of the shell, and it is clearly variable in some of the taxa that have it. Two specimens of Endocostea sp. from the Santonian-Campanian, Haslam Formation of Brannan Creek, Nanaimo District, British Columbia, Canada, demon- strate this point; in BM LL 28194 the inner ‘rib’ is poorly developed, starting at approximately 6 cm from the umbo of the right valve and continuing for only 2 cm, whereas BM LL 28217, a slightly smaller individual from the same locality and horizon, developed the rib at 1-5 cm from the umbo of the right valve, from which position it continues for at least 4-5 cm towards the posterior ventral margin. In three well-preserved specimens from the late Campa- nian, Fort Pierre Group (two examples of ‘/noceramus? aff. barabini Morton, BM L 21569 from the Cheyenne River Section, BM L 7577 from the Powder River Range, Montana, and a single example of ‘/noceramus’ tenuirostris Meek & 258 Hayden, 1862, BM L 21571, from the Cheyenne River), no fully developed internal rib is present. In its place are raised ridges on the innermost shell following the track of the normal position of the inner rib. These end distally as tangents to the 1 cm wide muscle scar that has been referred to as the posterior adductor. Clearly, inoceramids have an unusual arrangement of their soft parts, but it would appear the inner rib is formed as a sensitive emplacement of shell material to enhance strength in the part of the shell under stress from muscular shell closing. The internal rib is also present on some Jurassic species; a well-preserved specimen of Parainoceramus ventricosus (J. de C. Sowerby, 1823) from the Lower Jurassic, Pliensbachian, at Brockthorpe, near Gloucester, shows precisely the same details as ‘Inoceramus’ barabini, with an internal rib running down to the adductor. Some Campanian specimens of Endocostea, however, seem to show the internal rib running very close to the ventral margin, apparently past the position of the adductor. There are also preservational problems in observing the internal rib. Early diagenesis of the aragonite shell interior can destroy the rib, and there are many examples of inocera- mids preserved only as internal moulds of the outer calcitic shell layer. The rib is nacreous, is often hollow and has an arch-shaped section. In all the examples that I have examined where two valves are preserved together, one rib is an almost perfect mirror image of the other. The rib is in a morphologically comparable position in all species in which it occurs. It shows distinct evolutionary changes, one of the most notable being extreme broadness in some species of Seleniceramus (Seitz 1967), though it is usually distinctly narrow. The internal rib may be close to a muscle scar at its distal end. The claim that the internal rib is formed by, or in response to, a parasite may explain why it is sometimes filled with matrix. This can, however, be equally well explained if it is due to the activity of shell borers: if a hole is bored from the outside of the shell into the hollow of the rib, the damage cannot be repaired by deposition of more shell material as it is separated from the outer surface of the mantle by the thickness of the rib itself. The same would be the case if the inoceramid was already dead when its shell was drilled. All the cases of matrix- filled inner ribs that I have examined show evidence of drilling from the shell exterior. The inner rib is clearly of some importance for taxonomy at both the species and higher levels. Its variability in some species and its possible diagenetic loss mean that it needs to be used with caution. Genus ENDOCOSTEA Whitfield, 1877 TYPE SPECIES. E. typica Whitfield, 1877, by original designa- tion. The lectotype (USNM 12261) was selected by Seitz (1967: 54-55, pl. 2, fig. 4). It is the example ‘c’ of Whitfield (1880: pl. 9 fig. 3) and is from the Pierre Shale at Old Woman Fork on the Cheyenne River, Black Hills, South Dakota, which is of late Campanian age according to Cobban & Reeside (1952: 1011) (non Inoceramus cripsi var. typica Zittel, 1866: 98, from the late Cretaceous of Grunbach in Neuen Welt and the Gosau Valley, Austria). DisCUSSION. The generic name was introduced for relatively small inoceramids with a well-developed inner rib radiating N.J. MORRIS from the umbo towards the posterior ventral margin but stopping well short of that margin. The species is otherwise very similar to Whitfield’s own interpretation of Inoceramus barabini Morton (Whitfield, 1885: 75-76), except that the latter species apparently does not have the internal rib. Subgenus ENDOCOSTEA Whitfield, 1877 A number of species or subspecies of Endocostea are rather convex with prominent or enrolled anterior umbones. These occur in the late Campanian and Maastrichtian and resemble Endocostea (Endocostea) coxi (Reyment). They include Jnoc- eramus balticus pteroides (Giers, 1964: pl. 1, fig. 6) from the Upper Campanian, Polyplocum Zone of Haldem, Westfalia, and according to Sornay (1976) from Dau, Charente. They possibly also include Inoceramus borilensis Jolkicev, said to be from the Maastrichtian in Bulgaria. However, specimens that are very similar to J. bakalovi (said by Jolkicev (1961) to occur at the same horizon) occur with ammonites of the Upper Campanian, Donetzianum Horizon, in Nigeria. They also include Inoceramus impressus d’Orbigny (lectotype, Museum d@ Histoire Naturelle, Paris, 7592a, figured Sornay, 1957: 129) from the ?Upper Campanian of Royan, and ‘Inoceramus impressus’ d’Orbigny (pars, but not including the lectotype) from the Maastrichtian of Cotentin, Manche, France. These are provisionally included in the subgenus Endocostea sensu stricto, although their relationship with the less convex type species remains to be confirmed. Endocostea (Endocostea) coxi (Reyment, 1958) Plate 1, figs 24 1958 Inoceramus coxi Reyment: 140, pl. 3, figs 4-6. MATERIAL. The type material comes from Auchi in north- central Nigeria and is preserved in a medium grained, slightly ferrugenous sandstone (holotype BM L 82963). Some better preserved specimens occur on the [kot Ekpene Road in the Calabar area associated with magnificent specimens of Tro- choceramus ianjonaensis and are therefore of ‘mid’ (probably low Upper) Maastrichtian age. Another specimen in the BM(NH) collection comes from Madagascar. Three speci- mens were collected by Nolan and Skelton from the south- west face of Jebel Rawdah, BM LL 4164749. DESCRIPTION. Convex, equivalved species with strong radiat- ing sulcus posterior to the umbones, delimiting a much less convex posterior area. Somewhat quadrate, but with umbones well to the anterior and prosocline. Strong radiating internal rib well to the posterior of the convex part of the shell. Strong but low rounded comarginal ribs, separated by wider interspaces on anterior and flank, ie. the convex part of the shell, fading posteriorly. Smoother in later growth stages. | COMMENTS. This species was included in Cordiceramus by Dhondt (1983a) following Seitz (1967), a genus that was included as a synonym of Haenleinia by Cox (1969). Cordice- ramus Seitz (1961: 110, ex. Heinz, 1932) has a very distinctive wide external radiating furrow from the posterior of the — umbo to the posterior ventral margin and is more or less equivalve. The type species, C. cordiformis Sowerby, is a relatively tall species with a short hinge line, whereas Haen- leinia is an elongate genus related to Endocostea, with a posterior shell twist, resulting in an inequivalve shell in that = t : = ed ora & “7 i = - heb i ® od =—_-V \ - i Te — y leo - = i = ie - , z « = ‘ i > ° ne yaad agen rss Z a mr —_ oe 4 = P tae De and = = {at Dea Bi yess _ PREAS a roma Sarena farmer ACOCTV AR WATTEEAATEA AM OF 7 ie eee re eo aN \ n us wean, bal ga 2, tne a ae =e ar “ reser Ee : yp oe - a x =! i+ A ——- oe ee (tte) Os ange a Re lb ie geet Wi ela f FI te ‘ahi ete e. Pedi taP ety! eadeed cae oe | c An ae Y Vee Kk. ave Ph aa pa/ oe iN ered Oe tai a Bm “= ‘s \e = Y oe o = a re ead 7 od — e ttoo D._ f-be Aa ‘eevee & 1a : ree spoutpy= & me) TR Leia Hs ® > doe hh & — ripe Fai & “i = i 9 Mee Bee hie o-—, DB &. esa a ak F, ari, Be, —— sag eens er » otng ‘gat 0 iar abel Lares 1 Ss. inw a thoi Ae) a Th? Gis ey ies ji 2 ( Wie Jie Gog yew eS = i <->" 2 6 tae, & D po Bull. nat. Hist. Mus. Lond. (Geol.) 51(2):275-276 Issued 30 November 1995 Maastrichtian brachiopods from the United Arab Emirates-Oman border region ELLIS F. OWEN Department of Palaeontology, The Natural History Museum, Cromwell Road, London SW7 5BD INTRODUCTION Brachiopods were particularly rare in the Maastrichtian sedi- ments of the United Arab Emirates-Oman borders region. Despite intensive collecting only four specimens were found, all coming from the lower Loftusia-rich beds of the Qahlah Formation exposed at Jebel Huwayyah (see introduction for locality details). These four specimens belong to a hitherto undescribed genus of terebratulid. SYSTEMATIC DESCRIPTIONS Class TEREBRATULIDA Waagen, 1883 Suborder TEREBRATULIDINA Waagen, 1883 Superfamily TEREBRATULACEA Gray, 1840 Family TEREBRATULIDAE Gray, 1840 Genus PSEUDOGIBBITHYRIS nov. TYPE SPECIES. Pseudogibbithyris arabica sp. nov. DIAGNOsIS. Medium-sized uniplicate terebratulid, slightly longer than wide. elongate-oval in general outline and evenly biconvex. Umbo short, beak suberect, foraman small, perme- sothyrid. Cardinal process present, brachial loop simple. REMARKS. Pseudogibbithyris differs from Gibbithyris and Concinnithyris from the European Upper Chalk (Senonian), which it resembles in external morphological features, in its Fig. 1. Pseudogibbithyris arabica gen. et sp. nov., holotype, BMNH BF47; Jebel Huwayyah, section 2, beds 3-6. A, dorsal; B, lateral; C, anterior views: all x 1. © The Natural History Museum, 1995 distinctly flat bifid cardinal process, deep hinge-trough and short, triangular hinge-plates. Pseudogibbithyris arabica sp. nov. Figs 1, 2 DIAGNOsIs. As for genus. Types. Holotype, BMNH BF47; paratypes BMNH BF4446. DESCRIPTION. The dorsal valve is dominated by a low median fold bounded by faint carinae originating from a point midway between the umbonal area of the valve and the anterior margin of the shell. A corresponding shallow sulcus in the ventral valve forms a wide, shallow uniplication occupying the whole width of the anterior commissure. Internal characters include a well-developed flat, but dis- tinctly bifid cardinal process, and a deep hinge-trough. The hinge-plates are short, triangular in outline with a slightly concave ventral surface, and are deflected towards the dorsal valve. The triangular shape of the hinge-plates is maintained in the development of the horizontally placed bands of the descending branches of the brachial loop, which is uncompli- cated and terminates in a very high arcuate transverse band. REMARKS. The only morphological features which this spe- cies has in common with other Terebratulidae of the Upper Cretaceous are the distinctly oval general outline and unipli- cate anterior commissure. The internal structure, notably the distinctive cardinalia and short triangular hinge-plates, are features more typical of terebratulid species of the Upper Jurassic to Lower Cretaceous, the closest comparison being to species of Nucleata and Pygites. However, it is not sug- gested that there is any direct relationship between the species described here as Pseudogibbithyris arabica gen. et sp. noy. and the Tithonian genera mentioned above. As no reference to any forms which could be confidently compared to this species are known, it is treated as a previously undescribed genus and species. It would be unwise at this stage to draw any firm conclu- sions about the phylogenetic relations of this taxon. The simple cardinalia and brachial loop structure seen in the transverse serial sections (Fig. 2), is unusual and suggests a late Jurassic or early Cretaceous ancestry. However, the taxon should remain broadly assigned until more material is obtained, allowing further investigation. OCCURRENCE. All specimens came from the Loftusia-rich facies (beds 3-6) Jebel Huwayyah, section 2 (see volume Introduction for locality details). They are Maastrichtian in age. 276 E.F. OWEN Fig. 2. Sixteen transverse serial sections through the holotype of Pseudogibbithyris arabica gen. et sp. nov. (BMNH BF47) from Jebel Huwayyah, section 2, beds 3-6. Note the bifid cardinal process (sections 5 & 6), the short triangular hinge-plates, and high, arcuate transverse band of the brachial loop. The numerals denote the distance in mm between each section. All x 3. . Bull. nat. Hist. Mus. Lond. (Geol.) 51(2):277-305 Issued 30 November 1995 Late Campanian-Maastrichtian rudists from the United Arab Emirates-Oman border region N. J. MORRIS Department of Palaeontology, The Natural History Museum, Cromwell Road, London SW7 5BD P.W. SKELTON Department of Earth Sciences, The Open University, Walton Hall, Milton Keynes, Buckinghamshire MK76AA CONTENTS Sia CLAP MV GAN Cea PCa ten annccviacsnceeceiie deere snserlacceemiae ae