Cox 1988 English Callovian Perisphinctid Ammonites

P f R I O D i Q U E S 6 5 9 0

M O N O G R A P H OJK T H E P A I . A F O N T O G R A P I I I C A L S O C I E T Y

ENGLISH CALLOVIAm ^ fe. (MIDDLE JURASSIC)

PERISPHINCTID AMMONITES

B. M . C O X

P A R T 1 Pages 1-54: Plaies 1 - 2 3

© T H E P A L A E O N T O G R A P H I C A L S O C I E T Y - L U . N D O . N February 1 m

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Publication No. 575. issued as pare of Volume: HI) for 1986

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ENGLISH CALLOVIAN (MIDDLE JURASSIC) PERISFHINCTID AMMOM I ES

INTRODUCTION AND ACKNOWLEDOEMENTS

In England. theCallowan Stagenfrhe Middle Jurassic is represented by marine arenaceous to argillaceous clastic. and carbonate sediments. lr contains a clivara* ammonite fauna compr ising die Boreal Kosmoccratidae Ktppirri!*A, SignU>u?a$. K&snwcaas) and (iardiocrraliriae (Cadoctras* PscudoicuhiCTos* LftyigWincfirtut r t e o a w / w , ikteMitdtoww). and the Terhyati MacrocepludUidae {Mtzcmxphniitis). Pachyceratidae ; Erynwotwa*. P/iikyaru.*]. Oppehidao (/Vtf.VWw. W«:/irw>mv, Disiiihoaras). Reiner kcii<h-.i: Aspidocera tidar. {PôAxtpcii^yas. Pf\loc<r<ti. EuQUftUloc.txas) and Perisphinctidac (described herein). Alrhough r.hc Boreal families dominate and arc used a^ the principal biostraiigraphiial indices, ai some horizons ilie- Teihyan families constirurr a significant pari of the ammonite assemblage. Tint Perisphinctidac include forms assigned by Arkcll (195 7 si} to the PseudupcrHphincrinac. Perisphinctinac and Pruplanulitinar. In the English Callovian* these ammonites are common only at particular hari/.uns which arc discussed brlow.

This monograph is based on a Ph.D. ihcîs prepared under the scheme of co-operation between the University ofLoudon and government or public research inxiiiuuujis. The research was directed by Dr P. F. Rawson (then of Department nl'Gcology, Queen Mary College;. Official training aid was> received from the Hrirish Geological Survey (Natural F.nvironment Research Council). The constructive criticism of Professor ). H. (Jallomon (University College London) and Dr J . C. YV. Cope ! University College of Swansea) is acknowledged.

Dis P F. Rawson, S. R. A. Kelly ;L niver^jiy uf Cambridge) and ProfcssorJ. H. Cat lorn on donated material (now at BGS). Mr P. Hn.s<irn and Dr J . K. Wright supplied inlhruiatioii on collections in the Dorset (.nun.) MuKCum, Dorchester, and on specimens from the Yorkshire Gallovian respectively. Photographs, casts or information on specimens in foreign collections were supplied by Dr II. Lobhzcr and Dr F. Stojaspal :GBV:. Dr D Pajaud (LPP;u Dr M. Sastrv (GS1), Dr G. Sdiairer :BSM)> Professor Dr J . YVcndt (GPI) and Di R. Werner (SM F;. Mi C. Wood helped wkh German translations. Photography was carried out by the Photographic Section, BGS. This monograph is published with the approval of the Director, British Geological Survr.v (N.K.R.C.i.

RLPOSI I ORIES

BGS: British Geological Survey, Koywonh, Nottingham; BM: British Museum (Natural History;, London; BSM: Baycrischc Staatssanmiluii^ fur Palaontologie usul hisiorische Geologic, Munich: CNIGR: C M G R Museum. VSKCKL Leningrad: DM: Devizes Museum, Wiltshire; DCM: Dorset County Museum. Dure Lester; ETII : Eidgenossisc he I'cchnischc Ilochsc h u K Zurich: FSL: Department des Sciences de la I K:T<\ Faodte des Scicnccs de Lyon; FW: Private collection of Mr Felix YVhitham. Hull; GBV: Geologiscrhe BundesanÛll, Vienna; GPI: Geolopischpalaontologisrhrs lmtitut, Tubingen: GSI: Geological Survey of India. Calcutta; H U : Hull University: JKWs Private «>IUi:tiori of Dr. J. K. Wright, Chelsea College. Ixmdon; LIS: Services Gcologioo* de Portugal, Lisbon; LPP: Laboialoire de Paleontologie des Invern:bn:s% (Jiiivrrsiu* Pierce et Marie Curie, Pans: MM: Manchester Museum; M N H N P : Museum National d'l l istoirr NsitiirHle* Paris, NMW: National Museum uf Wales, Cardiff; OUM: University Museum, Oxford; PM: Pavlov Museum, MCRI, Moscow: SM: Sedgwirk Museum. Cambridge; SMF: For*cJiungsinstiuK Scnckcnbire, Krankfur.-am-Majn: WMS: YVoodcnd Museum. Scarborough; YM: The Yorkshire Museum, York.

ENCI.ISH CALI.OVLW PERISPHINCHI) AMMOXITKS

HISTORY Ob RESEARCH Although most oJ the English (Jallovian ammonite fauna is well known ami has been used as

tin: ha-us ol detailed biostratigraphical arc! rhioriostratigrapliical schemes* litcle systematic work has been published on the Pcrisplunctidac, which have liule primary slraligraphiral value in the Knglish sequence. In contrast, detailed descriptions of thi: abundant perisphirictids from the Bathouiari and the Coralliau Beds (Oxfordiari) were included in Arkelfs (1935-18; 19ol-58) monographs

The first monographic treatment of the Perisphinetidae was by Sieiriir.'.dzki (1008) who placed Callovian forms in G'wsouvrio. Pa-uphiacUs, Prwerito and Cfwffetw. Together with "jBiplica" and AtQ*ioccr<K, the classification of these perisphinctid groups was based on morpho-logical features such as size, ribbing, suture line and tKtfly-chamber, as well as on stratigra pineal distribution. Buckman (1909-30) in his Type Anwwmtes erected ten additional Callovian peri sphinctid genera; Kz)ikdih<wr<ti> ObtosicosliUs% PocvPufihinctts. Biw/JUpkmtss. GMSSipfantik'U*. Hamulisph incici, naphwjiilf <. HoMMQpiamtfues* Priuisphsnctes and AidgaliceMi, the last eight of which were based on Knglish material. However, his text was disjointed, with the classification di*-petted throughout the volume?, and not all groups were discussed in the same detail. The classification was based on the alleged irreversible phyletic. appearancc of new morphological characters: ;M: is by analyses of characters in this maimer that the descent and biological position of families and genera are to lie worked out -not by rash assumptions of affinity from general similarity, which* too often, may be merely homoeomorphic deceptions" (Buckman 1924, p. 9).

Spaih (1924; 1927-3 jj. erected further Callo\iau pcrisphincud yeucra from Gulch, India: Sub£WSS0Wri<iy Sv.ajuYic*. IhibtritceTus. Judos fifani'lts. ('uUhisphiucis.i* Orionoidw, and dismissed many ol Buck man's conclusions which he considered to he allectcd by too naYvr. au adherence ID the "law of earlier inheriT/iiice" JSpath 1931a, p. 2H0). Spath's genera were intentionally defined it; a somewhat 'clastic* manner, anil be tried co evolve a more practical classification of the peri-sphinctids* based on a worldwide study of thousands of specimens. He wrote (1931a, ft. 39?) "I am not satisfied with the classification her< adopted; and it is commonly admitted chat the Perisphinctids are perhaps the mosr dillicult group of Ammonites to divide* largely on ;irc ount of t he diversity of local faunas: but the presnr. schcmc at least embodies good deal of practical knowledge obtained since I used HicmiradzkPs classificanon in 1913 or tried my best to adopt Buck man's two families ten years later". Aikcli (1958. p. 163) commended S p a t h \ work, which he believed to represent the only atrempt at an orderly classification char took account of the Rule* of Nomenclature and showed any grasp of the cmiltiplicicy of forms of perisphinctids, and their scracigraphic«*l i mplicacions.

Arkell (1931 W ; used more or less arbitrary subdivisions in his classification of the Peri sphinciidac. "largely determined by systematic convenience so tar as is compatible with priority of nomenclature" !p. IfiS). The subfamilies and genera tecognircd were based on a consideration of as many characters as possible, rejecting any approach, in which one character was given ar. overriding imporunee. and following as far as possible the methods used earlier ( Arkell 193.">—1-HJ in classifying Corallian perisubinetids. He discerned a monophvletie thread through the Peri-sphinetidae which he subdivided largely stiatigraphically, as had Buckman and Spa:h. Fui the iiathonian, he recognized three successive subfamilies—Lcptusphinc.tinae Arkell. 1950, ZigzaL'iccratinae Bucknian, 1920 and Pseudoprrssphiactinac Siliiudcwolf, 192").; the genera of this last subfamily "pass up into the < iallovian or become slightly modified and are calico by other names" (Arkell 1950, p. 104). This scheme was also adopted in Arkell's (I9.ri7a) Jurassic ammonite classification in the Trtahs? m InvtoUbriM Pay*.*::o!og)t Part I.. Mangold <197lb'} recognized three perisphinctid subfamilies within the Bathonian and Oallovian ol the l;rench Jura—Leptosphinctiuae Arkell, 19j0. Zig/agiceraunae Buckman, 1920 and (Irossouvriinê Sp.ith. 1931. He assigned Callovian forms to the /.igzagiccratinar :JIomoMpiwMiiiM and IndoipfoKctes) and the < i: ossouvriinae {C.koJJada. Phibdihphinrtts, Bhuttii>phinrt*j and

1'AKi.ft: I. Gene r a and subgiuc.-a o: ihc Pcmpl;inc r jdac published records in :hc Callovian

HISTORY OI- RKSKARC.H

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4 K N C L I S I I CALLOVIAN" PERISPHINCTID AMMONITES

Hamulisphinctes), and continued to use Buckman ' s Proplanul i t idae for Propla7u1lit.es and related forms which s tand aside as a distinct group.

S T R A T I G R A P H Y

In England, rocks of Callovian age crop out in an almost cont inuous strip f rom Dorset to North Yorkshire, broken only in west Dorset and across the Market Weighton Block in H u m b e r -side where they have been removed by erosion before deposition of late Cretaceous s t ra ta ( Text-figs 1-3) . T h e y underl ie younger sediments over much of the land area to the east of the outcrop, with the exception of a large area beneath East Anglia and the T h a m e s Valley, and a smaller area on the Marke t Weighton Block. Tota l thicknesses are difficult to establish because of inadequa te or imprecise da ta in many areas, and only est imates can be given. Where complete at outcrop in

southern and central Eng land , the total thickness is c. 45 m on the Dorset coast, c. 50 m in Oxfordshire and c. 50-55 m in the East Midlands . T h e sequence is c. 50 m in par ts of T h e Wea ld subcrop, but thins due to a t tenuat ion and later erosion as it approaches the L o n d o n - B r a b a n t Massif. In Humbers ide , the thickness is reduced to c. 15 m, but increases to as much as 40 m in North Yorkshire.

LITHOSTRATIGRAPHY From Dorset to Humbers ide , Callovian s t ra ta arc included in three main l i thostrat igraphical

STRATIGRAPHY 7

units - U p p e r Cornbras lu Kellaways Beds and Oxford (Hay (Lower and Middle)—which prob-ably represent an increasingly transgressive mar ine sequence of shallow water shell d e p o s i t s ranging from thin limestones through silly clays and sands, to a relatively thick clay uni t . Recent mapping by BGS in Lincolnshire and Humber s ide lias not differentiated the Oxford (!lay from the other U p p e r Jurassic clay format ions recognized elsewhere; in I hat. a rea , the Oxford d a y is represented in the lower part, ol the so-called Ancholrric d a y Group . In North Yorkshire, the argillaceous aspect of I his succession is replaced by an almost completely arenaceous sequence, the Osgodbv Format ion , represent ing a change to near shore mar ine environments . The Callo-vian is represented by lour main l i thostrat igraphieal units, the Cornbrash , Kellaways Rock, Langdale Beds and Hackness Rock (Text-fig. 4)

C a i R O N O S I KA I KrKAiM I V

T h e current s t andard Ju rass i c zones were established on the basis of par t icular ammoni t e faunas, and most fall broadly within the categories of assemblage or tota l - range biozones of current usage (Hoc! berg lc)7b: H o l l a n d ^ / / . 1978). In the Callovian Stage all the M sub /ones of the six zones (largely based on and recognized by the Boreal kosinoeeraiid and cardioeeral id ammoni tes) now have type and reference sections (Duff?/* Cope etal. I WO, Tab l e 2). '['lie Stage is divided into three subslages Lower, Middle and Upper—wi th two zones in each. Locally in the English sequence even finer chronosl ra l igraphical subdivisions (chronohorizons) may be dist in-guished, such as the Cornptoni Bed and Acut is t r ia tum Band.

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6 K N C L I S I I CALLOVIAN" PERISPHINCTID AMMONITES

L O C A L I T I E S

Callovian s t ra ta arc exposed in coastal sections in Dorset and North Yorkshire, and in brickpits and quarr ies along the outcrop. In Dorset, a complete Callovian sequence is exposed along the shores of T h e Fleet behind Chesil Beach (at Abbotsbury Swannery , and between East Fleet and Tidrnoor Point) but generally in low, badly slipped and overgrown cliffs. T h e Lamber t i Zone is again poorly exposed at the base of H a m Cliff to the E of Weymouth . O n the Nor th Yorkshire coast, there is a magnif icent section through the Callovian at Red Cliff at the southern end o f C a y t o n Bay, S of Scarborough, and these beds can also be seen far ther S at Gr is thorpe Cliff and Cuns tone N a b (Wright 1968). T h e famous exposures of Callovian stata. at Cast le Hill, Scarborough are now part ly obscured and relatively inaccessible. In land , there is a n u m b e r of

exposures in small quarr ies and s t ream sections between Hackness and Newtondale (where the Kellaways Rock forms almost cont inuous crags for more than 3 km) and in the T a b u l a r and Hamble ton Hills (Wright 1968). Between Dorset and North Yorkshire, there are fine exposures of Oxford Clay in working pits of the London Brick C o m p a n y at Calvert and Bletchley, Buckinghamshire; Bedford, Bedfordshire, and Peterborough, Cambr idgeshi re , where the work-ing faces show the J a s o n to Athleta Zones (Callomon 1968). Exposures of Kel laways Beds can be seen in the banks of the River Avon near Kellaways, Ch ippenham, Wiltshire, and in sandpi ts near South Cave, Humbers ide . T h e strat igraphical position of certain sections and localities referred to are shown in Text-fig. 4.

TEXT-FIG. 3. O u t c r o p of Cal lovian s t r a t a in no r the rn E n g l a n d .

STRATIGRAPHY 7

COKNKKASII The Cornbrash is t radi t ional ly divided into Lower and U p p e r divisions on the basis uf

lirhologv, bur the two are more easily dist inguished by their ammon i t e and brach iopod faunas (Douglas & Arkell 1928; 1932). T h e Lower Cornbrash , consisting of poorly bedded mar ly limestone with shell detr i tus, rubbly mar l and massive beds of blue-hear ted l imestone, belongs to the Bathonian and the Uppe r C o r n b r a s h , consisting of sandy l imestone 01 fer ruginous sandy marls with dogger-like l imestone beds and occasional massive beds of l imestone wea ther ing in to flags, belongs to the Cal lovian (Arkell 1933). Toge ther they total only a few metres in thickness at most, and ei ther division may be locally very thin or absent .

-jir o < H in CD D

CC LLI CL CL

LU _J Q Q

DC LU

ZONE

Lamberti

Ath le ta

Coronatum

Jason

Calloviense

M a c r o -ccphalus

DORSET TO H U M BERSIDE

( l o c a l n o n s e q u e n c e s n o t s h o w n ]

^ u p p e r c o i M x s ^ ^ a m IZcCuiU.J ' : ' i •.' r1.' 1'.' i11 g E a M m rCJ-U-i 1,1 ..I

NORTH

Y O R K S H I R E

HACKNfcSS / • ROCK•

O h-<

cc o

L A N G D A L E BEDS

>-CO

K ELLA WAYS . ROCK

SECTIONS A N D LOCALIT IES

Dorset

§ ! l 2 o

a uu U1 El

X

X

8" 9

a1 a l UJI

I

1 D <n

f i i |

i

N. Y o r k s h j p ,

I

I I I I

I •-I "l SI

H % I i

I l

zl

I >

> <\d LL LU

> -LU -J X .

H LU I —11 CD

i i i i si Q,

31 SI Si UjI Ei I

I i cn\

3

§1

4-

X CD

1° la: a s

i i

21 II o

i TEXT-FIG. 4. Main l i thostrat igraphical units of the English Callovian and stratigraphical range of certain sections and

localities.

Perisphinctid ammoni t e s occur in both divisions and care must be exercised therefore when examining old collections where the s t ra t igraphical informat ion is not explicit; in m a n y cases, it is not clear f rom which division (Lower or U p p e r ) specimens came and whether spccies are exclusive to one division or c o m m o n to both. There is an added discrepancy in sys temat ic t r ea tment—the Ba thon i an Lower C o r n b r a s h per isphinct ids and Clydoniceras were described by Arkell (1951-58) , bu t the U p p e r C o r n b r a s h ammoni t e s (perisphinct ids and Macrocephaliles) have never been treated systematical ly as a fauna . U p p e r Cornb rash per isphinct ids have been recorded from Long I I a n [ d ] b o r o u g h , Oxfordshire ; Berkley, Somerset ; and Sta lbr idge. Holwcll and Corscombe, Dorset (Douglas & Arkell 1928, pp . 129, 143, 147, 148, 152; Arkell 1954b, p. 119). Confusion may occur because Arkell (1958) cited and figured specimens from the Lower Cornbrash from localities at which he had previously recorded per isphinct ids f rom the Uppe r

10 KNCLISII CALLOVIAN" PERISPHINCTID AMMONITES

Corn brash only. The following notes may help to avoid confusion abou t (he p rovenance of specimens. Long Hon\d\borough, Oxjoukhhv: Perisphinctid material has been recorded from both the Lower and U p p e r Cornbrash . No specimen ha&> been traced 10 support Douglas & Arkell 's (1928. ]>. 120) I Jppcr Cornbrash record (neither O L M nor SM). Material f rom the Lower Corn brash was recorded by Pringle (1926) and later described and figured by Arkell (1958) (BGS GSM37364 , C S M 8 3 I 4 0 ; O U M J1370;. T h e r e is othci undesciibed maierial , without""stratigraphical information ( O I J M JSOOOa, b. J14010-1 . J14016a , b j 1 4 0 1 7 J 1 4 2 5 4 - 3 ) . Berkley. Somerset: Pcrisphinctids were recorded h u m a cream-coloured sandy limestone in a shallow quar ry at Berkley and assigned to the U p p e r Corn brash by Douglas & Arkell (1928, p. 143). Subsequently, reinterprcratinn o f t h i s section with reference to ones nearby a t Frome showed that it belonged to the Lower Cornbrash (Arkell 1933, p. 330, l % 4 b , p. 120; 1958b, p. 218): (BGS GSM95035, 95039).

ZONE suezoNt TYPF LOCALITY REFERENCE SECTION(S)

Lamberti Lamberti Woodham brick pit Buckinghamshire:

lamberti Lrniestono (Bed C}1 Rivp' H'ora section.Sutherland: Haccally Sandstonelupocr oaril ond Cly- elish Quart y.SutliHrl/inil C.lynpiish Quarry Sandetono Lamberti

Henrici Woodham brickoit. Btjr.kiiujli#insliire: Oxford Clay iBcd U2}4

Foreshore section,south of Bioid HslitHiy, SuihmtancJ: Pascal y Siltctonc \jddci parti 6 Faace'ly 5flnd3'.one ;ioy/«r aaril

Athlota

Spinosum Wood ham brickpit Bucking ham shire: Oxford Clav '.Bed3 E-D1)4

Foreshore section,j>ouir of Brora estudiv . Sutherland, rascally SillHiriiw lu|i|i«i par?}

Athlota Proniee Calvort brickpit. Buckinghamshire: Oxford Clav !0eds 13a-hl4

Fintfhliuu? suction ,south of Bro'a Ofilua y, Sutherland: Faacelly Sillatonc (niodlif iJdill Athlota

Phocinum Calvert brickpit. Qucfcinohnmshiie Oxford Clav IBeds 10-121*

Foroshoro KPction,south of U'ora octuary Suthorlond: Hrora 3nck Clay b Foacolly Siltatone {'ower niirtj

Coronatum Grossouvrci Peterborough pits.Carnbridoeahirc! :

Uxtord Clay IB«ds'l7a-22cl4

Coronatum Obductum Peterborough pits.Cambridgeshire:

Uxtord Clav IBeds 14-16bl4

Jason Jason KidlinotonOxfordsliire:

Oxford Clay IBeds 15 26}* Peterborough oits. Cambridgeshire: Oxford C\*v (Beds 10-13I4

Jason Medea Kidlington, Oxfordshire

Oxford Clav<Reds 9-14}* Peterborough pits.Cambridgeshire: Oxford Clav IBeds M l '

Calloviense

Enodaturn South Cave. Mumbnrsirie: Kellawavs Rock

Calloviense CalloviensR Kellaways ,nr Chippenham . Wiltshire: Kcllawavs Rock Calloviense

Kueniyi Chippenham. Wiltshire: Kcllnwova Clav

Macro - Kamptus KidlingtonOxfordshire: Kellaways Clav IBeda 3 4\s Peterborough plts,Cambridgeshire:

Kellaways Clay (Bed 2f cephalus Mnr.ro-

cophalu6 Sutton Bingham roservoir.nr Yeovil. Somerset: Cornbrash (Beds 4 1U)2

Shipmoor Point Chester's Hill. Abbotobury Swannery, Cornbrash (Bads 3-14lJ Oorspt

Bed no* of 1 AiJk«H 1A39 2Arke«M9W 3n llnmon 1956 4Callomon I968 5Uouglas it Arkcl'

T A B L E 2. Cal lovian c .hronostrat igraphic zones and subzones. with their type localities, rcfercncc sections and horizons.

Slalbihige, Dorset: Pcrisphinctids have been recorded from the U p p e r Cnrnhrash ofSta lbr idge and the Lower Cornbrash of Stalbridge Weston. T h e specimen from t he Upper Cnrnhrash ofSta lbr idge (Douglas & Arkell p. 147) is OU.Vl J12767 (PI. 1, fig. 3); both U p p e r and Lower Cornbrash were present at the locality. Occurrences in the Lower Cornbrash ofSta lbr idge Weston (notably the type of I loinoeoplanidllvx fwitweomorphus Buckman) were discussed by Arkell (1958b, p. 226). Only Lower Cornbrash is thought to have been exposed in the quar ry there, which was worked in the last century; (BGS G S M 0 6 5 4 - 5 , SM J20354) . ' Hot well. Dorset: There is some d o u b t about specimens from this locality. An ex situ specimen was recorded from the Upper Cornbrash by Douglas & Arkell (1928, p. 148). but no specimen has been traced (neither O U VI. S M nor DC M ). Arkell (1958b. pi. 32 fig. 1) figured a specimen { O t ' M J 13002) which he inferred to be from the Lower Cor nbrash . 1 his was one of a n u m b e r of specimens formerly held by the Sherborne School Museum. M u c h of the material displayed there (Torrens 1978) from the C o r n b r a s h of Holwcll (including the Rev. H. Wood collection probably made between 1860 and 1880} has apparen t ly been lost; only two ammoni tes (a Clvdoniceras and a Macrocephalites) a re preserved in the material transferred to B M , and the D C M has none. The re are two specimens (BM CI 1796, CI 1825; PI. 1, figs 1. 2)

D E I ^ GHOLUGIP- I O ;

STRATIGRAPHY 9

collected by Blake {1905, p. 7) from the " U p p e r Beds" ar Holwell; he recorded 23 ft 9 in f 7-2 m | of s trata, and the upper 2-9 m at least is apparent ly K p p r r Cornbrash (with Maaocephalitrs); the rest is Lower Cornbrash and 1 ores I Marb le (Arkell 1958b. p 226). T h e former specimen was figured by lllake (1905. pi. 5 lig. 5, wrongly numbered 4 on plate) and discussed by Cox & Arkell (1950, p. 94, incorrectly numbered BM CI 1795). ' Corscombe, Doisd: T h e specimen from the U p p e r Cornbrash (associated wiih Macrocephnlitcs) recorded by Douglas & Arkell (1928, p. 152) f rom a temporary exposure at Corscombe, has not been traced (neither O L ' M , SM nor D O M ) . Both Lower and U p p e r Cornbrash were quarr ied around this village.

KELLAWAYS B I D S T h e Kel laways Beds of sou the rn Britain, which belong to the M a c r o c e p h a l u s Zone (pars) and

the Calloviense / o n e , are best known in Wil tshi re ( the type area; Cave & Cox 1975) and Gloucestershire . The major i ty of m u s e u m spec imens come f rom these two areas, e.g. former brickpits at S idd ing ton , Glouces tersh i re , ra i lway cut t ings at South Ccrncy , Glouces tersh i re , Bincombe Wood . C h i p p e n h a m and Trowbridge, Wil tshire , and road cut t ings on the M 4 motor-way near Chr i s t i an M a l f o r d , Wil tshire (Mante l l 1850, Harke r 1881, W o o d w a r d 1895, Reynolds & V a u g h a n 1902. Ga l lomon 1961, Barron 1972, Cave 1977). 1'he two-fold l i thological division of Kel laways Clay below and Kel lawavs Rock a n d / o r Sand above, can be recognized sou thwards into Somerset a n d Dorset (Arkell 1932, 1947c, 1948b, Cope 1969, Cope & Cox 1969), and nor thwards into O x f o r d s h i r e (Arkell 1947d, Gal lomon 1955) and the East M i d l a n d s (Ca l lomon 1968) where they floor the extensive Oxford Clay brickpits.

In Hurnbers ide , the Ke l l aways Beds compr i se a thin Kel laways Clay overlain by a renaceous Kellaways S a n d a n d Rock. T h e Cal loviense Zone a m m o n i t e f auna , which occurs main ly in the Kellaways Rock, differs f rom that of this s t r a tum in southern Eng land and is assigned to a younger ( E n o d a t u m ) Subzone (Ca l lomon 1964, p. 275). These a renaceous s t r a t a are known f rom quarr ies at South Cave , Sou th Newba ld and Ket l l e thorpe , a n d f rom the rai lway cu t t ing at Drewton (Keep ing & Midd lemiss 1883, De Boer, Neale & Penny 1958, Brasier & Brasier 1978).

Kel laways Rock is also recognized within the completely a renaceous Cal lovian sequence of Nor th Yorkshi re a n d has yielded per i sphinc t ids at the coastal localities of C a y toil Bay, O s g o d b y Nab , Corne l i an Bay and at scat tered localities between there a n d Newtonda le (Wr igh t 1968).

O X F O R D CLAY T h e Oxford Clay is a laterally r a the r uni form argil laceous sequence rang ing in thickness f rom

!j0 to 15Um. O n the basis of gross lirhology and Fauna., it is divided into th ree—Lower , Midd le and U p p e r . T h e Lower Oxford Glav is a sequence of b i tuminous shaly clays wi th crushed ir idescent fossils often preserved in whi te a ragoni te . T h e following zones are represented: Cal loviense (pan), J a s o n , C o r o n a t u i n and Ath le ta (pars). T h e Midd le and U p p e r Oxford Clay are sequences of non-bi tuminous , pa le grey calcareous clays. T h e y are less fossiliferous t h a n the Lower Oxford Clay and the a m m o n i t e f a u n a is preserved commonly as uncrushed pyri t ic inner whorls or nuclei . T h e Middle Oxford Clay belongs to the A th le t a Zone (pars) and Lamber t i Zone, bu t the U p p e r Oxford Clay is entirely O x f o r d i a n ( M a r i a e and C o r d a t u i n zones) in age. T h e clays a re or have been worked in a n u m b e r of pits, fossils f rom which are well represented in m u s e u m collections, a l though the a t t rac t ive small pyritized ammoni t e s are rarely accompan ied by detai led s t ra t igraphica l in format ion . M u s e u m collections also include mater ia l f rom coastal ou tc rops . These Cal lovian localities are listed below (from S to N ) .

Tidmoor Point, Dorset; Lamber t i and PMariae Zones (Spath 1933, p. ttoG, Arkell 1947c. p. 30). Wolvcreote and Summer town , Oxfordshire; Athleta Zone (Pringle 192(i, pp. 31, 30; Arkell 1939b. p. 207: 1947d, p. 70). Wood ham. Buckinghamshire ; Athleta , Lamber t i and Mar iae Zones (Arkell 1939b, Cal lomon 1968, p. 288). St Ncots, Cambridgeshire,- Athleta Zone ( Roberts 1892. p. 11. Edmonds & Dinham 1965, p. 14).

T h e a m m o n i t e f a u n a of the Lower a n d Midd le (i.e. Cal lovian) Oxford Clay is d o m i n a t e d by Kosmoceras, which is used as the basis for the zonat ion. f t is a ccompan ied by ra re r card iocera t ids ( though these equa l Kosmoceras in n u m b e r by the end of the Cal lovian) , oppel i ids , per isphinct ids , pachycera t ids , reineckeiids and aspidocera t ids . Per isphinct ids are par t icular ly c o m m o n at three


horizons which are widespread th roughout southern and central E n g l a n d — t h e C o m p t o n i Bed, the Acut i s t r ia tum Hand and the Lamber t i Limestone. They also form a significant, p a n of the assemblage in the basal shell-bed of the J a s o n Subzone, well-exposed in the brickpits at Calvert , Buckinghamshire , forming the base to which the pits a re worked (Cal lomon 1968. p. 285).

