An integrated microcrinoid zonation for the lower Campanian chalks of southern England, and its implications for correlation

Accepted Manuscript An integrated microcrinoid zonation for the lower Campanian chalks of southern England, and its implications for correlation Andrew Scott Gale PII:

S0195-6671(16)30218-X

DOI:

10.1016/j.cretres.2017.02.002

Reference:

YCRES 3524

To appear in:

Cretaceous Research

Received Date: 23 September 2016 Revised Date:

1 February 2017

Accepted Date: 2 February 2017

Please cite this article as: Gale, A.S., An integrated microcrinoid zonation for the lower Campanian chalks of southern England, and its implications for correlation, Cretaceous Research (2017), doi: 10.1016/j.cretres.2017.02.002. This is a PDF file of an unedited manuscript that has been accepted for publication. As a service to our customers we are providing this early version of the manuscript. The manuscript will undergo copyediting, typesetting, and review of the resulting proof before it is published in its final form. Please note that during the production process errors may be discovered which could affect the content, and all legal disclaimers that apply to the journal pertain.

ACCEPTED MANUSCRIPT An integrated microcrinoid zonation for the lower Campanian chalks of southern England, and its implications for correlation.

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Andrew Scott Gale

School of Earth and Environmental Sciences, University of Portsmouth, Burnaby Building, Burnaby Road, Portsmouth PO1 3QL, United Kingdom

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ABSTRACT

Detailed logging, sampling and processing of Lower Campanian chalks in Sussex,

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Hampshire, Isle of Wight, Wiltshire and Dorset (United Kingdom) have revealed the presence of a distinctive succession of pelagic microcrinoid faunas, including 25 species and forms of the order Roveacrinida. These occur through the Offaster pillula and Gonioteuthis quadrata zones and provide the basis for a new zonation characterized as CaR1-CaR11. Together with horizons of abundance and

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distinctive forms of the holasterid echinoids Hagenowia, Echinocorys and Offaster and limited microbrachiopod data, these provide a detailed biostratigraphical framework which allows independent assessment of correlations based primarily upon marker beds (marls, flints etc). The new biostratigraphy confirms

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and further constrains the detailed correlation of the Offaster pillula Zone (Newhaven Chalk Formation), but proposes a new correlation framework for the

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overlying Gonioteuthis quadrata Zone (uppermost Newhaven, Culver and Portsdown chalk formations). A new genus of Saccocominae (Assericrinus gen nov., type species A. portusadernensis sp .nov.), two new species of Saggitacrinus (S. alifer sp. nov. and S. longirostris sp. nov.), a new form of Applinocrinus (A. cretaceus forma spinifer nov.), a new form of Stellacrinus, S. hughesae forma lineatus, and two new species of Hessicrinus (H. cooperi sp. nov. and H. apertus sp. nov.) are described. It is likely that the roveacrinid zonation will be applicable internationally, given that many of the species are also present in the Campanian of the Gulf Coast of the USA.

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ACCEPTED MANUSCRIPT *email address: [email protected]

keywords: Roveacrinida Late Cretaceous

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Europe Biostratigraphy Taxonomy

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1. Introduction.

The biostratigraphy of the southern English White Chalk Group has a long

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tradition, and following the early studies of Barrois (1876), Arthur Rowe applied the macrofossil zonal scheme originally developed in France by Hébert (1863) to the chalk of the English coast (Rowe 1902, 1903,1904, 1905, 1908; reviewed by Gale and Cleevely 1989). This classification comprised assemblage zones based upon occurrences of echinoids, brachiopods, crinoids, bivalves and belemnites.

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Many of these zones have long durations; for example, the Micraster coranguinum Zone is typically 60m in thickness and represents approximately 2myr; the Gonioteuthis quadrata zone is nearly 100m and represents about the lower half of the Campanian Stage, at least 5myr. After some modification by

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Brydone (1912) this zonation was used almost exclusively for subdivision of the White Chalk for the first 80 years of the twentieth century (Gale and Cleeveley

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1989).

Subsequently, microfossils zonations were developed in the White Chalk.

Following the PhD studies of Bailey (1978) on the Turonian-Santonian and Swiecicki (1980) on the Campanian-Maastrichtian a UK benthic zonal scheme was established by Hart et al. (1989). Nannofossils were studied by Burnett (1998) who provided a series of zones which have been used widely (Fig. 1). The extensive studies of chalk lithostratigraphy by Mortimore (1983, 1986a,b, 2011; Mortimore et al. 2001) provided a lithostratigraphical subdivision of the White Chalk, which was adopted by the British Geological Survey (BGS) as the basis for mapping (Bristow et al. 1997; Rawson et al. 2001).

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ACCEPTED MANUSCRIPT Additionally, Mortimore named and correlated numerous individual beds of marl, flint, hardgrounds and nodular chalks in the White Chalk Group across southern England. Although based primarily upon lithology, macrofaunal biostratigraphy (and to a lesser extent, foraminiferal data) provided support for these correlations (Mortimore 1986a). However, in the relatively poorly

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exposed lower Campanian Culver Chalk Formation (80m+), well-preserved macrofossils are rather uncommon, mostly long-ranging, and the benthic

foraminiferal zonation (Hart et al. 1989) recognizes only three subdivisions,

based upon a single study of one section (Swiecicki 1980). Therefore, there is

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only limited biostratigraphical support for the correlation framework proposed for the Culver Chalk by Mortimore (1986a,b, 2011).

In 2014, I commenced extensive sampling and processing of Campanian

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chalk in southern England using Glauber Salt in 2014, and discovered the presence of a diverse, abundant and largely undescribed fauna of pelagic microcrinoids belonging to the Roveacrinida (Gale 2016). These are too small (0.5-3mm) to be found by field collecting and have been entirely overlooked by micropalaeontologists seeking foraminiferans and ostracods. Their potential for

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biostratigraphy was first recognized by Gaster (1924), who named a Saccocoma cretacea Subzone in the lower Campanian Gonioteuthis quadrata Zone in Sussex. It became apparent that the lower Campanian represents a significant evolutionary diversification of the families Saccocomidae and Roveacrinidae

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(Gale 2016), and 25 species and forma are recognized in this paper. The presence of a number of these taxa in the lower Campanian sediments of the USA

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Gulf Coast (Gale 2016) and elsewhere in Europe (e.g. Destombes and Breton 2001) offers considerable potential for interregional correlation. The zonal scheme presented here was developed by logging all available

sections of the upper Newhaven, Culver and Portsdown formations in southern England, and collection and processing of approximately 250 chalk samples. The residues also yielded abundant remains of the highly specialized echinoids Hagenowia (Gale and Smith 1982; Smith and Wright 2003) at a limited number of levels, and microbrachiopods throughout. Preliminary data from these two groups is incorporated into the study, and a review is provided of the taxonomy and distribution of the holasteroid echinoids Echinocorys and Offaster, which

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ACCEPTED MANUSCRIPT have been widely used in correlation of Campanian chalks (Griffith and Brydone 1911; Brydone 1912, 1914, 1939; Mortimore 1986a,b).

2. Methodology.

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All available sections were logged in detail, macrofossils collected, and samples were taken at intervals of approximately 1 or 2 metres in important

sections (see appendix Figs). One kg of each sample was processed using Glauber Salt, a process modified from Surlyk (1972) following Gale (2016). The residues

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were screened and mesofaunal elements were picked. The hard chalks of the Isle of Wight did not process successfully with Glauber Salt, and material was obtained by crushing samples, and treating finer grades of residue with 10%

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acetic acid. Although brutal, this provided some limited but useful information on the distribution of taxa. The pickings, the residues, and samples of the original unprocessed chalks will be deposited in the Natural History Museum, London (NHMUK). Grid references for all localities (Fig.2) are given in Table 1.

3.1 Previous work

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3. Biostratigraphy.

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Traditionally, following the work of Rowe (1902, 1903, 1904, 1905, 1908), the portion of the chalk of southern England now ascribed to the lower

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Campanian was identified as the Zone of Actinocamax quadratus, lying between those of Marsupites testudinarius and Belemnitella mucronata (Fig. 1, two left hand columns). Brydone, in a very detailed study of both faunas and lithostratigraphy, from Sussex to Dorset (1912, 1914), separated the lower portion as the Zone of Offaster pillula, characterized by the frequently abundant occurrence of the eponymous echinoid. He further divided the O. pillula Zone into a lower subzone of Echinocorys depressa, and an upper one of abundant O.pillula (Fig. 1, third and fourth columns). Gaster (1924) subsequently added an horizon of Hagenowia rostrata to the top of the O. pillula Zone, and characterized the lower portion of the quadrata Zone as the Subzone of Saccocoma cretacea

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ACCEPTED MANUSCRIPT (Fig. 1, fifth and sixth columns), neither of which emendations meet with the approval of Brydone (1939). Brydone’s zonation has been widely accepted (e.g. Rawson et al. 1978; Fig. 1, seventh column) and extensively used in both BGS sheet memoirs and independent publications through much of the twentieth century.

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It was apparent from Brydone’s work that the O.pillula Zone could be further subdivided using successive morphotypes of Echinocorys, and details of these on the Sussex coast were provided by Wood and Mortimore (1988).

However, the Zone of Gonioteuthis quadrata (uppermost Newhaven, Culver and

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lower Portsdown formations) represents a considerable thickness of chalk

(100m+) which remains rather intractable by macrobiostratigraphy, although Mortimore (1986b) provided some information on the Echinocorys variants

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which occur within this unit and on the occurrence of other fossils.

3.2 Microcrinoids

The first finds of chalk microcrinoids were made by the amateur geologist

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A.W. Rowe, who obtained material from the washed soft cores of flints (“flint meal”) from the Coniacian of Seaford Head, Sussex. His material was described by Douglas (1908) under the name Roveacrinus, a latinised form of Rowe. Subsequently, Bather (1924) described material collected by Gaster from the G.

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quadrata Zone of Sussex as Saccocoma cretacea, and Gaster (1924) erected a S. cretacea Horizon based on its stratigraphical distribution. Peck (1955)

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redescribed Douglas’material from the English chalk, and described a further three species from Cenomanian and Coniacian chalks, based on material in the NHMUK collections and the Sedgwick Museum, Cambridge. Rasmussen (1971) described microcrinoids from the Turonian of Hampshire. Gale (2016) described extensive new material from the Santonian-Campanian chalks of southern England, including six new genera and eleven new species. Microcrinoids are abundant in Campanian chalks in the United Kingdom, and their taxonomy was described by Gale (2016) who recognized 13 species belonging to the roveacrinid subfamilies Saccocominae, Applinocrininae, Roveacrininae and Hessicrininae. Further taxa are described below (see

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ACCEPTED MANUSCRIPT Systematic palaeontology). Their stratigraphical distribution (Gale 2016 fig. 2; herein Figs 9-16) is very distinctive, and permits the recognition of 11 zones, here named CaR1-11, through the Lower Campanian chalks of southern England. Examples of key taxa are illustrated in Figs 3, 4, ,5 and 6. The forms are all pelagic, and many occur in the Gulf Coast states of the USA (Gale 2016), although

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presently without a detailed stratigraphical context. The zonation is based upon the detailed distributions identified in the main localities, studied, the Sussex coast (Fig.9), North Lancing (Fig.10), East

Grimstead (Fig. 11), West Harnham (Fig. 12), Paulsgrove, Portchester east and

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Candy’s Pit, Portsdown (Fig. 13), Sussex pits nos. 10 and 26 (Fig. 14), Warren

Farm, Hants (Fig. 15) and Sussex pits nos. 17 and 24 (Fig. 16). The congruence of the distribution in these localities permits the recognition of a series of zones

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based on the microcrinoid first occurrences, last occurrences and acmes, defined below and summarized in Fig. 17. In the lower part of the succession, the O. pillula and lower G. quadrata Zones, a number of sections are available in Sussex, Hampshire, Wiltshire and Dorset, and permit the identification of microcrinoid

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Bioevents 1-7 (Figs 9-11). Fewer sections are available at higher levels.

These zones are as follows:

CaR1. LO of Uintacrinus anglicus Rasmussen, 1961 to FO (= First Occurrence) of

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Stellacrinus hughesae forma cristatus Gale, 2016. The former species has a worldwide distribution (Gale et al. 2008). Type section; Black Rock Brighton,

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0.5m above Friar’s Bay Marl 2. Also identified at Margate, Kent. Characterised by low diversity, and rather low abundance, with common S. h. forma hughesae Gale, 2016 (Fig. 5A-D,K), and uncommon Applinocrinus cretaceus cretaceus (Bather, 1924; Fig. 3I,J,N.O) continuing from below, and above, the successive appearance of Hessicrinus cooperi sp. nov. (level of Friar’s Bay Marl 3; Fig.4G,L,P) at the base of the zone, Sagittacrinus torpedo Gale, 2016 (Fig. 3 L,S) which appears between Friar’s Bay 3 and Ovingdean Marls) and rare S. hughesae forma cristatus Gale, 2016, immediately beneath the Rottingdean Pair of Marls (see Appendix Fig. 1 for details of sample levels).

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ACCEPTED MANUSCRIPT CaR2. Flood occurrence of Stellacrinus h. forma cristatus, to FO of Cultellacrinus gladius Gale, 2016 (Fig.5E,H,J), marking Bioevent 1. Type section: Friar’s Bay Steps, base at level of Old Nore Marl (Fig. 9, Appendix Fig. 1). Also identified in Paulsgrove, Hampshire (Appendix, Fig. 3), East Grimstead, Wiltshire (Fig. 11, Appendix Fig. 5) and Middle Bottom (Dorset). The flood abundance of S.

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hughesae forma cristatus (Fig. 5F,G,L) at the level of the Old Nore Marl and in the overlying few metres is a highly distinctive regional marker, here called Bioevent 1 (Figs 9,11). The upper part of CaR2 is characterized by a large increase in

abundance of A. cretaceus (up to 100 plates/kg) and the FO of Costatocrinus

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brydonei Gale, 2016, (Fig. 3A-E,M; Bioevent 2 – Sussex, West Harnham, East Grimstead, Figs 9-11) both just above the Peacehaven Marl. The FO of

Lucernacrinus woodi Gale, 2016 (Fig. 4O,S,W) and Hessicrinus scalaensis Gale,

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2016 (Fig. 4T,U) 2m above Peacehaven Marl, both uncommon, but laterally persistent, marks Bioevent 3, found on the Sussex coast and at East Grimstead (Figs 9,11).

CaR3. FO of Cultellacrinus gladius Gale, 2016 (Figs 5E,H,J) at the level of lowest

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marl of the Meeching Triple Marls, to FO of Hessicrinus filigree Gale, 2016 (Fig. 5H,M), beneath the Castle Hill Marls, which is Bioevent 4, found on the Sussex coast, East Grimstead and West Harnham (Figs 9-11). Type section; Friar’s Bay Steps, Peacehaven, Sussex (see Appendix Fig. 1). The occurrence of C. gladius

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increases rapidly up from the FO (rare) to abundant above, and persists into the

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overlying two zones.

CaR4. FO of Hessicrinus filigree (Fig. 5H,M) beneath the Castle Hill Marls, to FO of Platelicrinus longispinus Gale, 2016 (Fig.6P), 1m above Castle Hill Flint. Type section; west cliffs, Newhaven, Sussex. Bioevent 5, found in the Sussex coast sections, West Harnham and East Grimstead (Figs. 9-11).

CaR5. FO of Platelicrinus longispinus, 1m above Castle Hill Flint 4 (Fig. 6P) to FO of Platelicrinus campaniensis (Destombes and Breton, 2001)(Fig. 6F-L), Assericrinus portusadernensis gen. et sp. nov. (Fig. 4A-E,I), Sagittacrinus longirostris sp. nov. (Fig. 3K, P-Q ,X) Type section; west cliffs, Newhaven, Sussex,

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ACCEPTED MANUSCRIPT also well developed at Paulsgrove, Hampshire (Fig. 13). Characterised by the occurrence of P. longispinus and Costatocrinus mortimorei Gale, 2016 (Fig. 3.G,H). Bioevent 6 is marked by the FO of P. longispinus and is found on the Sussex coast, North Lancing, Paulsgrove, West Harnham and East Grimstead (Figs. 9, 11,12,13). Bioevent 7 is the FO of Costatocrinus mortimorei Gale, 2016, seen in

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Sussex (Seaford and North Lancing) and Paulsgrove, Hampshire.

CaR6. FO of Assericrinus portusadernensis sp. nov. (Fig. 4A-E,I), Sagittacrinus

longirostris sp. nov. (Fig. 3K,P,Q,X), Applinocrinus cretaceus forma spinifer nov.

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(Fig. 3V,W), Stellacrinus hughesae forma lineatus nov. (Fig.5I) and Platelicrinus campaniensis (Fig. 6F-L) at 44-45m in the Paulsgrove pit, Hampshire, together marking Bioevent 8 (see Appendix Fig. 3). Sagittacrinus alifer sp. nov. (Fig.

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3R,T,U) appears 2m higher (Bioevent 9) and is also characteristic of the zone. The top is taken at LO of S. longirostris sp. nov., and base of flood abundance of C. gladius at base of Cliff Dale Gardens section, Hampshire (Fig. 13, Appendix Fig. 4). Type section: Paulsgrove pit, Hampshire, level of sample PG14.

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CaR7. Last Occurrence (LO) of Sagittacrinus longirostris sp. nov., and base of higher flood abundance of C. gladius, base of Cliff Dale Gardens section, Hampshire (Fig. 13, Appendix Fig. 4, sample CD-3). Characterised by abundant C. gladius associated with less common C. mortimorei and P. longispinus. The top is

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taken at the last occurrence of C. gladius, Cliff Dale Gardens, Hampshire (Fig. 13).

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CaR8. LO of Cultellacrinus gladius to acme of Hessicrinus filigree and Platelicrinus campaniensis, 0.5-I metre beneath Charmandean Flint. Type section, Cliff Dale Gardens pit, Cosham, Hampshire (Fig. 13, Appendix Fig. 4, sample CD4). Also exposed in Charmandean Lane, Worthing, Sussex (Fig. 14, Appendix Fig. 1).The zone has a relatively low diversity and abundance of microcrinoids.

CaR9. Base of acme occurrence of Hessicrinus filigree (Fig. 4H,M) and P. campaniensis (Fig. 4F-L), 0.5-1 metre beneath the Charmandean Flint, to LO of Costatocrinus brydonei. Type section, Warningcamp, pit no. 24 of Gaster 1924 (Fig. 14, Appendix Fig. 2). Also identified at Charmandean Lane, Worthing,

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ACCEPTED MANUSCRIPT Sussex, pit no. 10 (Fig. 14, Appendix Fig. 1), Whitecliff, Isle of Wight (Appendix Fig. 10) and Mottisfont, Hampshire (Appendix Fig. 5).