Systematic palaeontology .................s.ceeeeeeeeeeeeeeeeeeees References Se cee ea ee GME Oe See ic eld oleae ctu elas a cluninis wis ec derteloster ogee 277 Pomp Geb aT RRR aDcOGe ae cOc oD ean eben cncascce qa concaC cnn eMeRseHonn tn acts 278 ap sjopebec candle soi aeaale deta sie face eee erie ea ee nteeteaaae a seats 303 Synopsis. About 30 species of Upper Campanian and Maastrichtian rudist bivalves from the Qahlah and Simsima Formations of the United Arab Emirates-Oman border region are described. They include Semailia smithi gen. et sp. nov. of the Dictyoptychidae, Glabrobournonia arabica gen. et sp. nov. of the Radiolitidae (Biradiolitinae), and Pseudosabinia gen nov. of the Radiolitidae (Joufiinae). The geological succession at Qarn Mulayh (Mileih), 7 km west of Jebel Buhays, is described. STRATIGRAPHY AND AGE In addition to the stratigraphy outlined by Smith, Morris and Gale at the beginning of this volume we include here strati- graphical details of the successions at Qarn Mulayh and Qarn Murrah (the ‘red qarns’) situated to the west of Jebels Buhays and Faiyah, UAE (Smith ef al., this volume, fig. 4). At these localities, limestone platform sedimentation on the serpenti- nized front of the Semail Ophiolite started earlier than in the jebels to the east, and is equivalent in age to the arenaceous facies of the Qahlah Formation. A section at Qarn Mulayh is given in Fig. 1. Similar beds also form the majority of the outcrop at Qarn Murrah, to the north. On the basis of the rudists, we suspect that the succession at both qarns is of late Campanian age, and may possibly extend into the earliest Maastrichtian. These lime- stones correlate laterally with the sandy facies of the Qahlah Formation, with only a few of the species present above in the basal beds of the Simsima Formation. Table 1 gives a full list of the rudists collected, and their distribution is shown in Fig. 2. The oldest fauna (1 on Fig. 2) occurs at Qarn Mulayh and Qarn Murrah, and also in the lower Qahlah Formation gravels at Jebels Huwayyah and Bu Milh. It is probably Campanian in age (the M1 fauna of Skelton et al., 1990). The middle fauna (2 on Fig. 2) occurs in the upper Qahlah Formation Loftusia-Beds at Jebel Huwayyah, and the top Qahlah Formation gravels (with acteonellids) at Jebel Bu Milh. It is Campanian/early Maas- trichtian in age (the intermediate M1/M2 fauna of Skelton et © The Natural History Museum, 1995 al., 1990). The upper fauna (3 on Fig. 3) occurs in the main part of the Simsima Formation, at Jebels Faiyah, Buhays, Rawdah and Huwayyah. It is Maastrichtian in age. The level 1 fauna compares closely with the so-called Pironaea-Pseudopolyconites Senonian fauna of Sladic¢- Trifunovic (1989), for which the type area is the “Vrbovac Beds’ of eastern Serbia. Diagnostic novel taxa (not present in the older Gosau Beds) are: *Pironaea, Yvaniella, *Pseudopolyconites, Joufia, Neoradiolites, Branislavia (?=*Colveraia), Sabinia |= * Pseudosabinia here), and Mitro- caprina (asterisks denote those also in the UAR/Oman fauna). In addition, the following were noted by Sladic- Trifunovié: *Vaccinites loftusi, V. ultimus, V. orientalis, V. bacevicensis, Hippurites colliciatus, ?*H. lapeirousei, * Hippu- ritella cornucopiae. Moreover, Sladic¢-Trifunovic (1989: 149) observed: ‘The species *Vaccinites oppeli Douvillé, also found in the Vrbovaé Reef (Baéeviéa), is certainly of a special biostratigraphic importance, since it was previously believed to exist only in Early Senonian’. Sladié-Trifunovié (1989: 153), following the earlier works of Milovanovic and Grubi¢, regarded the assemblage as Upper Campanian/Maastrichtian, mainly based on the sup- posed evolution of Pironaea. She did, however, acknowledge the arguments of others (for an earlier-extending range), based on orbitoids, and agreed that the fauna was probably absent from the Upper Maastrichtian. Pejovi¢ & Radoicic (1987) for example, had revised the age of the ‘Bra¢ Marbles’ (with Pironaea etc) to the Lower-Middle Campanian. Sladic¢- Trifunovié (1989: 154) responded: ‘If the ‘Bra¢ Marbles’, which include P. milovanovici [Kitihn’s species, regarded as 278 Table 1 Systematic list of the rudists found in the Qahlah and Simsima Formations. Family CAPROTINIDAE Gray, 1848 Genus GYROPLEURA Douvillé, 1887 Gyropleura sp. Family PLAGIOPTYCHIDAE MacGillavry, 1937 Genus PLAGIOPTYCHUS Matheron, 1843 Plagioptychus cf. toucasianus Matheron, 1843 Family DICTYOPTYCHIDAE Skelton in Skelton and Benton, 1993 Genus DICTYOPTYCHUS Douvillé, 1905 Dictyoptychus morgani (Douvillé, 1904) Genus EODICTYOPTYCHUS Skelton & El-Asa’ad, 1992 Eodictyoptychus aff. arumaensis Skelton & El-Asa’ad, 1992 Genus SEMAILIA Morris & Skelton, gen. nov. Semailia smithi Morris & Skelton sp. nov. Semailia sp. Family HIPPURITIDAE Gray, 1848 Subfamily TORREITINAE Grubic, 1979 Genus Torreites Palmer, 1933 Torreites sanchezi (Douvillé, 1927) milovanovici Grubic, 1980 Subfamily HIPPURITINAE Gray, 1848 Genus VACCINITES Fischer, 1887 Vaccinites loftusi (Woodward, 1855) Vaccinites vesiculosus (Woodward, 1855) Vaccinites oppeli (Douvillé, 1892) Genus HIPPURITES Lamarck, 1801 Hippurites aff. lapeirousei Goldfuss, 1841 Hippurites cornucopiae Defrance, 1821 Hippurites aff. cornucopiae Defrance, 1821 Genus PIRONAEA Meneghini in Pirona, 1868 Pironaea cf. polystyla Pirona, 1868 Family RADIOLITIDAE d’Orbigny, 1847 Subfamily RADIOLITINAE d’Orbigny, 1847 Genus Praeradiolites Douvillé, 1902 Praeradiolites cf. subtoucasi Toucas, 1907 Genus Radiolites Lamarck, 1801 ?Radiolites sp. Subfamily PPEUDOPOLYCONITINAE Sladi¢-Trifunovic, 1983 Genus Pseudopolyconites Milovanovi¢, 1937 Pseudopolyconites aff. parvus Milovanovic, 1935 Subfamily BIRADIOLITINAE Douvillé, 1902 Genus Biradiolites d Orbigny, 1850 Biradiolites aff. baylei Toucas, 1909 ? Biradiolites aff. baylei Toucas, 1909 Genus Glabrobournonia Morris & Skelton gen. nov. Glabrobournonia arabica Morris & Skelton sp. nov. Sub-family SAUVAGESIINAE Douvillé, 1908 Genus Durania Douvillé, 1908 Durania cf. gaensis (Dacqué, 1903) Durania cf. apula (Parona, 1900) Durania form A Durania form B Durania spp. Subfamily LAPEIROUSIINAE Kihn, 1932 Genus Lapeirousia Bayle, 1878 Lapeirousia sp. Genus Osculigera Kiihn, 1932 Osculigera cf. vautrinioides Vogel, 1970 Subfamily JOUFIINAE Karacabey-Oztemiir 1981 Genus Colveraia Klinghardt, 1921 Colveraia aff. variabilis Klinghardt, 1921 Genus Pseudosabinia Morris & Skelton gen. nov. Pseudosabinia aff. klinghardti (Boehm, 1927) N.J. MORRIS AND P.W. SKELTON ‘advanced’], were accepted as being of the Upper Campanian age, there would be no sense in talking about the evolution of the genus Pironaea’. The Pironaea story has, indeed, been shown to be incorrect by J. M. Pons & E. Vicens (unpub- lished), who suggested that the different forms reflect ontog- eny, rather than phylogeny, while Swinburne (1990: 27) dated the Bra¢ Marbles (= Pucis¢a Formation) as early- middle Campanian, based on Sr isotope correlations, in agreement with Pejovic & Radoicic. Moreover, by the same means, she and others (Swinburne et al., 1992) re-assigned some Pironaea beds in Bulgaria, which had previously been placed at various levels in the Maastrichtian, to the Campa- nian and lowest Maastrichtian. Thus the ‘Pironaea- Pseudopolyconites Senonian’ fauna seems to have ranged from the early Campanian to the earliest Maastrichtian. Additionally, Philip & Platel (1987) assigned their Torreites beds in Dhofar to the Campanian, based on orbitoids, while in the Caribbean the range of Torreites sanchezi (in our level 1 fauna) is also restricted to the Campanian (Rojas, Iturralde- Vinent, & Skelton, in press). In summary, a Campanian age for the level 1 fauna seems the most plausible (though on the rudist evidence alone, we still cannot yet exclude the earliest Maastrichtian). On the other hand, the level 3 fauna presents a marked contrast, having only V. oppeli, H. cornucopiae, Colveraia and Pseu- dosabinia in common with the classic Pironaea- Pseudopolyconites Senonian fauna. H. cornucopiae, though, is also well known from the Maastrichtian of Sicily and elsewhere. As noted in Skelton et al. (1990: 545), this younger fauna is also characterized by some distinctive Arabian/Iranian endemics (e.g. Dictyoptychus). The Level 3 fauna is independently dated by ammonites (Kennedy, this volume) as late Early to early Late Maastrichtian. SYSTEMATIC PALAEONTOLOGY Most of the described and figured rudists are in the collec- tions of The Natural History Museum, and are cited with the prefix BM; many of them have the additional prefix LL, and unless otherwise stated the specimens are in the Morris, Gale & Smith Collection; most of the other material is in the Skelton collection, collected by Skelton and Nolan (Skelton et al, 1990). The morphotype nomenclature used, eg. ‘eleva- tor’, ‘clinger’, ‘recumbent’, is according to the scheme of Gili & Skelton (1994), summarised by Skelton (1991) and Ross & Skelton (1993). Family CAPROTINIDAE Gray, 1848. (emend. Skelton, 1978; =Monopleuridae Munier-Chalmas, 1873) Genus GYROPLEURA Douvillé, 1887 TYPE SPECIES. Requienia cenomanensis d’Orbigny, 1850. REMARKS. There are no accessory cavities. Pl. 1, fig. 2 MATERIAL. Four small specimens from top bed 4 or basal bed 5, Simsima Formation Jebel Rawdah, section 1, ammo- nite and inoceramid horizon; two are attached to the upper valve of an ‘Umbonium’, BM LL41767-69. Gyropleura sp. LATE CAMPANIAN-MAASTRICHTIAN RUDISTS Metres 50 40 30 Texture > re > D ro ne) — fo) > = g =) WY Comments - Some 30 m, exposed only sporadically through dunes: red-stained bioclastic packstones to wackestones, with large gastropods (Campanile), irregular echinoids and small radiolitid rudists. - Localized clusters of moderate to large-sized elevator to radial clinger rudists, in life position: Pironaea, Vaccinites, Torreites, Osculigera. Shelly packstone matrix. - Bioclastic packstone with scattered flakes of radiolitid left valves. - Dense radiolitid flake floatstone. - Packstone with Vaccinites and rare Torreites. Basal lag with Pseudosabinia. - Biostrome of Osculigera and others. - Fine grainstone to packstone, with drifts of radiolitid flakes, and isolated Pironaea, Colveraia and Pseudosabinia. - Grainstone with Pseudosabinia. - Grainstone with toppled Pironaea and Torreites, and radiolitid floatstone cap. - Nodular shelly packstone with burrowed top. Rare small radiolitids, burrowing bivalves and Cyclolites. - (sand-covered) - Bouquets of small radiolitids - Sugary dolomitised calcarenite with limonitic (?ex-serpentinite) grains. Scattered Osculigera, small radiolitids, Plicatula, cyclolitid corals and regular echinoids. Omphalocyclus present. - (some 20 m sand-covered) - Coarse serpentinite and limestone clast conglomerate. Deepening ?Beach Shallow carbonate bank with mobile lime sands and rudist meadows Interpretation KEY Bioclastic limestone Shell-rich bioclastic limestone Orbitoid-rich bioclastic limestone Conglomerate Elevator rudist Recumbent rudist Radiolitid flakes Cyclolitid coral Burrowed surface Serpentinite pebble 279 ‘Fig. 1 Measured section at Qarn Mulayh (Mileih), 7 km west of Jebel Buhays; section logged at the north end of the western flank by P W | Skelton. | | N.J. MORRIS AND P.W. SKELTON 280 tees Bi omelu ta .Os Rk Ric Gon NEO Se sag ail yates Vale nV Sagal lel ee ct 3 ee e« e @ ? ¢ e oo e@ e@ ee e e 2 e e e e @ ] e e ? e e«© ee @ e e @ e Fig. 2. Stratigraphical distribution of the rudists. Ut = stratigraphical units: 1 = lower part of the Qahlah Formation, 2 = upper part of the Qahlah Formation (the gravels and the Loftusia-Beds), 3 = Simsima Formation. Ps = Pseudopolyconites, Bi = Biradiolites, Bo = Glabrobournonia, Du = Durania, La = Lapeirousia, Os = Osculigera, Pk = Pseudosabinia aff. klinghardti, Pr = P. ‘rtanjica’, Co = Colveraia, Eo = Eodictyoptychus, Se = Semailia, Di = Dictyoptychus, P| = Plagioptychus, Pi = Pironaea, V\ = Vaccinites loftusi, Vv = Vaccinites vesiculosus, Vo = Vaccinites oppeli, Hc = Hippurites cornucopiae, H = Hippurites aff. lapeirousei, T = Torreites. DESCRIPTION. All four specimens have the valves conjoined, with the upper left valve slightly exogyriform and the lower valve rather longer but with considerable variation in size of attachment surface; line of commissure slightly sinuous. Outer shell surface of both valves with fine, evenly and closely spaced, radiating striiform ribs. Shell margins crenu- late. Adductor attachments visible in the left valve of speci- men LL41767 on concave surfaces of the hinge plate, a little below the plane of commissure. Gyropleura is a clinger. Family PLAGIOPTYCHIDAE MacGillavry, 1937: 105, 152 (ex. Plagioptychinae MacGillavry, 1937 (plagioptychinés Douvillé, 1888: 729)) Genus PLAGIOPTYCHUS Matheron, 1843. TYPE SPECIES. Plagioptychus paradoxus Matheron, 1843, subsequently designated by Kutassy (1934: 172). REMARKS. Species of Plagioptychus are clingers to low eleva- tors. Plagioptychus cf. toucasianus Matheron, 1843 PI. 1, fig. 1 cf. 1843 Plagioptychus toucasianus Matheron, 117. MATERIAL. Two small specimens from near the base of the Simsima Formation at Jebel Faiyah, BM LL41765 (centre), LL41766 (from section 2), Skelton Collection. DESCRIPTION. Inequivalve, right valve smaller than free left valve, exogyriform with sinuous commissural margin with large attachment area, surface slightly rugose with prominent commarginal growth lines. The rather short exterior dorsal surface has a sub-vertical ligament groove and the shell surface bulges to the posterior of this line. Outer calcitic shell layer thin to medium, with thin inner layer without pallial canals. Prominent tooth and attached myophore projects into upper valve. Free left, upper valve globose, regularly coiled, gryphaeate with smooth surface; outer calcitic shell layer thin, inner layer recrystallized to calcite but medium to thick, with radially elongated narrow canals that do not pass into polygonal PLATE 1 Fig. 1 Plagioptychus cf. toucasianus Matheron, from Jebel Faiyah, lower part of Simsima Formation. 