T h e C o m p t o n i Bed is a thin (usually less than 0.3 m) shell-bed occurr ing at the top of the Corona t ion Zone in the Lower Oxford Clay. It is crowded with pcr isphinct ids , accompan ied by Erymnoceras, Hecticoceras and Kosmoceras, and m a y be associated with a band of hard calcareous shale or fissile l imestone rich in nuculoid bivalves and Mele.agrinella., with locally-developed l imestone lenses. T h e Acut i s t r ia tum Band is a highly calcareous muds tone or concret ionary l imestone at the base of the overlying Athleta Zone. T h e type locality is (Calvert, Buckinghamshi re (Buckman 1025) ; bu t most museum collections a rc from Chris t ian Malford , Wil tshire , where the horizon was exposed d u r i n g the construct ion of the Grea t Western Rai lway (Pra t t 1841; Cal lomon 1968. p. 274). T h e Lamber t i Limestone, at the top of the Midd le Oxford Clay, is a thin (generally less than 0-3 m), pa le grey, siltv. calcareous muds tone or marly l imestone, packed with ammoni tes which are usually dis tor ted and often f ragmentary; they a re preserved as internal clay moulds with a black or brown powdery coat ing of pyrite. This bed belongs to the uppe r part of the Lambert i Zone and is best developed in the Buckinghamshire area, where it was exposed at W o o d h a m brickpit (Arkell 1939b).

H A C K N E S S R O C K

T h e I laekness Rock was identified and defined as a discrete unit within the so-called Kel loway Rock [ = Osgodby Forma t ion , Text-f ig. 4] of the Nor th Yorkshire coast by Arkel I (1933, 1939b). 1 r is a thin (c. 2 0m) sequence of poorly sorted, sandy limestones and fine-grained calcareous sandstones, in both of which chamosi te ooliths may be common, together with silts and marls (Wright 1968, H e m i n g w a y 1974); it belongs to the Athle ta and Lamber t i zones. W h e n wea the red , some beds have a distinctive purpl i sh-red colour mottled with green. Its fossils were first described by Leckenbv (1859)1 including the type specimens of the pcr isphinct ids Attigaticeras Buckman , 1923, Binatisphinctes Buckman , 1921, Hamulisphinctes Buckman , 1921 and Poculisphinctes Buckman, 1920. T h e s e occur with kosmocerat ids , cardioccrat ids , oppeliids, aspidoccrat ids , pachycera t ids and rcineckeiids. Ca re m u s t be exercised in the examinat ion of old m u s e u m collections because Hackness Rock specimens arc included in, and must be identified from amongst , mater ia l marked "Ke l loway Rock" . T h e Hackness Rock has yielded pcr isphinct ids at the coastal localities o f C a y t o n Bay, Cornel ian Bay, Cuns tone Nab , Gr i s thorpe Cliff. O s g o d b y Nab , and a t Cast le Hill, Scarborough and Hackness Q u a r r y (Wright 1968).

C R I T E R I A F O R C L A S S I F I C A T I O N

T A X A OF T H E SPECIES G R O U P Only in a few cases (e.g. Ilomoeoplanulites difficilis and Binatisphinctes comptoni) is mate r ia l

sufficiently plent iful , well-preserved and wi th sufficient s t ra t igraphical control (and hence assumed to be isochronous) for biospecific assemblages to be m a p p e d out with any confidence. I n most cases, specific classification has to be based upon morphospecies , and it is p robab le tha t con temporaneous ones will p robablv be merclv var iants of a single biospecies (cf. Ca l lomon 1981, p . 2 6 0 ) .

M a n y ammon i t e workers have used Arkell 's (1936, p. x) in t roductory notes on general ' John T.eekenhy (1814—77) wa.s a hank manager in Scarborough and one time T rea su re r of the Borough of

Scarborough (Ob i tua ry in Proc. gtol Soc. Lonxt. 1878, 34, p 35}. He was a contemporary of William Bean (1787— I8t>6)—a local lossil collector of some renown. Bean himself did not publish many scientific papers describing the specimens which he collected but many new species were described by others, using his manuscr ip t (often specimen label} names . Bean's own fossil collection was sold to the British Museum in 1859 for £500 (McMi l l an Green wuod 1972). T h e Leckenby Collection was purchased, following an appeal bv Professor Adam Sedgwick, by the University of Cambr idge in 1871 for £800 {Nature, Dec. 1871, p. 151).

CRITERIA FOR CLASSIFICATION 11

classification, par t icular ly in the case of perisphinct ids. Arkell (1951a, p. 23; 1958b) later added only a few qualif icat ions to these. H e considered four main criteria—shell d imensions , whorl shape and coiling, septal sutures, and r ibbing; he (Arkell 1958b, p. 235) concluded that rib-style had proved to be the most valuable basis tor classification amongs t perisphinct ids . Since Arkcll 's work on Bathonian and Oxford ian forms, there have been two o ther significant m o n o g r a p h s (Enay I960; Mango ld 1971 b), in both of which Arkcll 's classification criteria were used as a basis. Mangold used fur ther morphological features to deline the var ious ranks in the hierarchy of taxa: sutures and o r n a m e n t on the early whorls for subfamilies, sequence of successive stages of ornament for genera and (in order of priority) mode ot division of pr imary ribs, average d iamete r for appearance of adul t charac ters , s t rength and n u m b e r of ribs, index of rib division, whorl section, and shell d imens ions and coiling, for species. ArkelPs four main cri teria a re each discussed below in the light of more recent work, including the approach adopted in this monograph.

T h e morphological terms are those used by Arkell (1935; 1957a, pp. L3, L81). Any qual i fy ing remarks or addi t ional terms, most of which reflect cur ren t common usage, a re given where appropriate .

Shell dimensions Species arc defined on the basis of complete adul t individuals , the criteria for which are:

(i) contract ion of the body-chamber towards the aper tu re with uncoiling of the umbil ical seam. (ii) modificat ion of the shell form near the per is tome (coarsening and regenerat ion or degenera-

tion of r ibbing, often with terminal constrictions, ventral collars, flares, horns , ros t ra , lateral lappets etc.).

(iii) approximat ion and degenera t ion of last few septal sutures (Cal lomon 1963, p. 25).

The m a x i m u m size of a pcr isphinct id species is a well-defined charac te r , with a variability from the mean of c. 10% (Cal lomon 1955, p. 237; 1963, p. 26). In reasonably numerous assemblages, assumed to follow a normal d is t r ibut ion, the range between the smallest and largest individuals will usually encompass a range abou t the mean of plus or minus two s t anda rd deviations (Howar th in Ca l lomon 1963, p. 28).

A widely used convention for expressing shell d imensions is the formula of Buckman (1912, p. viii):

(i) d iamerer (d) in mill imetres (h) whorl height (wh) as fraction of (i)

(hi) whorl thickness (wt.) as fraction of (i) (iv) umbilical width ( u w ) as fraction of (i)

to which mav be added

(v) d iameter at last s ep tum in specimens with some or all of the body-chamber preserved (vi) length of b o d y - c h a m b e r preserved, as fract ion of a whorl .

These pa rame te r s give some indicat ion of size, whorl section and coiling (see below). Whor l thickness is a meaningless p a r a m e t e r in crushed specimens but d iameter , whorl height and umbilical width m a y be only slightly dis tor ted. I n this monograph , pa rame te r s (i) to (iv) a re given for figured specimens, bur (ii) to (iv) a rc not expressed as fract ions of (i). Whor l thickness m a y be measured between or on the ribs; the. difference in per isphinct ids is usually only smal l , and the measurement is taken on the rib. Whorl height is measured in the plane of coiling, not obl iquely. The known m a x i m u m d iamete r for a species and pa rame te r (vi) arc given in cach species description.

Whorl shape and coiling The whorl section (Text-fig. 5a) may be described as compressed (higher than wide) or

depressed (or inflated) (wider than high) and expressed by the ratio:


wh'f

TKXT-FU;. 5a (left) Description of whorl section as ratio of whorl height (wh) and whorl thickness fwt) .

1'EXT-FIG. 5b (above) Coiling as ratio of whorl height (wh) and diameter (d).

(i) w h ; w t ( the greater the ratio, the more comprcssed the whorl)

T h e style of coiling (Text-fig. 5b) , is expressed by the following ratios:

(ii) w h : d (the greater the ratio, the more involute the shell) (iii) u w : d (the smal ler the ratio, the more involute the shell) (iv) d : n u m b e r of whorls ( the greater the ratio, the more evolute the shell)

T h e terms involute and evolute a re relative. All the Cal lovian per isphinct ids are more or less evolute with wh:d ratios ranging from 2 0 - 3 0 % (evolute) to 30 -40% (moderate ly evolute). T h e Proplamliles f auna is the only involute group, with wh:d ratios ranging above 4 0 % . Whor l sections (wh:wt ratios) r ange f rom compressed (140-150%)—modera te ly compressed ( 1 1 5 - 1 4 0 % ) — slightly compressed ( 1 0 5 - 1 1 5 % ) — r o u n d e d / s q u a r i s h (95-105%)—sl ight ly depressed ( 8 5 - 9 5 % ) to modera te ly depressed ( 6 0 - 8 5 % ) .

T h e combina t ion of style of coiling and degree of compression may be described by te rms such as p lanula te , p la tycone, serpent icone, and expressed by the ratio

(v) u w : umbilical dep th (ud) the greater the ratio, the more serpenticone-l ike the shell.

Each of these five ratios can be shown graphical ly in a bivariatc plot. In the ideal case, such a plot would show a positive correlat ion represented by a straight line (perfect funct ional relation-ship) or an even curve. T h e wh;wt ratio in per isphinct ids tends to increase dur ing ontogeny (i.e. inner whorls tend to be more or less depressed and outer whorls tend to be more or less compressed) , bu t it m a y decrease.

Septal sutures T h e general form of the suture has been the basis of many ammon i t e classifications, e.g.

Buc.kman (1909-30) . As internal shell features, they were thought to be of more f u n d a m e n t a l significance than external features such as r ibbing tha t might be influenced by envi ronmenta l condit ions. Spa th (1930, p. 7) considered the role of sutures in classification to be m u c h overra ted ,

c o m p r e s s e d

d e p r e s s e d

CRITERIA FOR CLASSIFICATION 13

and many authors have followed Arkell 's (1957b, p. 246) conclusion tha t they const i tute only one of a number of at least equally significant morphological characters ; he contended tha t the form of the suture was influenced strongly by the shape of the whorl and that var ia t ion within a species or an individual might be great . Al though Arkell 's views have been challenged, for example by Schindewolf (1957) and W e s t e r m a n n (1958), sutures have since assumed a less significant role in generic and specific classification. They are, however, f u n d a m e n t a l cri teria for recognizing the major (suborder) divisions of Palaeozoic and Mesozoic ammono ids ( W i e d m a n n & K u l l m a n n 1981). In this monograph , only a s u m m a r y of sutural charac ters is given for each genus.

Ribbing T h e ontogenetic deve lopment of rib style is generally useful for the definit ion of species and

where possible tha t lor the inner , middle and outer whorls (including body-chamber ) is descr ibed separately. All per isphinct ids tend to have similar inner whorls and these are in terpre ted by-association with accompany ing larger forms where available. Constr ic t ions are more c o m m o n in some groups than others and vary f rom broad and shallow to deep and nar row; they arc included as a feature of rib-style in the descript ions.

As well as style, there a re variat ions in r ib n u m b e r which Arkell and others though t also reflected specific differences. These are shown by plotting r ib-density curves (d iamete r against number of ribs). Ideal ly, each species is thought to have its own character is t ic c u n t , bu t Arkell (1935, p. xviii ) emphas ized that compar i son of r ib-density curves for ident ifying species should always be subord ina te to a considerat ion of the shape and style of the individual ribs. A line enclosing all the points in a plot of as many specimens as possible f rom one popula t ion shows the extent of var ia t ion; over lap between species may be considerable. Es t imated d iameters , which will have only a small marg in of error , can be used for perisphinct id inner whorl r ib counts . Rib-density curves are given in the species descript ions only where a reasonable n u m b e r of ma tu re individuals are available.

When rib counts are m a d e , intercalatory ribs are included with the secondary ribs. Simple, unbranchcd ribs m a y be included in pr imary and secondary r ib counts; in the lat ter case, the more accurate collective n a m e 'ventral r ibs ' should be used. Strong, widely spaced, b lunt ribs are called massive. T h e general style o f o r n a m e n t m a y be described as pcr isphinctoid , that is mainly regular bifurcate r ibbing with constrict ions.

Dimorphism Dimorphism has been a m a j o r considerat ion in ammoni te taxonomy since the idea was

revived by Ca l lomon (1963) and by Makowski (1963). D imorph ic pairs are des ignated mac.ro-conchs [M] and microconchs m] (Cal lomon 1955, p. 238). By analogy with living cephalopoda, and because it develops both ontogenctieally and phyletically (Cal lomon 1981, p. 260) in ammonites, this d imorph i sm is considered to be sexual.

The Per isphinctaceae provides some of the most consistent and impressive examples of dimorphism (Cal lomon 1963, p. 37), part icular ly those from the Ba thon ian , Callovian and Oxfordian. D imorph i sm in Ba thonian pcr isphinct ids was recognized by Arkell (1951 58). H a h n (1969) and Mango ld (1971b). Arkell (1951, p. 2) drew at tent ion to the presence of forms of large size (diameters greater than 400 m m wi thout the body-chamber as in Procerit.es) and smaller forms (diameters of only 100 m m ) with body-chambers bear ing lappets such as Siemimdrkia. bur he did not formally recognize d imorph i sm in his classification which used well-established generic names for g roups defined mainly on the basis of rib-style. However , he concluded that the persistence of the two main morphological types over a long period (Bathonian to U p p e r Oxfordian) lent a t t rac t ion to the hypothesis of sexual d imorph i sm. An examina t ion of phylogeny taking account o f m a c r o c o n c h and microconch forms led H a h n (1969) to recognize a phylogenet ic tree of paired d imorph ic lineages evolving in parallel . He a t t empted to express this systematical ly by dividing each genus into a microconch subgenus and a macroconch subgenus . Th i s a p p r o a c h was also adopted by Mangold (1971b) lor rhe Bathonian and Cal lovian per isphinct ids of the

14 ENGLISH CALLOVIAN PERISPIIINCTID AMMONITES

southern J u r a . T h e d imorph ic relat ionships amongs t existing Callovian perisphinet id genera were out l ined by Ca l lomon (1963, p. 37). Similar d imorph ic relat ionships within Oxford ian forms were recognized by Enay (1966).

In Callovian per isphinct ids d imorph i sm can be recognized ai most s t ra t igraphical levels. Although it is generally believed that macroconchs are females and microconchs males , the s t anda rd zoological symbols a re not used because we cannot be absolutely cer ta in . However , in allowing that d imorph i sm is sexual, ii does lead TO a rationalization of the t axonomy. T h e large n u m b e r (<?. 360) of described Callovian perisphinetid species were based on morphological criteria, and it m a y nor he possible in many cases TO tell whether r.he type specimen of a particular-one is a microconch or a macroconch .

In this monograph , a species is taken to include both the macroconch and microconch; where possible the type specimen is classified as [MJ or [m j. When the mater ia l is sufficiently preserved, both forms of each species are described separately; otherwise the descript ions are combined , with approp i i a le q uali li cal ion.

T A X A O F T H E G E M ' S G R O U P In a specific classification based on morphological differences, genera are merely well-

established, t radi t ional ly used " superg roups ' ' into which morphological ly similar species a re collected, r a the r than phylet ic units compr is ing a succession of biospecies. T h e subgenus is not used herein, because there is some ambigui ty in its usage either as a taxon of infra-generic, supra -specific rank, or as Lhe nominal taxon for incorpora t ing " the morphological manifes ta t ion of d i m o r p h i s m " into a morphological classification (Cal lomon 1969, p. I 16). As most genera were established before the significance of d imorph i sm was apprecia ted widely, and recent studies have dealt with d imorph i sm at generic and subgener ic level, the procedure adopted here has mean t that a n u m b e r of generic and subgener ic names have become jun io r synonyms. These are discussed under the app rop r i a t e genus .

T A X A OF T H E F A M I L Y - G R O U P T h e classification o f j u r a s s i c ammoni t e s adopted in the second edition of the Treatise (Part L)

favours the use of the subfamil ies Pseudoper isphinct inae , Proplanul i t inae and Per isphinct inac for the Cal lovian per isphinct ids , thereby following Arkell (1957a) except that the Zigzagicera t inae are restricted to the Lower and Midd le Ba thonian (Donovan el al. 1981). Th i s classification is followed herein, but Binalisphinclet> is included with the Pseudoper isphinct inae ra the r than with the Pe l toce raunae <>1 the Aspidocera t idae .

S Y S T E M A T I C D E S C R I P T I O N S

Fa mil v P E R I S P H I N C T I D A E S te inmann , 1890 Subfami lv P S E U D O P E R I S P H I N C T I N A E Schindewolf, 1925

Genus H O M O E O P L A N U L I T E S Buckman , 1922

(Synonyms. Anaplanulites Buckman , 1922 (type species A. difficiiis Buckman . 1922) [m] , Parachoffatia Mangold , 1971b ( type species Ammonites subbakeriae d ' O r b i g n v , 1850) [ M ] ,

?Loboplanulites Buckman , 1925 (type species L. longilobatus Buckman , 1925))

Type spmes. Homoe.oplamlitf.s hamMommphus Buckman, 1922 [m Diagnosis. Modera te ly evolute to evolute, with compressed ink die-outer whorls (Text-fig. 6);

lappered microconchs r ibbed th roughou t with ra ther dense, recl i radiate o r p rors i rad ia te pr im-aries. somet imes with rurs i radia te aspect to secondaries (Text-fig. 7). Macroconchs modera te ly and progressively var iocostate , with weakening of secondaries and finally loss of all r ibbing towards aper ture . W e a k parabol ic ribs or constr ict ions on inner-middle whorls. External su tu re has first lateral saddle with well-marked accessory lobe, a t r i f id , slightly asymmetr ica l lateral lobe about as deep as ventral lobe and a suspensive lobe with some well defined auxiliaries.

Discussion. W h e n Buckman (1922, pi. 328) created IIomoeoplanulit.es, he figured only one.

HOMOKOPLANLI.ITKS If)

specimen of the type species with a brief capt ion. He later (1924, pi. J15) figured an addi t ional new species. II. stabilis. T h e s e figures show medium-sized (100-140 m m m a x i m u m diameter ) evolute forms, r ibbed to the ape r tu re which bears lappets. T h e r ibbing consists of fairly dense, long, slender, s t ra ight , rec t i radia te or prors i radia te pr imaries which split into two or three secondaries with intcrcalatorics, and there are shallow constrictions. As originally proposed, the genus thus accommoda ted microconch forms. Buckman included the genus in the oldest hernera of his Macrocephal i tan age. which succeeded the Clvdonicera tan age, and thereby implied that it came from the U p p e r C o r n b r a s h (Callovian) ra ther than the Lower C o r n b r a s h (Ba thon ian) .

Arkell (1958. p. 225) believed that the holotypc o f / / , homoeomorphus was a s tunted individual whose growth may have been inhibi ted by injury, and tha t the holotypc of II. stabilis was a more normal, fully-grown individual (with a d iamete r of c. 135 m m ) . Spath (1931, p. 353), Arkell

TEXT-FIG. 6. Whorl sections of species Homoeoplamlites and Indospkincies: (a) BM CI 1825 at 110 mm; (B) BGS GSM /070 at (JO mm; (c) FW T N N 5 2 8 at 66 mm; (d) FW T N N 1 4 5 at 75 mm: (e: F W TN.Y173 at 87 m m ; (f) BM CI 17% at 230 mm: (g) BGS GSM7680 at 136 mm; (h) FW T N N 3 0 6 at 160 mm; fi) BGS /.t 167.5 at 10.5 mm: (j) GBV 1870/04/1 at 210 mm; (k) BGS 7682 at 65mm.

(1958, p. 22J) and Mango ld (1971 b, p. 61) synonymizcd these two species, and Homoeoplanufiles is now considered to include relatively large microconch forms of u p to 150 m m diameter . T h e following 'species' are also included: aculicosta Roemer , aequalis Roemer , balinensis Ncurnavr , bugesiacus Doiriinjon. couxi Domin jon , furcula N c u m a y r , leptns Cemmel l a ro , mangoldi Domin jon , pseudoannulum Lissajous, rambertensis Mangold andybbsensis Ju s scn : they are all based on Euro-pean forms and are recorded from the Upper Bathonian a n d / o r Lower Cal lovian. Both Arkell (1950b, p. 211) and Mango ld (1971b, p. 49) included Buckman ' s o ther Lower Cal lovian perisphinctid genus Anaplunulites, based on an Knglish form, as a j un io r synonym. Th i s la t ter genus has never really found accep tance and has also been included in Proplamtliles (Spath 1933; Arkell 1945) and Choffatia (Arkell 1957a). Buckman named only one species, A. difficilis, based on a single specimen (PL 1, fig. 6); discussion of the genus is therefore to a large extent a discussion of the species. Unti l M a n g o l d ' s (1971b) work, Homoeoplanulitcs was considered a microconch p a r t n e r and subgenus of Choffatia (Arkell 1957a, p. 317; 1958, p. 211; Cal lomon 1963, p. 37: H a h n 1969, p. 70). However. Mangold (1971b, p. 48) separa ted iUmocaplamdite.s as an independen t genus in


w h i c h he also included die larger forms belonging to the g roup of subbakeriae d ' O r b i g n y . H e removed this la t ter g r o u p from its t radi t ional place in Choffatia because of differences in o r n a m e n t and suture, and in t roduced a new taxon, Parachojfatia, for these macroconch pa r tne r s of the microconchs grouped in H amofoplamtliles s. s. As well a $ subbakeriae, Mangold based Pa/acfwfjatiaon Q[)[)vY s funalus and dis t inguished it f rom Proceriles by the more marked differentiat ion of pr imary and secondary ribs; also included were the species madam Spa th and arisphinctoides Arkell. Mango ld (197 I b, p. 73) likened ParachoffaUa to Buckman ' s (1925, pi. 596) genus Loboplanultle.s but concluded that (he type and only specimen of the type species of the latter, f rom the C o r n b r a s h of Long I l anborough . ( )xfbrdsh i re , was too f ragmenta ry and worn to be proper ly in terpre ted . Latei (1971b, p. 15/), he regarded it, as did Elmi (1962), a junior synonym of Subgmssouvria (now

Ti:x r-i-'iG. V. Ribbing ufsju-cics of Homotofilanvi'iHw and Indosbkinck.-..

HOMOEOPLANULITES 17

included in Choffatia). However . Ca l lomon (pers. c o m m . 1901) believes LoboplanulUes is "a lmos t certainly merely the inflated var iant of Homoeoplanulites".

The earliest known representat ives of Homoeoplanulites are from the Ba thonian , where it is thought to have originated f rom the d imorph ic g roup ofProcerites [macroconchs] and Siemiradzkia [microconchsj. Mangold (1971b, p. 13) included Homoeoplanulites with these two genera in the Zigzagiceratinae. m e m b e r s of which arc character ized by the presence of a var iab le "z igzag" stage of o rnament , result ing f rom a development of parabol ic ribs and nodes with s t rengthening of the ribs in the early ontogenet ic stages. Morphological in termedia tes in the Hodsoni and O r b i s (olim Aspidoides) zones of the U p p e r Bathonian make the dividing-line between the Pseudo-perisphinctinae and the ancestral Zigzagicerat inae arb i t ra ry , according to Ca l lomon (pers. comm. 1985).

Homoeoplanulites cf. subbakeriae (d'Orhigny, 1850) PI. 1. figs 1-3 Material. T h r e e poorly-preserved f ragments from the U p p e r C o r n b r a s h (Macroccpha lus

Zone) o fS ta lb r idge a n d U p p e r ? C o r n b r a s h of Holwell, Dorset. Discussion. Despi te a n u m b e r of records o \ H. suhhakeriae f rom the English U p p e r C o r n b r a s h

(Douglas & Arkell 1928), these are the only specimens that have been traced tha t can with any certainty be j u d g e d to have come from that horizon. They are too poorly preserved for detai led description and definit ive de te rmina t ion , but a p p e a r to he from moderate ly cvolutc forms with the following characterist ics; a m a x i m u m diameter of ?more than 250 m m (macroeoneh) ; moderately compressed whorl at least on middle and outer stages, with flat whorl sides and moderately broad venter . O r n a m e n t of ra ther dense and well-defined pr imar ies d iv id ing regularly into two or three secondaries with intercalatories which increase in n u m b e r with g rowth and fade on the venter .

Cox & Arkell (1950, p. 94) identified the specimen f rom Holwell , figured by Blake (1905) (see Introduction), as a f r agment of an outer whorl of Choffatia cf. lungilubala Arkell, but later Arkell (1958, p. 217) thought it p robably belonged with H. subbakeriae. a l though it was too small a fragment to be really identifiable. T h e former species, type of Lubup la nut ties, has s touter , r ounded , circular or depressed whorls and more massive r ibbing than the H. subbakeriae g r o u p (Arkell 1958, p. 212).

Data for figured specimens.

d wli wt uw pr imaries ventrals BM CI 1796 (PI. 1 fig. 1) est. 230 67 50 est. 85 (4) — BM CI 1825 (PI. 1 fig. 2) est. 110 c. 30 c. 20 est. 55 — (26) O U M J 1 2 7 6 7 (PI. 1 fig. 3) est. 60 — — est. 30 est. 40 —

Homoeoplanulites afT. balinensis (Neumayr , 1871) PI. 1, figs 4, 5 Material. T h r e e specimens (two completely septate , and one incomplete but with some body-

chamber), uncrushed (one in nodu la r cements tone), pinkish shell material part ial ly preserved; presumed microconchs. From Kel laways Clay (Calloviense Zone, Koenigi Subzonc) of Trowbridge, Wil tshire and Siddington, Gloucestershire.

Description. M a x i m u m d iamete r seen 95 m m , apparen t ly including c. 170° b o d y - c h a m b e r which begins a t e . 65 m m d iamete r . Five whorls visible. Moderate ly cvolutc ( w h / d = c . 3 0 - 3 5 % ) ; whorl section squar i sh Lo slightly compressed on inner whorls , becoming more compressed ( w h / w t = < 1 2 0 % ) with flat sloping whorl sides and ra ther flat venter (Text-fig. 6); steeply s loping umbilical wall. Ribs (Text-fig. 7) gently prors i radia te th roughout becoming rcct i radia tc near venter; on inner whorls b i furcate with occasional intercalatories and ra the r dense; on middle and outer whorls, t r i furcate with an associated intercalatory or bifurcate with two associated inter-calatories; pr imaries ra ther s t rong, sha rp and regular throughout , generally spl i t t ing above or about the middle of the whor l side; ribs pass unin ter rupted across venter . Occas ional weak constrictions on inner and middle whorls. Aper tu re not seen.



eh (septate to) wh wt uw primaries ventrals BGS G S M 7 6 7 6 (PI. 1, fig, 4) 66 (66) 22 20 27 34 c. 108 BGS G S M 7 6 7 9 (PI. 1, fig. 5) 47 (47) 17 — 20 40 est. 100

Remarks. T h e type specimen of H, balinensis comes from the richly fossiliferous Balin Ooli te of Balin in southern Poland which provided mater ia l for a number of monographs in the last century (Neumayr 1871; Sicmiradzki 1894) and f rom which Teisseyre (1887; 1888; 1889) described Proplamdites. T h e holotypc is incomplete but shows that its body-chamber begins at c. 60 m m diameter (cast f igured by Mangold 1971b, pi. 6 fig. 1). T h e present material is of comparab le propor t ions bu t . a s suming tha t the three specimens have been grouped together correctly, they show a ra ther coarser spacing of the p r imary ribs on the body-chamber (BGS Gcol. Soc. Coll. 3110) than generally accepted for this specics. However , with so little material it is not possible to assess how significant this difference is.

50-54 55-59 60-64 65-69 70-74 75-79 80-84 85-89 90-95 d i a m e t e r (inrri|

T E X T - F I G . Size distr ibution For adul t microconch Homoeoplanulites difficilis ;S. S. Buckman) [29 specimens including Buckman's type],

Homoeoplanulites difficilis (Buckman , 1922) PL l', fig. 6; PL 2, figs 1-6; PL 3, figs 1-3 ; PI. 4, f ig.l ; PL 5, fig. 2

1922 Anaplanulites difficilis. now; Buckman, pi. 229. 1933 Proptanuliles (Anaplanulites) difficilis. Buckman; Spath . p. 856. 1931 Proplamdites difficilis; [Spath] , p III 1 945 Prnpl.anuf.itts difficilis (Buckman); Arkell, p. 341. 1950 Anaplanulites \= Homoeoplanulites] difficilis S. Buckman; Arkell, p. 211. 1971b Anaplanulites [=HomueoplurtulilesJ difficilis Buckman; Mangold, p. 52.

Type specimen. Buckman ' s (1922, pi. 329) holotypc. from the Kellaways Rock of South Gave , Humbers ide is believed to have been kept at Hull M u s e u m (Frank Petch collection) and is presumed to have been destroyed dur ing the Second World W a r ; however, there is no proof of this because the whole of the Hull M u s e u m catalogue was lost at that t ime (P. Bovlan, pers. comm. 1976). In the present case, there is no "exceptional c i rcumstance" that warrants the designation of a ncotype at this t ime (1.C.Z.N., Article 75); there are a b u n d a n t topotypes.