CaR10. LO of Costatocrinus brydonei, Lucernacrinus woodi, and Sagittacrinus torpedo, at 4 metres above the Whitecliff Flint, to FO of Stellacrinus pannosus

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Gale, 2016, in lower of two Scratchell’s Bay Marls. Type section, Warren Farm Quarry (Hampshire, Fig. 15, Appendix Fig. 4), also identified in Cote Bottom, Sussex, pit no. 17 (Fig. 16, Appendix Fig. 2).

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CaR11. FO Stellacrinus pannosus Gale, 2016 (Fig. 5N,Q) in Scratchell’s Marl 1. Upper limit not presently defined. Type section: Warren Farm, level of

Scratchell’s Marl 1 (Fig. 15, Appendix Fig. 4). The abundance of the distinctive

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brachials (Fig. 5N,Q) makes this easy to identify in residues.

Some changes in the roveacrinid faunas through the Lower Campanian are evolutionary transitions, and therefore represent fundamental morphological shifts which have considerable potential for interregional

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correlation. These include: -

the transition between Costatocrinus laevis sp. nov. and C. brydonei.

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Stellacrinus hughesae forma cristatus appears to be a transitional form between S. hughesae forma hughesae and Cultellacrinus gladius (Gale 2016). Hessicrinus cooperi sp. nov. probably evolved into H. filigree by the

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-

development of radial/basal fenestrae (see Systematic palaeontology). Applinocrinus cretaceus forma spinifer nov., in which a substantial radial

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-

spine is developed, only appears at the level of the base of the zone CaR6 and evolved from the spineless A. cretaceus forma cretaceus.

Other changes in the faunas may represent cryptic speciation events, with

no evidence of intermediate forms (e.g. the replacement of S. hughesae by S. pannosus). Other taxa show levels of abundance, separated by units in which they are very rare or absent. In particular, Stellacrinus hughesae and Cultellacrinus gladius display recurring, mutually exclusive acme levels (Figs. 9-

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3.3 Holasteroid echinoids.

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3.3.1 Acme horizons of Offaster pillula (Lamarck, 1816)

Offaster pillula is a common, long-ranging, small species of holasterid which occurs in the chalk facies throughout the lower Campanian and the lower part of

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the upper Campanian across northwest and northeast Europe, eastwards to the Mangyshlak Peninsula in Kazakhstan (personal observation). It is present as discrete abundance levels, separated by intervals in which it is rare or absent.

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Ernst (1971, 1972) provided a detailed statistical analysis of the material from the UK and northern Germany, and identified a number of subspecies based upon size and shape, which were not recognized by Smith and Wright (2003) in their taxonomic revision of British material.

The stratigraphical usage of O. pillula in the UK chalk dates back to Rowe

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(1902, 1903, 1904, 1905, 1908) who used the species as an accessory guide to the zone of Actinocamax quadrata. Subsequently, Griffith and Brydone (1911) and Brydone (1912, 1914) erected a zone of O. pillula for the lower part of the original quadrata Zone, the upper part of which was identified as the “Subzone of

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abundant O. pilula”, and included two discrete levels of abundance of the species. The higher of these levels is terminated by a bed containing O. pillula of an

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exceptionally large size, delimited by a pair of marls with a centrally placed nodular flint which was called the Planoconvexa Bed by Brydone (1939). The lower part of the bed contains O.pillula planata Brydone, 1939, and the upper part O. pillula convexa Brydone, 1939. Brydone (1914) was able to trace this bed through Sussex, Hampshire, the Isle of Wight and Dorset. The bounding marls were later named as Telscombe Marls 2,3 by Mortimore (1986a,b). An additional (lower) horizon of abundant O. pillula has been discovered on the Sussex coast during this study, positioned between the Black Rock and Saltdean Marls (0.5-1.3m above Black Rock), well exposed in the western part of Friar’s Bay, Peacehaven and at Black Rock, Brighton (Appendix fig. 1). These are

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ACCEPTED MANUSCRIPT small-to medium-sized O. pillula, with rather variable morphology (usually with inflated oral surfaces), quite distinct from those occurring at the level of the Old Nore Marl. The material will be described at a later date. This horizon has not yet been found outside Sussex, but such is the lateral continuity of the other O. pillula occurrences that it is to be expected further west, if better sections of this

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horizon were available. Ernst (1972, fig. 9) undertook a biometrical analysis of O. pillula from the Planoconvexa Bed and underlying 2 metres of chalk in Sussex. This

demonstrates a progressive increase in size from a maximum length less than

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20mm, to over 30mm in the upper part of the Planoconvexa Bed. The

accompanying shape changes, which Brydone (1912) took to be significant, are probably an allometric consequence of increased size (Smith and Wright 2003),

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rather than an evolutionary development. Above the Planoconvexa Bed, O. pillula is rather uncommon, and usually small in size (O. pillula nana Ernst, 1971), and the species ranges up into the upper Campanian. These levels are:

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O.p. 1. Small O.pillula, <15mm length, abundant within the interval from 0.51.3m above the Black Rock Marl on the Sussex coast. Typically very inflated forms.

O.p.2. Small O. pillula, <15mm length, from beneath the Old Nore Marl to level of

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Peacehaven Marl.

O.p.3. Level of Meeching Paired Marls to Planoconvexa Bed. There is an overall

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increase in size of individuals through the interval.

Above this level, small O. pillula occur rather uncommonly in the G.

quadrata Zone (Brydone 1912; Mortimore 1986a).

3.3.2 The Hagenowia evolutionary lineage (Fig. 7)

The highly specialized holasteroid genus Hagenowia has the apical portion of the test extended into an elongated rostrum, the anterior surface of which forms a concave groove. The apical system is disjunct, and the number of plate

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ACCEPTED MANUSCRIPT rows present in the rostrum decrease through the evolutionary history of the genus (Gale and Smith 1982; Smith and Wright 2003; Fig. 7 herein). Entire specimens are exceptionally rare, but the rostra are common small fossils in Coniacian-Maastrichtian chalks. In processed chalk residues, rostra and rostral fragments are easily identified, and characterize discrete intervals of Campanian

can be very high (>50 fragments per kg).

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chalks, separated by units in which Hagenowia is entirely absent. Abundances

A continuous lineage (Fig. 7) leads from the Coniacian-Santonian species H. rostrata (Forbes, 1852), through the late Santonian H. anterior Ernst and

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Schulz, 1971, the Campanian H. blackmorei Wright and Wright, 1949 to the

Maastrichtian species H. elongata (Brünnich Nielsen,1942; compare Gale and Smith 1982; Smith and Wright 2003). In this, there is progressive narrowing of

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the rostrum, increased differentiation of the head of the rostrum, and reduction in number and elongation of the plate rows making up the rostrum (Fig. 7, bottom to top of page; see also figure caption). Finally, contact is made between genital plates 3 and 5, separating genital 2 from oculars II and IV (Fig. 7A-E), seen in Upper Campanian and Maastrichtian specimens. Hagenowia anterior has

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early and late subspecies (here referred to as A and B) characterized by the proportionate breadth/height of interamb rows 1b and 4a. In specimens from the M. coranguinum Zone (Fig. 7N) these are only slightly taller than broad, whereas in those from the Marsupites testudinarius Zone (Fig.7L,M) they are

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twice as tall as broad.

Jagt (2000, pl. 23 figs 3,4) demonstrated that the contact between

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genital plates 3 and 5 occurred low in the upper Campanian, because specimens from the lower Zeven Wegen Member of the northeast Belgium displayed the derived condition.

In Maastrichtian material of H. elongata only two madreporic pores,

situated vertically above each other, are present on genital plate 2 (Schmid 1972; Fig. 7D,E herein), whereas in Campanian specimens, a scatter of 6-8 pores are typically present (Gale and Smith 1982; Smith and Wright 2003, text -fig. 217B; Fig. 6C-D here). However, new material collected from both the lower and upper Campanian demonstrates that forms with only two vertically situated pores are present in about 25 per cent of rostra in H. blackmorei (Fig. 6A,B) These possibly

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ACCEPTED MANUSCRIPT represent a separate species or subspecies. I therefore propose that H. elongata is defined by the contact between genital plates 2 and 5, and its first occurrence is therefore close to the base of the upper Campanian.

The common presence of Hagenowia in chalks overlying the “Horizon of

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abundant O. pilula” led Gaster (1924) to erect a “Hagenowia rostrata Horizon”, identified in chalkpits around Worthing and on the Sussex coast. The

occurrences were confirmed by Gale and Smith (1982), Mortimore (1986a,b,) and Wood and Mortimore (1988). However, material picked from processed

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samples indicates that three levels of abundant H. blackmorei are present in

lower Campanian chalks. Rare individuals may be present at other levels, but

These levels are as follows:

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these records are unconfirmed.

H1. Uppermost U. anglicus Zone and immediately overlying chalk. Hagenowia

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anterior ssp. B. Sussex, Kent. Diffuse, specimens rather uncommon.

H.2. A level immediately overlying the Meeching Paired Marls, approximately 1.5m in thickness, in the lower part of O.p.2. Hagenowia blackmorei.

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H.3. A 7-m-thick horizon, in two parts, extending from 2m above the Castle Hill Marls, level of Castle Hill Flint 3, up to Castle Hill Flint 8 (Mortimore 1986a,b). An

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acme horizon of abundant specimens (up to 60 fragments per kg) is associated with Castle Hill Flint 4, and is identified in Sussex, Hampshire, Dorset and Wiltshire. H. blackmorei.

H.4. A 1m abundance level underlying the Portchester Hardgrounds. Probably H. blackmorei, but no complete rostra collected.

H.5. 1m thick, beneath the W Flint Weybourne Chalk, Norfolk (Peake and Hancock 1961). Hagenowia elongata ssp. A

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ACCEPTED MANUSCRIPT 3.3.4 Echinocorys scutata Leske, 1778 (Fig. 8)

Shape variants of the large holasterid E. scutata have played an important role in chalk stratigraphy, since A.W. Rowe (1902, 1903, 1904, 1905,

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1908) recognized their value in characterizing zones, and Lambert (1903) provided an hypothetical phylogeny. Griffith and Brydone (1911) and Brydone

(1912, 1914) named a number of Santonian and Campanian morphologies from Hampshire and Gaster (1924, 1937) applied these to the Sussex chalk.

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Whilst Echinocorys morphologies are undoubtedly of stratigraphical value, their use is hampered by a number of factors. Firstly, they are often

variable in form at any given horizon, so an assemblage of about 10 specimens is

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required to verify horizon. Secondly, they are relatively uncommon as wellpreserved specimens at some levels in the English chalk, including the G. quadrata Zone. Thirdly, the correct usage and validity of names for forms has never been satisfactorily agreed upon amongst workers; there are a plethora of names provided by English, French, and German authors which are sometimes

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used indiscriminately and informally. The monographic revision of British Cretaceous Echinoids (Smith and Wright 2003) did not really address these problems, but simply picked out a limited number of shape variants as named formae, stating as justification that (p.533), “However, these shape differences

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do not form an appropriate basis for various types, and all tests are constructed on the same architectural plan.” Forbes (1852,p. 4) thought along similar lines

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“……everyone of them is linked with the others by the most delicate shades of gradation.”

However, E. scutata morphotypes are actually highly restricted

stratigraphically, and some are extremely widespread geographically, with identical forms extending eastwards from the UK to Georgia, the Caucasus, and Mangyshlak, (Kazakhstan, central Asia; see Peake and Hancock 1961, 1970). They can be used to provide high-resolution interregional correlations, as demonstrated by Peake (in Hancock et al. 1993), who used species of Echinocorys to correlate between the Maastrichtian GSSP at Tercis-les-Bains in southwest France and Norfolk in eastern England. Although there appear to be

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ACCEPTED MANUSCRIPT continuous transitions between the main shape varieties (e.g. Forbes, 1852; Smith and Wright 2003), in reality “The characteristic shape of one horizon is probably never exactly mimicked at any other horizon.” (Peake in Hancock et al. 1993 p. 139). Additionally, other features of the test, in particular the size and shape of the apical system, provide independent and stratigraphically important

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data. Willcox (1953) showed that the shape and size of the apical system, proportional to the length of the test, changed significantly through the

Coniacian to Campanian interval in the English chalk – specifically, that there

was an increase in the proportional size of the apical disc, relative to the length

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of the test, up to E. s. cincta in the upper part of the O. pillula Zone, where a

dramatic decrease in disc size occurred (Fig. 8J). The shape of the apical disc followed a parallel trend, becoming shorter and broader up to the level of E. s.

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cincta, where it reverted to a narrow, elongated shape (Fig. 8). To illustrate the variation in apical disc morphology in Coniacan to Campanian Echinocorys, I here include drawings of the apical discs of Turonian and early Coniacian Echinocorys, which have a very elongated, narrow form (Fig. 8A,B; Olszewska-Nejbert 2007), and of the neotype of E. scutata from the upper Coniacian or lower Santonian of

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the UK (Fig.8C) which represents an intermediate morphology between early Coniacian and late Santonian types.

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The Echinocorys elevata-truncata lineage (Fig. 8F-I)

The successive forms E. s. elevata, E.s. tectiformis, E.s. depressula and E.s.

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truncata form a lineage in the Anglo-Paris Basin (Fig. 8), which evolved from a low variant of E. scutata, identified as E. s. striata (Ernst and Schulz 1972, no author given) in the coranguinum Zone (Fig. 8D,E). The overall characteristics of the group are the trapezoidal lateral profile, in which the apical disc forms a flattened, truncated apex to the test, the rather low position of the ambitus, the proportional increase in length of the apical disc, and the progressive shortening and broadening of the apical disc. The broadening of the apical disc is brought about by enlargement of genital plate 4 and to a lesser extent by genital plate 1 (Fig. 8D-H). Additionally, the ocular plates of the apical disc typically display irregular swellings, sometimes extreme, which superficially appear to be a result

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ACCEPTED MANUSCRIPT of parasitism, but are specific to this lineage. There are transitional forms between elevata, tectiformis and depressula, and possibly between depressula and truncata. Echinocorys s. truncata represents the final member of the lineage, which disappears just beneath the Peacehaven Marl.

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Echinocorys scutata elevata Brydone, 1912 (Fig. 8F)

Relatively large, tall, subpyramidal lateral profile, abruptly truncated aboral surface formed by relatively short apical disc. Typical form of the

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Marsupites laevigatus and M. testudinarius Zones (Wood and Mortimore 1988).

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Echinocorys scutata tectiformis Griffith and Brydone, 1911 (Fig. 8G)

Medium- to small-sized, relatively low trapezoidal profile, apical disc elongated, up to 50 per cent of total length. This occurs from the Friar’s Bay Marl 1 up to the level of the Saltdean Marl (Wood and Mortimore 1988).

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Echinocorys scutata depressula Brydone, 1939 (Fig. 8H)

Lateral profile low, aboral surface smoothly convex, ambitus rather high; posterior slope above periproct strongly and evenly reflexed. Base almost

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invariably inset by crushing. The subspecies only occurs between the Saltdean Marl and the Roedean Triple Marls on the Sussex coast, a thickness of

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approximately 5m, and is a valuable stratigraphical marker.

For Brydone (1912, 1914) the depressula form characterized the upper

portion of the Subzone of E. depressula of the O. pillula Zone, and its range in Sussex was correspondingly shown by Wood and Mortimore (1988, Fig. 18) as extending from the Saltdean Marl up to the Old Nore Marl. In contrast, Peake (in Hancock et al. 1993) viewed E. s. depressula as an ecological variant of E. s. tectiformis, common in the softer chalks of Hampshire, but rare in the higherenergy chalks of Sussex where hardgrounds are more commonly developed. New collecting by myself in Friar’s Bay, Sussex, shows that morphologically

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ACCEPTED MANUSCRIPT transitional forms between E. s. tectiformis and E. s. depressula are present in the 0.7-m-level overlying the Saltdean Marl, and E. s. depressula only occurs above this level, and E. s. tectiformis beneath it.

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Echinocorys scutata truncata Brydone, 1912 (Fig. 8I)

Small, lateral profile markedly trapezoidal, with a flat, short and very

broad apical disc, with irregularly swollen ocular plates; the disc represents up to 40% of the length (Willcox 1953). Early forms are larger (Wood and

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Mortimore 1988), and may be transitional from E. s. depressula. The form is

characteristic of the interval between the Old Nore and Peacehaven marls in Sussex, and elsewhere in southern England, over an interval of approximately 4-

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6m.

The Echinocorys scutata cincta- E.s. conica lineage

The Peacehaven Marl marks a major change in Echinocorys

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morphology, with the disappearance (probable extinction) of the E. s. elevata-E.s. truncata lineage, and the incoming of forms which possess a relatively elongated, narrow apical disc (Willcox 1953), lacking irregular ocular swellings of the disc and proportionately short in relation to test length. The shape of the test is

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variable; E. s. cincta in the sense of Brydone (1912) the name E. s. cincta is applicable only to tall, slightly gibbous forms in which the ambitus is high and

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the profile retracted beneath the ambitus (e.g. Brydone 1912, Pl. 1 figs. 3,4; Fig. 8J herein). This form is certainly characteristic of, and rather invariant morphologically, in the level between the Peacehaven Marl and the Meeching Marl Pair (e.g. Wood and Mortimore 1988), and is here distinguished as E. s. cincta A. Material from the level between the Meeching Paired Marls and the Planoconvexa Bed includes large, morphologically variable specimens which fall close to E. s. cincta, but include both tall, subconical and rather low, elongated forms. These are distinguished as E. s. cincta B.

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ACCEPTED MANUSCRIPT The level above the Planoconvexa Bed, up to the Castle Hill Marls yields large Echinocorys (“large forms” of Gaster 1937) of a very different type, which are subpyramidal in lateral profile (Fig. 8K E. s. new forma?) and of variable height. They appear to represent an entirely different group to that of E.s. cincta, and await detailed study.

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Gaster (1924) applied the name cincta to variable morphologies, often smaller in size and including common depressed forms, which occur at higher

horizons, notably the level of abundant Hagenowia in Sussex, between the levels of Castle Hill Flints 4-11 in chalkpits around Worthing, much to Brydone’s

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annoyance (Brydone 1939). Inspection of Gaster’s material in BGS and NHMUK collections supports his assignation of his “small forms” to the cincta group,

referred to E.s. cincta C.