1a, anterior view, 1b, section through structure and in that way differ from Mitrocaprina or the upper valve of Coralliochama. The narrow canals also do not bulge internally as is common in some forms of the genus. Anterior myophore robust and nearly level with the commis- sure. An oblique septum runs from the anterior tooth to the ventral margin. The cavity posterior to this septum houses the large central tooth and conjoined posterior myophore in a dorsal position. Family DICTYOPTYCHIDAE Skelton in Skelton & Benton, 1993 (ex. Trechmannellidae Cox, 1933: 65) DIAGNOSIS. Inequivalved rudists, attached by right valve. Valves uncoiled and ligament absent. Outer (calcitic) shell | layer compact. Inner (originally aragonitic) shell canaliculate throughout, in both valves. Two projecting teeth in left valve, straddling ridge-like central tooth in right valve. Posterior tooth dorso-ventrally flattened, flanking body-cavity, and separated from the dorsal margin by a small accessory cavity, which may be a relic of the ligamentary cavity. Anterior myophoral platforms extended both dorsally on hinge plate, around anterior tooth, and ventrally from hinge plate. Poste- rior myophore of left valve projecting, with adductor scar facing outwards, into recess, or socket, in posterior wall of right valve, and directly flanking body-cavity. REMARKS. The distinctive features of this family were dis- cussed, in relation to Eodictyoptychus, by Skelton and El-Asa’ad (1992). The dictyoptychid myocardinal apparatus differs from that seen in the Caprinidae (s.s.), the Plagiopty- chidae, and Sabinia (s.s.) in all of which the posterior tooth and posterior myophore (with inward-facing muscle scar) of the left valve are separated from the body-cavity by the combined central tooth socket and an accessory cavity,) extending ventrally from it, which receives the salient myo- phore of the right valve. The latter cavity is itself demarcated by a prominent lamina running from the anterior tooth to the postero-ventral margin of the valve. No equivalents of this) lamina, and the associated accessory cavity, are present in the) left valve of dictyoptychids. Pseudosabinia (Radiolitidae) differs in its possession of a | left valve approximately 5 mm from the commissure, BM LL41766, x 1, Skelton Collection 1c, dorsal view, BM LL41765, x 1. Fig. 2 Gyropleura sp., from Jebel Rawdah, section 1, loose from top of bed 2 or bed 3 of Simsima Formation; 1a, posterior view, 1b, dorsal view, Ic, view looking down on left valve, BM LL41768, x 2. Fig. 3 Dictyoptychus morgani (Douvillé), from Jebel Buhays, section 1, bed 11 of Simsima Formation, side view, BM LL41680, x 0-5. Fig. 4 Eodictyoptychus arumaensis Skelton & Al-Asa’ad, from Jebel bu Milh, section 1, top of Qahlah or basal Simsima Formation; 4a, top view, 4b, internal view, 4c, dorsal view, BM LL41927, left valve, x 1-5. 281 LATE CAMPANIAN-MAASTRICHTIAN RUDISTS 282 ligamentary invagination, and the presence of cellulopris- matic structure in the outer shell layer of the right valve. Despite the similarity to the Antillocaprinidae MacGillavry, 1937, in their lack of a ligament, and the presence of the small dorsal accessory cavity, which may be a relic of it, the dictyoptychids differ from Antillocaprina (at least) in having the posterior myophore of the left valve projecting, rather than parallel with the plane of commissure. Moreover, the outer shell layer of antillocaprinids is invariably very thin, whilst that of dictyoptychids is usually relatively thick in the right valve. Genus DICTYOPTYCHUS Douvillé, 1905. (nom. nov. for Polyptychus Douvillé 1904, non Huebner, 1816) TYPE SPECIES. Polyptychus morgani Douvillé, 1904, by monotypy. SYNONYM. Trechmannella Cox, 1933 (obj.). REMARKS. Attached right valve with large polygonal canali- cular structure. Outer layer of right valve thicker than that of left valve, with exposed growth margin; sharply peaked ridges on right valve growth margin in some specimens. We agree with Pons et al., 1992, who regard all described species as probably synonymous. Dictyoptychus morgani (Douvillé, 1904) Pipe ties'3 1904 Polyptychus morgani Douvillé: 248-51, pl. 33 bis. 1905 Dictyoptychus morgani (Douvillé); Douvillé: 198. MATERIAL. Jebel Faiyah, section 1, bed 3, BM LL41670-71; section 2, LL41659; Jebel Faiyah, centre, LL41683-—84. Jebel Agabah, Simsima Formation, loose, BM LL41676. Jebel Thanais, Simsima Formation, loose in scree, BM LL41668-69. Jebel Buhays, section 1, bed 11, LL41677, 41680; section 1b, LL41674; section 1a, Simsima Formation, BM LL41675. Jebel Rawdah, section 1, bed 3, BM LL41657-58, 41679, 41681; section 2, bed 19, LL41673; section 2, loose on scree, LL41667; section 4, bed 1, LL41672; Jebel Rawdah, north, LL41690. Jebel Bu Mihl, section 1, bed 3, BM LL41660-66. Jebel Huwayyah, section 2, beds 3-5, BM LL41678; Jebel Huwayyah, LL41685-89. Jebel Sa’ah, basal Simsima Formation, BM LL41682. REMARKS. Occurs widely in the lower half of the Simsima Formation, sometimes in great numbers. Dictyoptychus is a multigeniculate elevator (see Ross & Skelton, 1993: fig. 5.2). Genus EODICTYOPTYCHUS Skelton & El-Asa’ad, 1992 TYPE SPECIES. Eodictyoptychus arumaensis Skelton & El-Asa’ad, 1992, by original designation. PLATE 2 N.J. MORRIS AND P.W. SKELTON Eodictyoptychus aff. arumaensis Skelton & El-Asa’ad, 1992. Pl. 1; fig. 4; Pl 25 fig: 2 vy aff. 1992 Eodictyoptychus arumaensis Skelton & El-Asa’ad: 108-13, pl. 1, figs 1-6. MATERIAL. Jebel Bu Milh, section 1, Qahlah/Simsima For- mation Boundary, BM LL41927-8; section 2, LL41929; all are left valves. A possible fragment of a right valve from Qarn Murrah, BM LL41977, Skelton Collection. DESCRIPTION. Moderately inaequivalved, free left valve with thin outer calcitic shell layer, inner aragonitic layer recrystal- lised but formed of varyingly sized canals of polygonal section, even within the teeth and the massive myophores. Two subequal, projecting teeth in free valve, the anterior one below the umbo, the smaller, posterior one ventral to this, the two separated by a socket whose centre lies above and towards the umbo from the posterior tooth. The teeth lie on the body-cavity side of a massive plate which includes the attachment surfaces of the myophores, and takes up more than half of the area within the valve when looking down on the commissural plane. We have not been able to detect the accessory cavity dorsal to the posterior tooth, but this may be due to the difficulty of its preparation. The projecting posterior myophore is sepa- rated from the body-cavity by a narrow ledge which is not present in the type material from Khashm Buwaibiyat, approximately 50 km north of Riyadh. REMARKS. Eodictyoptychus varies from a lateral clinger to a recumbent. Genus SEMAILIA Morris & Skelton, gen. nov. TYPE SPECIES. Semailia smithi Morris & Skelton gen. et sp. nov. DIAGNOSIS. Bicornate and multicarinate, subequivalve with a thin outer shell layer in each valve. Inner shell canaliculate in both valves. Myocardinal arrangement typical of dictyopty- chids with two teeth in the left valve, the posterior tooth dorso-ventrally flattened and flanking the body-cavity with- out an intervening accessory cavity. No ligament present. The posterior myophore of the left valve projecting into a socket in the right valve, again flanking the body-cavity without an intervening accessory cavity. REMARKS. The absence of a posterior accessory cavity in the left valve excludes this taxon from the Caprinidae, Plagiopty- chidae and Sabinia s.s., and the outward facing, ie monopleu- riform, myophores, are typical of the dichtyoptychid plan. Unlike other members of the Dictyoptychidae this genus does not show the differentially greater thickening of the outer calcitic layer of the right valve. Fig. 1 Semailia smithi Morris & Skelton, gen. & sp. nov. Jebel Huwayyah, section 2, Loftusia-Beds; 1a, anterior view, 1b, posterior view, Ic, view of right valve, Id, dorsal view; holotype, BM LL41931, x 1. Fig. 2 Eodictyoptychus aff. arumaensis Skelton & Al-Asa’ad, Jebel bu Milh, section 1, top of Qahlah or base of Simsima Formation; internal view of left valve, BM LL41928, x 1. LATE CAMPANIAN-MAASTRICHTIAN RUDISTS 283 284 N.J. MORRIS AND P.W. SKELTON Fig. 3 Semailia smithi Morris & Skelton sp. nov. Four sections through the holotype approximately 10 mm either side of the commissure, BM LL41931; a, b, right valve, MC — accessory cavity for posterior myophore of left valve, PO — posterior tooth socket, AC — accessory cavity dorsal to tooth socket, CI — central tooth, AO — anterior tooth socket; ¢, d, left valve, with fine outer and large irregularly polygonal inner canals; X 1. PLATE 3 Fig. 1 Semailia sp., Jebel bu Milh,section 2, base of Simsima Formation; 1a, ventral view of broken right valve, 1b, dorsal view of right valve; BM LL41932 x 0-75. Fig. 2 Torreites sanchezi milovanovici Grubi¢; Haushi-Hugf Massif, Eastern Oman, BM LL41975, Samir Hanna Collection; 2a, view looking down onto commissural plane with broken left valve partly preserved in situ, X 1; 2b, dorsal view showing intucking of pillar L, 2c, posterior view showing infolds of pillars, 2d, ventral view; 2b-d, x 0-5. Fig. 3 Vaccinites aff. oppeli (Douvillé), Jebel bu Milh, section 2, Simsima Formation, bed 10, BM LL41730, marginal surface of outer shell layer of right valve, x 1. Semailia smithi Morris & Skelton sp. nov. PI. 2, fig. 1, Fig. 3 MATERIAL. A single specimen, the holotype, BM LL41931, from Jebel Huwayyah, section 2, Loftusia-Beds. DIAGNOSIS. Typical of the genus but with three strong cari- nae or flaring radial costae on each valve. DESCRIPTION. Both valves are preserved, closed, although the body-cavity is matrix-filled and includes larger foramin- ifera. The specimen has two serpulids attached, one to each valve and aligned radially with respect to the umbones, with their apertures close to the valve commissure, amid the plicae of the ventral margins. The shell is curvingly biconical, with the two valves sepa- rately following almost a semicircular direction of growth, in LATE CAMPANIAN-MAASTRICHTIAN RUDISTS 285 286 a single plane, so that the umbones or apices are beginning to approach each other; sub-equivalve with the right valve slightly ‘longer’ than the left. Three prominent carinae or flaring radial costae, skewed to the posterior, two at approxi- mately 180°, set anteriorly and posteriorly on both valves and the third normal to these on the ventral margin. Dorsal margin rounded. Shell surface otherwise relatively smooth on both valves. No ligament present. Very thin outer calcitic shell layer in both valves. Canals are present in the thick inner shell layer of both valves, round and capillary-like around the margins, becoming larger inside, irregularly rounded polygonal in section, especially in the left valve. Two teeth in the left valve are typical of the family, the anterior one is rounded and knob-like, the posterior one is dorso-ventrally flattened. One central tooth in the right valve. The posterior myophore of the left valve projects into an accessory cavity (socket) in the right valve. The anterior myophore of the left valve is a broad shelf and has large canals. There is an accessory cavity in the right valve which lies dorsally to the flattened posterior tooth socket. COMMENTS. The crescentic form of the shell suggests a recumbent life position, possibly reclining on the dorsal flank. It was found in a matrix of marly limestones with large specimens of Loftusia with a similar matrix filling the body- chamber, forming a loftusid packstone. COMPARISON WITH OTHER TAXA. There is a great similarity in the myocardinal arrangement with that of other dictyopty- ichids, but Semailia differs in having a much thinner outer shell layer in both valves and having strong shell carinae. Pl. 3, fig. 1 MATERIAL. A single specimen from Jebel Bu Milh, section 2, basal Simsima Formation, BM LL41932. Semailia sp. DESCRIPTION. The specimen is a large right valve, its anterior and posterior part nearly symmetrical about a dorso-ventral plane. Multicarinate