H O M O EG PL AN U1. II ES 19

Material. About 200 specimens , uncrushecl and generally undis ior ted , inc luding specimens that a rc comple te , with a p e r t u r e , or near ly complete , with b o d y - c h a m b e r and uncoil ing; pre-served in i ronshot oolite, usually as in te rna l moulds . Microconchs a n d macroconchs . From Kel laways Rock (Calloviense Zone , E n o d a t u m Subzone) , South Cave and Sou th Newba ld , Humbcr s idc .

Description. Mic roconchs (46 specimens) ; m a x i m u m d iamete r 95 m m , general ly 75 to 80 m m (Text-fig. 8). B o d y - c h a m b e r 220° to 250°. Five whorls visible. Modera te ly evolute ( w h / d = c.. 3 5 - 4 0 % ); whor l section rounded on inner whorls , compressed ( w h / w t = 140-150% ) on middle and outer whorls wi th flat whorl sides (Text-fig. 6); gently sloping and a lmost smooth umbil ical wall; umbi l icus c. 30 m m wide and up to 10 m m deep in adul t . Ribs p rors i rad ia te though secondaries may develop slight ru rs i rad ia te aspect; on inner whorls, ribs ra ther fine and dense , rarely weakening to lirac, b i furca te with occasional s imple ribs and parabolae ; on middle whorls, ribs t r i furca te or b i furca te wi th in tercala tor ies a n d rare s imple ribs, pa r abo l ae a n d rare weak constr ict ions; on ou te r whorls , long s t ra ight s lender pr imar ies which may be s t reng thened a t umbilical edge, spl i t t ing above or abou t the middle of whor l side, where they may weaken, into usually three secondar ies with some intercala tor ies (Text-fig. 7). Ribs pass across venter wi thout in terrupt ion a n d persist to, though m a y weaken towards , ape r tu re which bears l appe t s (up to c. 0-5X15 m m ) (Text-f ig. 9).

40

30

20

primary Er simple ribs

— i i i i i i i i 0 20 30 40 50 60 70 80 90

diametBr(mm) TEXT-KK;. 9. Rib density curves for adult mir rnronrh Hanwmplamdites (tijfialis (Buckman! .

Macroconchs (33 spec imens) : m a x i m u m d iame te r 180 m m , general ly 160 to 180 m m . Body-chamber 310° to 360°. AT least 5 whor ls visible. Modera te ly evolute ( w h / D = 3 0 - 3 5 % ) ; whorl section rounded on inner whorls , compressed ( w h / w t = 120-130%) on middle a n d ou te r whorls with flat whorl sides (Text-fig. 6): gently s loping to modera te ly steeply s loping and a lmost smooth umbilical wall; umbil icus 60 to 70 m m wide, and up to 25 m m deep in adul t . Ribs prors i rad ia te ; on inner whorls, r ibb ing dense with pa rabo l ae (only pr imaries visible); on middle whor ls , ribs t r i furcate or b i fu rca te with intercalator ies , r a re s t reng thened simple ribs or weak constr ic t ions, primaries generally s t ra igh t , long arid slender; on outer whorls , pr imaries m a y be s t reng thened at the umbil ical edge a n d gently prors i radia te . spl i t t ing abou t middle of whorl side where they appear to weaken in to usually three secondaries with some intercalator ies; r ibbing g radua l ly weakens towards b o d y - c h a m b e r which is almost completely smooth except for weak deve lopmen t of blunt p r imar ies which m a y persist par t icular ly a t the umbil ical edge (Text-fig. 7). Close to aper ture , which is s imple, these may be lost and growth lines m a y be visible. Ribs pass across the venter wi thout in t e r rup t ion .

• = start of body chamber

TNN 476 TNN 475

TNN 446 FW TNN 473

F W T N N 503

TNN 500

FW TNN 502


(septate to) wh wt n w pr imar ies ventrals 85 (c. 55) 29 24 35 33 —

78 (c. 50) 26 18 30 33 108 87 (c. 51) 30 22 33 31 e. 90 80 (c. 50) 26 e. 23 31 27 6. 105 72 {c. 45) 25 c. 19 27 33 c. 95 66 (c. 42) 21 18 28 33 est. 85 54 (c. 49) 18 est. 18 21 c. 45 —

02 ( - ) 35 25 30 25 —

c. 75 ( - ) 26 c. 18 29 180+ 180 (c. 95) c. 60 (40) 72 24 160 (c. 95) c. 55 46 64 24 c. 120 ( ) c. 43 c. 30 c. 40 26 est. 120 180 {c. 100) c. 55 45 70 21

Data for figured specimens, t = t y p e of H. spirorbis.

I lo lo tvpc (PI. 1, fig. 6) F W T N N 4 7 5 (PI. 2, fig. 1) F W T N N 4 7 3 (PI. 2, fig. 2) BGS Z k l 4 3 9 (PI. 2. fig. 3) F W T N N 5 2 7 (PI. 2. fig. 4) FW T N N 5 2 8 (PI. 2. fig. 5) F W T N N 5 1 0 (PI. 2. fig. 6) F W T N N 5 2 1 (PI. 3. fig. 1) BGS G S M 117460 (PI. 3. fig. 2) F W T N N 5 0 8 (PI. 3, fig. 3) F W T N N 5 0 6 (PL 4, fig. 1) t (PI. 5, fig. 1) F W T N N 5 1 6 (PL 5, fig. 2)

Remarks. Buckman ' s (1922, pi. 229) original figure of this spccics showed a microconch with about five-eighths of a whorl of incomplete body-chamber , with an actual d iamerer o f 8 5 mm, and an est imated m a x i m u m d iamete r o f 9 5 m m . T h e "perfect topotypc" (SM J 4 7365) of Arkell (1958, p. 211) is smaller (d=c . 75 m m ) , but compared with other topotypes seems to be the more normal size (Text-fig. 8). Arkell (1958b, p. 211) believed that , except for the smaller size o f / / , dijjicilis, there was little to dis t inguish it from H. homoeomorphus.

Spath (1933) and Arkell (1945) compared the South Cave perisphinct ids with P. scldosseri and P. comptoni figured by Krenkcl (1915) from Li thuania . However, there is no s t ra t igraphical information or descript ion to accompany Krcnkel ' s two figures and their s t ra t igraphical position is doubt fu l . Mos t of the per isphinct ids which Krenkel figured are t rue Binatisphinctes of the Ath le ta Zone (i.e. a much younger horizon than that of H. dijjicilis) (Arkell 1956b, p. 473). Spath (1933) and Arkell (1945) also thought that the difficilis f auna included "undescr ibed species allied to P.ferisphictes] spirorbis N e u m a y r " . T h e type of the lat ter (Neumayr 1870, p. 148, pi. 7, fig. 2; B S M ASI838; cast figured herein PL 5 fig. 1), from the Marrocephalenschich ten of southern G e r m a n y , is an incomplete macroconch which compares closely with some of the dijjicilis macroconchs, and apparen t ly comes from a similar s t ra t igraphical level. Mangold (1971b, p. 240) believed tha t spirorbis belonged in Indosphinct.es} bu t it has usually been assigned to Proplanulites in compar i son with Ncumavr ' s (1871, pi. 11 fig. 4) figure. This smaller specimen, cf. spirorbis of Neumayr , f rom the Lower Cal lovian Balin Ooli te of Poland, is a moderately involute compressed form wi th thickened short pr imar ies , and is correctly placed in Proplanulites. Neumayr ' s two figured speci-mens arc not jus t microconch and macroconch of the same species, for the macroconchs of Proplanulites arc found in Crassiplanulit.es which also has character is t ic thickened pr imary ribs, in contrast to those shown on Neumayr ' s type specimen of spirorbis, Krenkel 's (1915, p. 244, pi. 23, figs 4, 9, 10) two inner whorl examples of P. spirorbis f rom Li thuania (probably from a younger s t ra t igraphical horizon) and Corroy ' s (1932, p. 155) record of incomple te examples from the Paris Basin, do not throw fu r the r light on the species; however, both these authors assigned it to Proplanulites and included N e u m a y r ' s second example in their synonymies. Specimens from C u t c h (Waagen , 1875, pi. 41 (wrongly numbered 40 on plate), figs 1, 2) were excluded from P. spirorbis by Siemiradzki (1899, p. 323) and assigned to a new species P. indicus. Spath (1931, p. 333) confined the latter to VVaagen's fig. 1 and used the second figure as the type of a new species / . peregrinus) he included both species in Indosphinctes.

The species spirorbis is thus clearly not well known and needs revision; Neumavr ' s type shows only one whorl . Therefore , until the G e r m a n and Polish faunas are described and figured more-fully, H. dijjicilis is re ta ined, a l though ult imately it m a y prove to be a synonym of spirorbis.

H. dijjicilis shows some variability . Al though usually preserved only as internal moulds, some

INDOSPH1 NOTES 21

variat ion in the s t rength of r ibb ing on the body-chambers is apparen t . PI. 2, fig. 6 (FW T N N 5 1 0 ) is one of a small n u m b e r of less compressed individuals which have a more rect i radia te rib style. T h e microconch figured in PI. 2, fig. 5 ( F W T N N 5 2 8 ) is slightly more cvolutc with ra the r delicate r ibbing on the inner and middle whorls , and the secondaries have a marked rurs i radia te aspect . Rurs i rad ia te r ibbing is character is t ic of the microconch g roup of Elatmit.es Shevyrev. I960, which, with the macroconch g roup of Indosphinctes Spath , 1930, is believed to have evolved f rom Homoeoplanulites at the beginning of the Cal lovian (Mangold 1971b). As probably the youngest Homoeoplanulites, H. difficilis may show close affinities with Indosphinctes (e.g. the microconch nikitinoensis Sasonov, i % 5 , Midd le Cal lovian of the Russian Pla t form). However, microconch Indosphinctes have a bolder r ib style, par t icular ly on their inner-middle whorls. Also, Mango ld (1971b, p. 52) believed that their lappets were long compared with those of Homoeoplanulites and that Indosphinctes had a more marked development of parabol ic r ibbing par t icular ly at the end of the ph ragmocone and beginning of the body-chamber .

C o m p a r e d with the other microconch Homoeoplanulites, H. difficilis is most like / / . balmensis but the latter is more cvolutc. T h e macroconchs m a y also show similarities with certain Indosphinctes because the o r n a m e n t on the outer whorls is usually lost, except for the pr imaries . However , in Indosphinctes the p r imary ribs arc typically thickened at the umbilical edge at all stages and are shorter than those o f / / , difficilis. T h e II. difficilis macroconchs are much smaller than the other described macroconch Home.oplanuliles such as H. subbakeriae and H. funalus (Oppe l ) .

Genus INDOSPHINCTES Spa th , 1930

(Synonym: Elatmites Shevyrev, I960 (type species Perisphinct.es submutatus Nikitin) | m | )

Type species. Ammonites calvus J . de C. Sowcrbv, subsequent ly designated by Roman (1938. p. 275); lcctotypc Spa th 1931, pi. 52, fig. 1 ( B M C52288) [M] .

Diagnosis. Modera te ly cvolutc to cvolutc forms with compressed to rounded whorl section (Text-fig. 6); lappcted microconchs strongly ribbed throughout , usually with marked rurs i radia te aspect to secondaries; macroconchs show weakening of secondaries and development of short , b lunt pr imar ies which fade towards aper tu re (Text-fig. 7). I n n e r / m i d d l e whorls may have weak constrictions; parabol ic r i b s /nodes (par t icular ly near venter) not u n c o m m o n . External su tu re has first lateral saddle with well marked accessory lobe, a trifid. slightly asymmetr ica l lateral lobe slightly deeper t h a n ventra l lobe, and a suspensive lobe with some well defined auxiliaries.

Remarks. Indosphinctes was proposed by Spa th (1930, p. 36) for a group of per isphinct ids f rom the Callovian o f C u t c h , Ind ia . H e included the species calvus ). d e C . Sower by, indicus Sierniradzki, patina. Neumayr , abichi N e u m a y r &. I Ihlig, c.hoffdti Parona & Bonarelli and subpatina Petitclerc, and later (1931. p. 329) six new species, errans, nalmaris. patiniformis, peregrinus. mslicus and urbanus. H e noted par t icular ly the following characterist ics: a tendency to i rregular r ibbing on the inner whorls with inconspicuous p r imary ribs, often bundled or confined to a blunt node at the umbilical edge; a subsequen t loss of r ibbing with growth and its re turn, in modified form, on the body-chamber ; projection of the ribs on the venter , at least in later growth stages; and a highly-complex su ture (Spath 1931, p. 32!)). Spa th clearly considered the style and deve lopment of the r ibbing to be an impor t an t generic charac te r which he referred to as '"the Indosphinctes type of cos ta t ion" (1931, p. 331). M a n y of Spa th ' s described or figured specimens were incomplete , with d iameters rang ing generally f rom 115 to 150 m m ; when complete, specimens had d iamete r s of the order of 250 m m . F rom this descript ion, it seems certain that , as originally defined, the genus accommoda ted macroconch forms.

Indosphinctes was discussed by Mango ld (1971b, p. 84) who included in the genus microconch forms previously assigned to Elalmites. H e confined the macroconchs , including besavoensis Collig-non, brenoni Col l ignon, cesaredensis Mango ld , lancharesi Elmi & Mango ld , linaresi Elmi & Mangold . luceyensis Mango ld , pseudopatina Pa rona & Bonarelli and roberti Petitclerc, to Indosphinctes s.s., and used Elalmites to a c c o m m o d a t e the microconchs.


In proposing Elatmites, Shcvvrev (1960) discussed only ontogenetic development , part icular ly of the suture , of the type species {Perisphinctes submutatus Nikitin 1881b; lectotype (Niki t in 's pi. 1. fig. 4) designated and rcfigured by Sasonov (1965, pi . 2, fig. 3) [ C N I G R Nikitin coll. 6 /1344] ) . A fuller diagnosis given by Sasonov (1965, p. 10) and subsequent ly used by Mango ld (1971b, p. 110) is as follows: compressed forms with wide umbil icus and gently sloping umbil ical wall; last whorl covering 2 5 - 3 0 % of preceding whorl : whorl section circular in young, becoming oval in adul t : size between 65 and 80 m m ; o r n a m e n t of p r imary and secondary ribs, the lat ter rurs i radi-atc on phragmocone ; pr imaries well developed and prors i radia te on body-chamber , spl i t t ing a t middle of whorl side into two or three secondaries, somet imes with intercalatories; mouth border with lappets . L'sing the second and new species nikitinoensis assigned to Elatmites by Sasonov (1965), Mango ld (1971b, p . 110) pointed out addi t ional characters , part icular ly the presence of simple ribs and parabol ic tubcxcles (the lat ter being especially well developed at the end of the phragmocone and the beginning of the body-chamber ) ; he explained the lack of pa rabo lae on Sasonov's figure of the holotypc oCE. submutatus as being probably clue to preservat ion. T h e r e may also be weakening of the secondaries or a sulcus on the venter. As well as the two Russ ian species, Mangold (1971b) included in Elatmites the species arcicosta Petitclerc non VVaagen, calloviensis Loczy, cheyensis Peti tclerc, atrvicosta Oppel , gr ados us Siemiradzki. lobatus Buckman, prahecquensis Mangold , revili Mango ld , steinmanni Parona & Bonarell i and subrjasanensis Petitclerc, and Car iou (1974a) added revolt Petitclerc 1918.

Spath (1931, p. 325) suggested tha t Indosphinctes was connected with the Ba thonian genera Proce.ril.es | m a c r o c o n c h s | and Siemiradzkia [microconchs]. These have since been taken as the pa ren t groups of Hom.aeoplanuli.tes from which genus Indosphinctes is now thought to have evolved at the beginning of the Cal lovian (Mangold 1971b, p. 85). T h e dividing line between Homoeoplamdites and Indosphinctes is a rb i t ra ry and likely to be d r a w n differently by different au thors . Both genera persist into the Midd le Cal lovian.

Indosphinctes lobatus (Buckman . 1922) PI. 6, figs 1 -3

1922 Proplanulites lobatus nov.; Buckman, pi. 330. 1958 Proplanulites lobatus Buckman; Arkell, p. 235. 1971b Indosphinctes {m. Elatmites) lohatns (Burkman) ; Mangold , p. 115, tig. 83. 1975 Choffatia sp.; Cave & Cox : p. 40. '

Holotype. BGS C S M 7 6 8 2 , PI. 6, fig. 3, figured Buckman 1922, pi. 330. Microconch from "Oxford Clay" |Ke l l aways Clay] , Calloviense Zone, Kocnigi Subzonc, R a m p i s h a m , Dorset . (Blake (1905, p. 7) described this locality as "a deser ted quar ry in Oxford Clay but any C o r n b r a s h seen there would he avai lable" . Wilson el al. (1.958. p. 112) gave the horizon as Kellaways Clay) . Uncrushed specimen in cements tone with pink shell mater ia l partially preserved.

Material.. T w o uncrushed macroconchs with white chalky body-chambers and pink shell material part ial ly preserved. From Kel laways Clay (Calloviense Zone, Kocnigi Subzone) of Sutton Benger ( M 4 Motorway cut t ing) , Wil tshire and Siddington, Gloucestershire.

Description. O n e microconch: m a x i m u m d iamete r 65 mm. Body chamber 220°. Six whorls visible. Modera te ly evolute ( w h / d = 3 2 % ) ; at least outer whorl section moderately compressed ( w h / w t = r . 125%) (Text-fig. 6); modera te ly steeply sloping and almost smooth umbil ical wall; umbilicus 27 m m wide and 5 m m deep at 65 mm diameter . Ribs (Text-fig. 7) gently prors i rad ia te with slight rurs i rad ia te aspect, near venter , less so on body-chamber . Primaries s t rong and r a the r widely spaced th roughout (Text-fig. 10) spli t t ing about the middle of whorl side (nor seen on i n n e r / m i d d l e whorls) . O u t e r whorl has bifurcate ribs each with an associated intercalarory, or irregular t r i furcate r ibbing. Occas ional weak constrict ions on inner whorls. Ribs generally pass unin ter rupted across venter , though weakened in par t , arid continue to aper ture which bears lappets (up to c. 4X 14 m m ) .

T w o macroconchs : m a x i m u m d iamete r c. 150 mm. Body-chamber at least 220°. At least five *

(probably six) whorls visible. Modera te ly evolute to evolute ( w h / d = 3 0 - 3 5 % ) ; whorl section

IX DOS PI 11NCTES

(Text-fig. 6) rounded ( w h / w t = 1 0 0 115%) on inner whorls, becoming more compressed ( w h / w t = 115-130%) on middle and outer whorls, with flat whorl sides converging on fairly broad and ra ther flat venrer; modera te ly steeply sloping and almost smooth umbilical wall; umbil icus c. 65 m m wide and c. 15 m m deep at 135 m m diameter . Ribs (Text-fig. 7) gently prors i radia te th roughout , somet imes becoming rect i radia te near venter; on middle and outer whorls , t r i furcatc with an associated in tercala tory , or bifurcate with two associated intercalatories; pr imar ies r a the r strong, sha rp and regular th roughout generally splitt ing above or about the. middle of the whorl side where they weaken: on body-chamber , pr imaries shorter and more b lunt ; ribs pass across venter without in te r rupt ion . Shallow, weak constr ict ions (maybe two per whorl) on inner and middle whorls. Aper tu re not seen.

40

30

20

primary & simple ribs OUM J 309181ml

OUM J 30912 Iml OUM J 309131ml

K, BGSGSM /6821m

OUM J 30911

Im] — • K-

OUM J 30917 [ml

BGS GSM 7680

1M1

- start of horiy chnmbcr

10 20 30 40 50 60 70 80 90 diameter (mm)

T E X T - F I G . 10. Rib-density curVR.S for /ndnsphi.Mlt'i lobatus (Buckman) [broken line] and I. patina (Neumayr) [solid line].


BGS G S M 7 6 8 0 (PL 6, fig. I) BGS Zi 1639 (PI. 6, fi^. 2) BGS G S M 7 6 8 2 (PI. 6, fig. 3)

d (septa te to) 136 (c. 100)

t. 115 (c. 100) 65 {c. 40)

wh wt uw pr imaries 42 32 63 30

est. 40 est. 35 c. 55 21 17 27 31

128

est. 85

Remarks. Mango ld (1971b, p. 115) considered I. lobatus to be monotypic . He thought that the holotypc possessed all the charac te rs ofElatmites , in par t icu lar the rurs i radia te ribs on the end of the phragmocone , and that it was manifest ly not a Proplanulites to which genus Buckman (1921) originally assigned it. Proplanulites was one of the genera for which Buckman gave some notes for each species to a c c o m p a n y the type figures, but pe rhaps significantly, P. lobatus was the one exception. N o fur ther microconch specimen has since been found, but the two addi t ional specimens descr ibed here (PI. 6, figs I, 2) arc possibly the macroconchs . It is not certain how far from complete these m a t u r e specimens are (PI. 6, fig. 1 has at least 130° body-chamber missing), but they do not show any signs of loss of secondary r ibbing on the bodv-chambcr which is normal for Indosphinctes; however , they arc here assigned to the genus on the basis of the microconch. T h e microconch lobatus appea r s close to Sasonov 's (1965) probably slightly younger E. nikitinoensis from Yela t 'ma , U S S R (Zone of Cadoceras milaschevici and Kosmoceras jason). However , I. lobatus has more widely spaced pr imaries , at least on the last whorl of ' the phragmocone , and a p p e a r s to be more compressed. Mango ld (1971b, p. 114) believed that his species I. revili, f rom the Koenigi-Pat ina Subzoncs of M o n t du C h a t in the F rench J u r a , was very close to I. lobatus, but the former species was more involute and had a smaller whorl height, with less " i m p o r t a n t " spacing of the primaries on the body-chamber .

24 ENGLISH CALLOVIAN PKR1SPHINCTI1) AMMONI TES

I n d o s p h i n c t e s p a t i n a ( N e u m a y r , 1870) PI. 6, fig. 4; PI. 7, fig. 1; Pi. 8, figs 1-4 1870 PerisphincUspatina now sp.; Neumayr , j>. 140, pi. 8 fig. I.

turn 1871 PerisphincUs patina Ncumavr : Neumayr , p. 41, pi. 13 figs 2a, b [ = / . pseudopatina (Parona & Bonarelli)! . 1097 PerisphincUs patina Xcuni.; Fa rona & Bonarelli, p. 174.

? I 8 9 7 PerisphincUs submutatus N i k . , P a r o n a & B o n a r e l l i , p p . 181, 182; pi. 10 Iig. 2. 1898 PerisphincUs Comptoni Pratt ; Siemiradzki. p. 135 \Jpars\ 1899 Perispkinctes Patina Neum.; Siemiradzki, p. 297. 1910 PerisphincUspatina Neum.; von See. p. 698. 191 1 PerisphincUs patina X n i m . ; T i l l , p. 35 , pi. 4 (8) f ig . 11 [ s u t u r e o n l y ] .

?1913 PerisphincUs patina Neum.; Lowe. p. 154. 1915 PerisphincUs patina Neum. ; Loczy, p. 409. 1915 PerisphijuUs puiina .Ncuinayi', Pelitclcn;, p. 73. 1918 PerisphincUs arcicosta Waagen , sp.; Petitclerc, p. 23, pi. 18 (4), fig. 1; pi. 19 (5), fig. 14. 1930 Indosphinctes patina (Neumayr) ; Spath, p. 36.

?1931 Indosphinctespatina (Neumayr) ; Spa th , p. 338, pi. 102, figs 3a, h. 1932 PerisphincUs patina Neumayr ; f lorroy. p. 137, pi. 12. fie;. 3.

?1935 Choffatia patina Neum; Model & ICuhn. p. 473. I9i>b Grossouvria cf. comptoni (Pi all}; Cal lomon, p. 233. 1955 i Binatisphinctes cf. Jluciuosus ( Pratt}; Cal lomon. p. 233.

? 1961 PerisphincUs (Indosphinctes) gr .patina (Neumayr) ; Ruget-Perrot, pp. 38, 58, 101, 153. ? 1967 Indosphinctes patina; Elmi, p. 460.

1968 Choffatia (Elalmites) suhmutata Nikitin; Callomon, p. 286. 1968 Choffatia cf. patina (Neumayr) ; Cal lomon. p. 286. 1971b Indosphinctes (M Indosphinctes) patina (Neumayr) ; Mangold , p. 100, fig. 74; pi. 13, fig. 2 [XU'5]. 1971b Indosphinctes (m Etat mites) pruheiyuensis uuv. num.; Mangold, p. 121, tigs 91, 94; pi. 12, lig. 3. 1971 ChoJJutiu (Etuimiles) cf. suhmutata (Nikitin); Callomon & Cope, p. 168.

Holotypc by monotvpy; G B V 1870 /04 /1 , PI. lig. 1; PI. 7, lig. 1 (original of N e u m a y r 1870, pi. 8, lig. I which was reversed) , f rom Macrocepha len-Kalk (Lower Gal lovian), Brielral. n e a r G o s a u . Sa lzkammergut , Austr ia . Macroconch . All previous au thors have referred to Neumayr1?; original and only figure as " t y p e " lor the species.

Material. Nine crushed and mainly incomplete, pyritized microconchs and one macroconch uncrushed body-chamber in cemenrstone.

Description. Microconchs: m a x i m u m d iamete r 90 m m . Body chamber c. 180°. Six whorls visible. Modera te ly evolute ( w h / d = 1 5 0 - 4 0 % ) ; whorl section rounded at least on middle whorls; umbilicus c. 25-30 trim at 50—b 5 m m diameter . Ribs (Text-fig. 7) gently prors i radia te or rec-t i radiate with marked rurs i rad ia te aspect near venter. Ribs on inner and middle whorls s t rong, blunt but not swollen, even, moderate ly dense (Text-fig. 10) with long pr imaries spli t t ing into two ra ther high on whorl side, with occasional simple ribs and rare intercalatories; ou ter whorls develop t r i furcate , or b i furca te with an associated intercalatory, r ibbing which remains even and blunt to ape r tu re which bears lappets . Occas ional weak constrictions. Ribs apparent ly , though not certainly, pass un in te r rup ted across venter.

Macroconch : m a x i m u m diameter 265 m m . Body-chamber 210°. Evolute ( w h / d = 2 4 % ) ; whorl section compressed on outer whor l ( w h / w t = c . 100%) (Text-fig. G); r ibbing (Text-fig. 7) on body-chamber reduced to short , blunt pr imaries developed at umbilical edge, evenly spread c. 10 m m apar t , spli t t ing low on whor l side in to secondaries; all r ibbing fades as s imple a p e r t u r e approached .

Data for figured specimens. + = type of II. patina; *on body chamber .

d (septate to) wh wt O U M J30911 (PI. 8, fig. 2) 89 (c. 60) 28 — O U M J30912 (PI. 8, fig. 4) 67 (—) c. 21 — O U M J30918 (PL 8, fig. 3) est. 55 {—) est. 16 BGS Z t l 6 7 5 (PI. 8, fig. 1) 265 (—) 65 c. 40 t (PI. 6, fig. 4 & PI. 7, fig. 1) 210 (c. 145) c. 55 est. 36

uw primaries vent ra ls 35 c. 32 c. 84

31 c. 40 est. 90 29 —

c. 130 20* —

110 ± 3 2 —

INDOSPHINCTES 25

Remarks. I. patina was amongs t the species listed by Spa th (1030) in his proposal of Indosphinctes. It has been widely recorded and the macroconch form is one of the best known Indosphinctes in Europe. In France , it is used as a subzonal index in the Lower Cal lovian Gracilis Zone .

Specimens of macroconch I. patina were figured by Spath (1031, pi. 102, figs 3a, b), Corroy (1932, pi. 12, fig. 3) and Mango ld (1971b, pi. 13, fig. 2). Spa th ' s figure showed an i m m a t u r e example and his descript ion was based on a specimen that is much larger than the lcetotypc. T h e ass ignment of these specimens to I. patina is open ro doubt a view shared by Mangold (1971 b, p. 101). Corroy ' s figure is one of the best i l lustrat ions. Parona & Konarelli (1897, p. 177) excluded N e u m a y r s second publ ished figure o f / , patina because ir showed more involute coiling and gently curved ribs compared with the s t ra ight ribs typical of the species. T h e y assigned it to a new species I. pseudopatina, and this specific distinction has been accepted by all subsequen t au thors .

N o microconch species has previously been formally linked with the macroconch patina but , following M a n g o l d (1971b), the former are believed to he found in the suhmutatus Nikitin, 1881 group. Mangold did not a t t e m p t to pair up individual macroconch and microconch species but Cal lomon (1968, p. 286), having used both species for microconchs and macroconchs f rom the basal shell bed of the J a s o n Subzone at Calvert , believed tha t suhmutatus (Nikit in)[m]—patina (Neumayr ) [M] m a d e a plausible pair (Cal lomon, pers. comm. 1975).

The early Russ ian au thor s , including Nikitin (1881) assigned the macroconchs associated with suhmutatus xofunatus Oppe l , 1857 and mutatus T rau t scho ld , 1862. These records o f funa tus arc now included in Choffatiapseudofunata (Teisseyre, 1889) sensu Mangold 1971 b. p. 154. T rau t scho ld (1862, p. 211, pi. 6, figs 1,2) figured two specimens of mutatus, the second and smaller of which was included by Nikitin (1881b) in his original proposal of submutalus. T rau t scho ld ' s species was thereby restricted to his first figure and this usage has been followed by subsequent au thors (e.g. Lahusen 1883; Nikitin 1885; Sicmiradzki 1899: Stoll 1934; K u h n 1939). It is clearly a macroconch form (d—c. 240 m m ) which is dist inct f rom I. patina, par t icular ly its more involute coiling. It is reasonable to suppose that suhmutatus (Nikitin)[m]—mutatus (Trautschold) [M] are a d imorph ic pair and , a s suming the microconchs of mutatus and patina a re distinguish*!) e, the microconch of the latter must he sought elsewhere.