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because they have a high ambitus and a sub-ambital constriction. They are here

Above this level, well-preserved Echinocorys are uncommon in the G. quadrata Zone, and most of the material in collections is unhorizoned and therefore of limited value. However, it is evident that tall, domed morphologies, with rather rounded bases, occur throughout the interval between the Castle Hill

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Marls and the level of the Whitecliff Flint. These co-occur with more gibbous forms, which are derivatives of the cincta morphology. It therefore appears that the E. s. cincta group gives rise to the tall domed forms. Peake (in Hancock et al. 1993) applied the name E. turrita Lambert, 1903 to these tall, round based forms

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(Fig. 8L), and thought this species to be geographically widespread and highly distinctive. However, there are problems with the use of this name, because the

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author of E. turrita did not designate a holotype, but simply referred to outline lateral profiles given by Forbes (1852, pls 6,9) and Wright (1882, pl .77 Fig. 7), of which the horizon is completely unknown. Indeed, a very similarly shaped tall form, with an evenly rounded aboral surface, also occurs in the upper Coniacian of Germany (Ernst and Schulz 1974, p. 38, pl 4 fig. 2) who referred to this as the ‘E. ex. gr. scutata, “ planodoma” Typ’, without an author attribution. It will be necessary to study details of the test, particularly the apical system, to define these forms accurately. In the higher part of the G. quadrata Zone, Echinocorys were abundant at Downend Quarry, Hampshire (Gale 1980; Gale et al. 2015; Mortimore et al.

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ACCEPTED MANUSCRIPT 2001), and in the late 1970s I made extensive collections of material from this site, now at BGS, Keyworth. These include specimens which are close to E. s. turrita, which vary continuously with subconical forms in which the posterior slope of the lateral profile is strongly declined (Fig. 8M, E.s. undescribed). These are intermediate between E. s. turrita and E. s. conica. The latter becomes

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common a short distance beneath the Farlington Marls (M9,10; Gale et al. 2015).

3.3.5 Microbrachiopods

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Microbrachiopods are common in chalk mesofaunas and have been

widely used in biostratigraphy of upper Campanian and Maastrichtian chalks in Germany, Denmark and East Anglia, UK (Surlyk 1984, Johansen and Surlyk

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1990). They are abundant in lower Campanian residues collected in this study, and comprise about 10 species, most of which appear from an initial study to be long ranging. However, two species at least provide useful stratigraphical information. Terebratulina rowei Kitchin, in Rowe, 1902 (Fig. 6M), ranges up from the Marsupites Zone as a common fossil, to disappear in the lower G.

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quadrata Zone, approximately at the level of Castle Hill Flint 11. Additionally, the distinctive small terebratellid Leptothyrellopsis polonicus Bitner and Pisera, 1979 (Fig. 6N,O; see also Johansen and Surlyk 1990) is abundant in the highest Culver and Portsdown chalks, with up to 50+individuals per kg of chalk in some

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samples. This level can be traced from Sussex to Hampshire and the Isle of Wight.

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4. Correlation.

5.1 Upper Newhaven and basal Culver Chalks – O. pillula and lower G. quadrata Zones (Figs 18,19).

The biostratigraphy fully supports the correlations of marl seams and faunal horizons originally suggested by Brydone (1912, 1914) and developed and extended by Mortimore (1986a,b), with the addition of numerous named marker beds, from the Friar’s Bay Marls up to the Pepperbox Marls or a correlative level. A succession of flints, marl seams and beds of fossils can be

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ACCEPTED MANUSCRIPT correlated accurately from south Dorset (West Bottom), through Scratchell’s Bay, Isle of Wight, Wiltshire (East Grimstead), to Portsdown (Paulsgrove) and into the Sussex cliffs, supported by detailed faunal evidence from echinoids and microcrinoids. The lithological and faunal succession is similar across the region, as concluded by Brydone (1914) and Mortimore (1986a,b). The exceptions to

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this pattern are few but striking, such as the development of hardgrounds and phosphatic chalks at Whitecliff in the condensed upper O. pillula Zone, and at a scatter of other more poorly exposed localities such as Stoke Clump near

Chichester where a phosphatic cuvette is developed at approximately the same

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level (Gale 1980).

However, there is significant lateral variation in the presence/absence and local development of marls, in particular. Middle Bottom, in Dorset, has a

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virtually marl-free Newhaven Chalk, full of burrow flints, whereas there are very numerous marls in the equivalent levels in the less flinty Scratchell’s Bay succession. In Scratchell’s Bay, the numerous marls are rather evenly spaced (0.4.-0.7 metres) and appear to define precession cycles. Individual marls or groups of marls can change markedly laterally, such as the Meeching Triple

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Marls, which are triple in East Grimstead and Sussex, but pass into a single marl at Paulsgrove, Hampshire and are absent at West Harnham. The disappearance of marl seams is not related to condensation; for example, the succession represented by zone CaR4 is slightly thicker in Sussex than that at East

Sussex.

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Grimstead, but the Pepperbox Marls are present at East Grimstead but absent in

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However, a number of sections incorporated into the correlation

framework by Mortimore (1986a,b) are anomalous, and do not conform to the regular pattern shown in Fig.18. These include West Harnham, Salisbury, and Mottisfont, Hampshire (Fig. 19). The log of West Harnham presented (Fig. 12, Appendix, Fig. 6) here differs significantly from that of Mortimore (1986a fig. 19). The only marl seam I have identified in the lower part of the succession lies about 1.5m above the base; this is identified as the Peacehaven Marl, because it lies at the level of Bioevent 2 (compare with Sussex, Fig. 9 and East Grimstead, Fig. 11). Above this, the Meeching Triple Marls are not developed, and the Meeching Pair appear to be cut out on a nodular surface at 10.8m. The

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ACCEPTED MANUSCRIPT Telscombe Marls 1-3 and the Planoconvexa Bed are well developed. The single marl, resting on an omission surface at 21.2 m, lies above Bioevent 5 and is identified as a single Castle Hill Marl. I am unable to find the Castle Hill and Pepperbox Marls shown by Mortimore (1986a; Mortimore et al. 2001). However, the level of the double flint at 29.5-30 m yields abundant Hagenowia blackmorei

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and is identified as Castle Hill Flint 4. Bioevent 6 is present 1 m above this flint. The base of Mottisfont (Fig. 20; Appendix Figs. 5,6) appears, from the

faunal evidence presented here, to fall at a significantly higher level then shown by Mortimore (1986b). Rather, the microcrinoid evidence suggests that the

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section falls some way up in the Culver Chalk, with a condensed development of the Portchester Hardgrounds in CaR6 comparable to that seen on Portsdown, but with better-developed Solent Marls. Hardgrounds and marls are also present in

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CaR7 in Mottisfont which are absent elsewhere.

5.2 Correlation of the Culver Chalk (Figs 20, 21)

Correlation of the Culver Chalk in southern England has involved

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understanding the relationship between the complete section at Whitecliff, Isle of Wight and the incomplete, rather short mainland sections in Sussex and Hampshire. The correlation proposed by Mortimore (1986a,b, 2011) and Mortimore et al. (2001) was supported largely by the identification at Whitecliff

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of distinctive flints, originally identified and named in Sussex and Hampshire chalk pits. However, there is a dramatic change in flint morphology between the

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mainland (south Hampshire-West Sussex) and the Isle of Wight, with numerous lensoid tabular flints appearing in both Whitecliff and Scratchell’s Bay sections. A presumption was also made that the Sussex succession of the Culver Chalk is significantly thicker than that in the Isle of Wight (Mortimore 1986b). The Portsdown sections have been critical in the revised correlation, because they provide the only nearly continuous exposure of the Culver Chalk north of the Isle of Wight (Figs 19, 20; Appendix Figs 3,4). The Charmandean Flint (Mortimore 1986a,b) is a highly distinctive marker bed in Sussex (Figs 14, 21 ), comprising 15-20 cm dense flint lenses of up to 1 m diameter, set in a succession dominated by widely spaced layers of flint

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ACCEPTED MANUSCRIPT with burrow morphology following Thalassinoides systems. One to two metres beneath its base, in Lambley’s Lane (pit no. 7), Charmandean Lane (pit no 10) and Warningcamp (pit no. 24; Appendix Figs 1,2) bryozoan debris becomes abundant, and microcrinoids of CaR9 appear shortly beneath the flint, and continue up into the overlying beds. An identical succession can be observed in

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Candy’s Pit, Cosham (Hampshire); here the lenses of flint making up the Charmandean Flint are thinner, and the bed is less conspicuous than in Sussex

(Fig. 21; Appendix Fig. 4). At Candy’s Pit, it underlies a distinctive 30-cm-thick level of marl wisps, which afford a direct correlation to Whitecliff and

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Scratchell’s bays on the Isle of Wight, the Whitecliff Wispy Marl of Mortimore et

al. (2001) and Mortimore (2011). On the Isle of Wight, the Charmandean Flint is still present, but rather less conspicuous, and numerous other, stronger lensoid

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and semi-tabular flints appear. The identification is confirmed by the presence of the CaR8/9 boundary at Whitecliff. This placement of the Charmandean Flint at Whitecliff is 25m higher in the succession than that of Mortimore et al. (2001). At Whitecliff, Mortimore (1986a,b) named the Whitecliff Flint, a very strong, lensoid semi-tabular flint, overlying a triplet of marls, and approximately

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20m beneath the Portsdown Marl (Fig. 21; Appendix Fig. 10). In Candy’s Pit, Cosham, on the mainland, a strong flint layer, comprising closely spaced, often interconnected dense masses of flint, 15-20cm thick, is found 18m above the Charmandean Flint and was called the Cosham Flint by Mortimore (1986a). This

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overlies a distinctive triplet of thin marl seams, of which the lower two form a pair, 20cm apart (Fig. 21). This arrangement is identical to that seen in the lower

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part of Warren Farm Quarry (Gale et al. 2015), where the marls were called the Warren Farm Marls 1-3, and to the marls beneath the Whitecliff Flint at Whitecliff itself (Mortimore et al. 2001). The base of microcrinoid zone CaR10 lies 3-4m above the flint, in both Warren Farm and Candy’s Pit. In Sussex, the base of CaR10 lies 3 m above the Cote Bottom Flint in Cote Bottom Pit (no. 17 of Gaster 1924). The Whitecliff, Cosham and Cote Bottom Flints are morphologically identical, and fall at the same biostratigraphical level; they are therefore all the same bed, which is here referred to as the Whitecliff Flint (Figs 20, 21).

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ACCEPTED MANUSCRIPT The result of this new evidence is a dramatic reduction in the thickness of the upper part of the Culver Chalk on the mainland, in West Sussex and Hampshire, largely on account of the synonymy of the Whitecliff, Cote Bottom and Cosham Flints (Fig. 21). The succession developed in Sussex is remarkably similar to that seen on Portsdown Hill, Hampshire, in terms of thickness, marker

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beds and flint development, but marl seams are invariably absent above the level of the Castle Hill Marls in Sussex. Conversely, the lower part of the Culver Chalk is significantly thicker on the mainland than previously thought, and very

incompletely exposed above the level of the Castle Hill Flints in most areas.

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There appears to be a significant non-exposure of about 20m in Sussex,

representing all of microcrinoid zones CaR6 and CaR7 and the upper part of CaR5.

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The succession on the Isle of Wight (Whitecliff and in Scratchell’s Bay) is very different to that of the mainland in the massive development of thick semitabular flints in the Culver Chalk, which replace flints representing silicified Thalassinoides burrow systems which are dominant in Hampshire and Sussex (Fig. 21). The “Lancing Marls” and Solent Marls of the Isle of Wight (Mortimore et

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al. (2001) are not well represented in the mainland successions, but the latter are probably present at Mottisfont in Hampshire, and may correlate with the Portchester Hardgrounds on Portsdown Hill (Fig. 20). However, this part of the succession is very poorly exposed on the mainland, and the lateral variation is

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not well known.

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5.3 Uppermost Culver and Portsdown Chalk (Fig. 20)

The base of the Portsdown Chalk is marked by the Portsdown Marl

(Mortimore 1986a,b; M1 of Gale et al. 2015), which lies within a highly distinctive microbrachiopod level, marked by an abundance of the small, smooth terebratellid Leptothyrellopsis polonicus (Fig. 6N,O). The presence of this level at the top of the upper Warningcamp pit (no. 24 of Gaster, Arundel, Sussex; Fig. 16) indicates that the West Sussex succession probably extends into the lower Portsdown Chalk, even if marls are absent there (Fig. 20). At the level of Scratchell’s Marl 1 (M3 of Gale et al. 2015) there is a major change in the

23

ACCEPTED MANUSCRIPT roveacrinids, with the appearance of Stellacrinus pannosus in abundance, marking the base of CaR11. This change in the microcrinoids is also present in the southeast Netherlands (Gale 2016).

The development of marls within the Portsdown Chalk is highly variable,

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even over short distances. Thus, on the west of Portsdown Hill, at Warren Farm, M1-4 (Portsdown and Scratchell’s Marls) are present, but higher marls are

absent. In the east of Portsdown, in Bedhampton Limeworks, M8,9 are absent, but M10-11 present (Gale et al. 2015). Thus, without biostratigraphical

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information, identification of the base of the Portsdown Chalk, and levels within it is often difficult. For example, Mortimore and Pomerol (1991) took the base of the Portsdown Chalk in Warren Farm at the Warren Farm Marls, presuming

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these to be the Portsdown Marl, but actually some 20m lower in the succession. This was corrected by Mortimore et al. (2001).

5. Significance for mapped boundaries

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The consequences of the revised stratigraphy presented here for boundaries drawn on BGS maps are significant. Since 1995 (Bristow et al. 1995, Shaftesbury Memoir) BGS has mapped the UK Chalk Group using topographical features (essentially, breaks of slope) aligned to lithostratigraphical units, either

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derived from Mortimore (1986a,b) or of their own creation. Justification for this approach was provided by Bristow et al. (1998), and chalks of early Campanian

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age in southern England are now mapped as the Newhaven, Culver and Portsdown formations (Rawson et al. 2001). The Culver Formation was originally mapped in some areas (Dorset, south Hampshire, Sussex) as separate Tarrant and Spetisbury formations, now considered as members of the Culver Chalk (Hopson 2004). The formations and members are defined at stratotype sections by individual marker beds (Hopson 2004). When plotted against the composite succession for the Lower Campanian for southern England, the members and formations, as identified in larger exposures, vary in relative stratigraphical positions. On Portsdown, Hampshire, the base of the Portsdown Chalk has been mapped at the level of the Warren

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ACCEPTED MANUSCRIPT Farm Marls, immediately underlying the Whitecliff Flint (BGS digimap, Cosham). This falls approximately 20m beneath the Portsdown Marl itself, the chosen boundary marker (Hopson 2004). In the Arundel-Worthing district, West Sussex, BGS appears to have locally mapped the base of the Spetisbury Chalk approximately at the level of the Cote Bottom Flint (=Whitecliff Flint), although

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Bristow et al. (1988) stated that the mapped Ft2 feature in Sussex falls within Gaster’s pit 24 at Warningcamp, at about 2-3m beneath the Warningcamp Flint. This is very high in the Sussex succession, probably within a few metres of the

base of crinoid zone CaR11. Thus, the mapped base of the Portsdown Chalk on

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Portsdown falls approximately 15m above the base of the Spetisbury Chalk in

6. Systematic palaeontology

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West Sussex, some 40km distant.

Order Roveacrinida Sieverts-Doreck in Ubaghs, 1953 Family Saccocomidae d’Orbigny, 1852

Diagnosis. Roveacrinida in which the basals and centrale are small and

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commonly fused. Cavity enclosed by basals absent. Subfamily Saccocominae d’Orbigny, 1852

Diagnosis. Saccocomidae in which the cup is open on the adoral side, and in

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which elongated arms made up of primi- and secundibrachials are present.

Genus Costatocrinus Gale, 2016

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Type species. Costatocrinus brydonei Gale, 2016, by original designation Diagnosis. Theca conical, highly elongated, made up of five tall, aborally tapering, thin and delicate radials and a robust, spike-like aboral projection formed from the fused basals; radials and basals with V-shaped interdigitating contact; radial cavity broad and deep; radials with a raised central ridge with numerous less prominent parallel minor ridges present on either side; these converge with the central ridge towards the base of the plates; radial facets large, U-shaped with tall, simple interradial projections; IBr1 rectangular, external face small, IBr2 triangular, variably axillary, with broad, thin lateral flanges.

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ACCEPTED MANUSCRIPT Included species. In addition to the type species, Costatocrinus mortimorei Gale, 2016 and C. laevis sp. nov..

Costatocrinus laevis sp. nov.

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Fig. 3F

Derivation of name. From Latin ‘laevis’, meaning smooth, in allusion to the absence of sculpture on the radials.

Material. Three radial plates from the O. pillula Zone, Friar’s Bay Steps,

above these (SH18; Appendix, Fig. 1).

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Peacehaven, Sussex, level of the Rottingdean Marls (sample SH19) and 1.6m

Type. The fragmentary radial figured here is holotype. NHMUK EE16222. Sample

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SH18, 1.6m above Rottingdean Marls, Friar’s Bay Steps, Peacehaven, Sussex. Description. The radial plates are all incomplete, lacking the adoral articular region. They possess a single, centrally placed, narrow aboral-adoral ridge, which has a rounded top. The surfaces lateral to the ridge are quite smooth. The interradial contacts are strongly and rather coarsely sutured.

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Remarks. Costatocrinus laevis sp. nov. differs from both C. brydonei and C. mortimorei in the absence of any sculpture on the radials, except the central ridge. It occurs stratigraphically lower than C. brydonei, which appears a short distance above the Peacehaven Marl, and is probably ancestral to it. The rather

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large amount of material of C. brydonei (200+radials) invariably possess vertically oriented ridges and grooves lateral to the central ridge (Fig. 3A-C),

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which suggests that it is not simply be a variant of that species.

Genus: Assericrinus nov.

Derivation of name. From Latin, ‘asser’, meaning stave or beam, alluding to the shape of the vertically oriented portion of the elongated processes on the radials. Type species. Assericrinus portusadernensis sp. nov. Diagnosis. Saccocominae in which the adoral margin of each radial bears a process, comprising a short lateral spur and a tall, vertically oriented bar with a distinctive triradiate cross section.

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ACCEPTED MANUSCRIPT Remarks. Amongst the material from the UK lower Campanian chalks are numerous three-lobed, elongated bars, with a flattened process set at right angles to the axis of the bar. Although originally identified as Echinodermata problematica, well-preserved material from Waxahachie, Texas (USA), demonstrates that these are radial spines, laterally directed extensions from a

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thin, fragile radial plate. A reconstruction of Assericrinus gen nov. is provided (Fig. 26), based on new material from Texas.

Assericrinus portusadernensis sp. nov.

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Fig. 4A-E,I; Fig. 22B

Derivation of name. Roman name for Portchester, Hampshire, UK, Portus Aderni,

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from which the type material originated.