but otherwise smooth. Two wide ante- rior and posterior carinae form a wide kite-shape in dorsal view. Dorsal surface flattish near the commissure, but umbones incurved and separating two gently concave ante- rior and posterior areas with a low mid-dorsal carina that is prominent at the umbo, but faces towards the mid-dorsal margin. Ventral part of shell has three sub-equal strong carinae, which are equidistant from the dorsal margin and inclined towards the anterior. Outer shell layer thin and now formed of structureless recrystallized blocky calcite. Thick canaliculate inner shell layer with capillary-like polygonal canals throughout, includ- ing the teeth, fine at the margins becoming larger inwards, now also recrystallized to calcite. Right valve has part of projecting stout central tooth preserved behind base of the socket for the anterior tooth of PLATE 4 Fig. 1 N.J. MORRIS AND P.W. SKELTON the left valve and in front of the socket of the posterior tooth of the left valve, containing a fragment of that dorso-ventrally flattened tooth. COMPARISON WITH OTHER SPECIES. Semailia sp. differs from S. smithi in having two additional ventral carinae. There is not enough material to know if this difference is significant in discriminating separate species. Family HIPPURITIDAE Gray, 1848 Subfamily TORREITINAE Grubic, 1980 Genus TORREITES Palmer, 1933 TYPE SPECIES. Hippurites (Vaccinites) sanchezi Douvillé, 1927, by original designation. DESCRIPTION. Outer shell layer of right valve with tight marginal infoldings, giving rise to radiating crests (Douvillé, 1894) around the shell margin of the right valve, but variable in extent and number. Outer shell layer of left valve thin, postero-dorsally digitiform, smooth except for fine growth lines on the upper surface, but becoming overgrown with epibionts radiating from its low apex. Inner shell layer of both valves partly canaliculate, canals of right valve relatively large, sub-radial and sub-rectangular in section, in the area of the anterior muscle attachment. We have observed a similar reticulate network of vertical ridges separating tabulate canals in the anterior myophoral ledge of Vaccinites gosavien- sis (BM 33972, a specimen with the original aragonite pre- served). The canals of the left valve are narrow and radiate from the apex. REMARKS. Philip & Platel (1994) pointed to the similarity of Torreites with their new genus, Praetorreites, from the Lower Campanian, Samhan Formation of south-east Oman. The latter has canaliculate structure of the inner shell layer of the left valve and regular pedunculate radial structures in the outer shell layer of the right valve, which, they claim, are comparable with the radiating marginal crests of Torreites. They compare the subfamily, raised to family rank, with both the Plagioptychidae and the Hippuritidae, concluding that the similarity is greater with the Plagioptychidae. If Philip & Platel are correct in their suggestion that the Torreitinae are not hippuritids, then it follows that the three large infolds of the outer shell layer of the right valve are not homologues of the hippuritid pillars. The origin of two of the pillars of Torreites is claimed to be from the two small ‘pillars’ of Praetorreites and considered by Philip & Platel to be ana- logues of pillars ‘E’ and ‘S’. Philip & Platel’s plate 8, fig. 1 shows the position of these ‘pillars’ in relation to the teeth and myophores of the upper valve. If this arrangement is compared with the hinge structures of Dictyoptychus (Dou- villé, 1904: pl. 33 bis, fig. 4), the alignment of the sockets and teeth is very similar, although they are at a more obtuse angle Torreites sanchezi milovanovici Grubi¢, Haushi-Hugf Massif, Eastern Oman (also figured Skelton & Wright, 1987: pl. 67, fig. 1), BM LL28004, view looking down onto right valve (note coarse cellular structure of myophore in lower part of figure and tooth sockets to the right of pillar ‘L’), x 1. Fig. 2 Vaccinites aff. oppeli (Douvillé), Jebel bu Milh, section 2, Simsima Formation limestone, BM LL41733, view of naturally broken section of right valve, x 0-4. Fig. 3 Vaccinites vesiculosus (Woodward), Jebel Huwayyah, Loftusia-Beds, BM LL41716, Skelton Collection, 3a, view of partly eroded left valve, 3b, view of right valve, x 0-5. renee eee aaa iii eee inated aeeeeiena eked piaiinas = eeeaieaaedin aiTiaieaan aioe Ela EtEaiSeEe aE ceREGE eA enE — — LATE CAMPANIAN-MAASTRICHTIAN RUDISTS 287 288 N.J. MORRIS AND P.W. SKELTON to the ventral shell margin. This suggests a relationship between Dictyoptychus and Praetorreites. It is difficult to relate the anterior infolding of Torreites to the position of ligamentary invagination of the plagiopty- chids, but much easier if Torreites is interpreted as a hippu- ritid. Details of the morphology are illustrated by Skelton & Wright (1987: fig. 2). In Skelton & Wright’s interpretation of Torreites loss of the hippuritid canal system of the outer shell layer of the left valve is by recurvature of the shell margin to expose the mantle margins. The umbo of the operculiform left valve is close to the ventral margin. Philip & Platel (1994: fig. 4) claim a diphyletic origin for the ‘three-pillared’ genus Torreites and thus claim to have refuted the palaeobiogeographical interpretation of Skelton & Wright (1987). Their diagram shows Praetorreites occur- ring before Torreites and giving rise independently to the Caribbean and Arabian species. However, their date for the small Caribbean species Torreites tschoppi on this diagram does not show its full range. There is ample evidence to show that this species is best dated as Santonian in Cuba (Rojas et al., in press) and therefore pre-dates Campanian Praetorre- ites. Their analysis leading to the view that the torreitines are not closely related to the hippuritids relies heavily on the Fig. 4 Torreites sanchezi milovanovici Grubié; smooth top surface interpretation of the ancestral status of Praetorreites and the of left valve with advancing front of epibionts (arrowed), BM supposed homology of the pedunculate folds, with the mar- LL41975, x 2. ginal radiating crests of Torreites, which in the latter are a consequence of tight infoldings of the outer shell surface. A superficially similar structure in section may be seen in the outer shell layer of the lower valve of Dictyoptychus striatus, but this is formed as a consequence simply of salient radial ridges on the growth margin of that shell layer, without any infolding of the outer surface. In Philip & Platel’s (1994: pl. 7, fig. 3) illustration of Praetorreites the similarity, if anything, seems greater with the radial ridges of the dictyophychid, than with the intuckings of Torreites. In all particulars Prae- torreites is similar to Eodictyoptychus and appears to have little in common with Torreites. The criticism of Skelton and Wright’s (1987) explanation of the distribution of Torreites is therefore unconvincing. We Fig. 5 Camera lucida drawing of Vaccinites loftusi (Woodward); maintain the view that the differences between the Caribbean from Qarn Murrah, BM LL41933, Skelton Collection, x 1. and Arabian Torreites do not warrant greater that sub- specific separation, and that their similarities do reflect that Torreites sanchezi (Douvillé 1927) subsp genetic interchange between the two areas of occurrence did OVA O MEAG Enbic 1980 ; F take place. ; ; a Pl. 3, fig. 2; Pl. 4 fig. 1 1927 Hippurites (Vaccinites) sanchezi Douvillé: 54, 55, pl. 4, fig. 1. 1980 Torreites milovanovici Grubic¢: 92, 93, pl. 1, fig. 1, fig. 4, PLATE 5 Figs 1,2 Hippurites aff. lapeirousei Goldfuss; 1, Jebel Faiyah, section 1, Simsima Formation, bed 6, BM LL41754, mass of variously orientated individuals, x 0-5; 2, Jebel Faiyah, section 1b, Simsima Formation, bed 2, BM LL41755, view looking down onto a pair of right valves, X 2. Fig. 3 Vaccinites vesiculosus (Woodward), Jebel Huwayyah, Loftusia-Beds, BM LL41973, V. Chalmers Collection, view looking down on eroded left valve exposing pillars of right valve, x 1. Figs 4-7 Hippurites cornucopiae Defrance; 4-6, Jebel Faiyah, loose from low in the Simsima Formation, Skelton Collection; 4, BM LL41747, view of left valve (pores appear polygonal and radially vermiculiform where less eroded), x 1; 5, BM LL41744, eroded left valve showing pores and canals, x 1; 6, BM LL41745, view of left valve with oscules and pores appearing vermiculiform or polygonal depending on the degree of erosion, x 1; 7, BM LL41737, Jebel Thanais, natural section through right valve showing slightly pedunculate pillars, x 1. Fig. 8 Hippurites aff. cornucopiae Defrance, Jebel Rawdah, southern flank, loose from Simsima Formation, BM LL41753, Skelton Collection, natural section of right valve, x 0-5. Fig. 9 Pironaea cf. polystyla Pirona, Qarn Mulayh, BM LL41938, Skelton Collection, section through right valve, x 1. 289 LATE CAMPANIAN-MAASTRICHTIAN RUDISTS 290 1994 Torreites milovanovici Grubi¢; Philip & Platel: pl. 8, fig. 4, fig. 4. (For further synonomy see Skelton & Wright, 1987) TYPE SPECIMEN. The holotype of the subspecies is BM LL27699, Iraq Petroleum Company Collection (CP 86), on weathered surface of section, 7-7 m from top at Qarn Mulayh. OTHER MATERIAL. Two specimens from the same horizon on Qarn Murrah, BM LL41761, 62; a well-preserved specimen showing details of the upper valve, from the Haushi-Huqf Massif, eastern Oman, S. Hanna Collection, BM LL41975; the material of Skelton & Wright (1987) from eastern Oman, BM LL28004; one specimen from Jebel Huwayyah, section 1, Qahlah gravels, BM LL42763; Qarn Mulayh, no specimens collected. COMMENTS. BM LL41975 shows a boundary between shiny outer shell material and encrusted surface on the outer surface of the upper valve which we interpret as representing the limit of encroachment upon the exposed left mantle surface by epibionts. BM LL28004 shows the loose canalicu- lar structure of the anterior muscle attachment area of the right valve together with the tooth sockets. Subfamily HIPPURITINAE Gray, 1848 Genus VACCINITES Fischer, 1887 TYPE SPECIES. Hippurites cornuvaccinum Bronn, 1931, by monotypy. Vaccinites loftusi (Woodward, 1855) 1855 1897 Fig. 5 Hippurites loftusi Woodward: 58, pl. 3 figs 2-3. Hippurites loftusi Woodward; Douvillé: 210, pl. 33 (17), figs 1, la, 1b. Vaccinites loftusi (Woodward); Toucas: 82, 83, figs WA, WF. 1904 MATERIAL. Three specimens from Qarn Murrah, BM LL41933, LL41969-70; another from Jebel Huwayyah, Loftusia-Beds, LL41934; Skelton Collection. REMARKS. These specimens have the typical coarse external ribs of Woodward’s species but are very poorly preserved internally. The pillars of a Qarn Murrah specimen resemble those of Woodward’s type but were not enhanced by section- ing. The pores of the upper valve are in the form of coarse polygons and the outer surface of this left valve has radial undulations. Woodward’s type material has incipient multiple infoldings of the outer shell layer of the right valve, remins- cent of Pironaea. PLATE 6 Fig. 1 concave smooth dorsal radial band to the right, x 1. Fig 2 ventral view, 2c, anterior view; all Xx 1-5. N.J. MORRIS AND P.W. SKELTON Vaccinites vesiculosus (Woodward, 1855) PI. 4, fig. 3; Pl. Satiens Hippurites vesiculosus Woodward: 59, pl. 4, fig. 6. Hippurites vesiculosus Woodward; Douvillé: 201, pl. 29 (13), figs 6, 7. Vaccinites vesiculosus (Woodward); Toucas: 110, 111. 