Most earlier au thor s recorded or figured only macroconch patina, and wi thout a d e q u a t e figured examples a n d / o r s t ra t igraphical control, it is impossible to be certain if associated microconchs are present and , if they arc, u n d e r what name . They may have been included in suhmutatus (e.g. Parona & Konarelli 1897 recorded patina and submutalus f rom the s ame locality in the J ura) , or they may have been confused with those of the younger Binatisphincles, par t icular ly B. comptoni (e.g. Siemiradzki 1898; Cal lomon 1955). In the key to his figure of submulatus, Petitclerc (1915, pi. 5 fig. 3) explained tha t compared with B. comptoni. the species had a thicker whorl with a more oval section, more involute coiling, s t ronger r ibbing and more obvious parabol ic ribs in the ventral area (these are emphasized in Parona & Bonare l l r s (1897) figure of submulatus); B. comptoni also has much finer and denser r ibb ing ai least on the inner-middle whorls. Cor roy (1932) may also have included the microconchs in "Perisphmct.es Comptoni Pratt s p . ' \ for he recorded this species f rom both Lower and Uppe r Cal lovian and from a Lower Callovian locality with I. patina. Von See's (1910) descr ipt ion of ano the r microconch, Binatisphincles mosquensis, f rom the Lower Callovian, may also be a rnisidentilication which could be related to macroconch patina. A possible microconch to a c c o m p a n y Loezy's (1915) patina f rom Villany is his "Parkinsonia callovien-sis no v. s p . " (1915, p. 379, pi. 4, fig. 11; pi. ft. fig. 11: text-fig. 88) which Mango ld (1971b, p. 127) believed belonged to the submutalus g roup .

Mango ld ' s (1971 b, p. 121) speciesprahecquensis f rom Deux-Sevres, figured by Peti tclerc (1915, pi. 18 (4) fig. 1) as Perisphinctes arcicosta VVaagen, is considered here to be the microconch pa r tne r of /. patina. It occurs at the same s t ra t igraphical level in France , and the spec imen of Mango ld ' s addi t ional figure (Mango ld 1971b, pi. 12. fig. 3) from the French J u r a compares well with the English microconch mater ia l figured herein (PL 8, figs 2—4). Mangold (1971b, p. 122) believed


prahecquensis to be close to suhmutatus, bu t dis t inguished from this and all o ther Elatmites by its large size (up to 100 m m d iamete r ) .

Distribution. Oxfo rd Clay (Jason Zone) of Calver t , Buckinghamshire ; Witney. Oxfordsh i re and W a r l i n g h a m Borehole, Surrey (Cal lomon & Cope 197L). Lower Callovian of France (Paris Basin, Poitou, Deux-Sevrcs, and Ardeche (Elmi 1967); Lower-Middle Callovian of G e r m a n y (Macrocepha lensch ich ten of Saxony (Lowe 1913; von Sec 1910); Calloviense Zone -Midd le Callovian o f F r a n c o n i a (Model <Sc K u h n 1935) and the Jura (Parona & Bonarelli 1897): Lower Callovian of the Alps ( N e u m a y r 1870), H u n g a r y (Viilany (Till 1911; Loczy 1915)), Poland (Siemiradzki 1898-9) and Portugal (Rugct -Pcrro t 1961); PCutch, India ('" diastema tus Zone*' Spa th 1931).

Genus CHOFFATIA Siemiradzki, 1898

(.Sy«0«yw.f. Grossouvria Siemiradzki , 1898 (rype species Ammonites sulciferus Oppe l , 1857) [m] , Poculisphinctes Buckman , 1920 (type species Ammonitespoculum Leckenby, 1859),

Trinisphinctes Buckman , 1922 (type species T. trim Buckman. 1922), Subgwssoiwria Spa th , 1924 (type specics Perisphinct.es aherrans Waagen , 1875) [M]) .

Type species. Perisphinctescobra W a a g e n , 1875, by subsequent designation of Buckman (1920, p. 29); I lo lotype (GSI T y p e no. 2056), cast figured herein PL 9 fig. 1 | M | (BGS GSM11744a ) .

Diagnosis. Modera te ly evolute. forms with modera te ly compressed, rounded or depressed whorl section (Text-fig. 11); strongly r ibbed , often with well developed constrictions and sometimes parabol ic ribs and nodes (Text-fig. 12); r ibbing usually rurs i radia te near venter, par t icular ly in microconchs , becoming coarse on later whorls of macroconch, where pr imar ies may be d is tant and blunt ; lappetcd microconchs. External su ture has first lateral saddle with a well marked accessory lobe, a trifid slightly asymmetr ical lateral lobe usually abou t as deep as the ventral lobe, and a more or less re t racted suspensive lobe with some well defined auxiliaries.

Remarks. Choffatia was described originally by Siemiradzki (1898. p. 79) as one of six subgene ra within Perisphinctes. Lie gave the diagnost ic features as a smooth protoconch, compressed form without any trace of la teral lappets , s imple mouth border (like Perisphinctes mar tins i d O r b i g n y ) , lateral sculpture similar to the martiusi g roup and simple su ture line. Siemiradzki 's figure (1898, p. 327, Fig. 83) of the ape r tu re of P. martiusi shows a typical macroconch form with a gently sinuous mouth-border . T h e g roup was thought to link rhe per isphinct ids with Proplanulites and included a single lineage, tha t of Perisphinctes cobra. Waagen . Siemiradzki (1898, p. 334) described this species as having a thick, disc-shaped shell form (d iameter 125 m m ) with a wide, deep, funnel -shaped umbil icus and thick rounded whorl with coarse widely spaced ribs; these d i sappeared before reaching the venter which remained almost entirely smooth. T h e inner whorls were finely r ibbed , the ribs usually bi furcate and constr ict ions were rare . Siemiradzki 's definition of Choffatia thus accommodated macroconch forms.

Choffatia. and its type species have been discussed by Spa th (1931, p. 344) and Arkell (1958, p. 211). The larrer concluded tha t the typespec ies belonged to a Lower and Middle Cal lovian g roup of large constricted forms with gradua l ly modifying ribs and smoothing venter , and no lappet . Following Spath (1931, p. 325), he also included Ilonioeoplanulites Buckman as a microconch subgenus which like Choffatia s.s. ranged f rom the U p p e r Ba thon ian into the Cal lovian (Arkell 1957a, p. 317; 1958, p. 211). However , Mango ld (1971b) removed Ilomoeoplanulites as a separa te genus. Elmi (1962) included Spa t l r s (1924) Subgrossouuria (based on Perisphinctes aberrans W a a g e n ) as an addi t ional macroconch subgenus , to a c c o m m o d a t e forms with depressed or circular adul t whorls and an o r n a m e n t of coarse (massive) and dis tant p r imary ribs suddenly replacing closely spaced fine r ibbing on the inner whorls (Spath 1931, p. 327). Both Elmi (1962) and M a n g o l d (1971b) included the Bathonian based genus Loboplanulites Buckman as a jun io r synonym of Subgwssouvria. T h e associated microconchs of both Choffatia s.s. and Subgrossouuria have been included in Grossouvria (Cal lomon 1963, p. 37; Mango ld 1971b) which is discussed below.

CHOFFATIA 27

T h e r e has been confusion about The interpreta t ion of the type species of G rosso uvria and its type specimen, following B u c k m a n ' s (1920, p. 28) designat ion of Perisphinctes subtilis N e u m a y r , 1870 with N e u m a y r ' s (18 /1) pi. 14fig. 3 a s lectorype (viz'. Spa th 1931, p. 326; Arkell 1939, p. 157; Arkell 1958, p. 214; M a n g o l d 1971b. p. 163). T h e s e au thors have shown that Perisphinctes subtilis N e u m a y r , 1870 is a junior objective synonym of Ammonites sulciferus Oppe l , 1857, which is therefore taken now as the type species of Grossouvria ( I .C .Z .N. Article 67(e)) . T h e figure designated by Buckman as lectotype ( N e u m a y r 1871. pi. 14 fig. 3) and cited by Siemiradzki (1898) when he proposed the genus (see below), is considered to be dist inct from Oppe l ' s A. sulciferus. It is sep ta te to the end and therefore incomplete (Spath 1931, p. 326); Spa th considered tha t it might be the nucleus of a Choffatia species. He advocated that Grossouvria should be used for forms for which Oppe l (1857, nun 1862) used the n a m e sulciferus and for which N e u m a y r (1870, non 1871) used subtilis. Th i s view was suppor ted by Arkell (1958, p. 214) who des ignated Q u e n s t c d t ' s (1849, pi. 13, fig. I) Ammonites convolutus orriati as type specimen (GPI u n n u m b e r e d ; cast figured

T E X T - F I G . 1 I. Whorl sections o f C h o f f a h c i spccics: (a! IJGS GSM49454 a i 193 mm: (b) F W T N N 5 3 7 at 90 m m ; (c.) FW T N X 3 2 2 at 57 mm: (d) GSI 20:>(> at 127 mm; (c) FW T N N 5 0 7 at 91 mm; (f) FW T N N 4 6 0 at 58 m m ; (g) O U M J 1 2 0 3 at 60 m m ; (h) O U M J 17870 at 19 mm: fi) O U M J2588 at 30 mm; (j) GPI (httlotypr) ar Til mm: fk) O U M J2255'J at 87 nun: ii) S \ 1 J3298 at 60 mm: (m) BGS G S M 6509 at no mm.

herein, PI. 12, fig. 1). Mangold (1971b) also adopted this definition. Siemiradzki (1898) in proposing Grossouvria listed Perisphinctes sulciferus as character iz ing on of its lineages. Its use as type species docs not conflict therefore with the original definition.

Siemiradzki (1898, p. 76) diagnosed Grossouvria as a perisphinctid of small to med ium size, mostly not exceeding 100 m m d iamete r ; mou th -borde r with well-developed, more or less con-stricted, sword-like or tongue- or spoon-shaped lappets; s t rong parabolae recognizable, when the state of shell preservat ion allows, right up to the mou th border ; b i furcate r ibbing in the young, in older stages f requent ly three or more branches ; r ibbing extending to the end of the shell, which is never smooth; pro toconch smooth ; funnel -shaped umbil icus with rounded , strongly depressed whorl section; from the beginning of normal r ibbing, firstly some occasional large folds, then hairline, forward sloping, b i furcate r ibbing, with a somewhat backwards swing in the ventral region: normal r ibbing occurs first at about 3 m m total d iameter ; su tu re line s imple with few ramificat ions umbil ical lobe extending only a little downwards , the auxiliary lobes weakly

ti su In ifera sulci (era


inner whorls

middle whorls

outer w h o r l s — body chamber

Chuff a tin cardor i m

wcll-deveh wide con

:>pcd. rather strictions

Chuff a tin cardor i m

wcll-deveh wide con

:>pcd. rather strictions

1

llml

Choffatia recuperoi

moderat devel

constr

ely we l l -oped ictions

f ( [ f f r Choffatia

recuperoi moderat

devel constr

ely we l l -oped ictions WW**

Choffatia sulcifer a weak

moderatel constr

yW to y strong ictions

|FMI

1 Choffatia sulcifer a weak

moderatel constr

yW to y strong ictions

parabolic ribs & nodes m L m Choffatia aff.

iahuseni

c

0T Choffatia aff. iahuseni

c shallow

onstriction

I

s L i

Choffatia pocutum HJ3

c

m nodes,

flared ribs and weak

onstriction!

«m/rm ijL^r i /„/ Choffatia pocutum HJ3

c

m nodes,

flared ribs and weak

onstriction! mllr iMJr* Choffatia trina 112? f W

moderate shallow cc

ily strono, i >nstrictions llml

T K X T - F I G . 12. Ribbing of Choffatia species.

developed, the second lateral lobe a lways present a l though twice as small as the first, easily dis t inguishable f rom the auxil iary lobe by its s t raight position. Siemiradzki recognized several lineages within four different morphological types ( "Muta t ions rc ihc" ) characterized respectively by Perisphinctes subtilis N e u m a y r . Perisphinctes alligatus Lcckcnby. Perisphinctes aurigerns Oppe l and Perisphinctes euryptchus N e u m a y r . Grossouvria as defined by Siemiradzki thus accommodated micro-conch forms.

Choffatia is used in preference to Grossouvria in accord with the recommendat ion of the I .C .Z .N. Article 24A. It includes the largest n u m b e r (over 80) of described species of any Callovian per isphinet id , having been recorded f rom nearly all areas of ammoni te -bear ing Cal lo-vian s t ra ta (worldwide except for some of the most Boreal areas) .

CHOFFATIA 31

Choffatia is thought to have its origins in the Bathonian . Spath (1931, p. 325) and Arkell (1958, p. 211) both suggested, though not explicitly, that Choffatia [M]-Cr0W0tf»r/d[m] or iginated from Procerites[\i]-Siemiradzkia[m]. Mangold (1971b, p. 219) believed ra the r tha t the lat ter pair gave rise to Homoeoplanulites and Indosphinctes, and that together these belonged to quite ano ther subfamily. H e believed that the ancestors of Choffatia were to be found in Wagnericeras, full g rown specimens of which were described by Arkell (1958, p. 176) as "massive smooth ammoni t e s indis t inguishable f rom Procerites".

C h o f f a t i a c a r d o t i (Peti tclerc, 1915) PI. 9 figs 2-3; PI. 10 fig. 1

1915 Perisphinctes Cardoti nov. sp., nobis; Petitclerc, p. 04, pi. 5, lig. 2. cf. 1932 Perisphinctes cardoti Petitclerc; Corroy, p. 141, pi. 24, ligs 5. 6.

non 1939 Grossouvria (Poculisphinctes) cardoti (Petitclerc); Arkell, p. 179 [ — C. poculum\. 1958 Grossouvria cardoti Petitclerc. sp.; Arkell, p. 222. 1971b P. cardoti PMilderc: Mangold , p. 162. 1974 Choffatia cardoti Petitcl ; Cariou, p. 307.

Holotypc, by monotypy . Specimen thought to be in the Labora to i re de Paleonrologie des Invertebr<5s, Univcrs i tc Pierre ct M a r i e Curie , Paris, but not traced ( P a j a u d , pers. comm. 1980). Original ly figured Petitclerc. 1915, pi. 5, fig. 2. Lower Callovian (probably Koenigi Subzone Cal lomon, pers. comm.) . qua r ry of P rahccq , Dcux-Sevres, Aqui ta ine , France . Macroconch .

Material. T h r e e unerushed and undis tor ted internal moulds preserved in coarse friable sandstone; well displayed sutures and siphnncles. Macroconchs , including one with body chamber .

Description. M a x i m u m d iamete r at least 200 mm. Body chamber at least 280°. More than five whorls visible. Kvolute ( w h / d ~ c . 2 5 % ) ; whorl section rounded on inner and middle whorls becoming compressed ( w h / w t < 150%); Hat whorl sides and rounded venter , broad on inner and middle whorls (Text-lig. 11); modera te ly to steeply sloping atid almost, smooth umbilical wall; umbil icus wide and modera te ly deep (up to 105 m m wide and 20 m m deep at 195 nun d iameter ) .

40-

30

20

primary & simple ribs • = s t a r t of b o d y c h a m b e r

FW TNN 460 ImJ

FW TNN 522 FW TNN 507

[ M l

FW TNN 512 [ml FW TNN 537

BGS G S M 49454

IMJ

JL 10 30 50 70 90 110 130 150 170 190

diameter (mm)

TF.XT-FIG. 13. Rib-density curves for Choffatia cardoti {Petilclcrc) | broken line] and C. muperoi (Gemrnellaro) I solid line].

Ribs (Text-fig. 12) general ly rcct i radiatc to gently prors i radia te th roughout , though where pr imaries more markedly prors i radia te , secondaries may develop slight rurs i rad ia te aspect . Pr imary ribs strongly developed and ra ther widely spaced th roughout (Text-fig. 13), par t icular ly on middle whorls where they are almost bulla.te; on outer whorls , pr imar ies weaken bur. persist: on middle and outer whorls , pr imar ies split into three, a little above middle of whorl side, m a y b e with an associated inrercalatory (sometimes this combinat ion appear s a lmost v i rga to tome) ; on inner


wh wl uw pr imar ies 6". 21 c. 21 — —

28 32 41 23 49 6-. 32 105 est. 28

whorls (up to c. 40 m m d iamete r ) , some similar style r ibbing but also some bifurcate ribs. Ribs pass across rounded venter without in ter rupt ion . Well developed, ra ther wide constr ic t ions on inner and middle whorls , m a y b e two per whorl .

Data for figured specimens. d (septate to)

F W T N N 5 3 6 (PI. 9 lig. 2) 64 (64) F W T N N 5 3 7 (PI. 9 fig. 3) c. 90 (—) 28 32 41 23 c. 62 BGS GSM49454 (PI. 10 fig. 1) 193 (c. 123)

Remarks. T h e holotvpe was alleged (Petitclerc 1915) to be from the anceps Zone [ = Midd le Callovian |. However , little s t ra t igraphical informat ion was given and it possibly came instead from the Lower Cal lovian, because in that area of F rance it has not always been possible to separa te Lower and Middle Cal lovian faunas , which may occur in one unit (Couffon & Dollfus 1928, p. 406). Mangold (1971b, p. 162) believed C. cardoti belonged in the Lower Cal lovian . Corroy (1932) figured a specimen from the H a u t e M a r n e that appear s to be less coarsely r ibbed than the holorype. a l though he was clearly in no doubt abou t character is t ic features of the lat ter . Unpubl i shed work by Car iou on the type mater ia l from Aqui ta ine indicates that C. cardoti belongs to the Macrocepha lus / o n e , Rehmarmi S u b / o n e which is equivalent to the Calloviense Zone, Koenigi Snhzone of Grea t Britain (Cal lomon, pers. comm. 1981).

Specimens f rom the Lamber t i L imes tone o f W o o d h a m ( O U M J22375-7) , identified by Arkell (1939, p. 1 79) as C. cardoti. and described by him as "like a very coarse-ribbed poculum" a rc herein referred to C. poculum. T h e macroconch of this species appear s to be more involute than C. cardoti, with blunter r ibbing and only a weak deve lopment of constrictions; it also occurs at a younger horizon (Lamber t i Zone) . Later , Arkell (1958, p. 222) described C. cardoti as "a lmos t a homoco-rriorph or pe rhaps a direct descendent" of Choffatia (Loboplanulites) cerealis of the Ba thon ian , but with a less inflated whorl section. Loboplanulites was subsequent ly taken by Elmi (1962) and Mango ld (1971b, p. 157) as a j un io r synonym of Subgrossouvria.

C. cardoti also appea r s to have affinites with Sub grossouvria euryptchus (Ncumavr , 1871) f rom the Balin Ooli te of Poland (?Lower Cal lovian) (Spath 1931, p. 375; Gera rd & C o n t a u t 1936, p. 56 (al though their record, wi thout figure, is s t ra t igraphical ly less appropr ia te ) ) . N c u m a v r s (1871, pi. 12 lig. I, wrongly given in the text (p. 38) as pi. 14) figure shows a generally more evenly ribbed form whose r ibbing docs not become as coarse until a later stage, (cf. figure of S. euryptychus f rom Hungary : Loczy 1915, p. 403 fig. 119). However , wi thout bet ter i l lustrat ions of the type mater ia l , fur ther c o m m e n t canno t be made (see also Choffatia recuperoi below).

Distribution. Ke l laways Sand (Calloviense Zone , Calloviense Sub /one ) of South New bald and South Cave , Humber s ide . Lower — ?Middle Callovian of France ( l )eux Sevres. Aqui ta ine (Macrocepha lus Zone; Car iou 1974) (see above) ?Haute Marne ) .

Choffatia recuperoi (Gemmel laro . 1872) PI. 11, figs 1 4

aff. 1872 Perisphinctes Sciutoi Gemrn.; Genimellaiu, p. 25, pi. 4, ligs / - 9 . 1872 Perisphinctes Recuperoi Comm. , Gemmellaro, p. 26.. pi. 5, ligs 9-11.

nan 1875 Perisphinctes recuperoi Gemmellaro; Waagen, p. 172, pi. 43, fig., I \ = C. soorkaensis Spaih] . 1801 Perisphinctes Recuperoi. Comm. ; Hang, p. 99.

? 1090b Per. Recuperoi Gemm, ; Siemiradzki, p. 175. all". 1894 Perisphinctes etegans n. sp., Siemiradzki. p. 517, pi. 40. fig. aff. 1894 Perisphinctes crussus ci. sp., Siemiradzki, p. 518, pi. 40, fig. 5.

1894 Perisphinctes Recuperoi Gemmellaro: Siemiradzki, p. 525 pars. |~pars=C. soorkaensis]. non 1895 Perisphinctes Recuperoi Gemmellaro; Noeding, p. 20, pi. 13, lii». 4.

1897 Perisphinctes Recuperoi Gemm.; Parona & Bonarelli, p. 173. ff. 1898 Perisphinctes Sciutoi Gemm. ; Siemiradzki. p. 128 pars \pars~C. soorkaensisJ.

aff. 1898 Perisphinctes erassus Sic.rnir.; Siemirad'/.ki. p. 129 pars 1890 Perisphinctes Recuperoi Gemm. ; Siemiradzki. p. 295, fig. 68pars [/tars— C. soorkaensis],

all". 1913 Perisphinctes of. etegans Siem.; Lowe. p. 153. 1913 Perisphinctes Recuperoi Geniellaro Lowe. p. 15'1 pars

CHOFFATIA 31

non 1915 Perisphinctes recuperoi Gcmmrl la ro ; Petitclerc, p. / ' / , pi. 8, fig. 2 [— C. dumortieri Mangold & Elmi] . aff. 1915 Perisphinctes Sciutoi. Gemmellaro; Petit de rc , p. 82. non 1915 Perisphinctes Recuperoi Gcmm. ; Loczy, p. 420, pi. 13. fig. 8 | = C. dumortieri Mangold & Elmi].

1919 Perisphinctes Recuperoi Gemmellaro; Coufion, p. 215, pi. 14, figs 2. 2a, 2b pars, non 1924 Perisphi/uiei Recuperoi Gemmellaro: Roman, p. 99. pi. 11. fig. 3 [ = C. dumortieri Mangold & Elmi] . non 1930 Perisphinctes Recuperoi Gemmellaro; Roman, p. 181 pi 18, fig. 1 f = C. dumortieri Mangold & Klriiij.

1930 Choffatia aff. recuperoi (Gemmellaro):. Spath. p. 38. pi. 4, fig. 13; pi. 5, lig. 6. aff. 1930 Grossouvria cf. elegant (Sicmiradski); Spath, p. 40. pi. 4, ligi. 4. 11; pi. 5, fig. 4.

1931 Choffatia aff. recuperoi (Gemmellaro); Spath , p. 358. aff. 1931 Perisphinctes sciutoi; Spath, p. 303.

1932 Perisphinctes Recuperoi G e m m d l a r o ; Corroy, p. 129. pi. 16. fig. 3 pars fpars=C. mark ante mis and C. dumortieri). aff. 1932 Perisphinctes Sciutoi G e m m d l a r o ; Corroy, p. 112, pi. 23.. fig. 9. all'. 1932 Perisphinctes Sciutoi Gemmel la ro var. crassus Sicmiradski [sir]; Corroy. p 143, pi. 17. figs /, 0. non 1938 Choffatia Waageni (Teisseyrc) var. Recuperoi (Gemmellaro); Pfachler-Eiall i , p. 15, pi. 4, figs 2a, b. [fide

Mangold 1971b]. aff. 1939 Perisphinctes (Grossouvria) sciutoi Gemm.; Kuhri, p. 500, pi. 10, fig. 2.

1939 Perisphinctes recuperoi Gcmm. , Kul in , p. 512, pi. 4. fig. 2. non 1952 Perisphinctes (Perisphinctes) recuperoi Gemellaro |«c j ; Makowski, p. 33. pi. 9, fig. 3 [ = C. dumortieri Mangold &

Elmi]. 1958 Choffatia (Loboplanuiites) recuperoi (Gcmmo.lla ro); Arkell, p. 220, fig. 80.

?1961 Perisphinctes (Choffatia.) recuperoi (Quenst . ) ; Rugc t -Pcno t , p. 39. 1965 Subgrossouuria rf. recuperoi (Gemmellaro); Busnardo, Elmi & Mangold, p. 78. 1971b Choffatia (M Subgrossouma) ntuperoi (Gemmellaro); Mangold, p. 161. fig. 117. pi. 10 figs 2. 3.

a IT. 1971b Choffatia (m Grossotwrm) sautoi [ Gemmellaro) ; Mangold, p. 181, figs 122. 3, pi 7, fig 7 P1971 Chojjuliu {Choffatia) cf. recuperoi (Gemmellaro); H a h n & Roomer , pi. I 1.

all: 1974 Grossouvria sciutoi Gem.; Car iou, p. 307. all". 1974 G. crassa Siem.; Car iou, p. 307.

Type specimens. Gemmel la ro ' s (1872) specimens are t hough t to be ei ther in the Museo di Geologia e Mine ra log i ade l l a R. Universi ta di Palermo or the M u s e o di Palaeontologia e geologia, Palermo, Sicily, bu t have not been t raced. From the Rocea chi p a r r a ( 'zone of Stephanoceras macrocephalum') Ca la ta f imi , T r a p a n i , Sicily. Macroconch .

Material. Seven uncrushed and generally nndis tor ted specimens preserved in ironshot oolite, usually as in ternal moulds . Microconchs (complete with aper ture) and macroconchs ( incomplete); one pyriti&ed microconch.

Description. Microconchs ( three specimens) : max imum diameter 60 m m . Body-chamber at least 220°. Six whorls visible. Modera te ly evolute ( w h / d = 3 0 - 3 5 % ) ; whorl section rounded on inner and middle whorls , later becoming more compressed ( w h / \ v t = 110-130%) (Text-fig. 11); modera te ly steeply sloping and a lmost smooth umbilical wall; umbil icus c. 25 m m wide and u p to 8 m m deep in adul t . P r imary ribs prors i radia te , secondaries rnrs i radia te ; inner whorls have bi furcate ribs with occasional intercalatories; middle and outer whorls have ra the r long, s t ra ight pr imaries spli t t ing, somet imes irregularly and always with a backward swing, above the middle of the whorl side into two with an associated intercalatory to each pair , or more rarely into three (Text-fig. 12). Inne r and middle whor ls with moderately well developed constr ict ions. Ribs pass across venter wi thout in te r rupt ion . Body-chamber with terminal constrict ion; ape r tu re bears lappets (up to c. 5X15 m m ) .

Macroconchs (one wholly sep ta te specimen) : m a x i m u m d iamete r c. 90 m m . At least six whorls visible. Evolute ( w h / d = c . 3 0 % ) ; whorl section rounded or slightly compresscd on inner and middle whorls , becoming slightly depressed on outer whorls ( w h / w r = c . 8 5 % ) with ra the r flat whorl sides, gently a rched towards moderate ly broad venter (Text-fig. 11); steeply sloping and almost smooth umbil ical wall; umbil icus c. 40 m m wide and up to 12 m m deep at c. 90 m m diameter . Ribs (Text-fig. 12) gently prors i radia te but on inner whorls, secondaries have rurs i radi-ate aspect. Inner and middle whorls have bi furcate r ibs with occasional intercalator ies; outer whorls have t r i furca te ribs and more rarely bi furcate with an associated intercala tory, p r imar ies spli t t ing at abou t rhe middle of the whorl side. Inner and middle whorls have modera te ly well developed constr ict ions (c. 2 per whorl) . Ribs pass across rounded venter wi thout in te r rup t ion .


d (septate to) wh wt uw primaries vcntra ls 91 (91) e. 25 c. 29 40 33 est. 85

58 (c. 38) 19 17 24 38 90 c. 60 (c. 35) 21 c. 20 24 27 c. 75

57 (c. 36) 19 14 26 32 est. 80

Rib density curves for figured specimens are given in Text-fig. 13. For any one individual , f rom at most 20 m m d iamete r , the n u m b e r of pr imar ies per whorl apparen t ly varies r emarkab ly little.


F W T N N 5 0 7 (Pi. 1L fig. 1} F W T N N 160 (PI. 11, lig. 2) F W TNN.512 (PI. 11, fig. 3) F W TNN.522 (PI. 11, fig. 4)

Remarks. Perisphinctes recuperoi was proposed by Gemmel l a ro (1872) for macroconch mater ia l from Sicily. Arkell (1956, p. 207) believed, on the basis of field evidence, thai it c ame f rom a condensed bed represent ing the Lower and possibly the lowest pa r t of the Midd le Cal lovian. It has since been widely used, par t icular ly by French au thors , for several species from various horizons in the Cal lovian. T h e records of Petitclerc (1915), Loczy (1915), Roman (1924; 1930) and Makowski (1952) from the Middle Cal lovian were separa ted by Mangold & Elmi (I960) (who saw most of the figured and cited mater ia l ) into a new species Choffatia dumor fieri. Th i s is dist inguished f rom C. recuperoi by a less depressed whorl section and less thick p r imary ribs, but most not iceably by the straight and close r ibbing of the inner whorls , which in C. recuperoi persists to a comparat ively large d iamete r a feature a l ready noted by Spath (1930, p. 38; 1931, p. 359). Spath retained Gcmmel l a ro ' s species for " the more doubt fu l serpcnt icone Perisphinct ids of the macrocephalus beds" o f C u r c h and based his new species C. soorkaensis on W a a g e n ' s (1875) main example of C. recuperoi which was dis t inguished from Sicilian specimens by the ( lose r ibbing on the periphery of the earlier whorls and comparat ive ly simple su ture line (Spath 1931, p. 360). S iemiradzkrs (1891; 1898) records of the species included W a a g e n ' s example in their syn-onymies. Corroy ?s (1932) record also included this form as well as specimens subsequent ly referred to C. dumortieri. He (1932, p. 130) gave the horizon as U p p e r Callovian, but in the key to his plate 16. a t t r ibuted ir to the Lower Cal lovian; the s t ra t igraphieal section agrees with the text. T h e division is also included in the Midd le Cal lovian of current usage; the localities given by Corroy showed both Lower and U p p e r Cal lovian, and the record must therefore remain open to doubt .