Material. 50 radial spines from the lower Campanian G. quadrata Zone, Paulsgrove (Portchester) and Cosham, Hampshire, UK.

Type specimens. The radial spine figured (Fig. 4D) is holotype, NHMUK EE 16224, from sample PG14. The other specimens figured are paratypes (Fig. 4A,B,

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NHMUK EE 16225, sample PG8; Fig. 4C, EE 16226, sample PG13; Fig. 4E, EE 16227, sample PG13; Fig. 4I, EE 16228, from sample PG8, all from the G. quadrata Zone of the upper part of Paulsgrove pit, Hampshire (see Appendix Fig. 3 for detailed section).

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Diagnosis. As for genus.

Description. Only radial spines are known. The radial spines were attached to the

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body of the radial plate by an elongated, laterally flattened shaft-like process (Fig. 4A-E,I), which was set at an oblique angle to the exterior surface of the radial. The vertical portion of the spine is at right angles to the shaft, and is a robust structure with a three-cornered cross section (Fig. 4I), which tapers both ab- and adorally to blunt points. The lateral surface forms a shallow groove (Fig. 4I), bordered by two solid, rounded ridges. What appear to be immature individuals (Fig. 4C) possess only a very short vertical extension. Remarks. Since the original submission of this paper, I have collected complete but crushed cups of A. portusadernensis gen et sp. nov. from the Ozan Formation at Waxahachie dam spillway, Texas (details of site on Gale et al. 2008). A drawing

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ACCEPTED MANUSCRIPT based on this new material is included (Fig. 22B), and shows that the aboral portion of the cup is extended into a very elongated, narrow aboral process

Subfamily Applinocrininae Peck, 1973 Diagnosis. Theca conical to fusiform, delicately constructed, consisting of a basal

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circlet, usually fused, made up of five basals and a small centrale; five convex, trapezoidal radials; arms reduced to a single brachial, flattened, highly modified, triangular, imbricating; the five brachials form a cap over the cup (Gale 2016).

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Genus Applinocrinus Peck, 1973

Type species. Saccocoma cretacea Bather, 1924, by the original designation of Peck (1973).

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Included species. In addition to the type species, A. texanus Peck, 1973 and A. russelli Gale, 2016.

Diagnosis. Applinocrininae in which the brachials are equilaterally triangular, imbricated, and form a precisely articulated low, conical cap above the theca. The double articular structure comprises a larger, adoral, oval flange which

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imbricates a corresponding facet on the adjacent plate, and a lower small, low process which fits into a notch close to the base of the adjacent plate.

Fig. 1I,J,N,O

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Applinocrinus cretaceus forma cretaceus (Bather, 1924)

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Diagnosis. Applinocrinus in which the relatively low, broad radials are smooth, as are the brachials.

Type. A perfect theca with brachials in place, from the G. quadrata Zone of Durrington in Sussex, pit no. 17 (Cote Bottom Pit) of Gaster (1924), NHMUK E 24767 (figured by Bather 1924, p. 113, figs. 8, 9; Rasmussen 1961, pl. 57, figs. 9, 10; Hess and Messing 2011, fig. 108/2a, b; and Smith and Wright 2002, pl. 45, fig. 14).

Applinocrinus cretaceus forma spinifer nov. Fig. 3V,W

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ACCEPTED MANUSCRIPT p.2016 Applinocrinus cretaceus (Bather), Gale p. 12 (part) Fig. 5A,D; 7A,I,J only.

Diagnosis. Applinocrinus cretaceus in which an elongated, adorally curved, laterally compressed, tapering spine is present on the radial plate.

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Material. Over 100 radial spines from the G. quadrata zone of Sussex, Hampshire and Wiltshire. Material also from the Ozan Formation, Waxahachie dam spillway, Texas, and Ivõ Klack, southern Sweden (Gale 2016 fig. 8A,I,J).

Types. The elongated spine with a preserved radial facet figured in Fig. 3W is

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holotype, NHMUK EE 16228, Paulsgrove pit, Hampshire, sample PG14

(Appendix, Fig. 3). The other figured spine is paratype, EE 16229., sample WF8, Warren Farm, Hampshire (see Appendix, fig. 4).

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Remarks. Gale (2016) considered that the development of a centrally positioned, single tapering spine on the radial plates of Applinocrinus was highly variable, and forms with no spines intergraded with forms possessing short spines (e.g. Gale 2016 fig. 7A) and those with longer spines. This is true, but spines are entirely absent from radial plates found within the lower part of the range of the

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species; they appear very rarely a short distance above the Pepperbox Marls (e.g. sample PG16, Appendix, Fig. 3) and at this level the spines are short and rounded. They appear as a common element of the fauna at a higher level (PG13, Appendix Fig. 3), and here the spines are highly elongated and often recurved

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adorally. These forms continue into the highest levels sampled, and are here distinguished as forma spinifer nov.

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Description. The radial spines are preserved with the robust radial articulation at the base (Fig. 3W), but the remainder of the radial plate is invariably broken away. A single, centrally placed, elongated spine extends in an oblique, adoral direction from the surface of the radial, immediately aboral to the articular facet. The spines are 3 to 4 times longer than the width of the radial plates, and very variable morphologically. Some are straight, evenly tapering, and rounded in cross-section (e.g. Gale 2016 Fig. 5A, fig.7I,J), but many are laterally compressed towards the tip (Fig. 3V-W). The tip of the spine is often flexed adorally (Fig. 3.V) to a variable extent.

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ACCEPTED MANUSCRIPT Genus Saggitacrinus Gale, 2016

Type species. Saggitacrinus torpedo Gale, 2016, by original designation.

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Diagnosis. Brachials flat, shaped like arrowheads, with an oval adoral margin and three pointed aboral processes. The central process is tall, robust, and externally bears three ridges separated by deep grooves. Internally, it carries an adorally tapering flattened surface. The lateral processes are short, flat and pointed. A

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raised, trapezoidal articular structure for the radial facet is present on the inner face close to the adoral margin. No specialized articular structures are present between brachials, which simply overlapped.

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Remarks. Further collecting has demonstrated the presence of two additional species of early Campanian Sagittacrinus, characterized by the morphology of the brachial plates.

Sagittacrinus alifer sp. nov.

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Fig. 3R,T,U; Fig. 22A

Derivation of name. From the Latin ‘alifer’, meaning wing-bearer. Diagnosis. Sagittacrinus which possess a single, flattened, elongated, slightly

central ridge.

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aborally directed process on the left side of the brachial, and a raised narrow

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Material. Over 50 brachial plates, mostly from CaR6 in the upper part of Paulsgrove pit, Hampshire, but the species occurs at a higher level (Whitecliff Flint) as a rarity (Fig. 17). Types. The brachial figured here (Fig. 3R) is holotype, NHMUK EE 16230, from sample PG8, Paulsgrove pit, Hampshire; the other two brachials (Fig. 3T,U) are paratypes (NHMUK EE 16231; also sample PG8; EE 16232 sample PG13). G. quadrata Zone, see Appendix Fig. 4 for details. Description. The brachials are strongly asymmetrical, with a flattened, parallel sided process on the left side, and a short right margin which diverges slightly adorally (Fig. 3R,U). The process, which is four to five times broader than the

30

ACCEPTED MANUSCRIPT body of the plate, is deflected aborally. The exterior surface carries a swollen, narrow, central fridge which is vertically oriented, and flanked by two grooves. The radial margin is rounded. The brachials are never complete, and the adoral projection is broken away in all specimens. The interior surface (Fig. 3T) carries a rectangular process close to the radial margin which stands proud from the

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flattened interior surface. This bears a series of three, paired invaginations, which possibly carried podia (see Gale 2016 for discussion of the related S. torpedo).

Remarks. Sagittacrinus alifer sp. nov. is distinguished from the other two species

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of the genus by the elongated lateral process. A reconstruction (Fig. 26A) shows the very distinctive form of the species, in which the lateral processes project divergently from the margin of the cup. The striking asymmetry of this species,

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with a single elongated wing-like projection on the brachials, means that it would have spiraled in the water column. As these Roveacrinida were pelagic (Gale 2016) this means that it would have sampled a larger water volume as it ascended and descended, by comparison with graptolites (Rigby 1992).

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Sagittacrinus longirostris sp. nov. Fig. 3K,PQ, X.

Derivation of name. From the Latin, meaning long-nosed, with reference to the

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extended adoral process.

Diagnosis. Sagittacrinus with a very elongated, narrow, adoral process which

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bears a deep groove on the interior surface. Material. 46 brachials, mostly from the CaR6 zone on Portsdown Hill, Hampshire, G. quadrata zone.

Types. The brachial figured (Fig. 3P) is holotype, NHMUK EE 16233. Paulsgrove pit, Hampshire, sample PG10. The three other figured specimens (Fig. 3K,Q, X) are paratypes (Fig. 4K, NHMUK EE 16234, sample PG10; Figs Q,X, EE16235, 16236, both from sample PG14 – see Appendix Fig. 4 for details). Description. The brachials consist of an aboral flattened portion which is arrowhead shaped, and an elongated, narrow, adoral projection, which is 3 to 4 times the height of the basal part (Fig. 3P). A raised, narrow ridge runs from the radial

31

ACCEPTED MANUSCRIPT margin to the tip of the plate, and this broadens adorally (Fig. 3Q). The sides of the projection are raised and rounded, and separated by a median groove on the interior of the plate (Fig. 3K). The basal region is flattened, and raised into a barb-like projection on one side of the plate. The interior of the basal region carries a projecting trapezoidal structure, the base of which articulated with the

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radial facet (Gale 2016). The structure bears pits and grooves.

Remarks. Sagittacrinus longirostris sp. nov. is distinguished from the other two described species of the genus known by the elongated, three-sided adoral

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process.

Family Roveacrinidae Peck, 1943

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Diagnosis. Roveacrinida in which a double thecal cavity is present, the lower of which is enclosed by basals or basals and radials. Subfamily Hessicrininae Gale, 2016.

Diagnosis. Roveacrinidae in which the basals are enlarged and form the aboral portion of the theca. The basals are triangular, often fused, and possess adoral

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processes which contact the radials. The basal cavity is separated from the radial cavity by a basal web comprising a complex pentagonal structure made up of fine calcite struts, the corners of which are situated interradially. Large, radially positioned fenestrae are present between the basal/radial contacts, and

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interbasal fenestrae are often present. The lateral parts of the radials are perforated by small fenestrae and foramina, and from the interior; the cup has a

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reticulate appearance. The proximal brachials possess sheet-like lateral flanges.

Genus Hessicrinus Gale, 2016. Diagnosis. Hessicrininae in which the cup has a large adoral opening and a deep radial cavity. The height of the radials is twice that of the basals; the radials carry very elongated, laterally flattened spines, and are perforated by numerous foramina, which give a reticular appearance. The basals are rectangular, and form the base of the cup; two radial/basal foramina are present, of which the more aborally positioned one is the larger. IBr2 is not axillary. The brachials possess delicate, wing-like lateral processes.

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ACCEPTED MANUSCRIPT Type species. Hessicrinus filigree Gale, 2016 by original designation. Included species. In addition to the type species, H. scalaensis Gale, 2016, H. cooperi sp. nov. and H. apertus sp. nov.

Hessicrinus cooperi sp. nov.

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Figs 4G,L,P, 22D

2016 Hessicrinus aff. filigree Gale, 2016, p. 26-7, Fig. 2K, Fig. 11A,B.

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Diagnosis. Hessicrinus in which large radial/basal fenestrae are not developed, but a number of small perforations are present at the adoral margin of the

radial/basal contact. The basal plates form a star-shaped, conical, aboral base to

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the cup, perforated by numerous small foramina.

Derivation of name. Named in honour of John Cooper of the Booth Museum, Brighton, who has done much to help my work on chalk fossils. Material. 25 cups and fragmentary cups from the O. pillula Zone of Sussex, from the Friar’s Bay Marl 3 to about 3m above the Planoconvexa Bed.

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Types. The cup also figured by Gale (2016, fig. 11A,B; Fig. 4L,P herein) is holotype. NHMUK EE EE 16111, 4m above the Peacehaven Marl, Newhaven (sample SH13, Appendix Fig. 1) the other figured cup (Fig. 4G) is paratype, NHMUK EE 16236, Sample SH9a, Newhaven (Appendix Fig. 1).

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Description. The cup is rhomboidal in lateral aspect, with prominent, sharp radial ridges, each of which bears an elongated, laterally compressed spine

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immediately aboral to the radial facet (Fig. 4G,P). Raised ridges are present at the junction between adjacent radials. The interradial processes are tall, but broken adorally in most specimens. Numerous deep foramina are present between the radial and interradial ridges, and extend onto the interradial processes. An aboral process from each radial descends between each basal pair (Fig. 22D), and the basal/radial contact is marked by small fenestrae. The basals form a low, gently convex aboral margin to the cup, star-shaped in aboral aspect, which is perforated by tiny foramina (Fig. 4L). Each basal bears a single obliquely directed, laterally compressed spine, the tips of which are broken.

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ACCEPTED MANUSCRIPT Remarks. Hessicrinus cooperi sp. nov. lacks the enlarged radial/basal fenestrae of the descendant species H. filigree, rather it possesses small perforations at the radial/basal contact (Fig. 22). Additionally, the basals are more robust, and perforated by numerous tiny foramina. The transition evidently took place between a level 3m above the Planoconvexa Bed and the overlying Castle Hill

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Marls, from which interval complete cups needed for identification have not been collected in Sussex. However, both H. cooperi sp. nov. and H. filigree and

transitional forms are present in sample WH1 at West Harnham (Appendix Fig.

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5), at a level 2m above the top of the Planoconvexa Bed, Telscombe Marl 3.

Hessicrinus apertus sp. nov.

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Figs 4Q,R,V; Fig. 22C

Diagnosis. Hessicrinus in which the base of the cup is broad and stellate with five basal spines and radially arranged, irregular, elongated foramina; single interbasal fenestrae, single radial: basal fenestrae, and single large, deep interbasal foramen in the centre of each basal plate.

fenestrae.

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Derivation of name. Latin ‘apertus’ meaning open, in reference to the large

Material. Four incomplete cups from the G. quadrata Zone of Portsdown Hill,

Sussex.

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Hampshire, one from the G. quadrata Zone of Warningcamp pit no. 24, Arundel,

Types. The cup illustrated in Fig. 4V is holotype, NHMUK EE 16237, from sample

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1b, Warningcamp pit no. 24, Sussex (Appendix Fig. 2); the other two cups (Fig. 4Q, Fig. 4R) are paratypes, (Fig.4Q, EE 16237 is from sample PG8, Paulsgrove pit, Hampshire; Fig. 4R from the base of Cliff Dale Gardens pit, Cosham, Hampshire, sample CD-3, details in Appendix Figs 3,4). Description. The cups all lack the adoral portions of the radials, but the basal circlet and the aboral portions of the radials are well preserved. The aboral margin of the cup is gently convex and broad, with five rounded basal spines at the interradial extremities. The base is perforated by variably shaped and sized foramina, the largest of which are oblong and have a radial arrangement. Single, large, deep interbasal fenestrae are present at each basal contact, and a large,

34

ACCEPTED MANUSCRIPT rounded centrally positioned basal foramen is present on each plate. A pair of irregularly shaped radial/basal fenestrae are present, positioned between the adoral processes of each basal plate. The radials are perforated by foramina which decrease in size adorally. Remarks. Hessicrinus apertus sp. nov. is closest to H. scalaensis Gale, 2016 in the

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broad, pentagonal aboral base of the cup, with five prominent basal spines, and a single, large basal foramen in each plate, but differs in the single, large interbasal fenestra, the single pair of radial/basal fenestrae, and the radial arrangement of

elongated foramina on the base of the cup. It probably evolved from the older H.

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scalaensis, but there is a large gap between the occurrences.

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Stellacrinus Gale, 2016

Diagnosis. Derived Hessicrininae, in which the radials bear a centrally placed, elongated, and sigmoidally recurved spine. Radials weakly articulated, with one aboral and one adoral fenestra between abutting articular struts, and small central fenestrae adjacent to the radial spine. Basals form a fused ring which

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carries aborally deflected elongated blades, and tall, adorally expanding processes for articulation with the radials. Radial and basal cavities are separated by a thin, transverse, web-like complex network of fine, pentagonally arranged struts. Primibrachials and proximal secundibrachials carry irregularly

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shaped thin lateral projections.

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Type species. Stellacrinus hughesae Gale, 2016 by original designation.

Stellacrinus hughesae forma lineatus nov. Fig. 5I

Diagnosis. Stellacrinus hughesae in which the basal spines are elongated, daggershaped, and possess a low longitudinal ridge. Material. 30 basals from the G. quadrata Zone on Portsdown Hill, Hampshire, mostly from the upper part of Paulsgrove pit, samples PG13-11 (Appendix Fig. 3).

35

ACCEPTED MANUSCRIPT Types. The basal plate illustrated in Fig. 5I is holotype. NHMUK EE 16240. Sample PG13, G. quadrata Zone, Paulsgrove pit, Hampshire (Appendix Fig. 3). Derivation of name. Latin, ‘lineatus’, bearing a line, in reference to the ridge on the basal spine. Description. The basal plate is laterally compressed, elongated (up to 10 times

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longer then high) and tapers to a sharp point. The aboral margin is sharp and blade-like, the adoral margin is slightly rounded. A narrow raised ridge runs from the proximal portion to the tip along each side of the plate, set slightly above the mid-line. A short adoral articulation process is present.

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Remarks. Stellacrinus hughesae forma hughesae invariably possesses broad, rather blunt and short basal spines (Fig. 5A,D), and is characteristic of the

O.pillula Zone. In the lower G. quadrata Zone, from 12m above the Pepperbox

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Marl in Paulsgrove pit distinctive basals are found which clearly belong to Stellacrinus, but which are very elongated, sharp-tipped and dagger-shaped, and possess a low ridge running along the mid-height of the blade. These are one of the characteristic fossils of zone CaR6 (Fig. 17). The radial plates from the same samples do not appear to differ significantly from those of older material of S.

7. Conclusions.

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hughesae.