1855 1897 1904 MATERIAL. Qarn Murrah, BM LL41691-7, LL41721-26, Skelton Collection; Jebel Huwayyah, section 1, bed 9, BM LL41698-702; section 2, bed 7 (the main coral bed), BM LL41703+4; section 2, beds 2-7, BM LL41705-15 plus another 9 specimens; Jebel Huwayyah, loose, Skelton Collec- tion, BM LL41716—-20, V. Chalmers Collection, BM LL41973; three possible specimens from Jebel Thanais, loose in scree below lowest 3m of the Simsima Formation, BM LL41935-37. REMARKS. The specimens from the Loftusia-Beds at Jebel Huwayyah are larger than either those from Qarn Murrah or Woodward’s type material. They however have the same type of pillars in the right valve with a long thin aréte cardinal and pedunculate pillars. The complex pattern of the canals and pores of the left valve are also similar in each of the three groups of material. V. vesiculosus is the dominant rudist species in the Loftusia-Beds at Jebel Huwayyah. Vaccinites aff. oppeli (Douvillé, 1892) Pl. 3, fig. 3; Pl. 4 igs 2 aff. 1866 Hippurites dilatatus Defrance; Zittel: 142, pl. 24, figs 1-5. aff. 1881 Hippurites zitteli Munier-Chalmas in Zittel: 83, fig. 118 (non Matheron, 1880). aff. 1892 Hippurites oppeli Douvillé: 36, 37, figs 23, 24, pl. 4 (18), fig. 5. aff. 1897 Hippurites oppeli Douvillé: 203, pl. 31 (15), figs 1, la. aff. 1904 Vaccinites oppeli (Douvillé); Toucas: 109, 110, pl. 17 (18), figs 2, 2a. MATERIAL. Jebel Bu Mihl, section 2, common, many may be in life position, lower part of the main Simsima Limestone, BM LL41728-30, 41733; Hill to south of Jebel As-Saifr (east of Jebel Huwayyah), BM LL41731-32, Skelton Collection. REMARKS. In this very large form, the pores of the upper valve are present in one specimen and are similar to the pore system in topotypic material from Gosau, particularly on the outer third of the radius. The outer shell surface is rather smooth. The raised pattern on the growth surface of the outer shell layer of the right valve is extremely similar to the Austrian material (cf. Zittel, 1866: fig. 1). The pillars in the right valves of our material match very closely with those of Durania Form A, Jebel Huwayyah, section 1, from the Qahlah gravels, BM LL41948, part of outer shell layer of right valve with ?Radiolites sp., Jebel bu Milh, section 2, beds 7/8 at the Qahlah/Simsima boundary, BM LL41947, right valve, 2a, view of interior, 2b, Figs 3,4 Praeradiolites cf. subtoucasi Toucas, Jebel Rawdah, section 4, bed 1, basal rudist bed of Simsima Formation; 3, BM LL41941, section across lower part of right valve, x 1; 4, BM LL41942, view of anterior of right valve, x 1. Fig. 5 Pseudopolyconites aff. parvus Milovanovi¢, Qarn Mulayh, BM LL41974, Skelton Collection, x 1; 5a, section through right valve with aréte cardinale at top; 5b, surface of part of right valve with smooth ventral radial band centre and sediment with spines to right. Fig. 6 Durania cf. apula Parona, Jebel Rawdah, section 2, Simsima Formation, bed 10, BM LL41951; 6a, ventral view of right valve showing radial bands; 6b, part of upper surface of right valve with small holes near the radial bands, x 1. LATE CAMPANIAN-MAASTRICHTIAN RUDISTS 291 yf J ‘ yo 292 the Austrian material. The differences seem to relate only to phyletic size increase. Genus HIPPURITES Lamarck, 1801 TYPE SPECIES. Hippurites bioculatus Lamarck, 1801, by monotypy. Hippurites aff. lapeirousei Goldfuss, 1841 PI. 5, figs 1, 2 aff. 1895 Hippurites lapeirousei Goldfuss; Douvillé: 164, pl. 24 (11), figs 7-10. Orbignya lapeirousei Goldfuss; Toucas: 53, pl. 6 (12), figs 10, 11. aff. 1903 MATERIAL. Jebel Faiyah, Sharjah, section la, bed 6, BM LL41754; section 1b, bed 2b, BM LL41755; Jebel Faiyah, loose, BM LL41758, Skelton Collection; Jebel Rumaylah, BM LL41756, Skelton Collection; Jebel Mundassah, BM LL41757, Skelton Collection. REMARKS. This species is present in large masses of disorien- tated specimens close to coral patches, low in the Simsima Formation at Jebel Faiyah. It has broad and short pillars and lacks an aréte cardinale. It differs from typical specimens from Maastricht in having a rather smooth shell surface to its right valve. Pl. 5, figs 4-7 aff. 1821 Hippurites cornucopiae Defrance: 195, pl. 58, figs la, 1b (only). 1910 Hippurites (Hippuritella) cornucopiae Defrance; Douvillé: 79, pl. 7, figs 3-5. MATERIAL. Jebel Faiyah, section 1, BM LL41735; Jebel Faiyah, loose, BM LL41738, 4174447, Skelton Collection; Jebel Buhays, section 1, bed 15, BM LL41739-43; section 3, BM LL41736; Jebel Thanais, BM LL41737, Skelton Collec- tion; Jebel Rawdah, section 2, BM LL41734. Hippurites cornucopiae Defrance, 1821 REMARKS. The pores of the upper valve, together with the disposition and shape of the pillars of the lower valve, match those described by Douvillé from Sicily, which is the type locality of Defrance’s species. However, although the pores are polygonal and denticulate even when the surface is only slightly eroded, they are radially vermiculiform when the outer surface is intact. Hence the species should be assigned to Hippurites not Hippuritella (pace Douvillé, 1910). Rather conical specimens occur at the coral patch horizon at Jebel Faiyah, sometimes within the coral clumps. Hippurites aff. cornucopiae Defrance, 1821 PI. 5, fig. 8 MATERIAL. Jebel Faiyah, BM LL41750, LL41752, Skelton Collection; Jebel Rawdah, section 3, above Simsima conglo- morate, BM LL41748; Jebel Rawdah, loose, BM LL41749, 41751, 41753, Skelton Collection. PLATE 7 N.J. MORRIS AND P.W. SKELTON REMARKS. Specimens from Jebel Faiyah and Jebel Rawdah that occur well up the sequence of the Simsima Formation are of Upper Maastrichtian age. They are similar in general plan to H. cornucopiae but are much larger, always being more than twice the diameter of the latter. Genus PIRONAEA Meneghini in Pirona, 1868 TYPE SPECIES. Hippurites polystylus Pirona, 1868, p. 511. Pironaea cf. polystyla Pirona, 1868 Piles) ioe 9. 1868 Hippurites polystylus Pirona: 511. MATERIAL. Three specimens from Qarn Mulayh, loose from lower part of sequence, BM LL41938—40, Skelton Collection; the species was also seen at Qarn Murrah (see Skelton ef al. 1990: fig. 9a). DESCRIPTION. Outer shell layer of medium thickness with numerous secondary pillars. Inner shell layer of right valve slightly thicker than outer shell layer, recrystallised. COMMENTS. Swinburne et al. (1992) have shown that the supposed evolutionary sequence of Pironaea ‘species’ described as Maastrichtian in Serbia are more likely to be of Campanian to earliest Maastrichtian age. Family RADIOLITIDAE d’Orbigny, 1847 (as Radiolidae, emend. Gray, 1848: 439) Subfamily RADIOLITINAE d’Orbigny, 1847 Genus PRAERADIOLITES Douvillé, 1902, p. 467 TYPE SPECIES. Radiolites fleuriaui d’Orbigny, 1842, from the Cenomanian of Le Mans, by original designation. DESCRIPTION. Ligamentary invagination of right valve usu- ally present. Right valve elongate, left valve operculiform. Praeradiolites cf. subtoucasi Toucas, 1907 Pl. 6, figs 3, 4 cf. 1907 Praeradiolites subtoucasi Toucas: 31, pl. 3 (13), figs 8, 9. MATERIAL. Four specimens from Jebel Rawdah, section 4, from the Simsima Formation, bed 1, BM LL41941—-44. DESCRIPTION. A slightly distorted attached (right) valve, ‘D’-shaped in transverse section, which has developed a secondary bilateral symmetry about a dorso-ventral plane. Moderately wide radial bands are separated by a narrower sinus, on the ‘flat face’ which is formed at the postero-ventral margin. The two radial bands are wider than the central sinus and all three are deep below the plane of commissure. Internally the symmetry differs from the outside, with the ligamentary invagination set at about 30 degrees to the posterior of the axis bisecting the external ‘D’ shape. COMPARISON WITH OTHER SPECIES. The sinuses are deeper Figs 1-4 Durania cf. gaensis (Dacqué), Jebel bu Milh, section 1, Qahlah Formation; all x 0-75; 1, BM LL41922, individual from first clump, natural section showing shell layers and tabulae of right valve and part of surface and myophore of left valve; 2, BM LL41923, individual from second clump, commissural surface of right valve; 3, BM LL41924, individual with domed left valve from third clump, internal mould with some shell adhering; 3a, postero-dorsal view; 3b, basal view of right valve; 3c, anterior dorsal view; 4, BM LL41922, second individual from first clump; 4a, postero-dorsal view; 4b, postero-ventral view. 293 LATE CAMPANIAN-MAASTRICHTIAN RUDISTS 294 than those of P. aristidis (Munier-Chalmas, 1888) as figured by Toucas (1907, pl. 6 (16), figs 9, 10). They are more similar to P. boucheroni (Bayle) as figured by Toucas (1907, pl. 3 (13), fig. 10 only) and to P. subtoucasi Toucas (1907, pl. 3 (13), figs 8, 9). Genus RADIOLITES Lamarck, 1801 DESCRIPTION. Widely biconical with upper valve shorter than lower right valve. Outer layer of fixed right valve thick with calcite cellular structure radially stretched. Aréte cardi- nale usually present, usually short. Pl. 6, fig. 2 MATERIAL. Three specimens From Jebel Bu Milh, section 1, from the Qahlah/Simsima boundary, BM LL41945—46; Jebel Bu Milh, section 2, beds 7, 8, BM LL41947; one probable specimen from Jebel Rawdah, section 2, from scree, LL41980. ?Radiolites sp. DESCRIPTION. The simple smooth radial bands, the flaring growth rugae-like stacked projecting cones, and the radially aligned cell pattern of the outer shell layer of the right valve are similar to Radiolites, particularly the groups of Radiolites radiosus and R. sauvagesi (Toucas 1908). The radial bands are not thrown into stong folds and therefore this species does not belong to Praeradiolites. No aréte cardinale is visible and it may well be absent. This is a common trend in many rudist lineages and we suspect that it may happen independently in this form of Radiolites. Subfamily PPRUDOPOLYCONITINAE Sladi¢-Trifunovic, 1983b, p. 239 (ex. Pseudopolyconitidae Sladi¢-Trifunovic1983b, emend. herein) Genus PSEUDOPOLYCONITES Milovanovic, 1937 [Pseudopolyconites Milovanovi¢, 1935, was invalid because no type species was originally designated, in contravention of ICZN Rules, Art. 13A (b)] TYPE SPECIES. Pseudopolyconites parvus Milovanovic¢, 1935. Pseudopolyconites aff. parvus Milovanovic, 1935 PI. 6, fig. 5 Pseudopolyconites parvus Milovanovié: 188, 252 (nomen nudum). Pseudopolyconites parvus Milovanovic: 54-70, figs 1b-8. Pseuodopolyconites parvus Milovanovi¢; Milo- vanovic: 4-14, figs 2-9. 1934 aff. 1935 aff. 1937 MATERIAL. Two specimens from Qarn Murrah, BM LL41978-79, and one specimen collected loose at Qarn Mulayh, BM LL41974; all Skelton Collection. PLATE 8 N.J. MORRIS AND P.W. SKELTON REMARKS. The specimen LL41974 is well-preserved and shows the spines and a wide smooth ventral radial band. The aréte cardinale is long and narrow, with a typically rounded to ovoid distal end. The left valve LL41978 shows the aréte cardinale and the spine bases on its upper surface. Subfamily BIRADIOLITINAE Douvillé, 1902 Genus BIRADIOLITES d@ Orbigny, 1850 TYPE SPECIES. Biradiolites canaliculatus d’Orbigny, 1850 (ICZN Opinion pending). Biradiolites aff. baylei Toucas, 1909 Pl’ 8; hes 133 aff. 1909 Biradiolites baylei Toucas: 119, pl. 24 (9), figs 9, 10. ) Biradiolites royanus (d’Orbigny); Toucas: 103, pl. 19 (4), figs 34-38. Biradiolites aquitanicus Toucas: 107, pl. 20 (5), fig. 20. 21967 Biradiolites bulgaricus Pamouktchiev: 35, pl. 1, fig. 4; pl. 2, figs 3, 4. MATERIAL. Five specimens from Jebel Huwayyah, section 2, four from the top of the Loftusia-Beds, BM LL41799-41802, one from the Loftusia-Beds, BM LL41815, Skelton Collec- tion; one specimen from Jebel Bu Milh, loose, but probably from the basal gastropod bed of the Simsima Formation at Jebel Bu Milh, BM LL41805. 1909 cf. 1909 DESCRIPTION. Right valve with secondary approach to bilat- eral symmetry. Dorsal margin convex-alate, with a central (dorsal) raised portion, which is itself centrally grooved in a small specimen. Has strong ‘lateral’ carinae which curve downwards, giving the appearance of a stretched bow when the smooth dorsal surface is viewed. Ventral margin of right valve with wide radial bands separated by an acute carina that forms a prominent downward ‘V’ at about the same level as the ‘lateral’ carinae. REMARKS. B. baylei was a lateral clinger, on its broadly expanded antero-dorsal face. The latter feature, and the anterior-ward leaning of the interband, suggest Toucas’ group of B. fissicostatus, of which B. baylei is the Maastrich- tian representative. A specimen in the Trechmann collection, identified by Chubb (1971) as Bournonia thiadensi Vermunt from the Maastrichtian of Jamaica has similar plications on the ventral face of the lower valve, but a wider and flatter central area on the reverse side and in that way resembles the next species. ?Biradiolites aff. baylei Toucas, 1909 Pl. 8, figs 3-5 MATERIAL. Six specimens from the central eastern face of Jebel Faiyah, loose on lower part of the Simsima Formation, BM LL41793-98, Skelton Collection; one specimen from Jebel Rumaylah, from the lower Dictyoptychus level, LL41814, Skelton Collection; two specimens from Jebel Figs 1,2 Biradiolites aff. baylei Toucas, Jebel Huwayyah, section 2, Loftusia-Beds, beds 3-8; x 1-5; 1, BM LL41800; 1a, dorsal view; 1b, view of left valve; le, ventral view; 2, BM LL41799; 2a, view of left valve; 2b, ventral view; 2c, dorsal view; 2d, basal view of right valve. Figs 3-5 __Biradiolites ?aff. baylei Toucas, Jebel Faiyah centre, loose from lower part of Simsima Formation, Skelton Collection; x 0-75; 3, BM LL41794, 3a, ventral view; 3b, basal view of right valve; 3c, dorsal view; 4, BM LL41796; 4a, view of left valve; 4b, ventral view; 4c, dorsal view; 5, BM LL41795; 5a, view of left valve; 5b, ventral view. LATE CAMPANIAN-MAASTRICHTIAN RUDISTS 295 296 Faiyah, section 1, in hard limestone, ?from the lower Sim- sima Formation, BM LL41803-04; two specimens from Jebel Buhays, section 3, lower Simsima Formation, BM LL41810-11; three specimens (two conjoined) from Jebel Aqabah, BM LL41812-13; one specimen from Jebel Raw- dah, section 2, bed 13, BM LL41806; one specimen from the same section, bed 23, BM LL41807; two others from the same section, loose, BM LL41808-09; one specimen from Jebel Rawdah, BM LL41816, loose, Skelton Collection; one specimen from Jebel Ja’Alan, southern Oman Mountains, west side, lower Simsima Formation, BM LL41817, Skelton Collection. COMMENTS. Similar to Biradiolites aff. baylei but with an additional carina between the dorsal radial band and the posterior carina. The radial bands are relatively narrower than in B. aff. baylei. A number of species of this complex have been described by Pamouktchiev (1967); Pons et al. (1992) in a study of specimens from Somalia