O t h e r related forms are C. euryptychus (Neumayr ) and C. cardoti (Petitclerc) (see above); compared with C. recuperoi, the former is ra ther more irregularly r ibbed, and the latter is more strongly, sharply a n d irregular ly r ibbed. Both, bu t par t icular ly C. cardoti, lose the s t ra ight and dense r ibbing of the inner whorls at an earlier stage, and both have a less depressed whorl section.

Spath (1930, p. 39) believed C. recuperoi was in te rmedia te between Choffatia s.s. and Sub-grossouvria a n d more recently it has commonly been included in the latter (Busnardo , Elmi & Mangold 1965, p. 78; Mango ld 1971b). Arkell (1958) included it in Loboplanulites, since taken as a j un io r synonym of Subgrossouvria (see above).

The associated microconchs are tentatively recognized amongst and grouped with G c m -mellaro 's species Perisphinctes sciutoi: the type specimen (Gemmel laro 1872, pi. 4, figs 7-9) came from rhe same horizon and locality as C. recuperoi but it is much less well known. Mangold (1971b, p. 181) included it with the eve.xa-sulcifera g r o u p of Grossouvria (characterized by a regular style of o rnamen t ) which he used to a c c o m m o d a t e microconchs associated with Subgrossouvria. Peri-sphinctes elegans S iemiradzki and Perisphinctes erassus w:ere also included. Siemiradzki himself (1898) placed P. etegans in the synonymy with P. sciutoi'; P. erassus is clearly closely related and Cor roy (1932, p. 143) included it as a variety of P. sciutoi. Parona & Bonarelli 's (1897) Perisphinctes cf. subtilis listed by Siemiradzki (1898, p. 129) in the synonymy of P. erassus, is not inc luded. Perisphinctes gracilis is also excluded, a l though Siemiradzki (1898) subsequent ly g rouped it, together with P. etegans, in P. sciutoi; Mangold (1971b, p. 185) believed it belonged to a younger horizon. P. gracilis is less compressed than the otherwise similar but slightly younger Grossouvria

C H O F F A T I A - . ^ 33

vurvkosla (Oppel)-kontkiewczi (Siemiradzki) of Pfaehler Krar.h (1938. pi. 1. figs -4a, h) and those figured herein are less strongly constr icted, at least on the outer whorls, than some of the examples , including the types, ment ioned above.

Distribution. Ke l laways Rock (Calloviense Zone, Knodatum Subzone) of South Newbald and South Gave. Humber s ide ; basal Oxford Clav (Knoda tum Subzone) of the Hast Midlands : Lower

• / f

Callovian of Sicily; F rance ( H a u t e M a i n e , Ardennes . Aqui ta ine (Gracilis / o n e ; Car iou 1974a)); t h e j u r a ; southern Spain (Andalus ia ) ; ^Portugal; Poland ("Eiserioolithe" of Balin and Rudniki ) ; Kenya; Ind ia (Cutch) ; U S S R (Li thuania ; Siemiradzki 1890b); Macrocepha lus Zone of G e r m a n y (Saxonv (Lowe 1913), Swab ia (Ha lm & Koerne r 1971) and Franconia ( K u h n 1939)); and Lower Cal lovian of the Alps ( H a u g 1891).

Choffatia sulcifera (Oppel , 1857) PI. 12, ligs 1 (S 1840 Ammonites conuolutus ornati: Quens ted t , p. 169. pi. 13. fie:. 1857 Ammonites sulci ferns; O p pel. p. 555.

non Ub8 Amnmonites convolutus vrnati; Qucn.s(ed(, p. 544, pi. 71. fig. 9. | =e f . Choffatia aff. lahusent] 1802 Ammonites sulcijerus; Oppel . p. 155, pi. 49, figs 4a, I), c.

non 1932 Perisphinctes sulcijerus Oppel tip.; Corroy, p. 126. pi. 18., figs 8, 9. 1952 Perisphinctes (Grossouvreia) variabilis Lahusen; Maknw.ski, p. 32, pi. figs 4, 4a. 1956 Grossouvria aff. variabilis (I.ah ); Arkell, p. 482. 1957a Grossouvria (Grossouvria) sulcifera fOppel ) ; Arkell, p. LI519, Fig. 44)5, 12a. b. 1958 Grossouvria sulcifera (Oppel) ; Aikell, p. 214. 1971b Choffatia (in. Grossouvria) sulcifera (Oppel) ; Mangold, p. 189.

Holotype. By monotypy . Specimen figured Quens t ed t 1849, pi. 13, fig. 1, GPI (unnumbered ) ; f rom "Dogger zeta (Cal lov ium)" , Gammcl shauscn , W u r t t c m b c r g , W. G e r m a n v . Cas t , BGS G S M 117447, figured PI. 12, fig. 1.

Material. T e n uncrushed or part ial ly crushed specimens, small but nearly complete (adult) pyritized in ternal moulds , and one uncrushed incomplete specimen in a shelly calcareous dogger. Microconchs .

Description. Microconchs : m a x i m u m d iamete r c. 40 mm. Body-chamber p robably at least 250". At least 4 whor ls visible. Modera te ly evolute (wh/cl=c. 35%) ; whorl section generally modera te ly compressed ( w h / w t = 100-120%) with ra ther flat whorl sides and gently arched broad venter , but innermost whorls depressed (Text-fig. 11); umbilical wall steeply sloping and almost smooth; umbi l icus shallow and wide (c. 15 m m wide and 3 m m deep at 37 m m d iamete r ) . Ribs (Text-fig. 12) general ly prors i rad ia te on umbil ical edge and whorl sides, becoming rurs i radi-ate, often markedly , towards venter . Inne rmos t whorls smooth or with weak pr imar ies only, and weak constr ict ions (maybe two per whorl) . By 10 m m diameter , r ibbing becomes bi furcate with pr imaries spli t t ing into rwo on the outer par t of the whorl side and generally passing uninter-rupted across venter ; constr ict ions (maybe two per whorl) , weak to modera te ly s t rong—par -ticularly the one mark ing end of phragmocone . Somet imes weak nodes on the ventro- la tcral marg in , par t icular ly on the later whorls and body-chamber where r ibbing bccomcs a l together less regular with pa rabo lae and occasional s imple ribs. General ly 30-10 pr imaries for d iamete rs between 10 and 40 m m .


d (septate to) wh wt uw pr imaries ventrals Holo type (GPI , no number} (PI. 12, fig. 1) 31 (27) 9 9 1.5 29 est. 68 BGS GSM27932 (PI. 12. fig. 2) 32 (c. 24) 11 10 13 est . 35 O U M J2588 (PI. 12. fig. 3) 30 (18) 10 9 13 38 est. 66 J K W BH12G (PI. 12, fig. 4) c. 28 (—) 9 9 O U M J1218 (PI. 12, fig. 5) 37 (26) 13 (») 15 38 O U M J 2 5 8 7 (PI. 12, fig. 6) 27 (27) c. 9 8 11 c. 42 est. 65

Remarks. Al though C. sulcifera is the type species of (irassouvria and in that role has received


considerable coverage in the l i terature (see above, under Choffatia), it is not par t icular ly well known. T h e holotypc is a constricted form with prors i radia te pr imary ribs th roughout and a character is t ic ru rs i rad ia te twist in the secondaries. T h e r e may be the vestige of a ventral sulcus and pa rabo lae are confined to the body-chamber (PI. 12. figs l a - d ) . T h e mater ia l described herein includes nearly complete adult (microconch) individuals comparab le with the type material of de LorioPs (1808) species miranda which has been used previously to a c c o m m o d a t e the small pyritized representat ives of the 'ma ins t ream 1 per isphinct ids (Choffatia-Grossouvria) of the Athle ta to G o r d a t u m zones. Spath (1931, p. 394) restricted miumia to de LorioPs figures 15 (lectotype. Arkell, 1939) and 15a. However, compared with the present mater ia l , miranda shows a more i rregular rib style on the phragmocone with well developed parabol ic ribs and nodes, but with fewer constrict ions; the type also comes from a younger (Lower Oxford ian , Mar i ac Zone) horizon. Perisphinctes variabilis Lahusen (1883, pi. 10, fig. 4) is based on a complete Russ ian microconch f rom a condensed ironshot oolite possibly of late Callovian age (Lahusen 1883, p. 85) and inferred by Spath (1931, p. 328) and Mangold (1971b, p. 166) to come from the Lamber t i Zone. Tt differs from most of the mater ia l described here in having more p ronounced i r regular r ibbing on the ph ragmocone and no, or only a weak, development of constrictions. Makowski ' s (1952, pi. 8, figs 4a, h) figure of P. variabilis seems to fall within the range of var ia t ion here illustrated for C. sulci [era, and following Mangold (1971b), is included with it. Makowski ' s material from Poland ( U p p e r Cal lovian, Lamber t i Zone, fide Arkell 1956, p. 482) shows an a perl i i re heari ng lappets . 1 he A t hie la Zone material descri bed here occu rs wi t h A lligaticera.c rotifer (Brown) (see below). T h e few large pyrit ized, poorly preserved ( incomplete or with pyri te overgrowths) specimens in old collections (e.g. O U M J1211. J26254) , which come from similar s t ra t igraphieal levels or localities might be the associated macroconchs of C. sutcifera.

Distribution. Oxford Clay (Athleta Zone) of YVolvercotc and S u m m e r t o w n , Oxfordshire ; Hackness Rock of Hackness , Nor th Yorkshire (Bed 4 of Wrigh t 1968; Athleta Zone): Athle la-Larnberti Zones of Swabia , southern Ge rmany ( H a h n & Kocrner 1971); Lamber t i Zone of Lukow, Poland; Athleta Zone of the French J u r a (Mangold 1971a, b) and north-west Aqui t a ine (Cariou 1974).

C h o f f a t i a aff. l a h u s e n i (Sasanov, 1965) PI. 12, figs 7, 8 Material. T w o completely sep ta te nncrushed pyrit ized internal moulds f rom the Oxford Clay

(Athleta Zone) of Blctchley, Buckinghamshire (Bed 21 of Cal lomon 1968. p. 282) and Wolver-eote, Oxfordshire .

Description, M a x i m u m d iamete r seen c. 70 mm, completely septate; at least 4 whorls visible. Modera te ly evolute ( w h / d ~ c . 3 0 - 3 5 % ) ; whorl section rounded or slightly depressed ( w h / w t = 8 5 9 5 % ) with gently rounded whorl sides and broad venter (Text-fig. 11); modera te ly steeply sloping and a lmost smooth umbilical wall. Ribs (Text-fig. 12) prors i radia te th roughou t ; bifurcate with occasional intercalatories on inner whorls (up to r . 30 m m diameter) with shallow constrictions: on middle whorls, this style persists but t r i furcate ribs also common with some s t rengthened and flared pr imaries ; ribs split about middle of whorl side and pass un in te r rup ted across venter; 25 30 pr imaries at i. 20 mm diameter . O u t e r whor l s / body -chamber unknown .


d (septate to) wh wr uw pr imaries ventra ls O U M J17870 (PL 12, fig. 7) 19 (19 ) 7 8 7 30 — O U M J 1 2 0 3 (PI. 12, fig" 8) est. 68 (68) 21 22 est. 28 — —

Remarks. The specimens are comparab le with Neumayr ' s i l lustration of his species Perisphin ct.essubtilis (1871, pi. 14, fig. 3) (cast f igured PI. 12. fig. 10). The re has been confusion su r round ing the in terpre ta t ion of Choffatia lahuseni arising f rom N e u m a y r ' s (1870) original citation and his subsequent (1871) f igure and descr ipt ion of Perisphinctes subtilis. Th i s has been widely discussed and since resolved (see above) , a l though his later (1871, pi. 14, fig. 3) P. subtilis' is dist inct a n d requires a new name . N e u m a y r (1871, p. 38) m a d e it clear that this specimen (from Balin) was

CHOFFATIA 37

septate to the end and also noted that the species was "not r a re" in the upper part of the " O r n a t e n t h o n e n " of W u r t t e m b e r g (Franconia) (Uppe r Cal lovian) . His specimens from Wiirt-t cmburg apparen t ly showed body-chambers and had diameters up to 50 m m , but wi thout i l lustration, fu r ther compar i son is impossible. I favour the view that Neumayr ' s ' type ' specimen is an incomplete microconch, which remains finely r ibbed throughout ; because of its incomplete na tu re and uncer ta in age, I refrain from giving a new specific name. T h e two specimens described herein (PI. 12, figs 7, 8) a re c o m p a r a b l e with N e u m a y r ' s specimen (cast figured herein, PI. 12, fig. 10), but arc more coarsely r ibbed than this form at a comparab le d iameter which has c. 40 ribs at 20 m m diameter , the larger specimen ( O U M J1203; PI. 12 fig. 8), an incomplete macroconch . is larger than any specimen previously referred to N e u m a y r ' s P. sub lilts,

I.ahuserTs (1883) figured example of Perisphinctes sub til is f rom the U p p e r Cal lovian of the Russian Plat form, Peltoceras athleta and Qitensledtoceras keyserlingi Zone (Sasonov 1961), was m a d e the type oiOkaitespseudosubtilis Sasonov, 1965 (p. 24). However , in the same pape r (p. 26) he used the same specimen as the type of Loriolites lahuseni. These two species arc therefore object ive synonyms, and the lat ter n a m e is favoured because Sasonov gave a full descript ion. H e alleged that it showed an in ter rupt ion of the ribs on the venter , with a weakly developed sulcus. It is not possible to tell how signif icant or marked this feature is because Sasonov figured no addi t ional material , and gave no synonymy. It is certainly not well defined on Lahusen ' s figure. Unt i l more complete mater ia l becomes avai lable , the English specimens are referred to Choffatia aff. lahuseni.

Distribution. Possible records f rom Athle ta Zone of the western Alps ( I l aug 1891), Swabia (Ha lm & Koerner 1971), F rancon ia ( N e u m a y r 1871) and nor th-west Poland (Dayczak-Calikowska 1977).

Choffatia poculum (Leckenby, 1859) PI. 12, figs 9, 11-16; PI. 13 figs 1-5 ; Text-f ig. 14 1859 Ammonites pneubtm I.cckenby (Bean MS); Leckenby. p. 9, pi. 1. figs 4a-c .

non 1878 Perisphinctes poculum Lock. ?cf. cumcostu Oppe l ; L'lilig. pp, 613, 649, pi. 16, fig. 2. 1920 Poculisphinctes poculum Bean-Ldckcnby sp.: Buekman, pi. 185. 1920 Pvidhpfunites auricularis no v.; Buckman, pi. 186. 1931 Poculisphinctes poculus (Bean-Leckenby); Spath, p. 383. 1931 Poculisphinctes auricularis S. Buekman; Spath. p .393.

?1933 Poculisphinctes aff. poculum (Rcan MS) Leckenby sp..; Spath , p. 859. ? 1933 Perisphinctes of. poculus Leek.; Djanelidze. pp. 3. i , 15.

1939 Gwssouvrta (.Poculisphinctes/ poculum (Leckenby); Arkell, pp. 158, 178, 202. 1939 Grossotwria (Potulisphinctes) auruulare Buckman: Arkell, p. 179. 1957a Poiuiisphinci.espoculum (Leek.); Arkell, p. 319. 1971 Orionoides (Poculisphinctes) poculum Leckenby; Cariou, pp. 455, 458 1971 ; Poculisphinctes" auricularis Buekman; Cariou, p. 459 1971 Grossouvria poculum (Lorkenby}: Cal lomon & Cope, p. 166. 197-1 Pocuf.isphin.ctes cf. poculum Leek.; Car iou, p. 310.

Type specimens. Macroconch holotype, S M J3298, PL 12. fig. 9, designated B u c k m a n 1920, pi. 185 (original of Leckenby 1859, pi . 1, fig. 4b) . Microconch p a r a t y p e ^ S M J3299, PI. 12, fig. 11, designated Buckman 1920, pi. 186 (original of Leckenby, 1859, pi. I, fig. 4a) , — holotype of Poculisphinctes auricula Ruckman . Both specimens preserved in grey l imestone f rom "Kel loway Rock" [blackness Roek | . nea r Cr i s thorpc Bay, Nor th Yorkshire; probably Lamber t i Zone (see Wright 1968, p. 390, fig. 6).

Material. Abou t 105 specimens: the type specimens and about 25 others , unc rushed but mostly incomplete or i m m a t u r e (preserved in sandy limestone with ooliths); abou t 80 specimens, partially crushed or otherwise dis tor ted, some adul ts but mainly incomplete (many only whorl f ragments) o r i m m a t u r e (preserved in calcareous muds tone and marly l imestone as in ternal clay moulds with thin black powdery pyritic coat ings when fresh) and one specimen, uncrushed but incomplete pyritized internal mould . Microconchs and macroconchs .

Description. Microconchs (62 specimens) : m a x i m u m d iamete r 53 m m , general ly 40-50 m m . Body chamber 230°-240°. Al least tour whorls visible. Modera te ly evolute ( w h / d = 3 0 - 3 5 % ) ;


whorl section general ly rounded hut m a y be moderately compressed on outer whor ls ( w h / w t = 110-130%) with flat whorl sides (Text-fig. 11); gently sloping and a lmost smooth umbilical wall; umbil icus c. 20 m m wide and up to 8 mm deep in adul t . Ribs (Text-fig. 12) rectiraditc to prors i rad ia te but secondaries may have rurs i radia te aspect, par t icular ly on outer whorls; on inner whorls , weak, widely spaced prors i radia te pr imaries (sometimes very weak or replaced by lirae). i rregularly b ranch ing into two with intercalatories and occasional s imple ribs; nodes and flared ribs; on middle and outer whorls, the pr imaries are stronger, rect i radiare or weakly prors i radia te , spli t t ing low on the upper par t of the whorl side into three secondaries with an intcrcalatory or bundles of four, or more rarely bifurcate with two intercalatories; weak constrictions. Ribs pass across broad rounded venter without in terrupt ion. Aper ture , with associated s t rengthened rib and terminal constriction, bears lappets (up to c. i X 10 m m ) .

Macroconchs (46 specimens) : m a x i m u m d iamete r 180 m m . Body-chamber 270°. At least six whorls visible (shown by completely septate type specimen). Modera te ly evolute ( w h / d — 3 0 -40%) ; whorl section generally rounded, bu t may be slightly depressed on inner whorls ( w h / w t = 8 0 100% and modera te ly compressed on outer whorls ( w h / w t = l 10 130%) with flat whorl sides (Text-fig. I I ) : steeply sloping and smooth umbilical wall; umbil icus c. 45 m m wide and up to 15 nun deep in adul t (es t imated) . Ribs (Text-fig. 12) gently prors i radia te to rectiradi-ate; on inner whorls (up to c. 40 m m diameter ) , pr imar ies prors i radia te and ra the r weakly or irregularly developed and w idely spaced, with some flared ribs; on middle whorls ( f rom c. 40 m m to 80 m m diameter) , the pr imar ies s t rengthen and thicken, becoming blunt and widely spaced, spli t t ing abou t the middle of the whorl side into bundles of r a the r s lender secondaries (usually four with two intercalatories) . T h e secondaries become coarser with growth until on the outer whorl(s) (more than 80 m m d iamete r ) , the n u m b e r of secondaries decreases, usually to two with an inlercalatorv, or three, and the r ibbing is generally coarse and ra the r irregular , with only 16 pr imaries on the last whorl . Ribs pass across rounded venter wi thout in ter rupt ion .

Poor preservat ion makes it difficult to establish r ib density curves for individual specimens and only a few general observat ions can be made ; for macroconchs, the n u m b e r of p r imar ies ranges between 10 and 20 per whorl for d iamete rs between 60 and 100 mm; for microconchs. the number of pr imaries ranges between 25 and 10 for d iameters between 40 and 55 m m .


d (septate to) wh wt uw pr imar ies ventra ls SM J3298 (PI. 12, fig. 9) 60 (60) 22 21 21 12 c. 80 SM J3299 (PI. 12. fig. 11) •18 (c. 30) 15 14 20 c. 25 W M S J49 (PI. 12, fig. 12) 49 (c. 30) 15 est. 14 20 O U M J22362 (PL 12, fig. 13) 51 ( ) 16 est. 13 20 BGS FR1116 (PI. 12, fig. 141 11 ( ) 14 (8) 17 M M LL5664A (PL 12, fig. 15) 33 (33) O U M J22361 (PI. 12, fig. 16) 53 (c. 34) 16 14 21 est. 28 BGS Zt517 (PL 13, fig. 1) 80 ( ) 28 c. 25 33 16 c. 54 O U M J22553 (PL 13, fig. 2) 87 (87) 28 28 38 18 YVMSJ58 (PL 13, fig. 3) 95 ( - ) 30 38 40 18 ?c. 70 Y M 606 (PL 13, fig. 4) 51 (54) 20 21 19 c. 65 O U M J22375 (PL 13, fig. 5) 101 ( - ) 30 23 45 18 c. 61 Text-fig. 14 180 (—) 16

Remarks. Lcckcnby 's (1859, pi. 1, ligs 4a -c ) figures of specimens from the Hackness Rock show a d imorph ic pa i r—a microconch bear ing lappets (fig. 4a) and the inner whorls of a larger (macroconch) form (fig. 4h) . Buckman (1920) subsequent ly separated them on morphologica l g rounds into two species, poculum and aurkularis, and assigned both to a new genus Poculisphinctes, with poculum as type species. T h e holotypc o f / 1 , poculum (Lcckenby, 1859, lig. 4b) shows d is tan t , blunt pr imaries associated with close secondaries and a deep umbil icus with high smooth walls:

CHOFFATIA

T E X T - F I C ; . I I . Chojfalia poculum (Leckenbv) f M | ; L a m b c r l i Limestone; VVoodbam. Buck inghamsh i re ; L a m b e r t i Zone , Lamber t i S u b / o n e ; XI (J. II. C a l l o m o n collection and photoncgat ive) .

Spath (1931, p. 328) took these features to be character is t ic of the genus and added (p. 383) a fur ther species (Poculisphinctes kachhensis) based on a single completely septate specimen from the "athleta beds" of Clutch, Ind ia . Arkell (1939) included Poculisphinctes cardoti (Peti tclerc, 1915) in the genus, thereby suggest ing some similari ty with Subgrossouvria (macroconch Poculisphinctes show, after a d i amete r ofc . 40 m m , a change to a coarse r ib style (Spath 1931)) and also Poculisphinctes subpatina (Peti tclerc. 1915). Poculisphinctes trina Buckman . 1922 (see below) and Poculisphinctes


fascisculptus (I.oczy, 1915) (based on Ammonites cotwolutus piicomaphalus Quenstedt, 1886) have also been included (Sparh 1931. p. 328; Car iou 1971, p. 454). Poculisphinctes poculum is herein assigned to Choffatia; the other species listed above can also be included either therein, or in o ther well-established genera such as lndosphincles.

Uhlig ' s (1878) tentat ive de te rmina t ion of C. poculum from the nor thern C a r p a t h i a n Moun-tains, Poland is excluded as the s t ra t igraphical informat ion is inadequate ; Arkell (1956, p. 170) suggested that most of the ammoni t e s figured f rom tha t area came from the M i d d l e Cal lovian. Uhl ig 's figure shows a compressed form whose rib-style bears some resemblance to microconch poculum hut which, a ssuming the l i thograph is accura te , is ra ther too strongly r ibbed on the inner whorls. D j a n e l i d z r s (1933) record from the Callovian of Georgia , U S S R , based on two incomplete specimens and wi thout figures, or precise s t ra t igraphy, is open to serious doub t .

Car iou (1971, p. 459) though t that poculum (based on Buck man ' s type figure) showed str iking similarities with young Rugeticera-s and the middle growth stages of certain Orionoides such as 0 . monestieri (Gerard & Con tau t , 1936). He included it in Orionoides as a I .ambert i Zone form and considered that the microconch auricularis belonged to qui te ano the r genus. Orionoides (Spa th , 1931), based on Perisphinctes orion Waagen , 1875 [non Oppe l , =indicu.s Spa th] and Perisphinctes

pseudorion W a a g e n , 1875 (type spccies)seems bet ter restricted to involute, weakly constr icted forms with ii regular r ibbing ( including parabolae) on the inner whorls , which do not show early thickening and b luntness of the p r imary ribs on the ph ragmocone and which develop macroconch body-chambers "like Pseudopeltoceras. but not tubcrcu la tc" (Arkell 1957a).

In western France , C. poculum. a l though unf igurcd, is sufficiently well known to be used as a subzonal index in the lower par t of the Lambcr t i Zone.

Distribution. Rare in Athle ta Zone (Hackncss Rock of Hackncss, Nor th Yorkshire and Oxford Clay of Wolvercote, Oxfordshi re) . C o m m o n in I .amberti Zone (Lamber t i Limestone of W o o d h a m , Buckinghamshi re ; Hackncss Rock of Cayton Bay, Cuns tone Nab , Gr i s thorpe and Scarborough (Castle Hill) , Nor th Yorkshire; and ?Oxlbrd Clay of T i d m o o r Point , Dorset (Spath 1933)). Recorded f rom Lambcr t i Zone of Prance (north-west Aqui ta ine (Car iou 1974a), Poitou (Cariou 1971), N o r m a n d y (Arkell 1939, p. 203; Car iou 19/1)) .

Choffatia trina (Buckman , 1922) PI. 14, figs 1 - 5 1922 Trinisphinctex trims nov.; Buckman. pi. 332. 1931 T. trims; Spath , p 328. 1931 Poculisphinctes ("Trinisphtncfes') t.rinus S. Buckman; Spath, p. 383. 1939 Grossouvria fPoculisphinctes) tuna (Buckman); Arkclh pp. 159, 179, 203. 1947a Grossouvria (Poculisphinctes) trina (Buckman! (?); Arkell, p. 31 1971 Orionoides (Trinisphinctes) trim Buckman; Cariou. p. 455. 1971 Trinisphinctes tritius Buck.; Car iou, p. 310.

Holotype. BGS G S M 6 5 0 9 , PI. 14. fig. 1; completely septate, uncrushed limonitic cast from the Oxford Clay. Buckman (1922, pi. 332) gave the horizon and locality as Athle ta Zone . W e y m o u t h , Dorset , but Arkell (1939, p. 1 79; 1947a, p. 31) believed that the type of preservat ion indicared tha t it came from the Lamber t i Zone, T i d m o o r Point.

Material. Apar t from the type, 16 specimens, mostly incomplete (many only whorl f r agments ) , partially crushed or otherwise distorted (preserved in calcareous m u d s t o n c or mar ly l imestone as internal clay mou lds with thin black (when fresh), powdery pyritic coatings and part ial ly pyritized inner whorls) , and one completely pyrit ized specimen.

Description. M a x i m u m d iamete r (of macroconch) 110 m m . Body-chamber up to 360°. Al least five whorls visible. Modera te ly evolute ( w h / d = 3 0 - 3 5 % ) ; whorl section moderate ly compressed ( w h / w t = 100-130%) , with flat whorl sides and gently rounded venter (Text-fig. 11); steeply-sloping and almost smooth umbilical wall. Umbi l icus c. 20 m m wide and 7 m m deep at 55 m m diameter . Ribs (Text-fig. 12) prors i radia te , though less so on inner whorls, which may show

BINATISPHINCTLS 39

rurs i radia te aspcct near venter . Iiuierrnosi whorls smooth or weakly r ibbed, later developing bifurcate ribs with an associated inlercalalorv; middle whorls have t r i furcatc ribs with inter-calatories, with ra ther long straight pr imaries splitt ing at , or a little above, middle of whorl side into bundles of s lender secondaries; on later whorls, r ibbing coarsens with pr imar ies becoming more widely spaced and secondaries weakening. Modera te ly s t rong but shallow constrict ions, following course of r ibbing, par t icular ly on middle whorls. Ribs pass un in te r rup ted across venter . Poor preservat ion makes it difficult to establish rib density curves lor individual specimens, and only a few general observat ions can be made: from at most 10 m m diameter , the n u m b e r of primaries per whorl ranges between c. 28 and 36, arid remains fairly cons tant for any one individual .


vent ra Is c. 100

est. 105 c. 120

d (septate to) wh wt. uw pr imar ies BGS C S M 6 5 0 9 (PI. 14. fig. 1) 55 (55) 18 17 21 35 O U M J16729 (PI. 14, fi?. 2) 110 (c. 55) 29 21 49 28 O U M J22368 (PI. 14, fig. 3) 87 ( - ) ' f. 23 18 42 36 BGS Zr3766 (PI. 14. fig. 4) 74 ( - ) 24 20 31 O U M 122370 (PI. 14, fig. 5) 53 ( - ) 19 est. 16 21

Remarks. It is not certain whether the holotypc is a microconch or macroconch , because it is completely septate . Amongs t the mater ia l described herein, there is no definite microconch, and it seems likely tha t most of the larger specimens are incomplete macroconchs; specimen O U M J16729 (PI. 14 fig. 2) is near ly complete. T h e possibility that microconchs of (J. trim have been included with those of C. poculum cannot be ruled out.