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Microcrinoids of the order Roveacrinida are abundant and diverse in lower Campanian chalks of southern England, and some are relatively short-

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ranged. They progressively increase in diversity from the upper Santonian, and reach their diversity acme in the lower third of the G. quadrata Zone, above which the diversity decreases, but not the abundance (Fig. 17). New forms described herein (see section Systematic palaeontology) adding to the diversity described by Gale (2016). The total of 25 species and forms recognized here provide the basis for a new zonation, which includes zones defined as CaR1-11. The zones are largely based upon first occurrences, last occurrences and acmes. Together with evidence from levels of abundance of holasteroid echinoids (Offaster, Hagenowia) and shape variants of Echinocorys, these provide a new integrated biostratigraphical framework for the lower Campanian of southern

36

ACCEPTED MANUSCRIPT England which enables an evaluation of correlations based primarily upon lithological features. The correlation framework for the upper part of the Newhaven and lower part of the Culver formations (O. pillula and lower G. quadrata Zones) originally proposed by Brydone (1914) and extended by Mortimore (1986a)

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with the addition of extensive lithological detail, including numerous named marl seams and some flints, is largely supported by the new biostratigraphical

data (Figs 18,19). However, there is considerable lateral lithological variation in both the overall abundance of marly levels, and the development of individual

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marl seams. Thus, the chalk on the Dorset coast (Middle Bottom) is virtually clayfree, whereas that at Scratchell’s Bay in the Isle of Wight contains abundant thin marly beds. Individual marls and groups of marls vary significantly laterally. For

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example, the Meeching Triple Marls in Sussex and Wiltshire appear to merge into a single bed at Portsdown in Hampshire; the variably developed Pepperbox Marls are absent east of Arundel in Sussex (Mortimore 1986a) and represented by a single bed locally in Dorset. In spite of this, the major marker marl beds (Old Nore, Peacehaven, Meeching, Telscombe and Castle Hill) are remarkably laterally

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persistent, and synchronous biostratigraphically (Figs 18,19). For example, the acme of the crinoid Stellacrinus hughesae forma cristatus in and immediately above the Old Nore Marl provides supporting evidence for the correlation of this bed across the region.

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In contrast, the correlation framework proposed for the Culver Chalk (Mortimore 1986a,b, 2011; Mortimore et al. 2001), based largely upon the

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inferred correlation of flint beds from Whitecliff, Isle of Wight, to the mainland, is not supported at all by the biostratigraphy (Fig. 20). Correlation has been confused by several factors: 1) a significant change in flint morphology takes place between the Isle of Wight (lensoid and semi-tabular morphologies) and the mainland (predominantly silicified Thalassinoides system flints; Fig. 21). 2) the Sussex succession of the Culver Chalk is significantly thinner than proposed previously (Mortimore 1986b). 3) the complete absence of marker marl seams in the Culver Chalk in Sussex hampers correlation to the west.

37

ACCEPTED MANUSCRIPT The Portsdown, Hampshire, successions have been the key to understanding the correlation between the Isle of Wight and Sussex proposed here (Fig. 20), based upon flint and marl correlation supported by microcrinoids. This leads one to infer that the Cosham, Cote Bottom and Whitecliff flints

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(Mortimore 1986a,b) are synonymous, and that the Charmandean Flint is significantly higher in the Whitecliff succession than proposed by Mortimore et al. (2001). This has the effect of significantly decreasing the thickness of the

upper Culver Chalk, but the presence of a significant exposure gap in Sussex

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(perhaps 20 metres) is probable because microcrinoid zones CaR6 and 7 are not found there. The gap lies above the top of the North Lancing sections and the base of Charmandean Lane. In general, this interval appears to be very poorly

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exposed in mainland southern England.

The abundance of the small, smooth brachiopod Leptothyrellopsis polonicus in the uppermost Culver Chalk and the sudden appearance and abundance of Stellacrinus pannosus at the level of Scratchell’s Marl 1 and above provides further useful guides to correlation of the upper Culver and Portsdown

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chalks. It is necessary to calibrate these events with the detailed benthonic foraminiferal changes identified by Hopson et al. (2014) in Scratchell’s Bay (Isle of Wight).

It is probable that the microcrinoid stratigraphy described here is also

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present in other regions of Europe, and further afield, because ten of the taxa from southern England are present in the lower Campanian of Texas (Gale

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2016).

Acknowledgements

This paper is dedicated to the memory of Christopher John Wood, and I hope, probably vainly, that his exacting attention to detail and precise knowledge of chalk faunas are reflected in its contents. He was my first chalk mentor, who guided me through 25 years of my studies of the Cretaceous. I would like to thank Christine Hughes (University of Portsmouth), whose help with SEM

38

ACCEPTED MANUSCRIPT imaging has made my work over the past 10 years possible, and Hans Hess (Basle), whose extensive knowledge of the post-Palaeozoic crinoids has proved such a help and encouragement. I would also like to thank Professor Ian Jarvis (Kingston University) for the excellent photo of Scratchell’s Bay used in Appendix Fig. 20, and landowners and managers for permission to visit their

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properties, notably Mark Yorke (Mottisfont), Pat Riddell (Veolia Ltd, Warren Farm), George Vince (Charmandean Lane), and Mike Tristram, Lambley’s Lane (Sompting Estate). I thank John Jagt and Ian Jarvis for most useful reviews.

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genres, et des espèces de la classe des échinoderms. Annales des Sciences Naturelles, Troisieme Série, Zoologie, 6, (1846), 305-374, pls 15,16; 7 (1847) 129-168; 8 (1847), 5-35, 355-380.

Bailey, H.W. 1978. A foraminiferal biostratigraphy of the Lower Senonian of southern England. Unpublished PhD thesis, Plymouth Polytechnic. 328pp.

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Bailey, H.W., Gale, A.S., Mortimore, R.N., Swiecicki, A. and Wood, C.J. 1983. The Coniacian-Maastrichtian Stages of the United Kingdom, with particular reference to southern England. Newsletters on Stratigraphy, 12, 29-42. Barrois, C. 1876. Recherches sur le terrain Crétacé supérieur de l’Angleterre et

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de l’Irlande. Mémoires de la Société Géologique du Nord, 1, 234pp. Bather, F.A. 1924. Saccocoma cretacea n. sp., a Senonian crinoid. Proceedings of

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the Geologists’ Association, 35, 111-121.

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ACCEPTED MANUSCRIPT Brünnich Nielsen, K.B. 1942. Martinosigra elongata n.g. and n.sp. A new echinoid from the White Chalk of Denmark. Meddelelser fra Dansk Geologisk Forening, 10,2, 159-166. Brydone, R.M. 1912. The stratigraphy of the Chalk of Hants. Dulau and Co, London. 116 pp.

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Brydone, R.M. 1939. The chalk zone of Offaster pilula. Dulau and Co., London. 3-8. Brydone, R.M. 1914. The Zone of Offaster pilula in the south English Chalk, Parts I-IV. Geological Magazine, New Series, Decade VI, 1, 359-69, 405-11, 44957, 509-513.

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Burnett, J. A. 1998. Chapter 6, Upper Cretaceous. In: Bown, P.R. (ed), Calcareous Nannofossil Stratigraphy. Chapman and Hall, London. Pp, 132-199. Destombes, P. and Breton, G. 2001. Platelicrinus campaniensis nov. gen. nov. sp.

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Roveacrinidae (Echinodermata, Crinoidea) du Campanien des Charentes, France. Bulletin trimestriel de la Société géologique de Normandie et des Amis du Muséum du Havre, 87, 37-40.

Douglas, J.A. 1908. A note on some new Chalk crinoids. Geological Magazine, new series, (5)5, 357-359.

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Ernst, G. 1971. Biometrische Untersuchungen über die Ontogenie und Phylogenie der Offaster/Galeola-Stammesreihe (Echinoiden) aus der nordwesteuropäischen Oberkreide. Neues Jahrbuch für Geologie und Paläontologie Abhandlungen139, 169-225.

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Ernst, G. 1972. Grundfragen der Stammesgeschichte bei irrgulären Echiniden der norwesteuropäischen Oberkreide. Geologisches Jahrbuch A4, 63-175.

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Ernst, G. and Schulz, M.-G. 1971. Die Entwicklungsgeschichte der hochspezialisierten Echiniden-Reihe Infulaster-Hagenowia in der Borealen Oberkreide. Paläontologische Zeitschrift 45,120-143.

Ernst, G. and Schulz, M.-G. 1974. Stratigraphie und Fauna des Coniac und Santon im Schreibkreide-Richtprofil von Lägerdorf (Holstein). Mitteilungen aus dem Geologisch-Paläontologischen Institut der Universität Hamburg 43, 5-60. Forbes, E. 1852. Figures and Descriptions illustrative of British Organic Remains. Decade 4. Memoirs of the Geological Survey of the United Kingdom. Pls 110.

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ACCEPTED MANUSCRIPT Gale, A.S. 1980. Penecontemporaneous folding, sedimentation and erosion in Campanian Chalk near Portsmouth, England. Sedimentology, 27, 137-151. Gale, A.S. 2016. Roveacrinida (Crinoidea, Articulata) from the SantonianMaastrichtian (Upper Cretaceous) of England, the US Gulf Coast (Texas, Mississippi) and southern Sweden. Papers in Palaeontology doi:

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10.1002/spp2.1050, 44pp. Gale, A.S., Kennedy, W.J., Lees, J.A., Petrizzo, M.R. and Walasczyk, I. 2008. An integrated study (inoceramid bivalves, ammonites, calcareous

nannofossils, planktonic foraminifera, stable carbon isotopes) of the

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Waxahachie dam spillway section, Ellis County, north Texas, a candidate Global boundary Stratotype Section and Point for the base of the Campanian Stage.Cretaceous Research, 29, 131-167.

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Gale, A.S., Surlyk, F. and Anderskouv, K. 2013. Channelling versus inversion: origin of condensed Upper Cretaceous chalks, eastern Isle of Wight, UK. Journal of the Geological Society.170, 281-290.

Gale, A.S., Anderskouv, K., Surlyk, F. and Whalley, J. 2015. Slope failure of chalk channel margins: implications of an Upper Cretaceous mass transport

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complex, southern England. Journal of the Geological Society, London, http://dx.doi.org/10.1144/jgs2014-124 Gale, A.S. and Cleeveley, R. 1989. Arthur Rowe and the zones of the White Chalk

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ACCEPTED MANUSCRIPT Lamarck, J.B.P.M. 1816. Histoire naturelle des animaux sans vertébres, présentant les charactères généraux et particuliers de ces animaux, leur distribution, leurs classes, leurs familles, leurs genres, et le citation des principales espèces qui s’y rapportent; précédée d’une introduction offrent la détermination des charactères essentiells de l’animal, sa

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ACCEPTED MANUSCRIPT Orbigny, A.D. d’. 1850-1852. Prodrome de paléontologie stratigraphique universelle des animaux mollusques et rayonnés faisant suite au cours élémentaire de paléontologie et de géologie stratigraphique. Masson, Paris. volume 1. 1850, 394 pp.; volume 2, 1852, 427 pp.; volume 3 (1852), 196 pp.

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Peck, R.E. 1973. Applinocrinus, a new genus of Cretaceous microcrinoid, and its distribution in North America. Journal of Paleontology, 47, 94-100. Rasmussen, H.W. 1961. A monograph on the Cretaceous Crinoidea. Biologiske

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ACCEPTED MANUSCRIPT Rigby, S. 1992. Graptolite feeding efficiency, rotation and astogeny. Lethaia, 21, 51-68. Rowe, A.W.E. 1902. The Zones of the White Chalk of the English Coast, I. – Kent and Sussex. Proceedings of the Geologists’ Association, 16, 283-368. Rowe, A.W.E. 1903. The Zones of the White Chalk of the English Coast, II. –

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Dorset. Proceedings of the Geologists’ Association, 17, 1-76. Rowe, A.W.E. 1904. The Zones of the White Chalk of the English Coast, III Devon. Proceedings of the Geologists’ Association, 18, 1-52.

Rowe, A.W.E. 1905. The Zones of the White Chalk of the English Coast, IV.

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Rowe, A.W.E. 1908. The Zones of the White Chalk of the English Coast. V. Isle of Wight. Proceedings of the Geologists’ Association, 20, 209-352.

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Schmid, F. 1972. Hagenowia elongata (Nielsen), ein hochspezialisierter Echinide aus dem höheren Untermaastricht NW-Deutschlands. Geologisches Jahrbuch, 4, 177-195.

Smith, A.B. 1984. Echinoid Palaeobiology. George Allen and Unwin, London, Boston, Sydney. 190pp.

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Smith, A.B. and Wright, C.W. 2003. British Cretaceous echinoids. Part 7, Atelostomata, 1. Holasteroida. Monographs of the Palaeontographical Society, London, 156 (619), 440-568. Smith, A.B. and Wright, C.W. 2002. Echinoderms. In: Smith, A.B. and Batten, D.J.

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(eds), Fossils of the Chalk. Palaeontological Association Field Guides to Fossils, No. 2, Second edition, revised and enlarged. Pp. 251-295.

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Surlyk, F. 1972. Morphological adaptations and population structures of the Danish Chalk brachiopods (Maastrichtian, Upper Cretaceous). Det

Kongelige Danske Videnskabernes Selskab,19, 1-57.

Surlyk, F. 1984. The Maastrichtian Stage in NW Europe and its brachiopod zonation. Bulletin of the Geological Society of Denmark, 33, 217-23.

Swiecicki, A. 1980. A foraminiferal biostratigraphy of the Campanian and Maastrichtian chalks of the United Kingdom. Unpublished PhD thesis, Plymouth Polytechnic. 2 vols, 1, 358pp, 2, 155pp. Ubaghs G. 1953. Sous-Classe 4, Articulata J.S. Miller. In: Piveteau, J. (Ed.), Traité de Paléontologie 3, Masson, Paris. 756-765.

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ACCEPTED MANUSCRIPT White, H.J.0. 1913. The Geology of the Country near Fareham and Havant. Memoirs of the Geological Survey. England and Wales. Explanation of Sheet 316. HMSO, London. 96pp. White, H.J.O. 1921. A short account of the Geology of the Isle of Wight. HMSO, London. 219pp.

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White, H.J.O. 1924. The Geology of the Country near Brighton and Worthing. Memoirs of the Geological Survey. England and Wales. Explanation of Sheets 318 and 333. HMSO, London. 114pp.

Willcox, N.R., 1953. Zonal variations in selected morphological features of

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Echinocorys scutata Leske. Geological Magazine 90, 83-96.

Wood, C.J. and Mortimore, R.N. 1988, ch.6, Biostratigraphy of the Newhaven and Culver Members. In: Young, B. and Lake, R.D. 1988. Geology of the country

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around Brighton and Worthing. Memoir for 1:50 000 geological sheets 318 and 313 (England and Wales). HMSO, London,pp 58-64. Wright, T. W. 1882. A Monograph on the British Fossil Echinodermata from the Cretaceous Formations. 1. The Echinoidea. Part 10, pp. 325-371, i-xviii, pls 1-8, 76-80. Monographs of the Palaeontolographical Society, London.

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Wright, C.W. and Wright, E.V. 1949. The Cretaceous echinoid genera Infulaster Desor and Hagenowia Duncan. Annals and Magazine of Natural History,

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12, 454-474.

Figure captions.

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Fig. 1. Table to show bio- and lithostatigraphical classification of upper Santonian to upper Campanian chalks in southern England. From left to right, columns illustrate the development of a macrofossil zonal scheme through the the twentieth century (Rowe, 1902; Brydone 1912; Gaster 1924; Rawson et al. 1978. The benthonic foram zonation is after Swiecicki (1980) and Hart et al. 1989, the nannofossil zonation after Lees, 1998, the lithostratigraphy, after Mortimore (1983,1986a) and Rawson et al. (2001).

Fig. 2. Summary geological map of southern England to show major localities studied in the present paper. Modified from BGS online digimap. The two boxes

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ACCEPTED MANUSCRIPT show the positions of more detailed locality maps provided in the Appendix, Fig. 2 for detailed locations of Sussex pits and Appendix Fig. 3 for detailed locality map of Portsdown Hampshire. See Table 1 for details of localities. Numbers are as follows: 1. Seaford Head, Sussex, cliffs immediately east of town of Seaford. O. pillula and lower G. quadrata zones (Mortimore 1986a,b). 2. Newhaven, Sussex.

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Cliffs immediately west of town, beneath Castle Hill. O. pillula and lower G. quadrata zones (Mortimore 1986a,b). 3. Friar’s Bay Steps, Peacehaven, Sussex. 4, Black Rock, Brighton, cutting immediately east of marina. Marsupites and O.

pillula zones (Mortimore 1986a,b; Wood and Mortimore 1988). 5, North Lancing,

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Sussex, old chalk pit north of town, pit no. 2 of Gaster 1924. G. quadrata Zone. 6, Cote Bottom chalk pit, disused pit west of Durrington, Worthing, Sussex. Pit no.17 of Gaster (1924). Whitecliff Flint (= Cotes Bottom Flint) near base of

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section 7, old chalk pit north of village of Warningcamp, Arundel, no. 26 of Gaster (1924). G. quadrata Zone, Charmandean Flint near base. 8, Warningcamp, Arundel, Sussex. old chalk pit north of village, no. 24 of Gaster (1924). 9, Candy’s Pit, London Road, north of Cosham, Hampshire. Whitecliff Flint near summit. No. 1149 of Brydone (1912). G. quadrata Zone and adjacent Cliff Dale Gardens pit,

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London Road, north of Cosham, Hampshire, no. 1150 of Brydone (1912). 10, Paulsgrove pit, north of Portchester. O.pillula and G. quadrata Zones. 11, Portchester pits of White (1913), G. quadrata zone. 12, Warren Farm quarry, Fareham, Hampshire (Gale et al. 2015). Whitecliff Flint to Farlington Marls. 13,

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Whitecliff, Isle of Wight. Entire succession of Santonian-Campanian. 14, Scratchell’s Bay, Isle of Wight. Entire Santonian-Lower Campanian. 15, Middle

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Bottom, cliffs west of West Lulworth, Dorset (Brydone 1914). 16, Tarrant Rowston, Dorset. Lower part of G. quadrata Zone. 17, West Harnham chalk pit, Salisbury, Wiltshire. O. pillula Zone, and basal G. quadrata Zone.. 18, East Grimstead, Wiltshire, abandoned quarry. O.pillula and basal G. quadrata Zones (Mortimore 1986a,b). 19, working quarry, Mottisfont, Hampshire, no. 1067 of Brydone (1912).