suggested that the distinctions are unjustified. Genus GLABROBOURNONIA Mortis & Skelton gen. nov. TYPE SPECIES. Glabrobournonia arabica Morris & Skelton sp. nov. DIAGNOSIS. Small genus with a cornute lower valve and a very low, slightly convex upper valve. Right valve smooth except for fine growth lines and three major sinuses in the shell margin which leave sinusoidal traces, one at the ventral margin, one at the dorsal margin and one centrally on the posterior margin. Upper valve with reticulate sculpture of fine radiatig ribs and concentric growth laminae. REMARKS. At present we are aware of only one species. Glabrobournonia differs from Bournonia in being devoid of ribbing on the right valve. Glabrobournonia arabica Morris & Skelton sp. nov. Plo 9 igs 12 HOLoTyPE. BM LL41873 from Jebel Rawdah, section 1, the lower Simsima Formation, bed 2, just below low Upper Maastrichtian ammonites and inoceramids. PARATYPES. Jebel Rawdah, section 1, bed 2, LL41870-72; bed 3 and top bed 2, LL41874—95; loose, LL41869; section 4, bed 2, the basal rudist bed, LL41896-41916; Jebel Rawdah, southern flank, scree from Lower Simsima Formation, LL41818-22, Skelton Collection; Jebel Faiyah, section 2, lower Simsima Formation, LL41855—57; Jebel Thanais, lower Simsima Formation, LL41854; Jebel Buhays, section 1, LL41836-51, LL41917-—21; LL41825-27, Skelton Collection; PLATE 9 N.J. MORRIS AND P.W. SKELTON section 1, lower Simsima Formation, beds 4-10, LL41832-35; section 1b, loose, BM LL41852-S3. OTHER MATERIAL. Qarn Murrah, 50-60 m from bottom of section, LL41828 (16 specimens), LL41829-31, Skelton Col- lection; Jebel Huwayyah, section 1, beds 10-11, Loftusia- Beds, LL41858; section 2, beds 2-7, Loftusia-Beds, LL41859-68; LL41823-24, Skelton Collection. DIAGNOSIS. As for genus. DESCRIPTION. Mostly smooth horn-shaped lower valve with two bands representing a downward sinuosity where the cellular structure is exposed; one band is anterior, the other is ventral, and there is a further posterior sinuosity where growth increments are more obvious than the cellular struc- ture. Upper valve has exposed cellular structure radiating from excentric umbo, otherwise smooth, gently convex with a sinuous margin to fit lower valve. Subfamily SAUVAGESIINAE Douvillé, 1908 Genus DURANIA Douvillé, 1908 TYPE SPECIES. Hippurites cornupastoris Des Moulins, 1826, from the Turonian of France. REMARKS. The specimens of Durania from the Qahlah For- mation of Jebel Bu Milh are well-preserved, but even this material does not give us sufficient information concerning ecophenotypic variation. Different ‘morphs’ from different horizons are listed separately but we do not know whether they are different species. Durania cf. gaensis (Dacqué, 1903) Pl. 7, figs 14 cf. 1903 Radiolites ga’ensis Dacqué: 374, pl. 35, figs 7-9. cf. 1909 Sauvagesia gaensis (Dacqué); Toucas: 85, pl. 16 (17), figs 3-5. 21909 Sauvagesia flicki Toucas: 84, 85, pl. 16 (17), figs 6-8. cf. 1910 Durania gaensis (Dacqué); Douvillé: 50. LOCALITIES OF PREVIOUSLY FIGURED MATERIAL. Dacqué’s type material came from Ga’a near Abu Roash, Egypt, and was said by Douvillé (1910) to be Turonian in age; Toucas’ material occurs with Lapeirousia and was said to occur from Coniacian to Maastrichtian in Tunisia. MATERIAL. Three clumps from Jebel Bu Milh, section 1, from the upper part of the Qahlah Formation, LL41922-24; two further doubtful clumps from Jebel Faiyah, one loose from section 1, the second from section 1b, bed 2b, BM LL41925-26. DESCRIPTION. Rather large, outer shell layer of nght valve Figs 1,2 Glabrobournonia arabica Morris and Skelton gen. nov., sp. nov.; 1, Jebel Rawdah, section 2, Simsima Formation, bed 2, BM LL41873, holotype; la, ventral view; 1b, posterior view; 1c, dorsal view; 1d, anterior view; X 1-5; 2, south end of Jebel Buhays, lower part of Simsima Formation, BM LL41825, paratype, Skelton Collection; view of internal mould of left valve; x 1-5. Fig. 3 Osculigera cf. vautrinioides Vogel, probably from Qarn Murrah, BM LL22460, Iraq Petroleum Company Collection; polished section of right valve; x 1. Figs 4,5 Lapeirousia sp.; 4, Jebel bu Milh, section 2, Qahlah Formation, bed 5, the acteonellid gravels; BM LL41955, conjoined pair of right valves; 4a, ventral view; 4b, view of commissural surface; x 1; 5, Jebel Thanais, section 4, loose from lower part of Simsima Formation; BM LL41956, conjoined pair of individuals; view showing commissural surface with pseudo-pillars below internal mould of left valve; X 1. LATE CAMPANIAN-MAASTRICHTIAN RUDISTS 298 with thin-walled polygonal cells resembling those of Durania rather than Biradiolites. LL41922 includes three specimens with the upper valve in place, which is slightly concave and smooth in the centre but develops low rounded plicae towards the margins. LL41924 includes one specimen with the upper, left valve present but, in this case it is hemispheri- cally domed above the body-cavity of the right valve. We consider this to be a phenotypic variation, possibly related to the angle of the growth surface to the long axis of the right valve. Right valve steeply conical, ‘D’-shaped in section, with wide and smooth, somewhat indented radial bands on the straight part of the ‘D’. Radial bands separated by a large plication with two sub-plications at its crest. Rounded part of ‘D’ with approximately fourteen evenly distributed plicate ribs. Growing surface undulates with the plication of the outer shell sculpture. Durania cf. apula (Parona, 1900) Pl. 6, fig. 6 1900 Biradiolites apulus Parona: 21, pl. 3, figs 1-3.Q: 1909 Sauvagesia apulus Parona; Toucas: 97, fig. 65. MATERIAL. From Jebel Rawdah, section 2, bed 10, BM LL41951. DESCRIPTION. Medium sized species with approximately 40 narrow vertical ribs on lower valve. Wide growth surface of outer shell layer of right valve, which is not plicated. Ventral radial band relatively narrow and somewhat concave, with fine radial striations. Dorsal radial band narrow with fine riblets. Radial bands separated by a convex interband with a few N.J. MORRIS AND P.W. SKELTON sub-plicae. Growth surface of right valve has single narrow round holes approximately 2mm in diameter at the mid-point of its width near the position of the radial bands. Similar features have been found in Santonian hippuritids and are being described by Skelton & Vicens (in prep.), who regard them as the crypts of parasitic or commensal organisms. Pl. 6, fig. 1 MATERIAL. From Jebel Huwayyah, section 1, Qahlah gravel, BM LL41948; two specimens from Qarn Mulayh, BM LL41949-50, Skelton Collection. DESCRIPTION. Large form with approximately 70 ribs on the external surface of the right valve matched by undulations on the wide growth surface, and prominent down-twisted con- cave smooth dorsal radial band. Durania form A Fig. 6 MATERIAL. From Jebel Rawdah, section 1, bed 4, BM LL41952; section 2, bed 6, BM LL41953; three specimens, section 2, loose, BM LL41954. Durania form B DESCRIPTION. Small species with approximately 40 evenly spaced bifurcating ribs. Outer shell layer of right valve relatively thin with a folded growth surface, each upward fold in the position of the interspace between the ribs. REMARKS. The outer shell surface of the right valve and the growth form are similar to Durania cornupastoris as figured by Cobban, Skelton & Kennedy (1991: pls 1-3). Fig. 6 Durania form B; a group of three conjoined individuals with the remains of a fourth on the under surface; BM LL41954, x 0-75. PLATE 10 Fig. 1 commissural surface of right valve; x 1. Pseudosabinia aff. klinghardti (Boehm), Jebel bu Milh, section 2, bed 7-8, ? uppermost bed of Qahlah sands; BM LL41964, view of Figs 24 Colveraia aff. variabilis Klinghardt; 2, Qarn Mulayh, LL41958, Skelton Collection, view looking down on rim of right valve showing the internal structure of the left valve; x 1; 3, Jebel Thanais, loose from basal beds of Simsima Formation, BM LL41961, dorsal view of the two valves, x 1; 4, central Jebel Faiyah, loose from lower part of Simsima Formation, BM LL41960, Skelton Collection; 4a, view looking down on internal structure of left valve; 4b, ventral view of right valve; x 1. Fig. 5 Osculigera cf. vautriniodes Vogel, Qarn Murrah, BM LL41770, eroded section of right valve; x 1. Fig. 6 Lapeirousia sp., Jebel Buhays, loose from basal beds of Simsima Formation, BM LL41957; minute pseudo-pillars just visible; in the lower part of the figure the emplacement of the last shell layer is divided below the pseudopillars; x 2. 299 LATE CAMPANIAN-MAASTRICHTIAN RUDISTS 300 Fig. 7 Lapeirousia jouanneti Des Moulins, from the Upper Campanian, near Cognac, France, BM LL41976; a, b, upper and ventral view of a flat-based individual with pseudopillars marked, x 0-4; c, close-up of ventral pseudopillar showing its spout-like shape and division of the last layer of outer shell around it, x 1-5. Durania spp. Fragmentary or poorly preserved specimens that we can attribute to the genus Durania occur at most levels in the Qahlah and Simsima Formations: at Jebel Huwayyah, section 1, from bed 13, the basal Simsima conglomerate, the top rudist bed (two silicified specimens), and from near the top of the Simsima Formation approximately 6 m above the top oyster bed; at Jebel Rawdah, section 1, bed 2, and section 2, bed 16; and at Qarn Mulayh, a large thick-celled form with strangely shaped cells. PLATE 11 Fig 1-5 Pseudosabinia aff. klinghardti (Boehm); 1, Jebel Fayah, basal Durania-facies of the Simsima Formation, BM LL41971, Skelton Collection; eroded valve showing striate appearance of the outer part of the inner shell layer; x 0-4; 2, Virovi (between Lanac and Vestari, near Rtanj mine), eastern Serbia; BM LL41652, paratype of Sabinia rtanjica Pejovic; section through right valvewith aréte cardinale and ligament (arrowed); x 1-5; 3, Qarn Murrah, BM LL41972, Skelton Collection; close up of cellular structure of outer shell layer of right valve; x 5; 4, 5, Jebel bu Milh, section 2, Qahlah Formation; 4, BM LL41963, eroded commisural surface of a right valve; x 1; 5, BM LL41962, an eroded left valve; 5a, oblique dorsal view; 5b, posterior view; x 0-6. N.J. MORRIS AND P.W. SKELTON Subfamily LAPEIROUSIINAE Kuhn, 1932 Genus LAPEIROUSIA Bayle, 1878 TYPE SPECIES. Sphaerulites jouanneti Des Moulins, 1826, Upper Campanian or Lower Maastrichtian, Aquitaine, France. Lapeirousia sp. PI. 9, figs 4, 5; Pl 10; fig. MATERIAL. Two small joined specimens from Jebel Bu Milh, section 2, from the acteonellid bed towards the top of the Qahlah Formation, BM LL41955; three specimens, two of which are joined, from Jebel Thanais, from scree below basal part of the Simsima Formation, BM LL41956; one from Jebel Buhays, section 1, LL41957. LATE CAMPANIAN-MAASTRICHTIAN RUDISTS 302 DESCRIPTION. Small ovoid conical species, with the right valve approximately 75mm high and its marginal surface an ovoid approximately 37x45 mm. The structure of the pseudo- pillars of Lapeirousia jouanneti is shown in Fig. 7. Genus OSCULIGERA Kuhn, 1932 TYPE SPECIES. Osculigera cleggi Kthn, 1932, by original designation. SYNONYM. ?=Vautrinia Milovanovié 1938 (type species, Lapeirousia syriaca Vautrin, 1933, by original designation). REMARKS. Species differ in degree of complication of undu- lation of growth surface of outer shell layer. Osculigera cf. vautrinioides Vogel, 1970 Pip shige. 3: PIP 10; figs cf. 1970 Osculigera vautrinioides Vogel: 69, pl. 7, figs 3, 4, 6; pl. 8, fig. 3. MATERIAL. 21 fragmentary specimens from Qarn Murrah, BM LL41770-80; BM LL41781—90, Skelton Collection; also from the same locality BM LL41791 (a block with six conjoined juveniles), and BM LL41792 (one specimen and some fragments), Skelton Collection; also BM LL22460-63 (identified by A. Grubi¢ as O. magna Kiihn) labelled Trucial Coast, but almost certainly the same locality, Iraq Petroleum Co. Collection (RN 39/2,4,5.). DESCRIPTION. Right valve varies from a low inverse cone to a subcylinder. Surface with fine, low, even vertical ribs. Pseudopillars project into body-cavity in low but obvious gentle curves. Outer shell layer of right valve very thick, about equal to the diameter of the body-cavity. Shell margin approximately planar. Cells sub-polygonal and rather short, inner shell layer very thin. Approximately 24 radiating undu- lations, each topped by a row of radially elongate, radiating projections (secondary pseudopillars of Kthn) on the secreted marginal surface, which occasionally branch as they approach the outer shell surface. Up to about 12 projections, regularly spaced within each line, which are not visible in very small individuals. COMMENT. It is very difficult to see consistent differences between the species described by Kuhn and Vogel. The shell thickness and form of the radiating tuberculate undulations seem to be very similar to the specimen figured by Vogel as O. vautrinioides. Subfamily JOUFIINAE Karacabey-Oztemiir 1981 GENERA INCLUDED. Colveraia (?