Spath (1931) included C. trim in Poculisphinctes and believed that the holotype represented the inner whorls of one of the more finely ribbed (? and undescr ibcd) forms of tha t genus and compared with it his species Poculisphincteskachhensis (Spath, 1931) from Cu tch , India , based on a single specimen which, like C. Irinus, was completely septate and finely r ibbed. Car iou (1971) included the form with Orionoides, p resumably par t ly following Spath (1931) and Arkell (1939. 1947a) and part ly following his discussion of the species poculum.

C. trim bears compar i son with Prope risp h inctes pseudobe mens is Sasonov, 1965 from the Midd le Callovian of the Russian Platform which, however, has a more rounded and depressed whorl (cf. part icular ly Sasonov 1965, pi. 18, figs 5a, b) and also with 4Grossouvria'peliklerct de Loiiol, 1898 from the Lower Oxfo rd i an (Mar iac Zone) of the J u r a which is, as Arkell (1939) pointed out , more involute.

Distribution. L a m b e r t i Zone (Oxford Clay of Weymou th [T idmoor Point) , Dorset and ?Millbrook, Bedfordshire : Lamber t i Limestone of W o o d h a m , Buckinghamshi re) . Recorded f rom the Lamber t i Zone of France . \ W Aqui ta ine (Cariou 1974a), Poitou (Car iou 1971), N o r m a n d y (Arkell 1939).

Genus BINATISPHINCTES Buckman, 1921

(Synonyms: Hamulisphinctes Buckman , 1921 (type species / / . hamulatus Buckman , 1921) [m] , Okait.es Sasonov, 1961 (type species Ammonites mosquensis Fischer, 1813) [m], ?Rugeticeras

Tin tanr , 1961 (type species R. cesaredense Chof fa t MS, T in t an t in Rnget-Pcrrot el ai, 1961 [ MJ , pars Pse.udopeUoceras Spath , 1928 (type species Ammonites chauvinianus d ' O r b i g n y , 1847) [ M ] ) .

'type species. Ammonites binatus (Bean MS) Leckenbv, 1859 Diagnosis. Evolute forms with rounded inner whorls and more compressed later whorls (Text-

fig. 15). Inner and middle whorls with ra ther dense, mainly bi furcate and s imple r ibb ing (Text-fig. 16). Weak pa rabo lac a n d / o r constrict ions. General ly flattened venter with smooth band or


sulcus. Dur ing later growth stages, macroconchs develop strong, thick, widely spaced p r imary then simple non-Luberculate ribs. Lappe ted microconchs. External su ture has first lateral saddle with a well marked and qui te deep accessor)- lobe, a trifid asymmetr ica l lateral lobe, generally deeper than the ventral lobe, and a more or less retracted suspensive lobe with some well defined auxiliaries.

TF.XT-FIO. If). Whorl sections ofBinatisphinctesspecies: (a) J K W A C . / M a t lit* nun: (b: SM J:530bat 75 mm; (r) Y\L 61 I at 70 mm; (d) S \ 1 J3304 al 82 mm: ;"e) S M J 3 3 0 5 at 63 mm: (f) Y M 61 J at 77 mm.'

Remarks. As with many o f B u c k m a n ' s genera published in Type Ammonites, no formal descrip-tion was given. Buckman used Binatisphinctes to accommoda tc the species binatus "Bean-Leckenbv from the Cas t le Rock [Hackness Rock; Ath lc ta -Lamber t i zones) of Nor th Yorkshire.

Jluctuusus Prat l and possibly comptoni Prat t , f rom the Oxford Clay | Aciit isrriatum Band; Athle ta Zone] of Wiltshire. Buckman considered tha t B. comptoni could be placed only tentatively in the genus because its su tu re line was not available for study. O u r under s t and ing of the original meaning of Binatisphinctes mus t rely solely on Buckman '* four figures of these three species, with their cryptic d imension details (Buckman 1021, pi. 261 A, B (binatus): 1924. pi. 435 (comptoni); 1925, pi. 615 (JJucluosus)). His designated lcctotvpe of the type species is an incomplete, evolute

outer whorls middle whorls

inner whorls body chamber

weak parabolae

occasional weak constrictions

occasional weak constrictions

parabolae and ventral nodes

Binatisphinctes comptoni

Binatisphinctes ham u fat us

Binatisphinctes binatus

1 EX' I ' -HC. 16. Ribbing o[ Binulhphincles species.

BINATISPHINCTLS 41

com pressed p lanula te with a ventral smoorh hand . Ribbing is dense with long s t ra ight pr imaries . T h e su ture line on both figures has been painted in (usual Buckman pract ice) , bu t not descr ibed. T h e inclusion of Pra t t ' s B. ftuctuosus and B. comptoni means that (he genus as unders tood by Buckman, included both macroconch and microconch forms.

T h e present definit ion and scope ol the genus was discussed by Mangold (1971 b, p. 198). H e considered tha i the B. binatus g r o u p was restricted to macroconch forms rela ted to the microconch group of Perisphinctes rjasanensis Teisseyre. This latter species, together with o thers from the Callovian of the Russian Pla t form and Poland (subaurigerus' I 'eisseyre and mosquensis Fischer) had already been associated with Binatisphinctes by Spath (1931, p. 327). Sasonov (1961, 1965) subsequent ly proposed Ohailes for the g roup of mosquensis and rjasanensis; Mangold (1971b) considered this a microconch subgenus of Binatisphinctes, and restricted Binatisphinctes s.s. to the associated macroconchs .

T h e type specimen o f B . binatus cannot conclusively be recognized as a macroconch, and to use Binatisphinctes s.s. exclusively for macroconch forms with binatus as the type species is ques t ionable . In accordance with the policy of associat ing macroconch and microconch forms at species level, the genus, wi thout subgenera , is herein taken to include both forms, thereby following Buckman and avoiding the problem of the d imorph i sm of the type specimen of the type species. T h e r e is a number of o ther closely similar generic g roups of approximate ly the same age as Binatisphinctes— Okaites Sasonov. 1961. Hamulisphinctes Buckman , 1921, Ruge tic eras T i n t a n t , 1961, and Pseudo-peltoceras Spa th , 1928.

O t h e r species assigned to Binatisphinctes include B. giganleus Gera rd & Con tau t , 1936, B. robauxi Ge ra rd & C o n t a u t . 1936 and B. welschi Gera rd & Con tau t , 1936 f rom the Ath le ta Zone of France, the Russ ian B. arlti (Krenkel . 1915) and the mainlv Russian B. rossicus (Siemiradzki, 1898).

Binatisphinctes is thought to have evolved from Choffatia du r ing the mid to late Cal lovian (Corona tnm-Aih le ta zones). Some au thors , e.g. Spath , thought the microconch of B. comptoni to he better placed in Grossouvria ( = m i e r o c o n c h Choffatia) and forms such as Q u e n s t e d t ' s (1886-7, pi. 81, figs 15 19) evexa arc certainly qui te close. Mangold (1971 b, p. 186) designated Q u e n s t e d t ' s fig. 16 as leerotvpc of evexa and concluded that it was a form of the C o r o n a t u m Zone. T h e macroconch may be derived from Subgrossouvria [ = Choffatia] which has a character is t ic sudden change in its r ibbing style to s t rengthened and thickened pr imaries on the outer whorls (cf. Marigold 1971 b, p. 202). 11 shows affinities with Peltoceras which later in par t s of the Athle ta Zone becomes the p r edominan t per isphinctaccan m e m b e r of the English Callovian a m m o n i t e fauna.

Notes on genera regarded as synonyms of Binatisphinctes. (i) Hamulisphinctes Buckman, 1921

This genus was publ ished without formal descr ipt ion—only figures of three specimens of two new species (11. hamulatus and H. auricula) from the Cast le Rock | Hackness RockJ of Yorkshire (Buckman 1921, pis 262A, B; 263). This material was included by Leckenbv (1859) in his species A. binatus. B u c k m a n ' s figures show evolute forms, r a the r coarsely r ibbed (ribs having rurs i rad ia te aspect towards the venter at least on the outer whorls) with ventral weakening of the ribs or smooth band, and rounded whorl section. T h e type specimen of auricula is complete (d iameter 66 mm) and its ape r tu re bears lappets . Mangold (1971b. p. 208) restricted Hamulisphinctes to microconch forms related to the macroconch taxon Pseudopeltoceras (see below). No fur ther species has been recognized within, or assigned to, the g roup , a l though Spa th (1924. p. 13; 1931, p. 371) quoted ano ther Bean manusc r ip t species name (marifimus) a t tached to a specimen in the British M u s e u m (Natura l His tory) (no. 39530: PI. 18, fig. 5). Cal lomon (pers. comm. 1985) believes this specimen to be "an excellent [ m | of Pseudopeltoceras. (ii) Okaites Sasonov, 1961

T h e original mater ia l of the type species was in the Fischer Collection, kept at the Pavlov M u s e u m , Moscow. N. T . Sasonov reports that it is lost. H e in tends to select a neorype from material described by Lahusen (1883) (G. Sasonova. pers. comm. 1978). Sasonov included in


Okaites regular , evolute forms with rounded inner whorls. flattened venter with smooth band or sulcus, s t ra ight ribs a l te rna t ing simple and bifurcate, with parabolae , a n d a body-chamber occupying th reequar te r s of a whorl and with lappets. Suture details were described by Sasonov (1965), a n d in t ranslat ion by Mangold (1971 b. p. 200). As well as the type species, Sasonov (1961. p. 28) included N e u m a y r ' s species Perisphinctes scopinensis and Perisphinctes eurvptychus and later (Sasonov 1905, p. 22), added Teissevrc 's (1881) suhauri.gerus and rjasauensis, sinzowi Nikolajcva & Rozhdesiwenskaya a n d C. lahuseni Sasonov. Mangold (1971b), who restricted the genus to microconch forms, included KrcnkcTs (1915) species popilanicus, bodeni and credneri and SieniiradzkTs polonicus. All these spccics arc based on Russian or Polish forms; the geographical d is t r ibut ion, according to Sasonov (1961), eenrres on the Russian Plat form, nor the rn Caucasus and Poland, and includes England and France. (iii) Rugeliceras T i n t a n t (in Ruget-Pcrrot el al.), 19bI

Th i s genus was proposed to accommoda te some 40 specimens of the species R. cesaredense f rom the Uppe r Cal lovian (Athleta Zone) of Portugal . It included evolute forms with a wide umbil icus; inner whorls with bi furcate ribs a re followed by a stage with s t rong, spaced-out pr imar ies each giving rise, on the uppe r half of the whorl side, to three or four secondaries often slightly rurs i radia te and weakened or in ter rupted on the venter. Adul t forms have lengthened, s t rengthened and spaced out pr imaries which become very strong, but never with tubercles , and the secondaries d i s appea r totally. There is no constrict ion. Suture line incised with short ventral lobe, a much longer lateral lobe and an even, well developed suspensive lobe. Choffa t described in manuscr ip t cesaredense in 1910 and assigned it to "Peltoceras (Perisphinctes?)". H e thus clearly thought that this spccics was t ransi t ional between the t rue per isphinct ids and Peltoceras, which led Tintant in the subsequent published descript ion to compare it with Pseudopeltoceras (see below). Without the inner and middle whorls on which there are some differences in o rnamen t , T i n t a n t concluded that dist inction between Rugeliceras and Pseudopeltoceras was slight. He also pointed out the close similarity with Subgrossouuria; young examples of the two genera are dillicull to dist in-guish, but Rugeliceras has no constriction at tha t stage and more marked tendency to have an interrupt ion of the r ibbing on the venter. In adul t forms, Sub grossouvria has a much shor te r ( abou t a half whorl) s tage of s imple raised ribs; in Rugeliceras, it s tar ts on the ph ragmocone and lasts for two whorls. 'Tintant believed that Choffatia, Subgrossouuria, Rugeliceras and Pseudopeltoceras con-stituted a con t inuous morphological series character ized by the progressive deve lopment on the adult of a "pe l tocera to id" o r n a m e n t of coarse, raised simple ribs and progressive d i s a p p e a r a n c e of secondaries. 'The type species of Rugeliceras was thought to be closest to Pseudopeltoceras rollieri J c a n n e t . Binatisphinctes was not discussed, but it clearly belongs ro the same group .

(iv) Pseudopeltoceras Spa ill, 1928 'This genus was proposed for offshoot forms from the main perisphinetid stock transi t ional to

Peltoceras. Spa th different iated Pseudopeltoceras from Suhgrossouvria (see Choffatia) in the. g r adua l deve lopment of ventral tubercles on an inflated Nequeniceras like body-chamber (Spath 1931, p. 557). He referred to undescr ibed mater ia l from Monireui l ttellay (Fiance) and Scarborough , North Yorkshire (Hackncss Rock) including the unpubl ished Bean manusc r ip t species famulus (figured Spa th 193 L pi. 103, fig. 4; pi. 105, figs 6a, b) and le.ckenbyi (PI. 20, fig. 1). I I cdesc r ibed the inner whorls leckenbyi as "closely costatc, per isphinctoid" showing a very g radua l changc to the tubercula te s imple r ibbing of the body chambcr . His one new species Pseudopeltoceras retrorsum f rom the Callovian of Cu tch has uniform simple and bi furcate rurs i radia te ribs on the depressed and later rounded inner whorls; these ribs become more dis tant and thickened until the s u b q u a d r a t e -sectioned bodv-chambcr which narrows towards the aper tu re and has secondary ribs and intercalatories confined to the flat venter . The 'holotype1 of the type species (svntypc according to Cal lomon, pers . c o m m . 1985) was examined and re figured by T i n t a n t (1961, p. 187, pi. 2 fig. 2). This shows that d ' O r b i g n y ' s original i l lustration exaggerated the development of ventral tuber-

cles. Nevertheless, the so-called "pel tocera toid" tubercula te o r n a m e n t on s u b q u a d r a t e whorls was a generic charac te r understood by Spa th . J e a i m e t (1951) added to the genus two new species.

BINATISPHINCTLS 43

P. rollieri and P. zuberi f rom the Cal lovian of the S w i s s j u r a . Mangold (1971 h, p. 208) subsequent ly restricted Pseudopeltoceras to macroconch forms related to the microconch Hamulisphinctes. He described three successive stages of o rnamen t : inner whorls with simple and b i furca te ribs, weakened or in te r rup ted on the venter , pa rabo lae and constrictions; followed by a middle stage with prors i radia te pr imar ies becoming more widely spaced, s t rong at their base, weakening on the whorl side high up on which they split into secondaries with intercalatories having rursiradi-ate aspect and with compressed whorls , constr ict ions but no parabolae: and then a final s tage with widely spaced p r imary ribs thickened but without tubercles part icular ly in the v e n t r o l a t e r a l area, and ventral ribs d i sappear ing and being replaced by a broad ventral "cush ion" . Until Mangold associated Pseudopeltoceras with Hamulisphinctes, the former genus had been grouped exclusively with Pe.ltnce.ras in the subfamily Peltocerat inae. Specimens of' Pseudopellocers Jama turn Spa th are not u n c o m m o n in the Hackness Rock (Athleta Zone) and two examples are figured (PI. 21, figs 3, 6). Spat lvs type specimen from the Hackness Rock of Scarborough which he figured is BM 39512 not 39312 {fide Phillips 1977, p. 156).

Binatisphinctes comptoni (Pra t t , 1841) PI. 14, figs 6. 7; PI. 15, (igs 1-8; PI. 16. fig. 1; PI. 17, figs 1, 2; PI. 18, fig. I

1041 Ammonites Comptoni; Pratt , p. 163, pi. 1 fig. 1. 1841 Ammonites jluctuosus; Prat t , p. 161, pi. 6 fig. 1 only. 1849 Ammonites Comptoni; Brown, p. 244, pi. <20*"fig. 7. 1849 Ammonites jluctuosus; Brown, p. 245, pi. 20**fig. I (J. 1898 Perisphinctes Comptoni Pratt; Siemiradzki, p. 13.0 pars 1808 Perisphinctes ftuctuosus Prat t , Siemiradzki, p. 293, mm pi. 25 fig. 13 pars

non 1913 Perisphinctes Comptoni PraLL; Lowe, p. 152 J1915 Perisphinctes Comptoni Pratt ; Krcnkel , pi. 23 fig. 5.

1924 Bmitisphiwtes? Comptoni, Pratt sp.; Buckman, pi 485 1925 Binatisphinctes jhictuosus. Pratt sp.; Rnckman, pi. 615. 1932 Perisphinctes comptoni Pratt sp.; Corroy, p. 127, pi. Hi figs 11, 12 pars

non 1939 Perisphinctes (Grossouvria) comptoni Pratt : Kulin, p. 504. pi. 10 fig. 17 non 1939 Perisphinctes ftuctuosus Pratt ; Kulm, p. 511 non 1955 Grossouvria cf. comptoni ! Pratt) ; Cal lomon, p. 233 [ = / . patina [mj ] non 1955 ?Binatisphirttles jluctuosus (Pratt}; Cal lomon. p. 233 ( = / patina f M | |

I9G3 Am. jluctuosus P ra t t [M] compUmi Prarr [mj ; Callomon. p. >H. 1969 Binatisphinctes comptoni and B. jluctuosus; Smith in Cope, p. A42. 1971b Binatisphinctes (m. Okaites) comptoni [Pratt); Mangold, pp. 123, 199, 202, 220. 19711> Binatisphinctes (M. Binatisphinctes) jluctuosus (Pia t t ) ; Mangold, pp. 199. 202, 220. 1971 Binatisphinctes comptoni (Pratt) and B.jluctuosus (Prat t) ; Callomon in Callomon &. Cope, p. 167.

Hololype. Bristol Ci ty M u s e u m Ca7088; Oxford Clay [Acut is t r ia tum Band, Athle ta Zone] , Chr is t ian Mai ford, Wil tshire . Figured Buckman 1924, pi. 485 (see Morley Davies 1924, p. 258). Microconch.

Material. About 70 specimens, mainly adu l t and complete, but crushed almost flat in shale, with white aragoni t ic shells, some part ial ly pyrit ized; 4 specimens (mainly f ragments) par t ia l ly crushed, in f i n e g r a i n e d sands tone . Microconchs and macroconchs .

Description. Microconchs (57 specimens) : m a x i m u m d iamete r 102 mm (holotype) , general ly between 60 and 80 m m (Text-fig. 18). Body-chamber 180° to 220°. Six whor ls visible. Evolute ( w h / d —c. 3 0 % ) ; whorl section compressed at least on middle and adult stages; venter na r row with weakening of ribs and smooth band , at least in par t . F rom at most 5 trim diameter , r ibbing (Text-fig. 16) general ly s t ra ight and dense, rect i radiate to prors i radia te becoming slightly rurs i radia te nea r the ventral marg in . P r imary ribs generally long and slender, spli t t ing on the outer half of the whorl side into two or three secondaries with intercalatories, interspersed with simple ribs. R ibb ing becomes more widely spaced on the body-chamber with increasing rur-siradiate aspect towards the ape r tu re which bears long, nar row lappets (t. 5X30 trim). Weak deve lopment of pa rabo lae on both inner and outer whorls. N u m b e r of pr imar ies increases with growth u p to a d i amete r of c. 30 mm (from c.. 30-50 at 10 m m . tor . 35-60 at 20 m m , toe. 40 65 a t


30 m m ) , thereaf ter decreases regularly, though some rimes abrupt ly , to between 30 and 40 on the last whorl (Text-fig. 17).

Macroconchs (18 specimens): m a x i m u m d iamete r 250 m m , generally between 200 and 235 m m . Body-chamber 180° to 210°. At least six whorls visible. Evolute ( w h / d = c . 3 0 % ) . O n first four whorls , p r imary r ibs generally long, s traight and dense, recti radia te to prors i radia te ,

Acutistr iatum Hand Comptoni Bed

1 - BGS FR970 2 = B G S F R 1 3 2 0 3 = BGS FR1045 4 = BGS FR 9 3 8

5 = YM INN 887

BGS „ FR 1020

BGS GSM

30348 OUM J17889

— B G S ^ ^ . GSM YM607 3 0 3 1 9

j 80

BGS GSM

' 30344

-L 9 0 1 0 0

diameterimm) T K X I -MC;. 17. Rib-density curves: I b r a d u h microconch Bmatisphindts rnmpkm.i (Pra t t ) .

splitting on the outer half of the whorl side into two or three secondaries which are scarcely visible; s imple ribs and intercalatories also present with weak parabolae (Text-fig. 16). Pr imaries then becoming (by 80 m m diameter) increasingly s t rong and somewhat thickcncd until on the last two whorls, r ibs a re s t rong, simple, massive, widely spaced (20 to 25 per whorl) ; they apparen t ly do not cross the venter which remains smooth. Aper tu re simple. Aptychi present .


d (septate to'l wh wt uw pr imaries ventra ls BGS G S M 3 0 3 4 9 (PL 14, fig. 6) 77 (c. 52) 22 32 36 102 BGS PR 1233 (PI. 14. fig. 7) 70 ( - ) 22 30 HGS PR 1297 (PI. 15. fig. 1) 67 (c. 44) 23 30 r. 40 c. 210 BGS FR970 (PI. 15, fig. 2) 55 (c. 33) 18 — 24 37 7 0 + O U M J 17889 (PI. 15, fig. 3) 75 (c. 50) 24 — 33 c. 38 c. 100 YM 608 (PI. 15, fig. 4) 52 (c. 38) 15 24 34 c. 94 YM 607 (pi. 15, fig. 5) 87 (61) 30 — 34 31 116 BGS GSM30342 (PI. 15, fig. 6) 52 (59) 15 — 27 34 c. 85 BGS G S M 3 0 3 4 j (PI. 15, fig. 7) 86 (59) 27 — 37 2 4 + 101 BGS F R 1020 (PL 15, fig. 8) 78 (50) 29 33 42 96 + D M 2143 (PL 10, fig. 1) 235 (—) 62 — 95 c. 26 O U M J30868 (PI. 17, fig. 1) c. 220 (—) 115 c. 22 BGS GSM30351 (PL 1 7, fig. 2) 133 ( - ) •10 — 60 24 B M 32343 (PL 10, fig. 1) 250 ( - ) 75 120 c. 20

Remarks. Al though B. comptoni (microconch) and B. finetuosus (macroconch) were the only species apar t from the type assigned to Binatisphinctes by Buckman, their position in this genus has been open to doub t . Spa th (1931, p . 237) thought that the comptoni g roup connected Binatisphinctes with Choffatia and belonged to the assemblage for which Grossouvria Siemiradzki was mainly in tended. H e thought tha t Binatisphinctes should he retained for the g roup of evo lu te ' Grossouvria' of the mosquensis g roup , which included B. binatus and Hamu/isphinctes. However , he stated (p. 392)

BIXA'1 ISPIIINCTES 45

that " m a n y . . . examples of Pra t t ' s Amm. lomploui. . . are undoubtedly fiinalispbinctesoC the same \=z-rjasattensis'Vctescyre] g roup . . Cal lomon (1955, p. 233) also doubted the sys temat ic position a\'comptoni and allied species, and thought they were probably assignable to (iro\\nu;;ria ra ther than to Binatisphinctes.

Siemiradzki (1898) compared the species comptoni with mosquensis. rjasanensis a nd scop in ens is, all since recognized as typical microconch Binatisphincles and formerly grouped together in Okaites. There is no doub t that B. comptoni is correctly grouped with these Russian and Polish species, but

the figured specimens of their type mater ia l a re all incomplete and vvithoui examinat ion of more complete mater ia l , a more explicit s ta tement of similarity cannot be made, liven their recognition as definite microconchs must be considered speculative.

Binatisphinctes comptoni I ml

• - — I W - • | • • - ~ I I I M I L I A I • • M I I • • I « • • • •• -*»•

4 0 - 4 4 4 5 - 4 9 5 0 - 5 4 55 -59 6 0 - 6 4 6 5 - 6 9 /0 74 7 5 - / 9 8 0 u 4 h5 h9 9 0 - 9 4 9 5 99 100 104 d id m e t e r (mm)

Compton i Bed

A c u t i s t r i a t u m Band

c u m u l a t i v e to ta ls

T E X T l i e . ,

typo]. 18. Size distribution lor adult microcoiidi BmatkfflwicM comptmi (Pra t t ' |")8 s p e d in ens including Prat t ' s

Siemiradzki (1098) and some later au thors (e.g. Corroy 1032, p. 127) wrongly included mbmutatus Nikitin, 1881 (now type species of 'Elalmites) with H. comptoni. T h e two species bear superficial resemblance because they are both based on lappeted microconchs of s imilar size, bu t the latter is an older form with a different r ib pa t tern . Some records of B. comptoni most p robably should be included with submutatus or its related species: for example Lowe's (1913) record from the Lower Callovian of Saxony ( i 4Macrocephalei t tonenM) and K u h n ' s (1939) record of comptoni and Jluctuosus f rom the Lower Callovian of southern G e r m a n y (Kranconia) K u h n ' s figure of comptoni. showed only inner whorls and he equaled Jluctuosus with leptus Gemmel la ro . now con-sidered to be a Homoeoplanulites (Mangold 1971 b, p. 184). Cal lomon 's (1955) record ocomptoni and jluctuosus from the J a s o n Zone of Kidl ington, Oxfordshi re also probably belong with suhmulatus or its related spef ics. Siemiradzki (1898) also included jliuluosus in his monograph and assigned it to Perisphinctes s.s. His record f rom the Eisenoolilh (Lower Callovian) of Krakow, Poland is now exc luded from I he species.


Within the assemblages of B. comptoni f rom the Compton i Bed and Acut i s t r ia tum Band, there appears to be a n u m b e r of individuals that , a l though having similar inner whorls to a d i ame te r of c. 30 m m (cf. r ib curves in Text-f ig. 17), ma tu re earlier and consequently have a smal ler adul t d iameter (40 (SO m m . generally 4 5 - 5 5 m m ) . However , forms in termedia te in size between them and the ' no rma l ' comptoni do occur and there appears to be no dispar i ty in the s t ra t igraphica l dis t r ibut ion. At least 12 specimens of this smaller form have been identified, and three arc figured (PI. 15, figs 2, 4. b). T h e r e is insufficient macroconch material to detect any significant or comparab le size var ia t ion therein. In general terms, forms from the Compton i Bed arc smaller (mean d i a m e t e r = 6 5 m m ) than those of the Acut i s t r ia tum Band (mean d i a m c t c r = 7 4 m m ) but there is complete over lap in the variat ion f rom each bed (Text-fig. 18). A real difference between the two popula t ions based on adul t d iameter is not proven by statistical methods . C o m p a r e d with B. binatus, B. comptoni [m] is smaller, does not have constrictions, and has generally more finely r ibbed inner whirls.

Distribution. Oxford Clay (Acut is t r ia tum Band (Athleta Zone) and C o m p t o n i Bed (Cor-onatuni Zone)) of southern and central England; I .angdalc Beds (Coronat ion Zone) of Gr is thorpe and Row Brow Q u a r r y near Scarborough, Nor th Yorkshire. Kopik (in ka rczewski 1970, p. 41) recorded the species f rom Poland, based on Sicmiradzki ' s (1898) record and Stoll 's (1934) record of B. comptoni in the Midd le Callovian of Pomeran ia .

Binatisphinctes hamulatus (Buckman, 1921) PI. 18, figs 2-fi; PI. 19, fig. 1 1921 Hamrdisphwrits hamulatus, no v.; Buckman. pi. 262 A, B. 1921 Hamuli sph inctts auricula, nnv.; Ruckman, pi. 263. 1924 IlamulisphincUs sp. [A maritimus, Bean MS.); Spath, p. 13. 1931 Biriutisphirtctes { — ' Hamidisphindes') maritimus (Bean MS}: Spath, p. 371. 1931 Binatisphinctes (—' Hamulisphinites') hamulatus S. Buckman; Spath , p. 371

non 1931 Pseudopeltoceras leckenbyi (Bean MS}; Spath. pp. 557, 572 [sec below] a IT. 1936 Binatisphinctes Welschi nov. sp.: Gera rd & Conta in , p. 61, pi. 11, tigs 2. 3.

1936 Pseudopeltoceras leckenbyi Bean sp.; Gerard & Contain , p. 68. pi. 17. 1951 Pseudopeltoceras cf f.eckeubyi Bean ap . ; Jcanne t , p. 176. pi. 85, text-fig. 411. 1971b Itamuiisphinctes (m. Ilamulisphincles) hamulatus Buekman, 1921; Mangold, p. 211. fi^s 155, 6; 158. 9. 1971b Hamulisphirutes (M. Pseudopeltoceras) leckenbyi (Bean MS); Mangold, p. 214. tigs 153, 157. 1971 Ilamulisphincles hamulus [sicJ Buckman; Car iou, p. 453.

Type specimens. Holotype. S M J3306, PI. 18, fig. 6, originally designated Buckman 1921, pi. 262A. Microconch with about 225° of body-chamber but without its aper tu re preserved. From

Hackness Rock], PAthleta Zone, Scarborough, North Yorkshire. Preserved in i black ooliths. T h e para type , S M J3307 , PI. 18, fig. 3, is preserved in grey shelly

l imestone with rhvnchoncl l ids , comparab le with the doggers of bed 4 (Athleta Zone) of Hackness Q u a r r y (Wright I960, p. 392).

Material. Eight microconchs including types, uncrushed . preserved in sands tone with chamosite ooliths, or l imestone; one possible macroconch .