Fig. 3. Microcrinoids from the Santonian-Campanian of southern England. A-E, M, Costatocrinus brydonei Gale, 2016. A-C, radial plates; D,1Br2; E,M, fused basal rings. A, holotype, NHMUK EE 16198, original of Gale (2016, fig. 4C). B, paratype,

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ACCEPTED MANUSCRIPT original of Gale (2016 fig. 4E), NHMUK EE 16203. C, paratype, original of Gale (2016 fig. 4F), NHMUK EE 16196. Sample WC2, 0.2m beneath Charmandean Flint, Warningcamp pit no. 26, Sussex (see Appendix fig. 2). D, original of Gale (2016 fig. 4O), NHMUK EE 16047, Sample NL3, Gaster’s pit no. 2, North Lancing, Sussex (see Appendix, fig. 1). E, aboral spike of fused basals, original of Gale

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(2016 fig. 4P), NHMUK EE 16048, sample WC4, 4m above Charmandean Flint, Warningcamp pit no. 26 (see Appendix fig. 2). M, spike of fused basals, NHMUK EE 16239, G. quadrata Zone, Paulsgrove pit, Hampshire, sample PG10 (see

Appendix fig. 3). G,H, Costatocrinus mortimorei Gale, 2016. G, radial plate, H,

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brachial 1Br2. G, holotype, NHMUK EE 16192, original of Gale (2016 fig. 4B). Sample CD1, 3m above base of section, Cliff Dale Gardens pit, Cosham,

Hampshire (see Appendix, fig. 5). H,1Br2, original of Gale (2016 Fig. 4L), NHMUK

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EE 16046. Sample NL3, Gaster’s pit no. 2, North Lancing, Sussex (see Appendix, fig.1). F, Costatocrinus laevis sp. nov. Holotype radial, NHMUK EE 16223, sample SH18, O. pillula Zone, Friar’s Bay Steps, Peacehaven, Sussex (see Appendix, fig.1). I,J,N,O, Applinocrinus cretaceus forma cretaceus (Bather, 1924). I, cup in oblique view, NHMUK EE 16056, Original of Gale (2016 fig. 7A). J, basal circlet with

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extended aboral process, NHMUK EE 16076, original of Gale (2016 fig. 8L), 4m beneath Portsdown Marl, Warren Farm, Hampshire (see Appendix, fig. 4). N, brachial, external view, NHMUK EE 16057, original of Gale (2016 fig. 7R) from Charmandean Lane, pit no. 10 of Gaster (1924), 8m beneath Charmandean Flint

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(see Appendix, fig.1). O, fragmentary radial plate, original of Gale (2016 fig. 8N), 4m beneath Portsdown Marl, Warren Farm, Hampshire, sample WF7 (see

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Appendix, fig. 4). K, P,Q, X. Sagittacrinus longirostris sp. nov. Brachial plates. K, internal view, paratype, NHMUK EE 16236. P, holotype, NHMUK EE 16233, external view of brachial. Both from G. quadrata Zone sample PG10, upper part of Paulsgrove pit, Hampshire (see Appendix, fig. 3). Q, lateral view of paratype brachial, NHMUK EE 16234. X, paratype, adoral tip of brachial NHMUK EE 16235, both from G. quadrata Zone, Paulsgrove pit, sample PG14 (see Appendix, fig.3). L,S, Sagittacrinus torpedo Gale, 2016, brachials. L, internal aspect, NHMUK EE 16066, original of Gale (2016, fig. 7P). S, external view, NHMUK EE 16067, original of Gale (2016, fig. 7T). Both from G. quadrata Zone, Cliff Dale Gardens pit, Cosham, Hampshire, sample CD1, 3m above base of section (see Appendix,

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ACCEPTED MANUSCRIPT fig. 4). R,T,U, Sagittacrinus alifer sp. nov. Brachial plates. R, holotype brachial, external view, NHMUK EE 16230. T, paratype, internal view of brachial, NHMUK EE 16231. Both from G. quadrata Zone, Paulsgrove, sample PG8 (see Appendix, fig. 3). U, paratype brachial, external view, NHMUK EE 16232. Sample PG 13, Paulsgrove pit, Hampshire (see Appendix, fig. 3). V,W, Applinocrinus cretaceus

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spinifer forma nov. Radial spines in lateral (V) and adoral (W) aspects. W, holotype, NHMUK EE 16228. Paulsgrove pit, Hampshire, sample PG14 (see

Appendix, fig. 3), lower Campanian, G. quadrata Zone. V, paratype, Warren Farm, Hants, lower Campanian, G. quadrata Zone, sample WF8 (see Appendix, fig. 4).

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NHMUK EE 16229.

Scale bars equal to 200u A,B,D-F,H,J,K,M,O-R,V,W; 500u, C,G,I,L,N,S,U,X,

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Fig. 4. Microcrinoids from the Campanian of southern England.

A-E, I, Assericrinus portusadernensis gen. et sp. nov. Radial processes, in lateral (B,C,D,E), adoral/aboral (A) and distal aspects (I). D, holotype, NHMUK EE 16224, sample PG14. A, B, paratype, NHMUK EE 16225, sample PG8. C, paratype, NHMUK EE 16226, sample PG13. E, paratype, sample PG13, NHMUK EE 16227. I,

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paratype, NHMUK EE 16228, sample PG8. All from the G. quadrata Zone, Paulsgrove pit, Hampshire (see Appendix, fig. 3). F, Jakeocrinus ellisensis Gale, 2016, abraded cup in lateral aspect, G. quadrata zone, sample PTE 5, Portchester east pit (see Appendix Fig. 3), NHMUK EE 16240. J, probable basal plate of

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undescribed saccocomid, original of Gale (2016, fig. 13R), NHMUK EE 16147, sample CD3, Cliff Dale Gardens Pit, Cosham (see Appendix, fig. 4).

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K, Roveacrinus cf. communis Douglas, 1908, abraded cup in oblique aspect, original of Gale (2016, Fig. 9O), NHMUK EE 16095, O. pillula zone, sample SH11, Newhaven, Sussex, level of Meeching Marl Pair (see Appendix, fig. 1). G,L,P, Hessicrinus cooperi sp. nov. L,P, holotype cup in lateral (O) and aboral (K) aspects respectively, original of Gale (2016 fig. 11A,B), figured as Hessicrinus aff. filigree. NHMUK EE 16111. 4m above Peacehaven Marl, Offaster pillula Zone, Newhaven, Sussex, sample SH13 (see Appendix, fig. 1). G, paratype cup, NHMUK EE 16236, sample SH10a, O. pillula zone, west of Newhaven, Sussex (see Appendix, fig. 1). H,M,N, Hessicrinus filigree Gale, 2016. H, cup in lateral aspect, NHMUK EE 16112, original of Gale (2016, fig. 11C), level of Castle Hill Marls, O. pillula Zone,

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ACCEPTED MANUSCRIPT Newhaven, Sussex, sample SH7 (see Appendix, fig. 1). M, paratype cup, NHMUK EE 16103, original of Gale (2016 figs 10G, 11E), 0.2m beneath Charmandean Flint, G. quadrata Zone, Warningcamp, Sussex, pit no. 26 of Gaster (1924), sample WC2 (see Appendix, fig. 2). N, 1Br2, original of Gale (2016, fig. 10H), NHMUK EE 16105. 4m above Charmandean Flint, G. quadrata Zone,

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Warningcamp, Sussex, pit no. 26 of Gaster (1924), sample WC4 (see Appendix, fig. 2). T,U, Hessicrinus scalaensis Gale, 2016. Holotype cup, original of Gale (2016, fig. 11J-L), NHMUK EE 16114, 4m above Peacehaven Marl, Offaster pillula Zone,

Friar’s Bay Steps, Peacehaven, Sussex, sample SH13 (see Appendix, fig. 1). Q, R,V,

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Hessicrinus apertus sp. nov. V, holotype cup, aboral aspect, NHMUK EE 16236, sample 1b, Warningcamp pit no. 24, Sussex (see Appendix, fig. 2) R, paratype cup, lateral aspect, NHMUK EE 16238, sample CD-3, Cliff Dale Gardens pit, G.

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quadrata zone,Hampshire (see Appendix, fig. 4). Q, paratype cup, aboral aspect, NHMUK EE 16237, sample PG8, Paulsgrove pit, Hampshire, G. quadrata zone (see Appendix, fig. 3). O,S,W, Lucernacrinus woodi Gale, 2016. O, isolated basal circlet, aboral aspect, original of Gale (2016 fig. 9G), NHMUK EE 16088. W, holotype cup in lateral aspect. NHMUK EE 16084. Original of Gale (2016 fig. 9A,B). O and W

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from G. quadrata Zone, North Lancing, Sussex, pit no. 2 of Gaster (1924), sample NL3 (see Appendix, fig. 1). S, adoral portion of cup in lateral aspect, original of Gale (2016 fig. 9I), NHMUK EE 16089. 2m above Peacehaven Marl, O. pillula Zone, Friar’s Bay Steps, Peacehaven, Sussex. Sample SH14 (see Appendix, fig. 1).

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Scale bars equal 100u (P), 200u (A,B,D,F,G,H,I,J,L-N,Q-U) and 500u (C,E,K,O,V)

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Fig. 5. Microcrinoids from the Campanian of southern England. A-D, K,O,P, Stelllacrinus hughesae forma hughesae Gale, 2016. A-C, holotype cup, in lateral (1), aboral (2) and adoral (3) aspects, original of Gale 2016 Fig. 14A-C,H NHMUK EE 16148, level of Meeching Triple Marls, sample SH13 (see Appendix fig. 1), Friar’s Bay Steps, Peacehaven, Sussex. D, isolated basal plate, original of Gale (2016 fig. 14J), NHMUK EE 16152, O. pillula Zone, West Harnham, Wiltshire, sample WH6 (see Appendix fig. 5). K, isolated radial plate showing form of elongated radial spine, original of Gale (2016 fig. 14F), NHMUK EE 16150, G. quadrata Zone, 4m above Charmandean Flint, Warningcamp, Sussex, sample WC4 (see Appendix, fig. 2). O, isolated radial plate, spine broken, original of Gale

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ACCEPTED MANUSCRIPT (2016 fig. 15B), NHMUK EE 16156, base of section, Candy’s Pit, Cosham, Hampshire, sample CP2 (see Appendix, fig. 4). P, axillary primibrachial (1Br2), original of Gale (2016, fig.15I), NHMUK EE 16159, sample WH6, West Harnham, Salisbury, Wiltshire (see Appendix, fig. 5). F,G,L, Stellacrinus hughesae forma cristatus Gale, 2016. F,G, radial and interradial aspects of holotype radial plate,

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original of Gale (2016 fig.15H,L), NHMUK EE 16162. L, lateral aspect of radial spine, original of Gale (2016 fig. 14G), NHMUK EE 16204. Both from O. pillula Zone, level of Old Nore Marl, sample SH17 (see Appendix, fig.1), Friar’s Bay

Steps, Peacehaven, Sussex. I, Stellacrinus hughesae forma lineatus nov. Holotype,

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NHMUK EE 16241, basal plate, lateral aspect. G. quadrata Zone, sample PG13, Paulsgrove pit, Hampshire (see Appendix, fig. 3). E,H,J, Cultellacrinus gladius

Gale, 2016. J, holotype radial plate, original of Gale (2016 fig. 16D), NHMUK EE

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16169. O. pillula Zone, West Harnham, Wiltshire, sample WH1 (see Appendix, fig. 5). H, paratype radial plate, to show articular processes, original of Gale (2016 fig. 16B), NHMUK EE 16167. West Harnham, sample WH0 (see Appendix, fig. 5). E, fused basal ring, original of Gale (2016 fig. 16M), NHMUK EE 16177, Cliff Dale Garden Pit, Cosham, Hampshire, sample CD3 (see Appendix, fig.4). M,

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Cultellacrinus labyrinthus Gale, 2016, holotype, fused basal ring, original of Gale (2016 fig. 17I), NHMUK EE 16187. Sample WF8, beneath Portsdown Marl, Warren Farm, Hampshire (see Appendix, fig. 4). N,Q, Stellacrinus pannosus Gale, 2016. N, primibrachial (1Br1), original of Gale (2016 fig. 15N), NHMUK EE

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16165. Q, holotype axillary primibrachial (1Br2), original of Gale (2016 fig. 15J), NHMUK EE 16163. Level of lower Scratchell’s Marl (M3), sample WF10a (see

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Appendix, fig. 4), Warren Farm Quarry, Hampshire. Scale bars equal 100u (E,M), 200u (D,F,G,H,N-Q), 500u (A-C, I,J-L).

Fig. 6. Echinoids, microcrinoids and brachiopods from the Lower Campanian chalk of southern England. A-D, Hagenowia blackmorei Wright and Wright, 1949, tips of rostra. A, West Harnham, Wiltshire, sample WH16, G. quadrata zone (see Appendix, fig. 5). B, East Grimstead, Wiltshire, sample EG7, O. pillula zone (see Appendix, fig. 5). C, Newhaven west cliffs, original of Gale and Smith (1982, pl.4 fig. 6), NHMUK E 76846, level of Castle Hill Flint, Newhaven, Sussex. D, West Harnham, sample WH16, G. quadrata zone (see Appendix, fig. 5). E, H. elongata

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ACCEPTED MANUSCRIPT (Brünnich Nielsen). Tip of rostrum, Keswick pit, Norwich, 1m beneath major semi-tabular flint. F-L, Platelicrinus campaniensis Destombes and Breton, 2001. F, cup, original of Gale (2016 fig.12A), NHMUK EE 16115. G, cup, original of Gale (2016 fig. 12I), NHMUK EE 16123. H, isolated radial plate, original of Gale (2016 fig. 12J), NHMUK EE 16124. I, cup in adoral view, original of Gale (2016 fig. 12D),

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NHMUK EE 16118. J, cup in aboral view, original of Gale (2016 fig. 12C), NHMUK EE 16117. K, primibrachial (1Br2), original of Gale (2016 fig. 12P), NHMUK EE 16202. L, primibrachial (1Br1), original of Gale (2016 fig. 13E), NHMUK

EE16136. F,G, 0.2m beneath Charmandean Flint, sample no. WC2 (see Appendix,

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fig. 2); K, sample WC4, pit no 26, Warningcamp, Sussex (see Appendix fig. 2). J,L, 1m beneath Charmandean Flint, sample CH6, pit no. 10 (see Appendix, fig. 1), Charmandean Lane, Worthing, Sussex. I, sample CB2, 0.5m beneath Whitecliff

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Flint, Cote Bottom pit, Durrington, Worthing, Sussex (see Appendix, fig.2). H, 10m beneath Portsdown Marl, sample WF5, Warren Farm pit, Hampshire (see Appendix, fig.4). P, Platelicrinus longispinus Gale, 2016, isolated radial plate. Paulsgrove pit, Hampshire, sample PG16 (see Appendix, fig. 4) NHMUK EE 16242. 13, Terebratulina rowei Kitchin, 1902, brachial aspect. Basal part of

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section, sample Wh0, West Harnham, Salisbury Wiltshire (see Appendix, fig. 5). N,O, Leptothyrellopsis polonicus Bitner and Pisera, 1979. Warnincamp pit no 24, sample WT0b (see Appendix, fig. 2). Note septum on brachial valve. Scale bars

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equal 200u (G-I,K,L,P), 500u (A-F,J,M-0).

Fig. 7. Evolution of the holasteroid echinoid Hagenowia, modified after Gale and

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Smith (1982) and Smith and Wright (2003), from the Santonian to the Maastrichtian, with addition of new data. In the Coniacian- lower Santonian H. rostrata (O-R) the rostrum is short and low (O,R) with a large coelomic space (P), and genitals 1 and 4 are still present (green), and amb plate rows II and IV are present throughout their length. In H. anterior (J-M) the genitals 1 and 4 have been lost, and plates of rows ambs IIa and IVb are greatly reduced and separated. In the early form, H. anterior A (N) from the upper part of the coranguinum Zone, the interamb plate rows 1b and 4a are still relatively broad, but in the late Santonian crinoid zones these are narrow and elongated (L,M). The rostrum is strengthened by buttressing at the corners (K). In the early Campanian H.

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ACCEPTED MANUSCRIPT blackmorei (F-I) all plates of the shaft of the rostrum are tall and narrow, and amb rows IIa and IVb are completely lost, the rostrum is more strongly buttressed, and the internal coelomic space is reduced but still triangular (G). In H. elongata from the upper Campanian and Maastrichtian, the oculars II and IV are separated from genital 2 by interambs 2a and 3b, and the coelomic space is

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further reduced and oval. In the form from the late Campanian (A-C), the number of madreporic pores on genital 2 are variable, but in the Maastrichtian form

these are invariably reduced to two vertically arranged pores. B,C, are after

Smith and Wright (2003); D,E, after Schmid (1972); G-I, K-M, P-R after Gale and

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Smith 1982; F,J,O after Smith (1984 Fig. 5.4).

Fig. 8. Stratigraphical variation in the holasteroid echinoid Echinocorys scutata

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Leske, 1778, from the Santonian-lower Campanian interval in the chalk of southern England. The main part of the figure (F-N) shows the succession of forms which characterize the upper Santonian to lower Campanian interval, with lateral profiles of typical specimens (left) and apical disc plating (right) Genital plate 4 is coloured green. The arrows indicate the presence of transitional forms.

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Echinocorys s. cincta B and C are not depicted. A,B are apical discs of stratigraphically early Echinocorys after Olszewska-Nejbert (2007) to show the narrow, highly elongated form found in Echinocorys gravesi Desor, in Agassiz and Desor, 1846. (A) from the upper Turonian or lower Coniacian of Petites Dalles,

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northwest France (her fig. 18A) and B is E. ex gr. scutata (her fig. 22b) from the Lower Coniacian of Mangyshlak, Kazakhstan. C, disc of the neotype of

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Echinocorys scutata from the Late Coniacian or Lower Santonian of Gravesend, Kent (NHMUK E. 8721). D, specimen from the Santonian upper coranguinum Zone of Broadstairs, Kent (Peake Coll). E, disc of E. scutata from the Uintacrinus Zone of Palm Bay, Kent (Peake Coll.). Note the distinctive enlargement of Ge4 in D-G and other members of the elevata-truncata lineage. F, E. s. elevata Brydone, 1912, from the base of the Marsupites laevigatus Zone, Margate, Kent (Peake Coll). G, E.s. tectiformis Griffith and Brydone, 1911, lower O. pillula Zone, Sussex coast (Peake coll). H, E. s. depressula Brydone, 1939, lower O. pillula Zone, 0.5m above Saltdean Marl, Friar’s Bay Sussex. A.S. Gale coll. I, E. s. truncata Griffith and Brydone, 1911. Lower level of O. pillula (O.p. 2 herein), Sussex coast. (Peake

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ACCEPTED MANUSCRIPT Coll). J.E. s. cincta Brydone, 1912, O. pillula Zone, above level of Peacehaven Marl, Sussex coast (Peake coll). K, E. s. new forma? “Large form” of Gaster (1937). Sussex, Gaster Coll., NHMUK, unregistered. L, E. s. turrita Lambert, 1903, from between the Charmandean and Whitecliff Flints, Mottisfont, Hampshire (A.S. Gale coll). M, E. s. undescribed form, Downend Quarry, Hampshire (upper Culver

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Chalk) BGS Coll, unregistered. N, E. s. conica Agassiz, 1847. Portsdown Chalk, Whitecliff Bay, Isle of Wight (Peake coll.). The N.B.Peake Collection is in Norwich Castle Museum, but remains unregistered. The drawings of his specimens are

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based upon casts made by Stuart Baldwin in the 1980s.