=Dechaseauxia) and others from Serbia and Romania, Joufia, Pseudosabinia gen. nov. and an undescribed genus for Radiolites albonensis Toucas, 1908. REMARKS. These are highly specialised radiolitids with pal- lial canals and well-developed aréte cardinale. They differ from the the subfamily Chiapasellinae Alencaster, which lack the ligamentary invagination. The aréte cardinale of one species of the family, Pseudosabinia rtanjica, has the split inner ligament preserved on its inner surface (PI. 11, fig. 2). They are fixed by the right valve. N.J. MORRIS AND P.W. SKELTON Genus COLVERAIA Klinghardt, 1921 TYPE SPECIES. Colveraia variabilis Klinghardt, 1921, by origi- nal designation. SYNONYMS. Branislavia Sladi¢-Trifunovicé, 1983a; ?Dech- aseauxia Tavani, 1949. Colveraia aff. variabilis Klinghardt, 1921 Pl. 10, figs 2-4 MATERIAL. One specimen from Qarn Mulayh, BM LL41958; one specimen from Qarn Murrah, BM LL41959; one specimen from Jebel Faiyah, north-west face, low in the Simsima Formation, low in the Dictyoptychus-facies, BM LL41960, all Skelton Collection; one specimen from Jebel Thanais, loose from basal Simsima Formation, BM LL41961. DESCRIPTION. Moderately thick calcitic outer shell layer of the right valve with a narrow elongate aréte cardinale. Canals obvious in the inner shell layer of both valves. In the right valve a single row of large, subquadrate canals separate the sockets and myophores from the outer shell layer. In the left valve the canals are somewhat narrower and radially elongate and penetrate the teeth and myophores. In one specimen (PI. 10, fig. 4a) the outer surfaces of both the teeth and the myophores are longitudinally ridged, with the ridges appar- ently formed by the canal walls, which interdigitate with ridges on the muscle attachment surface of the opposing valve. Genus PSEUDOSABINIA Morris & Skelton gen. nov. TYPE SPECIES. Pseudosabinia klinghardti (Boehm, 1927). INCLUDED SPECIES. Sabinia klinghardti Boehm, 1927, Sab- inia serbica Kihn & Pejovi¢, 1959, and ?Sabinia rtanjica Pejovic, 1967. DIAGNOSIS. Both valves extended, conical, gently curved. Outer shell layer finely cellulo-prismatic to compact in the right valve, thinner and smooth in the left valve. Well- developed aréte cardinale projecting between two close, well-developed teeth, flanked by radiolitiform myophores in left valve. Inner shell canaliculate throughout in both valves. REMARKS. Originally assigned to Sabinia, the species of this genus are recognized by the finely polygonal cellulo-prismatic structure of the outer calcitic layer of the right valve, and for their radiolitiform myocardinal apparatus. In P. serbica the cells appear to be present in some areas of the right valve outer layer (Kiihn & Pejovic , 1959: figs 7, 8), whereas they occur throughout the outer shell layer of the right valve of S. klinghardti. The thickness and surface sculpture of coarse, zig-zag growth rugae of this outer shell layer are similar in S. serbica and §. klinghardti. In the interpretation of Philip (1986) all species of Sabinia are considered to be related and are off-shoots of the Radiolitidae that have secondarily reduced the outer calcitic shell layer of the right valve. We consider that there may be at least two distinct genera within the genus Sabinia as presently constituted, and that these may be quite unrelated. The features of the true Sabinia with its type species, Sabinia anienis Parona 1908, may relate it to the Plagioptychidae. NOTE ON THE TYPE SPECIES OF SABINIA. Parona described three species: in sequence they were S. sublacensis, S. sinuata LATE CAMPANIAN-MAASTRICHTIAN RUDISTS and S. anienis from the Pietra di Subiaco, Late Cretaceous, of Monte Affilano, Valle dell’Aniene, Provincia di Roma (the type specimens are in Rome). All three nominal species may be merely varieties. In Parona’s material a septum cuts off the posterior accessory cavity from the body-cavity in the left valve (Parona, 1908: figs a, c). It runs between the anterior tooth and the postero-ventral margin, and could be equiva- lent to the septum in the left valve of the plagioptychids. Parona does not describe the structure of the outer shell layer of the right valve in any detail. Pseudosabinia aff. klinghardti (Boehm, 1927) Pl. 10, fig. 1; Pl. 11, figs 1-5 Sabinia klinghardti Boehm: 205, pl. 15, figs 1, 2; pl. 16, fig. 1. aff. 1927 ?aff. 1927 Schiosia bilinguis Boehm: 207, pl. 18, figs la—Ic. aff. 1967 Pseudosabinia rtanjica Pejovic : 295-97, pl. 1, fig. 1. aff. 1986 Sabinia rtanjica tunisiensis Philip: 248, 49, pl. 1, figs 1-6. Types. The holotype, BM L49455 from the Campanian- Maastrichtian, east of Hereke and west of Tauchanly, Bythinia, north-west Turkey, has both valves preserved; the paratype, BM 49454 from the same locality, is a much larger specimen of a crushed right valve. The holotype of Schiosia bilinguis, BM L50929, is from the ‘Upper Senonian’ between Herake and Tauchanly, Bithynia. BM LL41652 is an unfig- ured paratype of P. rtanjica Pejovic, from near Rtanj mine, eastern Serbia. NEW MATERIAL. From Qarn Murrah, a large crushed right valve with most of the outer shell layer missing except for a thin layer remaining, together with a second crushed valve which may be left or right, BM LL41972, Skelton Collection; four specimens from Jebel Bu Milh, section 2, Qahlah Sands, BM LL41962-65; two possible specimens from Jebel Raw- dah, section 4, bed 1, the basal rudist bed, BM LL41966-67, and another small left valve that shows well-developed pallial canals and a narrow aréte cardinale, BM LL41968; a large left valve from the north-west face of Jebel Faiyah, Simsima Formation, the Durania-facies, BM LL41971, and a small left valve from central Jebel Faiyah, BM LL41981, both Skelton Collection. DESCRIPTION. The material from Qarn Murrah is badly crushed but does show the cellular outer shell layer (PI. 11, fig. 3). The large, horn-like right valve (PI. 11, fig. 4), at least 16 cm long, has a long narrow aréte cardinale, a thick inner shell consisting of irregularly polygonal pallial canals, fine at the valve margin, becoming coarser inwards, and prominent cystose tabulae. Accessory cavities are apparently absent. There are two sockets for prominent prong-like teeth with longitudinal grooves and ridges either side of a low, ridge-like tooth, immediately ventral to the aréte cardinale. The myo- cardinal arrangement is not capriniform. The small right valve of Pl. 10, fig. 1 shows the aréte cardinale straddled by the ridged sockets for the teeth. Two equally prominent teeth are present in the fixed left valve of Pl. 11, fig, 5. REMARKS. The progression from small to larger pallial canals in the left valve of S. bilinguis Boehm is similar to the pattern in the present species, otherwise Boehm’s species has very few characters preserved. The outer shell layer of the right valve of Pseudosabina rjtanjica is thin (less than 1 mm, fide 303 Philip) and of dense fibrous calcite, ie not cellular as in P. klinghardti. This seems to be the only difference, but is confirmed by our material. Pejovi¢ (1967) and Philip (1986) suggest that a row of canals separating the myophores from the body-cavity distinguishes P. rtanjica from P. klinghardti. Careful inspection of Boehm’s holotype leads us to the conclusion that this difference does not exist. The myophores actually have a purely radiolitiform arrangement, albeit with contained canals, and face outwards opposing the inner surface of the right valve. Pejovic’s (1967) description of Sabinia rtanjica was based only on eight specimens of the upper, left valve, in which the myophores were stated to be separated from the living chamber by a number of pseudocanals, characterized by regularly spaced tabulae and not considered homologous with the canals of the Caprinidae. A well- preserved small left valve collected by one of us (PWS) from the ?Lower Campa- nian of Monte Kamilja, near Leposavi¢ in south-west Serbia, confirms the description of P. rtanjica in having large canals on inner sides of the myophores. However, this feature is in fact the coarse canaliculate structure of the myophoric apo- physes and is also present in the holotype of Pseudosabinia klinghardti. Philip’s (1986) Tunisian form differs from Pejov- ic’s original description in the length of the aréte cardinale which reaches well between the teeth, unlike that in the Serbian original which, it is claimed, only reaches the top of the teeth/sockets. Inspection of Pejovié ‘s 1967, pl. 1, fig. 1 suggests that this is incorrect. Secondly Philip stated that the pallial canals of the lower (right) valve are of oval section in the Tunisian form, but oval and polygonal in the nominal subspecies. He also claimed that the upper valve canals have a suboval section in the Tunisian form, but these are pentago- nal, hexagonal or triangular in the nominal subspecies. We suspect that these latter two distinctions reflect only the diagenetic growth of fibrous cement crust within the canals of the Tunisian material. Pseudosabinia rtanjica has a very thin, compact outer shell layer in the right valve, according to Philip (1986), which is in contrast to the much thicker outer shell layer of Pseudosab- inia klinghardti, although there are dense areas in the outer part of this layer in Boehm’s holotype. This seems to be supported by our rather poorly preserved material, although a specimen with part of both valves preserved, apparently of P. rtanjica, from Serbia, kindly donated from Mdm. Pejovic, shows a thin zone of cells on the innermost zone of the outer shell layer grading into dense structure in the outer part. This suggests the possibility of intergradation between the two taxa. Inspection of further material would be required to confirm this. ACKNOWLEDGEMENTS. We would like to thank Diana Clements for considerable help in both sorting the material and preparing the manuscript. We are indebted to Phil Crabb for taking the photo- graphs, and to Tony Wighton and his colleagues for cutting the sections of the rudists. REFERENCES Boehm, J. 1927. Beitrag zur Kenntniss der Senonfauna der bithynischen Halbinsel. Palaeontographica 69: 187-222, pls 11-18. 304 Chubb, L. J. 1971. Rudists of Jamaica. Palaeontographica Americana, 7 (45): 161-257, pls 27-58. Cobban, W. A., Skelton, P. W. & Kennedy, W. J. 1991. Occurrence of the rudistid Durania cornupastoris (Des Moulins, 1826) in the Upper Cretaceous Greenhorn Limestone in Colorado. Bulletin of the United States Geological Survey, Shorter Contributions to Paleontology and Stratigraphy, 1985 (D): DI-D8, 3 pls. Cox, L. R. 1933. The evolutionary history of the rudists. Proceedings of the Geologists Association, 44: 379-88. Dacqué, E. 1903. Mittheilungen uber den Kreidecomplex von Abu Roash bei Cairo. Palaeontographica, 30 (2) (4): 337-92, pls 34-36. Defrance, J. L. M. 1821. Dictionaires des sciences naturelles, 18-22. Paris. Des Moulins, C. 1826. Essais sur les sphérulites qui existent dans les collections de MM. F. Jouannet et C. Des Moulins, et considerations sur la famille a laquelle ces fossiles appartiennent. Bulletin de l’Histoire Naturelle Société Linnéenne de Bourdeaux, 5 (1): 141-43. Douvillé, H. 1888. Etudes sur les Caprines. Bulletin de la Société géologique de France (3) 6, 699-730, pls. 22-25. 1890-94. Etudes sur les rudistes. Révision des principales espéces d’Hippurites. Mémoire no. 6; Mémoires de la Société Géologique de France; Paléontologie, 1 (1890): 1-32, pls 1-3 (15-17); 2 (4) (1892): 33-56, pls 4-7 (18-21); 3 (4) (1893): 57-94, pls 8-15 (19-26); 4 (2) (1894): 95-138, pls 16-20 (6-10). — 1895-97. Etudes sur les rudistes. Distribution régionale des Hippurites. Mémoire no. 6; Mémoires de la Société Géologique de France; Paléontologie, 5 (1895): 139-188, pls 21-28 (8-15); 7 (3) (1897): 189-236, pls 29-34 (13-18). 1902. Classification des radiolites. Bulletin de la Société Géologique de France, (serie 4) 11: 461-482, pl. 15, figs 1-8. 