Description. Microconchs: m a x i m u m d iamete r c. 90 mm. Body-chamber ar least 225°. Six whorls visible. Evolute ( w h / d = c . 2 5 - 3 0 % ) ; whorl section generally rounded or slightly depressed ( w h / w t = 9 0 - 1 0 0 % ) but may be more depressed on inner whorls (wh /wr > 7 0 % ) (Text-fig. 15); umbil icus shallow and wide (c. 38 m m wide and 8 m m deep at 75 mm diameter ) with steeply sloping umbilical walls, smooth on the outer whorl , r ibbed to umbilical seam on the inner and middle whorls. O n inner and middle whorls , r ibbing (Text-lig. lb) mainly gently prorsiradiate becoming rurs i rad ia te nea r ventral marg in , bifurcate with intercalatories (pr im-ary: sc co ndarv ratio of 1:2-5); s imple ribs and weak constrictions may also occur; oil ou te r whorl , ribs become more widely spaced remain ing mainly bifurcate, with intercalatories and t a re s imple ribs bu t developing rurs i rad ia te aspect which may be marked. Ribs on venter a lmost invariably in ter rupted by smooth band . Aper ture bears lappets. N u m b e r of pr imar ies increases with g rowth up to c. 30 m m (from c. 30-35 at 10 m m , toe. 35 45 a t 20-30 mm) , thereaf ter decreases regularly to 25-35 on the last whorl .

Cast le Rock sands tone wit

BINATISPHINCTLS 47

d (septate to) wh wt uw pr imar ies ' 87 ( - ) ' 21 24 45 c. 36

30 ( - ) n / 10 16 39 c. 68 (—) t-. 16 6'. 18 a 40 80 (c. 55) 22 22 40 24

75 (54) 22 20 38 28 225 ( - ) 60 54 110 24

Possible macroconch: m a x i m u m d iamete r seen 225 m m . with 70° body-chamber . Evolute f * (wh/d=<?. 2 5 % ) ; whorl section squar ish to slightly compressed ( w h / w t — 1 1 0 % ) . Ribb ing on later whorls, s t rong, simple, s t ra ight becoming thickened and massive (24 per whorl) .

Data for figured specimens. ventra ls

Y M 614 (PI. 18, 2) 87 (—) 21 24 45 c. 36 est. 70 SM J3307 (PI. l / , fig. 3) J K W A C / 6 6 (-P1. 18, fig. 4) B M 39530 (PI. 18, fig. 5) 80 (c. 55) 22 22 40 24 est. 78 S M J3306 (PI. 10. fig. 6) BM C I 0 7 8 (PI. 19, fig. 1)

Remarks. Following Arkell (1957a), hamulatus is included in Binatisphinctes. Hamulisphincles auricula Buckman , 1921 and the specimen bearing the Bean M S n a m e maritimus (PI. 18 fig. 5) cited by Spa th (1924; 1931) are also included in B. hamulatus. T h e type specimen o f / / , auricula is a lappeted microconch (SM J3308) , and as no new record or informat ion abou t it has been added since the original citation, it is considered to be a small example of/?, hamulatus. T h e pinkish shell material of the II. auricula type is, however, reminiscent of Lamber t i Zone preservat ion, and the possibility tha t this form is a late variety of real genetic significance a n d / o r which is s t rat igraphical ly useful should not be overlooked. .4. maritimus is ra ther more involute than the type specimens of B. hamulatus; it may be more appropr ia te ly placed in Pseudopeltoceras Spa th when that genus is more comprehensively descr ibed.

T h e macroconch form of hamulatus was included previously in Pseudopeltoceras leckenbyi (Bean MS) Spath , 1931 (Mangold 1971b). However , the type specimen of P. leckenbyi, B M 46516 from the Hackncss Rock of Scarborough has never been figured and it has been mis in te rpre ted , following Cera rd & C.ontaut (1936) who based their identification on Spath ' s (1931, pp. 557, 572) few words of descr ipt ion. T h e type specimen (figured herein PI. 20, fig. 1) shows distinctive curved prors i radia te ribs each wi th an incipient tubercle at the vcntro-latcral margin . T h e macroconch hamulatus is therefore now rccognizcd as the spccics leckenbyi Gerard & Gontau t , non Spa th . Fur the r examples of this form were described and figured by J c a n n c t (1951) and Mango ld (1971b). J e a n n e f s species rollieri (1951. p. 176, pi. 87) and zuberi (1951, p. 176, pi. 86) a p p e a r to be o ther related non- tubercu la te Pseudopeltoceras which should be regarded as Binatisphinctes macroconchs . Mangold (1971b, p. 211) in fact grouped these three macroconchs together, wi thin Pseudo-peltoceras, point ing our the similari ty of at least the inner whorls to Binatisphinctes-Okaiies. H e did not recognize any macroconch Binatisphinctes in his J u r a faunas, but described five microconch species. Only one possible macroconch specimen of B. hamulatus from England is recorded here (PI. 19, fig. 1); its inner -middle whorls are very poorly preserved and therefore ir is only tentatively assigned to the species. Ca l lomon (pers. comm. 1985) views this specimen as a "splendid typical Pseudopeltoceras [ M J".

B. hamulatus is dis t inguished from both B. binatus and B. comptoni by its coarser and ra ther more irregular and rurs i rad ia te r ib style, and more depressed whorl section. B. welschi Gerard & Con taut appear s to be a closely related microconch form.

Distribution. Hackness Rock (Athleta-• 'Lambert i zones) of Cornel ian Bay. Osgodhy Nab and Scarborough (Cast le Hill), Nor th Yorkshire. Recorded from France (Athleta Zone of the southern J u r a and west-central F i a n c e (Cariou 1971)) and f rom the Uppe r Cal lovian of I Ie rznach , Switzer land.

Binatisphinctes binatus (Leckenby, 1859) PI. 21, figs 1, 2, 4, 5 1859 Ammonites binatus. Bran. MS; T.cc.kcnhy; p 10. 10/11 Rin/itisphindes binatus, Bean-Leckcnby sp.; Buckman, pis 261 A, B. 1931 Binatisphinctes' binatus (Bean M S ) Leckenby sp., Spath . p. 327. 1935 Pnisphindes binatus Leek.; Model & Kuhn , pp. 478, 9, 481.


1939 Grossouvria (Binatisphinctes) binata (Leckenbv); Arkell, p. 180pais 1957a B. binatus (Leek.); Arkell, p. 319, figs 405, 6a-c . 1971 Binatisphinctes (M. Binatisphinctes) binatus Buckman; Mangold , p. 198.

Types. Lectotypes S M J 3 3 0 4 (PL 21, fig. I), designated Buckman 1921, pi. 261 A. F rom Cast le Rock | Hackness Rock], /Lamber t i Zone. Scarborough. Nor th Yorkshire. ?Microconch preserved in sparsely oolitic l imestone with some pinkish shell material a t t ached . Wholly septate paralcc-totype, SM J3305 (PL 21 fig. 2), preserved in reddish stained sands tone with black chamosi te ooliths and Meleagrindla\ /Ath le ta Zone.

Material. 'The type specimens and nine others, incomplete, mainly inner whorls or juveni les but generally uncrushed , preserved in sandy l imestone usually with chamosi te ooliths. O n e addit ional f r agment of partially crushed internal clay mould. Microconchs .

Description. Microconchs (eight specimens) : m a x i m u m diameter amongs t incomplete mater ia l c. 85 mm. Body-chamber at least 170°. Six whorls visible. Evolute ( w h / d = c . 3 0 - 3 5 % ; for inner whorls 2 0 - 2 5 % ) ; whorl section rounded or slightly depressed ( w h / w t = c . 9 0 % ) becoming more compressed on later whorls ( w h / w t = 120-140%) and with flat whorl sides (Text-fig. 15); umbilicus shallow and wide (c. 35 m m wide and 6 m m deep at c. 75 m m diameter ) with ra the r steeply sloping umbilical walls and ribs extending weakly down to the umbilical seam. O n inner and middle whorls , r ibbing (Text-fig. 16) gently prors i radia te becoming rurs i radia te near ventra l margin, a l ternat ing b i furca te a n d simple with dense, long, slender, s t raight pr imaries; on outer whorls, ribs of s imi lar aspect , bu t n u m b e r of secondaries increases; pr imaries remain ra the r long and well defined, spl i t t ing into three, or two with intercalatories, on the outer half of whorl side; pr imaries become more widely spaced towards aper ture which is not seen. Weakly developed constrictions and parabol ic ribs, with nodes at ventral marg in , somet imes occur. Ven te r becom-ing nar rower with growth , with weakening of ribs and usually a ventral smooth band th roughou t . N u m b e r of pr imaries increases with growth up to a d iameter of c.. 40 m m (from c. 35 45 at 10 m m , to c. 40-50 at 20 m m , to c. 45 -55 at 30 mm, to c. 50-55 at 40 m m ) , thereaf ter decreases regularly, p robably to between 30 and 40 on last whorl.


d (septate to) wh wt uw est. 82 (—) c. 29 20 est. 35

63 (63) 23 c. 18 25 32 (—) 7 8 17 77 (58) 23 20 35

pr imar ies ventrals SM J3304 (PL 21. fig. 1) S M J3305 (PL 21. fig. 2) W M S J-19-1 (PI. 21, fig. 4) 32 (—) 7 8 17 45 Y M 615 (PL 21, fig. 5) 77 (58) 23 20 35 37 c. 103

Remarks. Despite be ing type species of Binatisphinctes, it is one of the least well known. It is unclear f rom Lcckenby's original descript ion and Buckman ' s .subsequent. (1921) figures whe the r the type specimens are macroconch or microconch. T h e lectotype is incomplete and broken, bu t septate to at least 70 m m ; the paralec.totypc is completely septate . Opin ion has varied; Spa th (1931, p. 327) considered them to be microconchs belonging with " t he similar "Hamulisphinctes to Grossouvria of the mosquensis g roup . T h e latter was used subsequent ly as the type of the microconch 'genus ' Okaites. Arkell (1957a p. 319) apparen t ly considered Binatisphinctes, based on binatus, to be a macroconch g r o u p but included Haniulisphinc.tes, which accommoda tes microconch forms, in synonymy. Mango ld (1971b, p. 199), following Gerard & C o n t a u t (1936 p. 60), considered binatus to character ize a macroconch group , but I do not consider that his a r g u m e n t s justify this. Although no conclusive s ta tement can be made , it seems more likely that the type specimens are microconchs. As type species ol Binatisphinctes. the mat te r is of some impor tance , part icularly for those who incorpora te d imorph ism at subgeneric level. If one supposes that as defined by Buckman, Binatisphinctes included both macroconch and microconch, a n d one restricts Binatisphinctes s.s. (with binatus as type) to macroconchs and B. (Okaites) to microconchs. as done by Mangold (1971b), then a new n a m e is required for the Hackness Rock microconchs; but this group) probably includes the type of binatus.

ALLIGATICERAS 49

T h e microconch o[ B. binatus resembles most closely the microconch of B. comptoni, but it is p robably slightly larger in s i ze—up to 120 m m diameter , compared with 95 m m ; B u c k m a n thought tha t the lec totype of B. binatus would be 120 m m diamete r if complete , and the para lec-totypc 110 m m . In addi t ion , it has constr ic t ions, albeit weak, whereas B. comptoni does not , a n d a l though the rib curves for the few avai lable spec imens fall wi thin the range shown for B. comptoni (Text-fig. 1 7), B. binatus r ibb ing seems to be slightly coarser on the inner whor ls relat ive to its la ter whorls . Xo mac roconch is known f rom Eng land . B. binatus, like B. comptoni belongs to the s a m e g roup as the Russ ian and Polish species mosquensisrjasanensis and scop mens is but more deta i led compar i son with these forms is not possible (see above) .

Distribution. Hackncss Rock (Ath lc ta -Lamber t i Zones) o f C a y t o n , Gr i s thorpc , Hackness and Scarborough , Nor th Yorkshire . Rare in Lambcr t i Limestone (Lamber t i Zone) of Wood h a m , Buckinghamshi re . Recorded f rom N o r m a n d v , F rance (Arkell 1939, p. 180) a n d sou the rn Ger-many (Athleta Zone) (Mode l & K u h n 1935,'p. 478).

Type species. Ammonites alligatus (Bean MS) Leckenby. 1859 Diagnosis. Evolute forms with squar i sh or r ounded whorl section (Text-fig. 19); regular , often

wiry, mainly b i furca te r ibb ing and s t rong constr ict ions (Text-fig. 20). Lappc t cd microconchs . Externa l su tu re has iirst lateral saddle with a wel l -marked and qui te deep accessory lobe, a trifid slightly a symmet r i ca l lateral lobe slightly deeper than the ventral lobe and a r a the r shor t slightly retracted suspensive lobe with some well defined auxiliaries.

Remarks. B u c k m a n (1923, p. 157) gave no formal descript ion when he created the genus to a c c o m m o d a t e the single species Ammonites alligatus described by Leckenby (1859, p. 9. pi. 2 : figs 2a, b) f rom the 'Kel loway Rock' a n d 'Sca rborough Cast le Rock1 | Hackness Rock; Athlera-Lamber t i zones] of Nor th Yorkshi re ( B u c k m a n 1921, pi. 212). A p a r t f rom this f igure with its

TEXT-HI,. 19. Whor l sections of Alligaticcrasspecies, (a) S M J 3 3 0 3 a t II m m ; <b) B G S Z t 5 1 6 a t 70 m m ; (c) O U . M J 2 5 8 6 at 19 m m : id) M M LI 1191 a v 3 3 m m ; (el BGS7.13495a t57 mm: (f) O U M J l 2 5 0 a t 19 mm: (g) Y M f> 13at 88 m m ; (h) BGS Z1.3 703 at 65 m m .

cryptic d imens ion detai ls , the only in format ion B u c k m a n gave abou t the genus was in dist in-guishing it f rom Dichotomoceuis to which genus he originally referred alligatus ( Buckman 1921, pi. 212, emend . Oc t . 1927). l i e descr ibed it as having "vers i - rad ia te" [recti r ad ia t e of cu r ren t usage | ribs, pa r abo l ae and a s u t u r e wi th only a small accessory lobe on the external (ventral) sadd le and the second lateral lobe smal ler ( ra ther than longer as in Dickolomoceras) t han the first auxil iary lobe. Arkell (1936, pp . xlii-xliii) believed this 'geno-holotypc ' to be apparen t ly adul t at a d i a m e t e r of 45 m m , with the last th ree -quar te r s of the ou te r whorl occupied by body-chamber ; the r ibb ing undergoes no modif icat ion towards the ape r tu re . Al though not preserved on the type spec imen ,

Subfami ly Pr .Kt.sni iNCTiNALStcinmann. 1890 G e n u s ALLICATICERAS Buckman , 1923

O r o t i f e r


Arkell thought lappets would be present on complete adult individuals. It seems cer ta in that Alligati.ee/as a ccommoda ted a microconch group.

Aliigaticeras is taken to include microconch and macroconch forms; a l though Mango ld (1971b, p. 221) did not descr ibe the systematics of Aliigaticeras he suggested that a t least some of its macroconchs had been included in Orionoides Spa th , 1931. Collections from west-central F rance might provide an answer , because the s t ra t ig raphy is well known, and there are records of alligatum and associated forms, and published il lustrations (cf. Gerard & C o n t a u t 1936) which include forms assigned to Orionoides. C c r a r d & Con ta in ' s (1936) species O.piveleaui, 0. raguini and 0. termieri could be Alligatoceras macroconchs .

Aliigaticeras alligatum

Aliigaticeras rotifer

Alligatfcoras a f f . rotifer

inner whorls

middle whorls

occas ional parabolae

outer whorls — body chamber

I 1MI

deep cons t r i c t i ons

deep cons t r i c t i ons

weak co istr ict ions

T E X T n o . 20. Ribbing of Alligathnos spccics.

Aliigaticeras includes the Knglish species alligatum and rotifer of the U p p e r Cal lovian; in the Lower Oxford ian pseudograciosus Arkell, 1939 belongs to this g roup and forms identified as bemensis de Loriol, the type species of Hroperisphincles (Spath , 1931), mus t be very closely related (see PI. 23, fig. 8; Arkell 1936,.pp. xli-xlii) , a l though this genus is based only on pyritic nuclei. Forms from India occurr ing a t a s imilar horizon and assigned to the genus by Spa th (1931) include A. jooraense (Waagen) , A. nbliqueiplicatum (Waagen) , A. polymorphum Spa th and A. indicum Spa th . Arkell (1936, p. xliii) added several o ther species from the Lower Oxfordian of the J u r a including ,4. birmensdor-

fen sis (Moesch) , A. noellutgi (de Loriol) and .4. mairei (de Loriol). Cal lomon (pers. comm. , 1985) considers A. bimensdorfemis to be f rom the Midd le Oxford ian T ransve r sa r inm Zone, and not closely related to the others.

Aliigaticeras is thought to have evolved from Choffatia du r ing the late Callovian ( Ath le ta Zone) ; .4. rotifer shows transi t ional charac ters (cf. Choffatia poculum) and is probably an early form. T h e g roup persists into the Oxford ian where it is c o m m o n amongst the per isphinct id f auna of the Lower Oxford ian in m a n y sub-Boreal and more southerly areas.

ALL IG ATI CE R A S

Alligaticeras rotifer (Brown, 1849) PI. 22, figs 1-6, 8

1841 Ammonites rotifer (nob.); Williamson, p. PI7 [nom. nud.]. 1849 Ammonites rotifer; Brown, p. 246. pi. 20* figs 14, 15. 1918 Perisphinctes rotifer Will iamson-Brown sp.; Buckman, pi 113, pp. 113b, c.

?1971 Grossouvria rotifer (Will iamson); I l a h n & Koerner , pi. 12.

Holotype. M M LI 1494 (PI. 22, fig. 1), des igna ted B u c k m a n 1918,pi . 113 f rom 'Ke l loway Rock ' [Hackness Rock: A t h l e t a - L a m b e r t i zones] , Cas t l e Hill, Sca rborough , Nor th Yorkshi re . Pre-served in a highly ca lcareous oolitic sands tone ; it a lmost cer tainlv comcs f rom the Ath le ta Zone (cf. Bed 3 at Cas t l e Hill; W r i g h t 1968).

Material. T h e type spec imen and a b o u t 18 others, unc rushed but incomple te or i m m a t u r e (completely sep ta t e ) , preserved in ca lcareous a n d / o r oolitic s ands tone or in shelly ca lcareous doggers; and a b o u t 35 spec imens , ma in ly small , unc rushcd but complete ly sep ta te ( inner whor ls ) , pyri t ized in ternal moulds .

Description. M a x i m u m d i a m e t e r a m o n g s t incomple te mater ia l 90 m m . B o d y - c h a m b e r at least 180". At least four a n d more often five whor l s visible. Modera te ly evolute ( w h / d = 3 0 - 3 5 % ) ; whorl section modera t e ly depressed ( w h / w t = 6 5 - 8 5 % ) with gent ly rounded whorl sides a n d broad venter ( Text-fig. 19); s teeply s loping and a lmost smooth umbil ical wall ; umbi l i cus modera te ly shal low and wide (c. 2.5 m m wide and 8 m m deep at c. 55 m m d i a m e t e r ) . Ribs (Tex t -fig. 20) general ly p ro r s i r ad ia te ( though inner whorls m a y show slight ru rs i rad ia te aspect near the venter ) , b i furca te on inner whor ls (up to 25 m m d iame te r ) , becoming b i furca te with intercala-tories and occasional s imple ribs (up ro 35 to 40 mm d iamete r ) and then (more than 40 m m d iamete r ) t r i furca te with in tercala tor ies giving a p r i m a r y to vent ra l ra t io of u p to 1:4. Ribs split high up on the whorl side (on the vent ro- la tera l shoulder of depressed whorls) and pass un in t e r rup t ed across the venter . Modera t e ly s t rong arid deep constr ic t ions are c o m m o n on the inner a n d middle whor ls (usually two per whorl) . N u m b e r of pr imar ies decreases wi th g rowth from 3 0 - 5 0 at 2 0 - 2 J irun d i ame te r , to 3 0 - 4 0 at 30 -50 m m d iamete r , to c. 3 J at 5 0 - 5 5 n u n d iamete r , to c. 30 at 90 m m d iame te r .

Data Jot figured specimens.

M M LI 1494 (PI. 22, fig. 1) Y M 613 (PI. 22. fig. 2) O U M J 1 2 0 4 (PI. 22, fig. 3) O U M J1250 (PI. 22, fig, 4) B G S Y2125 (PI. 22. fig. 5) BGS Zt3495 (PI. 22. fig. 6) O U M J 2 6 3 6 0 (PI. 22, fig. 8)

Discussion. B u c k m a n (1918, pp . 113b, c) discussed A. rotifer in cons iderably m o r e detai l t h a n any of his own species, quo t ing Wi l l i amson ' s (1841) original nomina l ci tat ion (in a f auna l list, wi thout f igure or descr ip t ion) a n d Brown ' s (1849) subsequen t descr ipt ion. As publ i shed in the former, the n a m e is a no men nudum, and the la t ter is the first valid publ ica t ion . Brown (1849) made-no ment ion of Wi l l i amson , bu t as indica ted by B u c k m a n (1918, p. 113c) the re is c i r cumstan t i a l evidence to suggest tha t Brown was descr ib ing Wi l l i amson ' s species. T h e ava i lab le a n d valid name is therefore Ammonites rotifer (Wil l iamson MS) Brown, 1049. Wi l l i amson (1841) gave its p rovenance as a 1-8 m thick ha rd i rons tone [p robably u p p e r p a r t j at Sca rbo rough Cas t l e Cliff, which B u c k m a n concluded to be the "grey sands tone of [the] Ath le ta Zone" . B u c k m a n gave his own descr ip t ion based on an examina t ion of the wholly septa te holotype, bu t evident ly could not relate it to any of the n e w generic g roups tha t he rccognizcd amongs t the Cal lovian per i sphinc t ids , and so assigned it to Perisphinctes. P e r h a p s bceausc of this, the spccics has since t ended to be overlooked. C a l l o m o n (pers. c o m m . 1985) believed tha t it was p robab ly the oldest avai lable n a m e for m a n y of the pyri t ized nuclei f rom the Brown J u r a zcta (Athleta Zone) of sou the rn G e r m a n y

d ( sep ta te to) wh wt uw pr imar ies veni ra l s 53 (53) 14 22 24 35 fc\ 100

88 (c. 65) 22 25 45 30 c. 75 <;. 40 (—) 13 c. 16 26 39 —

19 (19) 6 8 8 c. 36 c. 62 30 ( - ) 10 14 13 57 (57) 18 22 24 90 ( - ) (25) (16) 45


"described in the past , originally as Amm. convolutus of one sort or another , then r e n a m e d s tar t ing with Amm. convolutusgigas Q u . 1047. pi. 13. fig. 6, . . . Amm. conv. dilalatus, Qu . 1887, pi. 81 . . ."; he concluded that "rotiferdeserves more currency than it has had , and there has been no excuse post-Buckman ."

T h e species rotifer is thought to belong in Aliigaticeras not least because of the presence of the character is t ic s t rong deep constrictions, i ts more depressed whorl section and bundled rib-style suggest a position in te rmedia te between Choffatia and the typical alligatum g roup Aliigaticeras. I ts s t ra t igraphieal position suppor t s this in terpre ta t ion. It differs from Choffatia poculum (bu t the adul t microconchs seem very similar) in having the characteris t ic cleep constrict ions and more Strongly r ibbed inner whorls.

Distribution. Hackness Rock of Hackness ( including Bed 4 (Athleta Zone) of Wrigh t 1968), Cay ton and Scarborough Cast le , Nor th Yorkshire; Oxford Clay, Athleta Zone of S u m m c r t o w n and Wolvcrcotc , Oxfordshi re ; Athleta Zone, Proniac Subzone of Mil ton Keynes A 5 diversion, Buckinghamshire . Recorded f rom the Athle ta—lower Lamber t i zones of south-west G e r m a n y (Hahn & Kocrncr 1971).

Aliigaticeras aff. rotifer (Will iamson MS), Brown 1849 PI. 22, figs 7, 9; PL 23, figs 1, 2 1939 Perisphinctes (Proper isphificlei) tulitmgtiaius Noelling; Arkell, p. Hit), 195. pi. 9, ligs 3. 1969 Properisphwctes (?) sp.; Hudson & Palf raman, p. 414. pi. 19. lit*, f. 1969 Properisphinctes sp.: Hudson & Palf raman, p. 414, pi. 19, lig. h.

Material. T h r e e unc rushed , completely septate, pyritized specimens (presumed macro-conchs); 20 nuclei and inner whorls , similarly preserved and two similar specimens with outer whorls preserved as clay moulds . Microconchs and macroconchs .

Description and discussion. T h e inner whorls are indis t inguishable from those of .4. rotifer, bu t later whorls are less depressed ( w h / w t = 9 0 - 9 5 % ) (Text-fig. 19), more finely and densely r ibbed , and constrict ions arc less deep (Text-fig. 20). Microconch body-chamber (c. 55 m m diameter ) with bundled ribs (pr imary:vent ra l rat io about 1:4) and lappets (up to c. 5 x 2 0 m m ) resembles that of Choffatia poculum. N u m b e r of pr imaries decreases with growth from c. 40 at 20 -50 m m diameter , to 30-35 at 65-75 m m diameter , to c. 35 at 90-110 m m diameter .

Arkell 's (1939, pp. 160, 161, 195) records ( O U M J22503, 4 and possibly O U M J22L26, 7, Arkell 1939, pi. 9. figs 2. 3) of latilinguatus Noctl ing 1887, from the Athle ta Zone of W o o d h a m belong to this group. With their small d iameter , they might equally well be assigned to .4. rotifer, but because of the associated more complete material from this horizon and locality, they are included herein with A. aff. rotifer. Noet l ing 's material came from the Mar iae Zone of Moun t Hermon , Syria and the i l lustrat ions (Noet.Iing 1887. pi. 4, fig. 7 8c) show septa te f r agmen t s and inner whorls only. Arkell (1938b) also used this species n a m e for the pyritized per isphinct id f ragments and nuclei ( O U M J22837 collective) f rom the Lamber t i Zone clays, exposed in the excavations for the Bodleian Library extension in Oxford ; however, a l though these included forms of the rotifer-alligatum bemensis g roup , they are now considered to be inde te rmina te at specific level.

T h e relat ionships of .4. aff. rotifer are. not clear; its more, compressed whorl section, finer r ibbing and slightly younger s t ra t igraphiea l position compared with .4. rotifer, migh t indicate a position in te rmedia te between .4. rotifer and .4. alligatum, but it has only a compara t ive ly weak development of constr ict ions.

ventra ls Da ta fo r fig u red specim ens.

d (septate to) wh Wt 11 w pr imaries O U M J29909 (PL 22. lig. 7) c. 53 (—) c. 17 — —

BGS Z13703 (PL 22, fig. 9) 65 (65) 19 20 29 —

O U M J29910 (PL 23, fig. 1) 110 ( - ) 33 — 47 c. 36 O U M J1206 (PL 23, fig. 2) 92 ( - ) 34 35 35 c. 36

Distribution. Oxford Clay ( Athleta Zone of Summerrown, Oxfordshi re ; Athleta Zone, Proniae

AI.LICATICERAS 53

Subzone of Milton Keynes . Buckinghamshire ; Athleta Zone. Spinosum Subzone o f W o o d h a m , Buckinghamshi re ) .

Alligaticeras alligatum (Leckenby, 1859) PI. 23, figs 3-7 1059 Ammonites alligatus Leckenby (Bean MS); Leckenby. p. 0, pi. 9} fig.?, 2a, b.

i'1897 Perisphinctes alligatus (Bean); Parona & Rnnarrlli, p 178. pi. 9, figs 1, la . 1898-9 Perisphinctes alligatus Leek.; Siemiradzki, p. 86. pi. 10. fig. 0, ?pl. 17, lit;. 58. 1921 Dichotomoceras alligatum, Bcan-Leckcnby sp.; Buckman, pi. 212. 1927 Alligaticeras alligatum, Bcan-Leckenby sp.: Buckma.il [corrigenda slip]. 1931 Alligaticeras alligatum (Bean MS) Leckenby sp.; Spath , p. -107.

?1933 Alligaticeras all", ulligutum (Bean MS) Leckenby sp.; Spath, p. 859. 1936 Perisphinctes (Alligaticeras) alligatus Leckenby; Arkell. p. xlii, pi 100. figs 4a—c. 1939 Perisphinctes {Alligaticeras) alligatus (Leckenby); Arkell, pp. 161, 100, 2(13.

? 1939 Perisphinctes bermnsis Lor.; Knhn , p. 509. 1957a Alligaticeras alligatum (Leckenby): Arkell, p. 319. lii>. 407. 1971 Alligaticeras alligatum Leckenby; Cariou, p. 453. 1971 Alligaticeras alligatum Leek.; (Janon. p. 309.

Holotype. S M J 3 3 0 3 (PI. 23, fig. 3), designated Buckman 1921, pi. 212 from 'Cast le Rock' [Hackness Rock; A t h l e t a - L a m b c r t i zones], Scarborough, North Yorkshire. Microconch. T h e preservat ion of this specimen (l imestone with pinkish shell mater ia l ) s trongly suggests that it comes from the Lambcr t i Zone. O t h e r specimens suppor t this—they are associa ted with Qitensted-locems a n d / o r come from Cuns tone N a b (see Wright 1968).

Material. The type specimen and about 10 others, uncrushed but mainly incomplete or i m m a t u r e (preserved in sandy l imestone with ooliths) and abou t 30 specimens part ial ly crushed or otherwise dis tor ted, including at least two adults , otherwise incomplete or i m m a t u r e and all more ot less f r a g m e n t a r y (preserved in calcareous muds tonc or marly l imestone as internal clay moulds with thin black (when fresh) powdery pyritic coat ings) . Microconchs and macroconchs .