Fig. 9. Distribution of microcrinoids and holasteroid echinoid Hagenowia from the lower Campanian O. pillula and G. quadrata Zones of the Sussex coast. ONM,

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Old Nore Marl; PHM, Peacehaven Marl; MT, Meeching Triple Marls; MP, Meeching Paired Marls; P-C, Planoconvexa Bed; CHM, Castle Hill Marls (see Table for locality details and Appendix, Fig. 1 for detailed logs and sample positions). Note the Bioevents B1-B7 which are regionally extensive across southern England;

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compare with Figs 10-12.

Fig. 10. Distribution of microcrinoids and holasteroid echinoid Hagenowia from the Lower Campanian G. quadrata Zones of pit no.2 (Gaster 1924), North Lancing, Sussex (see Table for locality details and Appendix Fig. 1 for detailed

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logs and sample positions). The acme of H. blackmorei at 0.8 m provides evidence of the correlation of this flint with Castle Hill Flint 4 on the Sussex coast.

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Compare distributions with those on the Sussex coast (Fig. 9) and Wiltshire (Figs 11,12).

Fig. 11. Distribution of microcrinoids and holasteroid echinoid Hagenowia from the Lower Campanian O. pillula and G. quadrata Zones of East Grimstead, Wiltshire (see Table for locality details and Appendix, Fig. 5 for detailed logs and sample positions). The occurrences and distribution of taxa is very similar to that on the Sussex coast (Fig. 9), and Bioevents 1-6 can be identified. These confirm the correlation proposed by Mortimore (1986a,b fig. 19).

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ACCEPTED MANUSCRIPT Fig. 12. Distribution of microcrinoids and holasteroid echinoid Hagenowia from the Lower Campanian O. pillula and G. quadrata Zones of West Harnham, Salisbury, Wiltshire. (see Table for locality details and Appendix, Fig. 5 for detailed logs and sample positions). Although microcrinoids are less abundant here, the overall patterns of abundance and distribution permits the

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identification of Bioevents 2 and 4-6 also shown in Figs 9, 10 and 11.

Fig. 13. Distribution of microcrinoids and the holasteroid echinoid Hagenowia from the Lower Campanian G. quadrata Zone of sections on Portsdown Hill,

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Hampshire (see Table for locality details and Appendix Figs 3,4 for detailed logs and sample positions). Note particularly the simultaneous appearance of five

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taxa at 44.9 m in the Paulsgrove pit which mark the base of CaR6.

Fig. 14. Distribution of microcrinoids from the Lower Campanian O. pillula and G. quadrata Zones of Charmandean Lane (pit no. 10) and Warningcamp (pit no. 26) of Gaster (1924), West Sussex (see Table for locality details and Appendix Fig. 2 for detailed logs and sample positions). Although diversity is low, and some

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occurrences are sporadic, the significantly increased abundances of P. campaniensis and H. filigree at the level of the Charmandean Flint can be identified throughout southern England, and mark the base of CaR9. This

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coincides with a marked increase in coarser bioclastic debris, mostly bryozoans.

Fig. 15. Distribution of microcrinoids and the brachiopod Leptothyrellopsis

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polonicus from the Campanian G. quadrata and B. mucronata Zones of Warren Farm Quarry, Hampshire (Table for locality details and Appendix Fig. 4 for detailed logs and sample positions). Note the extinctions of three species (most notably Costatocrinus brydonei) at 6.5 metres (2-3 metres above the Whitecliff Flint) marking the base of zone CaR10, the disappearance of Stellacrinus hughesae at 21.5 metres, and the appearance of Stellacrinus pannosus, marking the base of zone CaR11 at the level of Scratchell’s Marl 1 at 27.8 metres. The abundance of Leptothyrellopsis polonicus in samples WF5-10 is matched by a similar occurrence in Warningcamp pit no 24 (Fig. 16) in West Sussex.

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ACCEPTED MANUSCRIPT Fig. 16. Distribution of microcrinoids and the brachiopod Leptothyrellopsis polonicus from the Lower Campanian O. pillula and G. quadrata Zones of Sussex, pits at Cote Bottom (no. 17) and Warningcamp (no.24). (see Table for locality details and Appendix Fig. 2 for detailed logs and sample positions). The extinction of Costatocrinus brydonei 2-3 metres above the Whitecliff Flint at Cote

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Bottom exactly parallels the distribution in Warren Farm, Hampshire (Fig. 15), as does the abundance of Leptothyrellopsis polonicus in the uppermost Culver Chalk at Warningcamp pit no. 24.

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Fig. 17. Compiled distribution of microcrinoids and holasteroid echinoid

Hagenowia from the lower Campanian O. pillula and G. quadrata zones of all studied sections in southern England, with proposed microcrinoid zonation (CaR

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Zones) and Echinocorys biostratigraphy. FB, Friar’s Bay Marls; BR, Black Rock Marl; RT, Roedean Triple Marls; ON, Old Nore Marl; P, Peacehaven Marl; MT, Meeching Triple Marls; MP, Meeching Paired Marls; P-C, Planoconvexa Bed; CH, Castle Hill Marls; P-B, Planoconvexa Bed; CH, Castle Hill Marls; PB, Pepperbox Marls; F10, Castle Hill Flint 10; PH, Portchester Hardgrounds; CHA,

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Charmandean Flint; WF, Whitecliff Flint; PM, Portsdown Marl; SM, Scratchell’s Bay Marls; FM, Farlington Marls.

Fig. 18. Correlation between upper Santonian to lower Campanian sections in

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southern England, based on lithology (black), microcrinoids, Offaster and Hagenowia. Note the lateral continuity of some lithological markers from east to

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west, but variation in the abundance and development of marl seams, almost absent in Dorset, but abundant in Scratchell’s Bay, Isle of Wight. Note also the significantly expanded succession in Scratchell’s Bay.

Fig. 19. Correlation between lower Campanian sections in southern England, based on lithology (black), microcrinoids, Offaster and Hagenowia. Note the strikingly similar lithological and faunal successions at East Grimstead and the Sussex coast (Seaford Head), and the laterally consistent occurrences of O. pillula and H. blackmorei. RTM, Roedean Triple Marls; ONM, Old Nore Marl; PM,

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ACCEPTED MANUSCRIPT Peacehaven Marl; MTM, Meeching Triple Marls; PC, Planoconvexa Bed; CHM, Castle Hill Marls; CHF, Castle Hill Flints.

Fig. 20. Correlation between lower Campanian G. quadrata Zone sections in southern England, based on lithology (black), microcrinoids and Hagenowia,

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upper part of succession. Note the revised correlation for the upper part of the succession, in which the Cote Bottom, Cosham and Whitecliff Flints are shown to be synonymous, and the inferred exposure gap in Sussex.

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Fig. 21. Bed-scale correlation between Culver Chalk successions across

Hampshire, the Isle of Wight and Sussex. Note the larger, lensoid or more tabular flints on the Isle of Wight (Scratchell’s Bay), which replace Thalsssinoides burrow

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flints on the mainland, and the disappearance of marker marls in Sussex.

Fig. 22. Drawings of Roveacrinida and reconstructions. A, Reconstruction of Sagittacrinus alifer sp. nov, in adoral aspect. Note asymmetrical wing-like flanges on brachials. The cup is unknown. B, Reconstruction of Assericrinus

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portusadernensis gen et sp. nov., reconstruction, in lateral aspect. The robust radial spines are the only known parts from the English chalk (fig. 4A-E,I). The cup reconstruction is based on entire specimens of A. portusadernensis from the Campanian Ozan Formation of Waxahachie dam spillway, Texas (Gale et al.

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2008). C-F, Lateral view of aboral cup in species of Hessicrinus, to show distinguishing features. C, H. scalaensis Gale, 2016; F, H. apertus sp. nov.; E, H.

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filigree Gale, 2016; D, H. cooperi sp. nov. H. scalaensis and H. apertus are closely related; note the large central basal foramen (bfo), the prominent basal spine, the flat stellate base to the cup (see also Fig. 5G,H,L,M,P-R,T-V). However, H. scalaensis possesses paired radial/basal fenestrae, as does H. filigree. Note that in H. cooperi sp. nov. radial/basal fenestrae are tiny and inconspicuous. Bfo, basal foramen; ibfe interbasal fenestra; r:bfe, radial/basal fenestra; rfo radial foramen. Scale bar equal to 1mm for A,B.

Appendix. Locality logs and descriptions.

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ACCEPTED MANUSCRIPT 1. Sussex coast (Fig. 9, Appendix Fig. 1)

The lower part of the succession (upper Newhaven and basal Culver chalks, Marsupites testudinarius to basal G. quadrata Zones) was studied on the coastal cliffs between Brighton and Seaford Head (Appendix, fig. 1), and used as a

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standard for the lower part of the microcrinoid succession (Fig. 9). Samples were obtained from Black Rock, Brighton, Friar’s Bay Steps, the cliffs west of

Newhaven, and Seaford Head. Lithostratigraphical nomenclature follows

Mortimore (1986a,b). Samples were taken approximately every metre; the

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detailed microcrinoid occurrences are plotted in Figure 9.

A taxonomic account of some of the material was provided by Gale (2016), with supplementary taxa described below, and detailed quantitative

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records are provided here (see section Systematic palaeontology). They demonstrate the presence of a succession of distinct roveacrinid faunas which are used to define successive zones, CaR1 to CaR5.

CaR1 extends from the last occurrence of Uintacrinus anglicus, 1metre beneath Friar’s Bay Marl 3, up to the Old Nore Marl, where the base of CaR2 is

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marked by a flood abundance of Stellacrinus hughesae forma cristatus (Fig. 9, Bioevent 1; Appendix Fig. 1 sample SH17). Numerous changes take place in the interval between the Peacehaven Marl and the Meeching Triple Marls. Firstly, Applinocrinus cretaceus forma cretaceus and Stellacrinus hughesae forma

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hughesae become abundant and Costatocrinus brydonei appears (Fig. 9, Bioevent 2; Appendix Fig. 1, sample SH14a, 1m above the Peacehaven Marl). Secondly,

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Lucernacrinus woodi and Hessicrinus scalaensis appear (Fig. 9, Bioevent 3; Appendix Fig. 1, sample SH14, 2m above the Peacehaven Marl). Finally, the FO of Cultellacrinus gladius (Fig. 9, Bioevent 4; Appendix Fig. 1, sample 13, base of Meeching Triple Marls) marks the base of CaR3. Cultellacrinus gladius becomes abundant (>50 plates per kg) 0.7m above the Meeching Triple Marls (Fig. 9; Appendix Fig. 1, sample SH12). The evolutionary transition between Hessicrinus cooperi sp. nov. and H. filigree takes place beneath the Castle Hill Marls; the latter species appears in sample SH8a (Fig. 9 Bioevent 5; Appendix Fig. 1), marking the base of CaR4. The first occurrence of Platelicrinus longispinus between Castle Hill Flints 4 and 5

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ACCEPTED MANUSCRIPT (Fig. 9, Bioevent 6; Appendix Fig. 1, sample SH4) marks the base of CaR5. Finally, the FO of Costatocrinus mortimorei (Fig. 9, Bioevent 7; Appendix Fig. 1, sample SH2a) is observed. These new discoveries augment the traditional macrobiostratigraphy for the Sussex coast chalk, based upon Echinocorys scutata variants and levels of

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abundance of Offaster pillula (Brydone 1914, 1939; Mortimore 1986a,b; Wood and Mortimore 1988), and the high-resolution microbiostratigraphy provided by benthonic forams and nannofossils (Hampton et al. 2007).

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2. North Lancing, Sussex (Fig. 10, Appendix Fig. 1)

In view of the importance of the work of Gaster (1924), who established

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the Saccocoma cretacea Subzone in the Worthing district, cuttings and old chalk quarries in the region of North Lancing were also studied. Pits nos. 3 and 4 of Gaster are now gone, but the lower part (5m) of the succession which he described is well exposed in a cutting behind an old factory in Halewick Lane, North Lancing, and the upper part of the succession is still seen in Gaster’s pit no.

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2 which was re-excavated and sampled. The results of this study permit precise correlation of individual beds with the coastal succession. Hagenowia blackmorei appears above the paired marls exposed in Halewick Lane (sample H4), interpreted as the Castle Hill Marls, and reaches an acme of abundance in the

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Thalassinoides burrow flint close to the base of pit No. 2 (sample NLB), which therefore correlates with Castle Hill Flint 4 of the coast (Mortimore 1986a,b).

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Samples from the highest few metres of pit no. 2 (NL1-4) yield a rich fauna of CaR5, including C. mortimorei and P. longispinus, demonstrating the presence of Bioevents 6 and 7. This part of the section here is evidently slightly more expanded than that at Seaford Head, where the mesofauna is fragmented and abraded at the level of Castle Hill Flints 9-11. Mortimore (1986a) referred to a flint in the upper part of pit no. 2 at North Lancing as the Lancing Flint, and identified a thin marl seam beneath this as the Lancing Marl. However, as shown by Wood and Mortimore (1988), this flint correlates with Castle Hill Flint no.9 on the Sussex coast, and the Lancing Flint is therefore not used in this study. I am unable to find a marl at this level.

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ACCEPTED MANUSCRIPT Gaster (1924) took his base of the Saccocoma cretacea Subzone a short distance beneath what Mortimore (1986a,b) subsequently named the Lancing Flint in pit no. 2. However, the range of A. cretaceus is now known to extend down to the Santonian (Gale 2016), so Gaster was in error here. In conclusion, Gaster’s (1924) North Lancing sections and inferred

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correlation with the Sussex coast was remarkably accurate, but he mistakenly believed that the section included the Planoconvexa Bed (i.e. Telscombe Marls

2,3) at the base, as did Wood and Mortimore (1988). In fact, the base of the North Lancing sections of Gaster are situated some distance (about 4m) above the

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Planoconvexa Bed, so his suggestion that part of the coastal succession described by Brydone is absent at North Lancing (Gaster 1924) is incorrect. White (1924) commented that he had never seen the bed with large O.pillula in the Worthing

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district.

3. Worthing – Arundel, Sussex (Figs 14, 16, Appendix Figs 1,2)

The succession in Sussex is continued up above North Lancing in the 17-

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m -section adjacent to Charmandean Lane, Worthing (Gaster 1924, pit no. 10; Mortimore 1986a,b; Appendix Fig. 1). Wood and Mortimore (1988) showed an overlap of several metres between the base of this section and the top of pit no. 2, North Lancing. However, pit no. 2 is still within CaR5 (see above), whereas the

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base of pit no. 10 is in CaR9 (Appendix Figs 1,2, samples CH1-4), so there is a significant biostratigraphical gap between the two sections, and the distinctive

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CaR7 and CaR8 faunas are not recorded from Sussex. It is therefore highly probable that there is a significant exposure gap in Sussex, which, by comparison with the Portsdown succession, represents about 20 m of chalk. It is noteworthy that Gaster (1924) implied the presence of this gap, remarking (p. 102) that pit no. 7, which exposes the Charmandean Flint, lies about 100 feet (30m) above the base of his Saccocoma cretacea subzone, i.e. close to the top of pit no 2. The upper part of Charmandean Lane pit provides an excellent exposure of the distinctive semitabular Charmandean Flint, which is immediately underlain (0.5m) by the base of CaR9, characterised by common Hessicrinus filigree and Platelicrinus campaniensis. Chalks immediately beneath and above

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ACCEPTED MANUSCRIPT this flint contain lenses of abundant bryozoan debris, and are technically wackestones. The Charmandean Flint is again exposed 1m above the base of the 13-m-section in pit no. 26 of Gaster (1924) at Warningcamp, near Arundel (Fig. 14, Appendix Fig. 2), a locality which is an important source of well-preserved roveacrinid material (Gale 2016), including a number of type specimens.

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The succession is continued in Cote Bottom Pit, Durrington, Worthing (Gaster’s pit no. 17; Fig. 16, Appendix Fig. 2), the type locality of Saccocoma

cretacea (Bather 1924). The lower part of the section yields abundant crinoids of CaR9 (samples CB1-3), and the double flint close to the base is taken to correlate

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with a similar flint at the top of Warningcamp, pit no. 26, supported by detailed comparison with the similar Candy’s Pit succession in Cosham, Hampshire. The base of crinoid zone CaR10 is found at 3 m above the semi-tabular Cote Bottom

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Flint (Mortimore 1986a,b; Fig. 16 herein, sample CB4), and is marked by the LO of Costatocrinus brydonei and Lucernacrinus woodi.

The Sussex section continues in the more southerly of the Cote Bottom pits, and in the upper Warningcamp pit, no. 24, which provide a composite section of the highest chalk which is exposed inland in Sussex (Fig. 16, Appendix

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Fig. 2). The highest 2m here (samples WT 2, 1b, 1 and 0) yield abundant small, smooth terebratellid brachiopods (Leptothyrellopsis polonicus), characteristic of the uppermost Culver and basal Portsdown chalks. However, the highest samples, 2.8m above the Warningcamp Flint, still yield a CaR10 fauna and

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indicate that the highest chalks probably fall a short distance beneath the level of

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the Scratchell’s Marls.

4. Portsdown Hill, Hampshire (Figs 13, 15, Appendix Figs 3,4)

Numerous disused chalk quarries are present on Portsdown Hill, to the

north of Portsmouth, Cosham, Portchester and Fareham, and these provide a nearly complete composite section from the upper part of the Newhaven Chalk, almost the entire Culver Chalk and the basal Portsdown Chalk. Macrofossil zonation of these sections was provided by Griffith and Brydone (1911) Brydone (1912) and White (1913) and skeletal logs of the lower part of Paulsgrove, The George, Cosham (Candy’s Pit herein), Downend and Warren Farm by Mortimore

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ACCEPTED MANUSCRIPT (1986a,b) and Mortimore et al. (2011). Gale et al. (2015) published a section of Warren Farm quarry, and proposed a detailed correlation with the Isle of Wight successions. The lowest part of the succession is exposed in the eastern part of the extensive Paulsgrove pit (Appendix Fig. 3), which includes a total of nearly 55 m

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of chalk. The lowest 5 metres lack marl seams and contain numerous small Thalassinoides burrow flints. Sample PG6a yielded a CaR1 fauna, and the base of CaR2, marked by flood occurrence of Stellacrinus hughesae forma cristatus was

identified in a prominent marl (sample PG6) which confirms the identification of

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this bed as the Old Nore Marl (see also Mortimore 1986b). The overlying beds contain a succession of marl seams, and samples PG0a, PG5, PG4a, PG3c and

PG3b yield a typical fauna of CaR3, including abundant Cultellacrinus gladius. The

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presence of abundant small O. pillula between 24 and 26m, and large O. pillula at 27m confirm identification of the Meeching Marl Pair at 24m and the Planoconvexa Bed at 27 m. The paired Castle Hill Marls (31m) yield the lowest Hessicrinus filigree (sample PG3) and overly the base of CaR4. The single marl seam at 33.1m yields abundant Hagenowia blackmorei

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(sample PG2a) associated with a CaR4 fauna and is the correlative of the Pepperbox Marls. The appearance of Platelicrinus longispinus in sample PG2 (34.6m) marks the base of CaR6 (Bioevent 6), and another typical species of the zone, Costatocrinus mortimorei, is present in sample PG16 (Bioevent 7, 37.3m).