1904. Etudes Géologiques ; part 4, Paléontologie. Mollusques fossiles in: de Morgan, J. Mission Scientifique en Perse, 3: 189-380, pls 25-50. Leroux, Paris. — 1905. Observations, in J. de Morgan. Note sur la géologie de la Perse et sur les travaux paléontologiques de H. Douvillé sur cette region. Bulletin de la Société géologique de France, (4) 5: 170-189. — 1910. Etudes sur les rudistes. Rudistes de Sicile, D’Algérie, d’Egypte, du Liban et de la Perse. Mémoires de la Société Géologique de France, 41: 1-84, pls 1-7. — 1927. Nouveaux rudistes du Crétacé de Cuba. Bulletin de la Société Géologique de France, (4) 27: 49-56, pl. 4. Gili, E. & Skelton, P. W. 1994. Classificacio paleoecologica de les formes dels rudistes — una eina per a l’analisi paleoambiental. Butlleti de la Instucio Catalana d’ Historia Natural, 61 (Sistemes I Processos): 97-116. Gray, J. E. 1848. On the arrangement of the Brachiopoda. Annals and Magazine of Natural History, (2) 2 (12): 435-40. Grubic, A. 1980. Torreites milovanovici sp. nov. iz Omana. T. coxi sp. nov it T. chubbi sp. nov sa Jamajke, nov prikaz roda Torreites Palmer i osvrt na znacaj njegovog paleogeografskog rasprostranjenja. Vestnik zavod za geoloska i geofizicka Istrazivanja, A 37: 81-99, pls 1, 2. Karacabey-Oztemiir, N. 1981. Three new species of the genus Miseia and proposal of a new subfamily of Radiolitidae. Bulletin of the Mineral Research and Exploration Institute of Turkey, 92: 40—46, pls 1-2. Klinghardt, F. 1921. Die Rudisten. Teil 1: Neue Rudistenfauna aus dem Maastrichtien von Maniago (Friaul) nebst stratigraphischem Anhang. Archiv fiir Biontologie, 5 (1; 1): 68pp., 13 figs, atlas. Kiihn, O. 1932. Rudistae from Eastern Persia. Records of the Geological Survey of India, 66 (1): 151-179. & Pejovic , D. 1959. Zwei neue Rudisten aus Westserbien. Sitzungsberich- ten der Osterreichischen Akademie der Wissenschaften, Wien, Mathematisch- Naturwissenschaftliche Klasse, (1) 168 (10): 979-989, pls 1-4. Kutassy, A. 1934. Pachyodonta mesozoica. (Rudistis exclusis.). Fossilium Catalogus, 1, Animalia, 68: 202pp. W. Junk, The Hague. Lamarck, J. B. de 1801. Systeme des animaux sans vertébrés. viii + 432 pp. Paris. MacGillavry, H. J. 1937. Geology of the Province of Camaguey, Cuba with revisional studies in rudist paleontology (mainly based upon collections from Cuba). Physiographisch-Geologische Reeks der Geographische en Geolo- gische Mededeelingen, Amsterdam, 14: 168pp., 10 pls. Matheron, P. P. E. 1842-43. Catalogue methodique des fossiles des Bouches du Rhone. 8vo, Marseilles. — 1880. 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Mémoire no. 36; Mémoires de la Société Géologique de France; Paléontolo- gie, 14 (4) (1907): 1-46, pls 1-8 (11-18); 16 (1) (1908): 47-78, pls 9-15 (1-7); 17 (1) (1909): 79-132, pls 16-24 (1-9). Vautrin, H. 1933. Sur quelques formes nouvelles de rudistes recueillis en Syrie Septentrionale. Notes et Memoires de la Haute Commissariat de la Syrie et du Liban, Republique Frangaise, 1: 29-43, pls 1-S. Vogel, K. 1970. Die Radioliten-Gattung Osculigera Kiihn (hohere Oberkreide) und die Funktion kennzeichnender morphologischer Eigenschaften der Rudisten. Paldontologische Zeitschrift, Stuttgart, 44 (1-2): 63-81, pls 6-8, 6 figs. Woodward, S. P. 1855. On the structure and affinities of the Hippuritidae. Quarterly Journal of the Geological Society, 11: 40-61, pls 3-S. Zittel, K. A. 1865-66. Die Bivalven der Gosaugebilde in den nordostichen 305 Alpen Beitrag zur Characteristik der Kreideformation in Osterreich. Denk- schriften der Kaiserlichen Akademie der Wissenschaften, Mathematisch- Wissenschaftliche Klasse, Wien, 24 (2) & 25 (2), 293pp. 27 pls. — 1881-85. Handbuch der Palaeontologie 2. Mollusca und Arthropoda. 900pp., 719 figs. Munchen und Leipzig. Note added in proof: One of us (PWS) has recently had the Opportunity to see some specimens of Durania mutabilis (Stoliczka, 1871) collected from the Maastrichtian of south- ern India by Professor Malcolm Hart. It is clear that the material described above (p. 296) as Durania cf. gaensis (Dacqué, 1903) can be attributed to D. mutabilis. Whether D. gaensis and D. mutabilis (which has priority) should be considered to be synonyms remains to be resolved. vila voit icine r* « “ i ] ols wha de, wai hp(hy kCahE nit Lhe | teat oak eels tole a Fs (FA 4p ORAK. Gee agi Wa ehrk A | iy etd yh sp Re ty s ’ ae i 4 1A %e@— AP Mla Laon bs Spalgiun Wre air oie (a Hay «ae i ie 7 my Cer ' al wry ~~ rt ‘Ta tale ; : at - hy ’ sneha) wale Se ee Comea)oh prt, disdald « L lit ive 2S) Wr ypohitgron «lapeiieee (oe ween bs Vandre 1.01 i ‘t , ee oe oe inoempel tt 4 ts nt ¥ , ; y dx) en Ath your iwi. y ; 7 ik == wo Pui oh al er Sve esiey ty oe ale \ 2 : : 7 ere, it f ey tie Me PPO e ‘ an t oper piety alipsieyn mele’ oy ake step iegepaienl sk yobs Dpto: ' r ‘n : ; ; 4 say ce SOE! Hy ae : : : é , F wy teased sie asain T fi ‘a ca peal = aan A Fd) ie pies: f i 4 ‘ ; i = 7 rr, vee byedoe 1 Toe . sci Ma : > ‘ee DL & weeteey PO |e Gee Be hie i 3 fi a ; , aa mig ew -96an s ane ot 8 Bes 7 part bet ; na ; : ee gal . mn a ae -— ca oF al e. Sie) a 2 Cie 1 i =: ee - legmemetiht 4 = me iad Aye i a ie > z , y +m) a a) Patera ear! R y : | seta wae . n , oe = oles ee i vi ey wal ths eal p Da ‘ oe, ay ten" Tae Te oe 7 a Ja? a ee ee ee ee. | Sint ys Sie et A Ol peg Paha l a) eS an & a oe eS led "ee iam eis f il “a. f eb ' *é,, a or 1 We , i “eile 4 i « = tliat 3s" sip ; ‘ Me = re nerd Actas + ¥ ate = fe s ae ‘ i ; 3 tie 7 gen Ph tae He A 9. wh! Pay) = s me ea iy i, nti fk Merc yg Mae = s, ii % ‘waaay are (tes arua eat iy yi , \ ! [So Te, ; ' \ a Se ee ey i } P ; awd Smid ¥L ow ; 7 — . eeeed ve mr tamed pat a ~ttemys ie OLS Tees ee ‘ : ¢. hem me ier wp RS oe fb Moench Mintle ages Tie & a4 - 6 eal, ny elt ee i = ee, ee) heaonaee — : a , ' Phin: Mab WH SS ee m 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 32 No. 1 No. 2 No. 3 No. 4 Volume 33 No. 1 No. 2 Miscellanea Observations on Cycloclypeus (Cycloclypeus) Carpenter and Cycloclypeus (Katacycloclypeus) Tan (Foraminiferida). C.G. Adams & P. Fame. 23 figs. The provenance of Sivapithecus africanus. P.J. Andrews & T.I. Molleson. 3 tables. A silicified brachiopod fauna from the Silurian of Iran. L.R.M. Cocks. 41 figs. Two new condylarths (Mammalia) from the early Eocene of southern England. J.J. Hooker. 15 figs, 3 tables. Miocene sharks’ teeth from Ecuador. A.E. Longbottom. 26 figs, 3 tables. A new fossil terrestrial isopod with implications for the East African Miocene land form. S.F. Morris. 12 figs. A re-evaluation of the fossil human calvaria from Singa, Sudan. C.B. Stringer. 2 figs, 1 table. New species of Protorthoptera and Protodonata (Insecta) from the Upper Carboniferous of Britain, with a comment on the origin of wings. P.E.S. Whalley. 5 figs. 1979. Pp. 1-90. £10.50 Palaeoenvironments and correlations of the Carboniferous rocks in west Fermanagh, Ireland. C.H.C. Brunton & T.R. Mason. 1979. Pp. 91-108, 6 figs, folded map. £4.00 The Ordovician trilobite faunas of the Builth-Llandrindod Inlier, central Wales. Part III. C.P. Hughes. 1979. Pp. 109-181, 177 figs. £10.00 The stratigraphy and brachiopods of the upper part of the type Caradoc of south Salop. J.M. Hurst. 1979. Pp. 183-304, 557 figs. £18.50 An account of the Ordovician rocks of the Shelve Inlier in west Salop and part of north Powys. W.F. Whittard, F.R.s. (Compiled by W.T. Dean). 1979. Pp. 1-69, 38 figs, frontispiece, coloured map, folded, in pocket. £10.00 Map available separately £1.00 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. 1 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. Pp. 165-235, 105 figs. £9.00 No. 4 Fossil insects from the Bembridge Marls, Palaeogene of the Isle of Wight, southern England. E.A. Jarzembowski. 1980. Pp. 237-293, 77 figs. £7.50 No. 5 The Yorkshire Jurassic fern Phlebopteris braunti (Goeppert) and its reference to Matonia R.Br. T.M. Harris. 1980. Pp. 295-311, 20 figs. £2.75 Volume 34 No. 1 Relative dating of the fossil hominids of Europe. K.P. Oakley. 1980. Pp. 1-63, 6 figs, 17 tables. £8.00 No. 2 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 No. 3 Ashgill Brachiopoda from the Glyn Ceiriog District, / north Wales. N. Hiller. 1980. Pp. 109-216, 408 figs. £14.75 No. 4 Miscellanea Type specimens of some Upper Palaeozoic Athyridide brachiopods. C.H.C. Brunton. 31 figs. Two new British Cretaceous Epitoniidae (Gastropoda): evidence for ev olution of shell morphology. R.J. Cleevely. 14 figs, 1 table. 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.P. Owen. 13 figs. Tupus diluculum sp. nov. (Protodonata), a giant dragonfly from the Upper Carboniferous of Britain. P.E.S. Whalley. 1 fig. Revision of Plummerita Bronniman (Foraminiferida) and anew 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. Pp. 79-107, 8 figs, 3 tables. £4.20 No. 3 Centenary miscellanea Reassessment of the Ordovician brachiopods from the Budleigh Salterton Pebble Bed, Devon. L.R.M. Cocks & M.G. Lockley. 35 figs. Felix Oswald’s Turkish Algae. G.F. Elliott. 3 figs. J.A. Moy-Thomas and his association with the British Museum (Natural History). P.L. Forey & B.G. Gardiner. 3 figs. Burials, bodies and beheadings in Romano-British and Anglo-Saxon cemeteries. M. Harman, T.I. Molleson & J.L. Price. 5 figs, 7 tables, VI appendices. The Jurassic irregular echinoid Nucleolites clunicularis (Smith). D.N. Lewis & H.G. Owen. 4 figs. Phanerotinus cristatus (Phillips) and the nature of euomphalacean gastropods. N.J. Morris & R.J. Cleevely. 12 figs. Agassiz, Darwin, Huxley, and the fossil record of teleost fishes. C. Patterson. | fig. The Neanderthal problem and the prospects for direct dating of Neanderthal remains. C.B. Stringer & R. Burleigh. 2 figs, 1 table. 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. Ensom. 23 figs. A possible non-calcified dasycladalean alga from the Carboniferous of England. G.F. Elliott. 1 fig. Nanyjinoporella, a new Permian dasyclad (calcareous alga) from Nanjing, China. X. Mu & G.F. Elliott. 6 figs, 1 table. Toarcian bryozoans from Belchite in north-east Spain. P.D. Taylor & L. Sequeiros. 10 figs, 2 tables. Additional fossil plants from the Drybrook Sandstone, Forest of Dean, Gloucestershire. B.A. Thomas & H.M. Purdy. 14 figs, 1 table. Bintoniella brodiei Handlirsch (Orthoptera) from the Lower Lias of the English Channel, with a review of British bintoniellid fossils. P.E.S. Whalley. 7 figs. Uraloporella Korde from the Lower Carboniferous of South Wales. V.P. Wright. 3 figs. 1982. Pp. 43-155. £19.80 No. 3 The Ordovician Graptolites of Spitsbergen. R.A. Cooper & R.A. Fortey. 1982. Pp. 157-302, 6 plates, 83 figs, 2 tables. £20.50 No. 4 Campanian and Mastrichtian sphenodiscid ammonites from southern Nigeria. P.M.P. Zaborski. 1982. 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. noy., 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. noy. 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. 1 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. 1 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. (IIT) 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 5. £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. 1 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. 1 The Ordovician graptolites of the Shelve District, Shropshire. I. Strachan. 1986. Pp. 1-58. 38 figs. 0 565 07010 X. £9.00 The Cretaceous echinoid Boletechinus, with notes on the phylogeny of the Glyphocyphidae and Temnopleuridae. D.N. Lewis. 1986. Pp. 59-90. 11 figs. 7 tables. 0 565 07011 8. 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 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. £19.00 No. 2 £5.60 No. 4 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. No. 5 £13.00 Volume 41 No. 1 The Downtonian ostracoderm Sclerodus Agassiz (Osteostraci: Tremataspididae), P.L. Forey. 1987. 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 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 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 No. 2 No. 3 No. 4 Volume 42 No. 1 Cenomanian and Lower Turonian Echinoderms from Wilmington, south-east Devon. A.B. SMith, C.R.C. Paul, A.S. Gale & $.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. Rickards. 1988. 394 pp., figs. 0 565 07020 7. £70.00 Volume 44 No. 1 Miscellanea: Palaeocene wood from Mali—Chapelcorner fish bed—Hererotheca coprolites—Mesozoic Neuroptera and Raphidioptera. 1988. Pp. 1-63. 0 565 07021 5. £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 1. £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 No. 4 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 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. 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 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. 1 ‘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. 2 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 Volume 50 No. 1 Systematics of the melicerititid cyclostome bryozoans; introduction and the genera Elea, Semielea and Reptomutltelea. 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. Pp. I-88. 89 90 91 121 241 251 257 267 275 277 Bulletin of The Natural History Museum CONTENTS Palaeontology of the Qahlah and Simsima_ Formations (Cretaceous, Late Campanian-Maastrichtian) of the United Arab Emirates-Oman Border Region Preface M.K. Howarth Late Cretaceous carbonate platform faunas of the United Arab Emirates-Oman border region A.B. Smith, N.J. Morris and A.S. Gale Late Campanian-Maastrichtian echinoids from the United Arab Emirates-Oman border region A.B. Smith Maastrichtian ammonites from the United Arab Emirates-Oman border region W.J. Kennedy Maastrichtian nautiloids from the United Arab Emirates-Oman border region N.J. Morris Maastrichtian Inoceramidae from the United Arab Emirates-Oman border region N.J. Morris Late Campanian-Maastrichtian Bryozoa from the United Arab Emirates-Oman border region P.D. Taylor Maastrichtian brachiopods from the United Arab Emirates-Oman border region E.F, Owen Late Campanian-Maastrichtian rudists from the United Arab Emirates-Oman border region N.J. Morris and P.W. Skelton GEOLOGY SERIES Vol. 51, No. 2, November 1995 C 1 a ‘ 1 _ es 4 We v oy! 1, et ty tae ‘on or | ’ | « a iy i 1s sos. aay vay: Ih raraeay Met le ircen ler mH rh Bye ace = pea rors — tie tt i i — Sree een See WH ee Saar asores gi