Description. Microconchs (29 specimens) : m a x i m u m d iamete r c. 55 m m , generally 4 0 - 5 0 m m . Body-chamber a t least 270". At least four whorls visible. Evolute ( w h / d —25-30%); whorl section squarish (wh/wr~~80 100%) with flat whorl sides and broad ra ther flat or gently rounded venter (Text-fig. 19); inner whorls may be depressed; gently sloping and almost smooth umbilical wall; umbil icus shal low and wide (up to 25 m m wide and 5 m m deep in adul t ) . Ribs (Text-fig. 20) mainly recl i radia te but prors i rad ia te on the inner whorls and gently prors i radia te on the venter; apa r t f rom this change in t rend, r ibbing remains remarkably constant at all stages of g rowth with sha rp regular p r imar ies spl i t t ing very high on the whorl side, often at the ventro-lateral margin , into two secondaries; occasionally two (or more rarely one) simple ribs with an associated intercalatory. S t rong deep constr ict ions common , usually two per whorl, and occasional nodes a t the ventro-lateral margin . Ribs pass across broad venter usually wi thout in te r rupt ion bu t somet imes wi th vestige of weak smooth ventral band . Aper ture , with associated s t rengthened rib and terminal constr ict ion, bears lappets (apparen t ly short) .

Macroconchs (six specimens) : m a x i m u m diameter amongs t incomplete mater ia l (no speci-men seen to aper tu re ) c. 100 m m . Length of body-chamber unknown. At least four whor ls visible. Evolute ( w h / d = 2 5 - 3 0 % ) ; whorl section squarish ( w h / w t = 9 0 - 1 0 0 % ) with flat whorl sides and broad ra ther flat or gently rounded venter (Text-fig. 19); gently to modera te ly steeply sloping umbilical wall; umbil icus shallow and wide (est imated up to 50 m m wide and 10 m m deep in adul t ) . Ribs (Text-fig. 20) mainly rect i radia te , becoming gently prorsiradi te on the venter. As in the microconch, the r ibbing style apparen t ly remains remarkably constant with s h a r p regular pr imaries spl i t t ing very high on the whorl side, often at the ventro-lateral marg in , into two secondaries; occasional simple ribs with an associated intercalatory; s t rong deep constr ict ions part icular ly on middle and probably inner whorls. Ribs pass across b road venter wi thout in ter rupt ion . For microconch and macroconch, the n u m b e r of p r imar ies increases wi th growth from 10-50 at 20 -25 m m d iamete r , to (40) -60 al 30 -35 m m d iamete r to (50)—65 at 70-75 m m diameter .

54 E N G L I S H C A L L O V I A N P E R I S P H I N C T I D A M M O N I TES

Data far figured specimens.

d (septate to) wh wl uw pr imar ies ventra ls SM J3303 (PI. 23, fig. 3) 44 (29) 13 14 21 58 c. 112 O U M J258b (PI. 23, fig. 4) 1 9 ( - ) 5 8 9 40 YVMS J46 (PI. 23, fig. 5) 91 ( - ) 23 f. 22 46 —

BGS Z t 5 l 6 (PI. 23, fig. 6) c. 70 (—) 6". 18 20 c. 33 c. 47 c. 90 O U M J22384 (PI. 23* lig. 7) 36 ( - ) 9 c. 10 18 50 c. 90

Discussion. A discussion of Aliigaticeras has almost always involved .4. alligatum and for this reason the microconch form, albeit based on the holotypc only, has become fairly well known. T h e macroconch is less well known, and no complete specimen is known from England . Mos t o ther records of the species are not suppor ted by il lustration or systematic descr ipt ion. A. alligatum is more compressed, with finer and more regular bifurcate r ibbing than A. rotifer.

Distribution. Lamber t i L imes tone (Lamber t i Zone) o f W o o d h a m , Buckinghamshi re ; Hackness Rock (Lamber t i Zone and possibly Athleta Zone) of Cuns tone Nab . Cayton , Hackness and Scarborough, Nor th Yorkshire; ?Oxford Clay (Athle ta Zone of Wolvercote, Oxfordshi re ; ?Lam-berli Zone of T i d m o o r Point , Dorset (Spath 1933)). Recorded from France ( A t h l c t a - L a m b c r t i zones of Ca lvados (specimen figured by Siemiradzki 1098, pi. 20 fig. 8), Poitou, nor thwes t Aqui ta ine) , the J u r a , possibly as bernensis in the Athleta Zone of Franconia ( K u h n 1939) and doubtful ly f rom the "Eisenool i the" (Jason Zone) of Poland (Siemiradzki 1898).

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westdeutschcn J u r a . Jah.rh. geol. Lmdesamt Baden-Wiirltemberg, I I , 29-86. pis 1-9. & KOERNER, L . 19V1. Die Aufschliisse im oberen Dogger (Baihoniuui-Cal lovium) im Albstollen der Boden-

seewasser-versorgung uuier der Zol lernalb (SW-Deutsch land) . Ibid., 13, 123-44. MARKER, A. 1884. O n a r emarkab le exposure of the Kel lawav ' s Rock in a recent cut t ing near Cirencester . Proc. Cott.eswold

Nat. Field Club, 8, 175-87. MACG, E 1891. T.es chaines suba lp ines en t re G a p et Dignc. Bull. Sen.. Carte geol. Fr., 3 , no. 21. 1-197. HEDBERG, I I . D. (eel.) 1976. International stratigraphic guide. xv i i+200 pp. New York. HEMINGWAY, J . E. 1974. Jurassic. Pp . 161-223. In RAYNER, D. H . & HEMINGWAY, J . K. (eds) The Geology and Mineral

Resources of Yorkshire. 405 pp. 7 pis. Leeds. HUDSON. J . D. & PAI.FR A MAN, I). F. B. 1969. T h e ecology and preservation of the Oxford Clay f auna at Wood h a m .

Buckinghamshi re . Q. JI geol. Soc. Loud., 124, 387-418. pis 19-20. HOLLAND, C . H . et al. 1978. .4 guide to stratigraphical procedure Spec. Rep. Geol. Soc. Land. No. 11, 18 pp. INTERNATIONA!. COMMISSION ON ZOOLOGICAL NOMENCLATURE ( I C Z . N . ) 1985. In terna t ional C o d e oi Zoological

Nomencla ture 3rd edition, x x + 3 3 8 pp. London. JEA\-NET, A. 1951. Die Kiscn- und M a n g a n e r z e der Schwei/.. S i ia t igraphic und palaeontologie des oolithischen

Eisener / lage is von I l e r z n a c h und seiner I Jmgcbung. Beitr. geol. Karte Schcveiz, Geotech. Ser , no. 13, 5. xiii + 240 pp . , 107 pis.

KARCZEWSKI, L. (ed.) 1970. Geology of Poland. I I Ca ta logue of Fossils, Pt 2. Mesozoic. 234 pp. Warsaw. KEFPTNO. VV. &. MLDDLEMLSS. C. S. 1883. O n some new raiKvav sections and o ther rock exposures in the distr ict of Cave.

Yorkshire. Geol. Mag., ( 2 ) , 1 0 , 2 1 5 - 2 1 . KRENKEL, E. 1913. Die Ke l loway-Fauna von Popilani in VVesu ussland. Palaeontographico, 6 1 , 191 3 6 2 , pis 1 9 - 2 8 . K U H N . ( ) . 1 9 3 9 . Die Ammoni t en des f rankisehen Cal loviums. Nova Ada Levpoidina, n.s., 6 , no. 4 3 , 1 5 1 - 5 3 2 . pis I—10. LAHUSEN, I . 1883 . Die F a u n a de r Jurassischcn Bi ldnngen des Rjasanschen Gouvernemcnts . Tr. geol. Kom.. 1 , no. 1,

1 9 4 , pis l - l 1. LECKENBY. J . 1859. O n the Kelloway Rock of the Yorkshire coast. Q.Jlgeol. Soc. Land., 15, 4-15. pis 1 -3 I .OOZY, L . VON LOG/. 1915. M o n o g r a p h i c dei Villanvcr Callovien Ammoni ten . Geol. Hungarica, 1 , 255-507, pis 1-14. LORIOL, P. DF 1898. fitudesur les mollusques et brachiopodes de l 'Oxfnrdien in fe r i eu rou Zone a Ammoni tes rengerri

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36, 113-213. MAKOWSKI, H . 1902. La I'auue cal lovienne de L.ukow en Pologne. Palaeontol. Pol., no. 4, x + 64 pp . , 9 pis

1963. Problem of sexual d imorph i sm in ammoni tes . Ibid., no. 12. viii + 92 pp., 20 pis.

/ u /

KKFKKKNCKS * ^ iii

MANGOLD, C. 1971 a. S t ra t ig raph ie des etages Bathonien et Callovien du Jura meridional Doc. I.ab. Geol. Fac. Sci. Lyon, no. 11, (1), 1 -376 .

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QUENSTF.DT, F. A. 1845-1849. Petrej'aclenkunde Deutschlands. Die Cephalopoden. 1845, 1 104, pis 1 6: 1846. 1 0 5 - 8 1 pis 7-14; 1847, 185-264. pis 15-19; 1848, 265 472, pis 20 29; 1849, 173 580, pis 30-36. Tub ingen .

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UHLIG, V. 1870. Beitrage zur Kenn tn i s der Jnraformat ion in den Karpa t i schen Kl ippen . Jahrh. Kais.-Kbnig. geol. Reiihsanst.. 2 8 , 0 4 1 - 5 8 , pis 10-17.

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W ILLIAMSON, VV. C . 1841. O n the dis t r ibut ion of organic remains in i he s t rata of the Yorkshire coast from the U p p e r Sands tone tu the O x f o r d Clay inclusive. Trans, geol. .Soc. Loud. (2) , 6 , 143-52.

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R E F E R E N C E S v

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PLATES

Unless otherwise stated on the plate, the figures are reproduced at natural size; most specimens were coated with ammonium chloride before photo-graphy. Start of the body-chamber is marked ( > ) where visible.

P L A T E 1 Fig. Page

Homoeoplanulites c f . subbakeriae ( c T O r b i g n y , 1 8 5 0 ) 1 7

la , b BM CI 1796 (J. F. Blake Collection); [M] body-chamber f ragment ; Upper? Cornbrash ; Holwell , Dorset .

2a, b BM C I 1825 (J. F . Blake Collection); whorl f ragment ; Upper? Cornbrash ; Holwell, Dorse t .

3 O U M J12767; Uppe r Cornbrash ; Stalbridge, Dorset ; Macrocepha lus Zone and Subzone.

Homoeoplanulites aff. balinensis (Neumayr , 1871) 17 4a, b BGS GSM7676 ; completely septate; Kellaways Clay; Siddington, Gloucester-

shire; Calloviense Zone , Koenigi Subzone. -5 BGS GSM7679; completely septate; Kellaways Clay; Siddington, Gloucester-

shire; Calloviense Zone , Koenigi Subzone.

Homoeoplanulites difficilis (S. S. Buckman, 1922) 18 6a, b Holo type (presumed lost) as figured by S. S. Buckman (1922, pi. 329); [m];

Kellaways Rock; South Cave, Humbers ide; Calloviense Zone , Eno-d a t u m Subzone.

M O N O G R A P H OF T H E PALAEONTOGRAPHICAL SOCIETY

Plate 1 COX, Callovian perisphinctid ammonites

PLATE 2 Fig. Page

Hamoeaplanulites difficUu (S. S. Buckman, 1922) 18 I 3: 6a, b Topotypes; [m]; Kellaways Rock; South Cave Station Quarry. South Cave.

Humbcrsidc; Calloviense Zone, Enodatuin Subzone. L FW TNN475; 2, FW TNN473; 3. IKiS Zkl439 fall complete adults with lappets); 6a. b, FWTNN510.

4a, b, 5a-c Complete adults (m) with lappets; Kellaways Rock; South Ncwhald Quarry, South Ncwhald, I I umherside; Calloviense Zone. Ivnodalum Subzone. 4. FW TNN527: 5, FW TNN528.

MONOGRAPH OF THE PALAEONTOGRAPHICAL SOCIETY

COX, Callovian perisphinctid ammonites Plate j2

PLATE 3 Fig. Page

Ilomoevplanulitesdifficilis (S. S. Buckman, 1922) IK I FW TNN521; [M]; Kellaways Rock: South Newbaid, Hiirabcrsicle; Callovi-

ense Zone, Lnodatum Subzone. 2% 3 Topotypes; Kellaways Rock; South Cave Station Quarrv, South Cave, Hum-

be rside; Calloviense Zone. Enodaium Sub/one. 2, BGS GSM 117460, [m]> septate inner whorls and pare of body-chamber; 3, FW TNN508, adult [M|.

M O N O G R A P H O F T H E PALAEONTOGRAPHICAL SOCIETY \

COX, Callovian perisphinctid ammonites Plate 3

PLATli 4

HomneopUiniilUes difficilis (S. S. Buckman, 1922) FW TNN506; topotype; adult [MJ: Kellaways Rock; South Cave Station

Quarry, South Cave, Humberside: Calloviense Zone. Enodalum Subzone.


COX, Callovian perisphinetid ammonites Plate 4

PLATE 5 I'if?. Page

Perisphinctesspirorbis Neumayr, IK'70 20 la. b Plaster cast of holotypc; BGS GSM 117448; according to Neumayr, last whorl

almost all body-chamber with some mouth border bul no suture visible; Macrocephalcn-Schichten; Vogi&heim im Brcisgau, Baden> W. Germany; original specimen is BSM AS1838.

HomoeopIanuMes difficilis (S. S. Buckman. 1922) 1X 2 FW TNN516; topotype; adult [MJ; Kellaways Rock; South Cave Station

Quarry, South Cave, Humberside; Calloviense Zone, Lnodatum Subzone.



PLATE 6 Fig. Page

Indosphinctes lobatus (S. S. Buckman. 1922) 22 la. I> ItOS GSM7680] |M|; Kellaways Clay, Siddington, Gloucestershire; Callovi-

ense Zone. Koenigi Subzone. 2 BGS Zrl639: [M]; Kellaways Clay; Sutton Benger (M4 motorway cutting),

Wiltshire; Calloviense Zone, Koenigi Subzone. 3a, h BGS GSM7682; holotypc; complete adult [m] with lappets: Kellaways ('lay;

Rampiskam, Dorset; Calloviense Zone. Koenigi Subzone. Indosphinctespaiina (Neumayr, 1870) 24

A GBV 1870/04/1; lectotype; original of Neumayr 1870, pi. 8, fig. I ; [M]; Macro-cephalen-Kalke; Rrictal, near Gosau, Salzkammcrgut, Austria; see PL 7, lig. 1 for lateral view.

MONOGRAPH O F THE PALAEONTOGRAPHICAL SOCIETY


Fig. PLATE 7

Page Indosphinctes patina (Neumayr, 1870) 24

GBV 1870/04; 1: leclolypc; original of Neumayr 1870. pi. 8, lig. 1; (Ml; Macro-cephalcn-Kalke; Brictal, ne-tir Cosau. Salzkammergut, Austria: see PI. 6, fig. 4 for apertural view.

M O N O G R A P H OF THE PALAEONTOGRAPHICAL SOCIETY


PLATE 8

Indosphinctes palifia (Neumayr. 1870) BGS Z1I675; [M] body-chambcr; Lower Oxford Clay; Witney. Oxfordshire;

Jason Zone; lb, x 0-8. OUM J30911; adult [m| with broken lappet; Lower Oxford Clay; LUC pit,

Calvert, Buckinghamshire; Jason Zone. Jason Subzone (Bed 2e of Callo-mon 1968).

OUM J309I8; [mj; Lower Oxford Clay; LBC pit. Calvert, Buckinghamshire; Jason Zone. Jason Sub/one (Bed 2c of Callomon I96K).

OUM J30912; [mj; Lower Oxford Clay; LBC pic, Calvert, Buckinghamshire; Jason Zone. Jason Subzone (Bed 2c of Callomon l%K).



PLATE 9 Fig. Page

Choffatia cobra (Waagen, 1875) 26 la, b Plaster cast of holotype; BGS GSM 117449; |M|; 'Beds with Per. aneeps\

south of Bhooj. Cutclu India; original specimen is GS1 Type no. 2056.

Choffatia cardoti (Petitclerc, 1915) 29 2a~e FW TNN536; completely septate; Kellaways Sand; South Newbald Quarry,

South Newbald, Humberside; Calloviense Zone, Calloviense Subzone. 3a-c FW TNN537; no suture visible; Kellaways Sand; South Newbald Quarry,

South Newbald. Ilunihersidc; Calloviense Zone, Calloviense Subzone.


PLATE 10

Choffatia cordon (Petitclerc. 1915) IKiS <SSM49454; (MJ; Kellaways Sand: South Cave, Humberside; Calloviense

Zone. Calloviense Subzone.

PLATE 11 Fig. Page

Choffatia recuperoi (Gemmellaro, 1872) 30 la c FW TNN507; completely septate |MJ; Kellaways Rock; South Cave Station

Quarry. South Cave, Humberside: Calloviense Zone. Enodatum Subzone. 2a-e FW TNN460; adult [m] with lappets; Kellaways Rock; South Cave Station

Quarry, South Cave, Humberside; Calloviense Zone, Hnodatum Subzone. 3a c FW TNN512; adult [m]; Kellaways Rock: South Cave Station Quarry, South

Cave, Humberside; Calloviense Zone, Hnodatum Subzone. 4a-c FW TNN522; adult [m]; Kellaways Rock; Soulh Newbald Quarry, South

Newbald. Humberside: Calloviense Zone, Enodatum Subzone.


COX, Callovian perisphinctid ammonites

PLATE 12 Fig. Page

Choffatia sulcifera (<)ppcl, 1857) 33 la d Plaster cast of holotype; HC.S GSM 117447; [m]; Brown Jura zcta; Gammel-

shauscn. Wurttetnberg, W. Germany; original specimen is GPI |no number] (Quenstedi 1X49, pi. 13, fig. I).

2 BCJS GSM27932; [in); Oxford Clay; Summer town, Oxfordshire; Athlcta Zone.

3fl-c OIJM J2588; [m]: Oxford Day; Wolvercote, Oxfordshire: Athlete Zone. 4 JKW BH/126; [mj; Ilacknesa Rock; Hackncss Quarry. Hackney North

Yorkshire; Athlcta Zone (Bed 4 of Wright 1968). 5 OLM J1218; [mj; Oxford Clay; Wolvercote, Oxfordshire; Athlcta / one . 6 OUM J25X7; completely septate; Oxford Clay; Wolvercote* Oxfordshire;

Athlela Zone.

Chojj'utia alT. lahuseni (Sa.sonov, 1965) 34 7a, b OUM J17870; completely septate; Middle Oxford Clay; I.BC pit, Bletchlcy.

Buckinghamshire: Athlela Zone, Proniac Subzone (Bed 21 of Callomon 1968).

8a t h OUM J1203; completely septate: Oxford Clay; Wolvercote, Oxfordshire; Athlela /one .

Choffatiapocutwn (Lcckcnby. 1859) 35 9a c SM J3298: holotype; completely septate | M); Hackneys Rock: near Gristhorpc

Bay, North Yorkshire; Lamberti Zone. Ila ? b SM J3299; para type; complete adult (mj with lappets; Hackncss Rock; near

Gristhorpe Hay, North Yorkshire; Lamhcrl.i Zone. 12 WMS J49; [mj; Hackness Rock; Cunstone Nab. North Yorkshire; Lamberti

Zone. 13 OUM .122362; complete adult (mj with lappets; Lamberti Limestone; Wood-

ham, Buckinghamshire; Lamberti Zone, Lamberti Subzone. 14 BGS FR1116; complete adult [in] with lappei.s; Lamberti Limestone; Wood-

ham, Buckinghamshire; Lamberti Zone. Lamberti Sub7one. 15 MM LL5664A; completely septate; Oxford Clay; Wolvercote, Oxfordshire;

Athleta Zone. 16 OUM J22361; complete adult fm| with lappets; Lnmbcrii Limestone; Wood-

ham, Buckinghamshire; Lamberti Zone, l amberti Subzone.

Perisphinctes suhtilis Neumayr, 1871 35 10a, b Plaster cast of Neumayr 187K pi. 14, fig. 3a c; BOS GSM 117446; according

to Neumayr, completely septate; 'oberer Dogger'; Balin, near Krakow, Poland; original specimen is BSM ASXXI2.


COX, Callovian perisphinctid ammonites P l a t e j2

PLATE 13 Fig. Page

Choffatia poculum (Locke n by, 1859) 35 1 IKiS Zt51? [M]; Lamberti Limestone; Wood ham, Buckinghamshire; Lam-

berti Zone, l.ambcrti Subzone. 2a, b OUM J22553; completely septate [MJ; Lamberti Limestone; Woodham.

Buckinghamshire; Lambcrli Zone. Lamberti Subzone. 3a. b WMS J53; [MJ; Hackncss Rock: Gristhorpe(7): North Yorkshire; Lamberti

Zone. 4 YM 606; completely septate [M); Ilackness Rock: near Scarborough, North

Yorkshire; Lamberti Zone. 5 OUM J223~5; [M]; Lamberti Limestone; Woodham, Buckinghamshire; Lam-

berti Zone, Lamberti Subzone.

PLATE 14 Fig.

Choffatia trina (S. S. Buckman. 1922) 38 la c BGS GSM6509; holotype; completely septate; Oxford Clay; Fleet Bridge

[? = Tidmoor Point]" Weymouth, Dorset; ?Lumberti Zone. 2 OUM .11 6729; complete adult [M] wilh one whorl body chamber; TWuodham,

Buckinghamshire; Lamberti Zone. 3 OUM J22368; [MJ; Lamberti Limestone, Woodham, Buckinghamshire; Lam-

berti Zone, Lamberti Subzone. 4 BGS Zr3766; 'Lamberti Limestone; Millbrouk, Bedfordshire: Lamberti

Zone. 5 OUM J22370; Lamberti Limestone; Woodham. Buckinghamshire; Lamberti

Zone. Lamberti Subzone. Binatisphinctes comptoni (Pratt, 1841) 43

6 BGS GSM30349; complete adult [m] with lappets; Oxford Clay [Acutistri-atum Bund]; Christian Mai ford : Wiltshire; Athleta Zone, Phaeinum Sub-zone.

7 BGS FR1233; adult [m] with lappds; Oxford Clay [Comptoni Bed]; LBC pit, Calvert, Buckinghamshire; Coronatum Zone. Grossouvrei Subzone.

M O N O G R A P H OF THE PALAEONTOGRAPHICAL SOCIETY \


PLATE 15 Fig. Page

Binatisphinctes comptoni (Pr&ii^ 1841) 43 1 BGS FR1297; adult |m| with lappets: Oxford Clay (Acutistriatum Bond);

LBC pit, Stewartby, Bedfordshire; Athleta /one , Phaeinum Subzone. 2 A small form; BGS FR970; adult |m) with lappets; Oxford Clav [Comptoni

Bed); LBC pit. Calvert, Buckinghamshire; Coronatum Zone, Grossouvrei Subzone.

3 OUM J17889; adult |m|; Oxford Clay (Comptoni lied): LBC pit, Blctehley, Buckinghamshire; Coronatum Zone, Grossouvrei Subzone.

4. 6 A small form: adults [nij with lappets; Oxford Clay [Acutwtriatuiu Band]; Christian Mai ford, Wiltshire; Athleta Zone, Phaeinum Subzone; 4. YM 608; 6, BGS GSM30342.

5*7 AdulLs (m] with lappets: Oxford Clay [Aeulislriatum Band]; Christian Mal-ford, Wiltshire; Athleta Zone. Phaeinum Subzone: 5, YM 607; 7. BGS GSM 30345.

8 BGS TR1020; adult ("ml with lappeta; Oxford Clay [Comploni Bed]; LBC pit, Calvert, Buckinghamshire: Coronatum Zone. Grossouvrei Sub7one.

M O N O G R A P H OF THE PALAEONTOGRAPHICAL SOCIETY \ 0

COX, Callovian perisphinctid ammonites Plate 15 A

PLATE 16

Bimiisphincies lompfoni (Pratu 1841) DM 2143; [MJ: Oxford Clay [?Acuminatum Band]: ?\ViJtshirc; ?Athlcta

Zone. Phacinum Subzone.


COX, Callovian pcrisphinctid ammonites Plate 16

PLATi: 17 Fig. Page

Bimtisphincies comptoni (Pratt, 1841) 43 1 OUM J 30868; [MJ; Oxford Clay I Acuminatum Baud]; LBC pit: Blctchley,

Buckinghamshire; Athleta Zone, Phaeinum Subzone. 2 BGS GSM30351; [M]; Ox lord Clay [AeuListriatum Hand]: Christian Mall'ord,

Wiltshire; Athleta Zone, Phaeinum Subzone.



PLATII IS Fig. Page

Biaatisphincfrs comptoni (Pra 11, IH41) 43 i BM 32343; [M] with aptychi; Oxford Clay (Aeutislriatum Band]; Christian

Malford. Wiltshire; Athleta Zone. Phacinum Subzone; xO-5.

Bimithphhw/as hamulatiis (8. 5. Buckman, 1921) 46 2a. b, Topotvpcs; [mj; Hackncss Rock; Scarborough. North Yorkshire; Athlera 5a, b Zone; 2a. b. YM 614; 5a, b. BM 39530. 3a, b SM J3307; paratype; Hackncss Rock; Scarborough [?1 lackness], North York-

shire; Athlcta Zone. 4a. b JKW AC/66: Hackncss Rock: Cornelian Bay, North Yorkshire; Athlela

Zone. fta-e SM .13306; holotype; adult [m|; Hackney Rock; Scarborough. North York-

shire: Athleta Zone.

MONOGRAPH OF THE PALAEONTOGRAPHICAL SOCIETY \

PLATE 19 Fig. Page

?Bina(isph'tncteshwnuUttus (S. S. Buekmun, 1921) 46 la, b BM C107X; adult [MJ; Hackncss Rock; Scarborough, North Yorkshire;

Alhlcla Zone.



PLATE 20 Fig. P a ?S

Pseudopeltoceras tecktwbyi Spath. 1931 47 la. b BM -'16516; holotypc; adult |MJ; Hackncss Rock; Scarborough. North York-

shire: Athleta Zone.


COX, Callovian pcrisphinctid ammonites Plate 20

RS

PI.ATE 21 Fig. Page

Binatisphinctes binatus {Leckenby, 1X59) 47 la, b SM J33G4; lcctotypc; ?[m]; Hackness Rock; Scarborough, North Yorkshire;

?Lamberti Zone. 2n c SM J3305: paralectotype; completely septate; Hackncss Rock; Scarborough.

North Yorkshire; YAthlcta Zone. 4a, b W.MS J-19-J; Hackncaa Rock; [?Casile Hill, Scarborough]; North Yorkshire;

?AthIeta Zone. 5a-c YM 615: Hackness Rock: Scarborough. Norlh Yorkshire; ?Athlcta Zone.

Psetuiopefloceras famulwn Spath, 1931 43 3a, b YM 612: Hacknc^s Rock; Scarborough NorLh Yorkshire; Athleta Zone. 6a, b WMS J4S; Hackncss Rock; [?Castle Hill, Scarborough], North Yorkshire;

Athleta Zone.



J

PLATE 22 Fig. Page

Alligaticeras rotifer (Brown, 1849) 51 la, b MM LI 1494; holotype;completely septate; Hackness Rock; Castle Hill. Scar-

borough, North Yorkshire; Athlela Zone. 2a. b YM 613; [M]; Ilackne&s Rock; Scarborough, North Yorkshire; Athlcta Zone. 3a, b OUM .112<W; Oxford Clay, Summertown, Oxfordshire; Athlcta Zone. 4a, b OUM J1250; completely scptale: Oxford Clay; Summcrtown, Oxfordshire;

Athlcta Zone. 5a, b BGS Y2125: I lackness Rock; Caylon Bay. North Yorkshire: Athlela Zone. da. b BGS Zt3495; completely septate: Middle Oxford Clay; A5 diversion, Milion

Keynes, Buckinghamshire; Athlcta Zone, Prouiae Subzone (Beds 5b 5c of Callomon MS).

X OUM J26360; Oxford Clay; Pear Tree Hill, Oxford, Oxfordshire; Athlcta Zone.

Altigaliceras all', rotifer Brown. 1849 52 7 OUM J29909; adult [ml with lappets; Middle Oxford Clay: Woodham, Buck-

inghamshire; Athlela Zone. Spinosum Subzone. 9a. b BGS Zi.3703; completely septate |M); Middle Oxford Clay; A5 diversion,

Milton Keynes. Buckinghamshire; Athleta Zone, Prouiae Subzone (Bed 7a of Callomon MS).



PLATE 23 Fig. Page

Alligatkeras aft. rotifer (Brown. 1849} 52 1 OUM .129910; | M|; Middle Oxford Clay: Woodham, Buckinghamshire:

AlhleUi Zone. Spinosuni fvuhznne. 2 OUM .11206; [MJ; Oxford Clay: ?Suramerto\vn1 Oxfordshire: Athleta Zone.

Aliigaticeras alligatum (I.eckenby, 1 &59) 53 3a c SM .13303; holotype; [ml; Hackness Rock; Scarborough, North Yorkshire;

Lamberti Zone. 4a-c OUM J2586; <>xford Clay; Wolvcrcotc. Oxfordshire; Athleta Zone. 5a, b WMS J-M>; [M]; Hackness Rock; Can stone Nab, North Yorkshire; Lamberu

Zone. 6a c BGS 7,1516; [MJ; Lamberti Limestone; Woodham, Buckinghamshire; Lam-

berti Zone, Lamberti Subzone. 7 OUM .I223K4; complete adult [m] with lappets; Lamberti Limestone; Wood-

hum, Buckinghamshire; Lamberti Zone, Lamberti Subzone. Propmsphinctes bernensis (de Loriol, I89K) 50

8a-c MM L6251A; Oxford Clay; St. Ives, Cambridgeshire; Lower Oxfordian, Mariae Zone.



Cox 1988 English Callovian Perisphinctid Ammonites

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