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Above this are 20 metres of soft, bryozoan-rich chalks containing widely spaced nodular flint layers, many representing silicified Thalassinoides systems,

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inaccessible until partial infilling of the quarry in the 1990s. A diverse microcrinoid fauna of CaR6 (Bioevents 8-10) is found in the uppermost 9m of the Paulsgrove pit, appearing in sample PG 14 and continuing to the top of the exposed chalk. This includes Saggitacrinus longirostris sp. nov., S. alifer sp. nov., Assericrinus portusadernensis gen. et sp. nov., and Applinocrinus cretaceus forma spinifer nov. Although White (1913) believed that there was an overlap between the top of Paulsgrove and the base of Candy’s Pit, to the east in Cosham, there is a significant faunal gap, represented by a 6-m-section north of Portchester (Portchester east, Fig. 13, Appendix Fig. 3) which was described as containing two nodular hardgrounds and a underlying marl (White 1913, fig. 3). The

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ACCEPTED MANUSCRIPT hardgrounds are clearly exposed, but the “marl” appears to be a soft fracture surface lacking clay content. The section was re-exposed in the present study and has yielded a CaR6 fauna, including S. longirostris sp. nov., and a level with abundant Hagenowia blackmorei (sample PTE2). There is clearly an unexposed gap between the top of Paulsgrove and the base of Portchester east, probably of a

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few metres magnitude. Portchester west continues the succession upwards, above an unexposed interval of about 10-12m. The lower part yields faunas of CaR8 (samples PTW1,2), the upper part CaR9 (samples PTW4-6). The

Charmandean Flint is present at 6.6m, overlain by a hardground on which the

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Whitecliff Wispy Marl is cut out.

The succession immediately above Portchester east continues up in Cliff Dale Gardens and Candy’s pits on the London Road at Cosham, for which a

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composite section is provided (Fig. 13, Appendix Fig. 4). The base of Cliff Dale Gardens pit (sample CD-3) yields a CaR7 fauna, with abundant Cultellacrinus gladius, mixed with late elements of the CaR6 fauna (S. longirostris sp. nov.) and probably lies a short distance above the top level exposed in Portchester east. The top of CaR7 (sample CD4, 5.3m) is marked by the extinction of C. gladius and

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P. longispinus. The overlying 11 m of chalk (samples CD5, CD6, CP2, CP2a, CP3, CP3a) yield a fauna of the CaR8 zone, typified by rather low abundance and diversity. The base of CaR9 is marked by the abundance of Platelicrinus campaniensis and Hessicrinus filigree, associated with bryozoan-rich chalks, in

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samples CP5, CP6, CPT, CPT1, CPT2 and CP1. The Charmandean Flint and overlying Whitecliff Wispy Marl are present just above the base of the zone. The

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Warren Farm Marls and overlying Whitecliff Flint are present in the higher part of the section.

Warren Farm Quarry, now a recycling centre owned by Veolia Ltd., still

provides an excellent exposure of the uppermost Culver and Lower Portsdown chalks (Fig. 15, Appendix Fig. 4). Gale et al. (2015) provided a detailed log, and proposed correlation with the Isle of Wight exposures. The base of the section exposes the Warren Farm Marls 1-3 and the overlying Whitecliff Flint. The base of the CaR10 zone is marked by the extinction of Costatocrinus brydonei in sample WF3(6.2m). The higher part of the Culver and basal Portsdown chalks yield abundant small terebratellid brachiopods, Leptothyrellopsis polonicus in

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ACCEPTED MANUSCRIPT samples WF8, WF9 and WF10. The base of CaR11, in the lower of the two Scratchell’s Marls (20.8m) is represented by the first occurrence and flood abundance of Stellacrinus pannosus (Gale 2016). The section at Farlington Redoubt (Appendix, Fig. 4) is also developed in CaR11.

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5. Mottisfont, Hampshire (Appendix Figs 5,6)

The large roadside chalkpit at Mottisfont, west of Winchester, is still

operated actively by Somborne Chalks, and provides nearly 40m of exposed

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chalk succession. A skeletal log was given by Mortimore (1986a,b), which

identified the Charmandean Flint in the upper part of the section, and the lower part of the section, incorporating other nearby pits was interpreted as

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representing the Old Nore to Pepperbox Marl succession by Mortimore (1986b, fig. 19). The section was relogged and 27 samples collected and processed. The lower part of the section shows considerable lateral variation from the north to the south of the section, and two sections are given with a proposed correlation (Appendix Fig. 6). The lowest beds in the northerly exposure are

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nodular, and contain three marl seams. A sample from the second marl (M0, 1.4m) yielded abundant Hagenowia blackmorei. The overlying succession comprises soft chalks with flints, overlain by a number of marl seams and nodular units, and terminated by a 0.9-m-nodular hardground. In the southerly

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succession the, upper nodular units have coalesced to form a 1.5-m-thick nodular hardground, within which the marls have been cut out. Samples M3 and

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M4 (southerly section) yield a distinctive microcrinoid fauna, including Costatocrinus mortimorei and Platelicrinus longispinus, together with abundant Cultellacrinus gladius. The highest record of Cultellacrinus gladius is in sample 4a, within the nodular complex. The lower part of the section, by comparison with Portsdown, is interpreted as representing the upper part of CaR7 (base to about 5m), and the lowest group of nodular chalks and marls are the Portchester Hardgrounds and Solent Marls. The flinty chalks and overlying nodular unit represent CaR7, including samples M3 and M4. The top of CaR7 is taken at the top of the nodular unit in the southerly section.

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ACCEPTED MANUSCRIPT Above, 16 metres of chalk with well-developed, evenly spaced nodular flints yield a fauna of the CaR8 zone (samples M5-18). The Charmandean Flint, at 21.8m is overlain by the Whitecliff Wispy Marl, and samples from this level to the top of the pit contain a CaR9 fauna. Dissolution seams, now full of soil, appear to represent the Warren Farm Marls, and large semitabular flint masses

6. East Grimstead, Wiltshire (Fig. 11, Appendix Fig. 5)

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immediately above the highest chalk are from the Whitecliff Flint.

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The disused chalk quarry adjacent to the railway line south of East

Grimstead sits on the southern limb of the Dean Hill Anticline, and exposes 48 metres of upper Newhaven and lower Culver chalks. A skeletal section was

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provided by Mortimore (1986b) and Mortimore et al. (2001). The section was cleaned and relogged for the present study and 25 samples were collected and processed (Fig. 16). As shown by Mortimore (1986b), the development of marl seams is strikingly similar to that on the Sussex coast, with minor differences. The prominent Old Nore Marl (sample EG0, 5.5m) yielded abundant Stellacrinus

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hughesae forma cristatus, indicative of the base of CaR2, and Bioevent 1 in Sussex (Fig. 9). Bioevent 3, in the upper part of CaR2, is marked at East Grimstead, as in Sussex, by the occurrence of Hessicrinus scalaensis and Lucernacrinus woodi at 17.2m (sample EG4), immediately beneath the Meeching Triple Marls. Bioevent

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3, which marks the base of CaR3 (Bioevent 4, FO of Cultellacrinus gladius) is present in sample EG18 at 19.9m. Hessicrinus filigree, indicative of CaR4, is

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present between the Castle Hill Marls (sample EG13). The large burrow flint overlying the Pepperbox Marls yielded abundant Hagenowia blackmorei (EG3, 39.9m). An unusual occurrence is the present of common Sagittacrinus cf. longirostris sp. nov. in the upper part of the succession (samples EG2, EG17, EG21), not found in Sussex.

7. West Harnham, Wiltshire (Fig. 12, Appendix Fig. 5)

The disused chalkpit at West Harnham to the southwest of Salisbury, an important source of fossils for H.P. Blackmore, was mentioned briefly by Reid

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ACCEPTED MANUSCRIPT (1903) and Mottram (1956) stated that it contained the level of abundant Offaster pillula. The first section was published by Mortimore (1986b), and more details were provided by Mortimore et al. (2001), including a review of the important type fossils collected at Harnham. Hopson et al. (2006) followed the stratigraphy of Mortimore et al. (2001).

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The section was logged (Appendix Fig. 5), and 22 samples collected and processed (Fig. 11). The base of the section, below and above a strong marl

seam at 1.5m (samples WH3, WH0, WH4, WH5) yielded abundant microcrinoids of zone CaR2, including Costatocrinus brydonei. Cultellacrinus gladius appears at

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8.5 metres (sample WH8), marking the base of CaR3. The Planoconvexa Bed, and associated Telscombe Marls 1-3, was identified from the presence of large Offaster pillula, and also yielded a CaR3 fauna. A single marl at 21.2 metres

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resting upon a nodular chalk, falls within the lower part of zone CaR4 and is therefore interpreted as a Castle Hill Marl. The Hagenowia blackmorei acme was identified in association with a prominent double flint (25.5m, sample WH16), which is therefore a correlative of Castle Hill Flint 4. Microcrinoids diagnostic of CaR5 (Platelicrinus longispinus) are present in sample WH20 (29.2m) and extend

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to the top of the section.

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8. Middle Bottom, Lulworth, Dorset (Appendix Fig. 7)

The cliff section beneath Middle Bottom, west of Durdle Door, is only

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accessible by boat in calm weather. The section was measured in the summer of 1996 with the aid of D.S. Wray. The locality was visited and zoned by Rowe (1905), and a quite detailed log was provided by Brydone (1914). The section extends continuously from the Upper Greensand at White Nothe to the lower part of the Culver Chalk. The Newhaven Chalk contains abundant nodular flints beds, mostly consisting of small flints which replace Thalassinoides systems. Although the chalk appears to be undisturbed tectonically, the few marl seams developed in the Newhaven Chalk are represented by minor thrust surfaces. The ranges of the crinoids Uintacrinus socialis, Marsupites laevigatus and M. testudinarius were

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ACCEPTED MANUSCRIPT determined, but U. anglicus could not be found, probably on account of the wavesmoothed surfaces. The Old Nore Marl is well developed (51.2-51.4 m), and a sample yielded Stellacrinus hughesae cristatus, indicative of the base of CaR2. The O.p.2 and O.p.3 levels of abundant Offaster pillula are well developed, and the Planoconvexa Bed is prominent, as described by Brydone (1914). A single Castle

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Hill Marl is present (81.4 m), as elsewhere in Dorset (see below); the Pepperbox Marls are not present, but the distinctive burrow flint, Castle Hill Flint 4, is well developed (85.3m), as inland in Dorset. A marl seam near the top of the

9. Tarrant Rowston, Dorset (Appendix Fig.7)

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accessible section may represent one of the Solent Marls of the Isle of Wight.

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The disused chalkpit situated at the base of the Cliff, south of Tarrant Rowston, exposes 10m of uppermost Newhaven and basal Culver chalks (Tarrant Chalk in BGS terminology). The section was briefly described by Bristow et al. (1995, p.132), who identified a prominent marl in the lower part of the section as the Pepperbox Marl. The base of the section exposes a lower marl,

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which is a single representation of the Castle Hill Marls. The lower sampleTR6) yielded a fauna of CaR4, and overlying samples TR4 and TR3 both yielded Hessicrinus filigree, indicative of CaR4.

The prominent nodular flint overlying the Pepperbox Marl at 5.5m yields

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abundant Hagenowia blackmorei (sample TR3), which is also present but less common in TR2 and TR4. The flint is therefore correlative with Castle Hill Flint 4

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in Sussex. Samples TR1 and TR2 yield microcrinoids typical of CaR5, including Platelicrinus longispinus.

10 Scratchell’s Bay Isle of Wight (Appendix Figs 8,9)

This section, accessible only by boat, was logged by ASG during numerous visits over the period 1994-2013. Major macrofaunal levels and marker beds have been identified, but the chalks are too hard to process with Glauber Salt. Marl samples were treated with acid and yield some limited microcrinoid data.

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ACCEPTED MANUSCRIPT Published studies of the Scratchell’s Bay succession were based almost entirely upon the log in Swiecicki’s PhD thesis (1980) which provided evidence for the assignment of benthic foraminiferal zones for the UK benthic scheme (Bailey et al. 1983; Hart et al. 1989). However, the log of Swiecicki (1980) is not sufficiently detailed to identify marker beds with any certainty. Hopson et al.

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(2014 fig. 3a) also provided a log of the Scratchell’s Bay succession as well, but this also contains insufficient detail to identify most individual beds. However, the distinctive marl/flint signature at the level which they identify as the

Scratchell’s Marls (M3,4) is significantly higher than the level identified by Gale

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et al. (2015), and actually represents M5-6. Hopson et al. (2014) additionally identified the boundaries of the BGS benthic foraminiferal zones.

A log of the upper Newhaven, Culver and lower Portsdown Chalk in

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Scratchell’s Bay is provided here (Appendix Fig. 8), with an accompanying annotated photograph (Appendix Fig. 9). The Newhaven Chalk contains very numerous thin marl seams, far more than in Sussex, and the fauna provides support for the proposed correlations of these. The occurrence of Uintacrinus anglicus (3.2-3.7m) permits identification of the Friar’s Bay Marls, and the

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second horizon of Offaster pillula (O.p.2) starts a short distance above a prominent marl, which yields abundant Stellacrinus hughesae forma cristatus, indicative of basal CaR2. The highest horizon of abundant O. pillula (O.p.3) is surmounted a typically developed Planoconvexa Bed at 47m (paired Telscombe

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Marls 2,3; centrally placed nodular flint), and the Castle Hill and Pepperbox Marls are well developed. The Solent Marls (1-3) are similarly well developed,

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and the Charmandean Flint is overlain by the distinctive Whitecliff Wispy Marl. In the interval above the Whitecliff Marl, numerous lensoid semi-tabular flints are developed, as on Whitecliff. A prominent semi-tabular flint overlying the Whitecliff Wispy Marl is present throughout the Isle of Wight and is here called the Vectis Flint.

The base of the Portsdown Chalk is marked by the Portsdown Marl (107.8m). The overlying Scratchell’s Marls, at 111.2-111.7m, yield Stellacrinus pannosus, indicating the base of CaR11.

11 Whitecliff, Isle of Wight (Appendix Fig. 10)

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ACCEPTED MANUSCRIPT General descriptions of the succession were given by Rowe (1908) and White (1921). A detailed log of the Campanian chalk succession in Whitecliff Bay, eastern Isle of Wight, was provided by Mortimore et al. (2001), and details of parts of the section were given by Gale et al. (2013, 2015). The chalks are too

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hard for Glauber Salt processing, but a small number of samples were treated with acetic acid (see above) and provided some limited microcrinoid data. The uppermost Santonian and lowermost Campanian are highly

condensed, with the development of nine mineralised hardgrounds and some

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phosphatic chalk which floor a complex channel system (Gale et al. 2013).

Occasional large Offaster pillula typical of the Planoconvexa Bed are present on the summit of H9 (13.2m). The overlying chalks are expanded and represent a

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channel fill (Gale et al. 2013). The Castle Hill and Pepperbox marls can be clearly identified (Mortimore et al. 2001), and are overlain at 43.4 and 46.2m by welldeveloped marl seams which they called the “Lancing Marls 1,2”, which are apparently not developed on the mainland. The Solent Marls and underlying chalks (WCL11, WCL10, WCLS2-3) yield elements of the CaR6 fauna, and

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Hagenowia blackmorei at 55.2m. The CaR8 fauna is present in a suite of samples (samples WCL4-6), and CaR9 was identified in the Whitecliff Wispy Marl at 76.8m. Chalks immediately beneath the Portsdown Marl yielded abundant Leptothyrellopsis polonicus (sample WCL14), and the Scratchell’s Marls contain

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numerous Stellacrinus pannosus, marking the base of CaR11 (sample WCL15).

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Appendix Fig. 1. Stratigraphy of Campanian sections on the Sussex coast (Black Rock, Seaford Head; all Sussex coastal samples bear the prefix SH), and the Worthing-North Lancing district. See Table for details of localities. Microcrinoid zonal boundaries are marked based upon data provided in Figs 9 and 10.

Appendix Fig. 2. Stratigraphy and correlation of lower Campanian sections in inland Sussex, pit numbers following Gaster (1924), with locality map and location of numbered samples. See Table for details of localities. Microcrinoid zonal boundaries are marked, based upon data provided in Figs 14 and 16.

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ACCEPTED MANUSCRIPT Appendix Fig. 3. Stratigraphy of lower Campanian sections on Portsdown Hill, Hampshire, with position of samples marked. Lower part of stratigraphy. See Table for details of localities. Microcrinoid zonal boundaries are marked, based upon data provided in Fig.13.

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Appendix Fig. 4. Stratigraphy of Campanian sections on Portsdown Hill, Hampshire, with position of samples. Upper part of stratigraphy. See Table for

details of localities. Microcrinoid zonal boundaries are marked, based upon data

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provided in Fig. 13.

Appendix Fig. 5. Stratigraphy of Campanian sections in Wiltshire (East Grimstead, West Harnham) and Hampshire (Mottisfont), to show position of

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samples. See Table for details of localities. Microcrinoid zonal boundaries are marked, based upon data provided in Figs 11 and 12.

Appendix Fig. 6. Correlation of the lower part of section, Mottisfont pit, Hampshire, showing lithological changes from the northern to the southern

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succession (approximately 150m apart) available. Marls present between 6 and 10m in the north are cut out by nodular hardgrounds in the southern section. Microcrinoid zonal boundaries are marked.

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Appendix Fig. 7. Stratigraphy of Lower Campanian sections in Dorset. Middle Bottom coastal cliffs, west of West Lulworth. Chalk pit at Tarrant Rowston, SE of

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Blandford Forum. See Table for details of localities. Microcrinoid zonal boundaries are marked.

Appendix Fig. 8. Stratigraphy of the Upper Santonian and Lower Campanian exposed in Scratchell’s Bay, Isle of Wight. See Table for details of locality.

Appendix Fig. 9. Photograph of Lower Campanian exposure in Scratchell’s Bay, Isle of Wight. PC, Planoconvexa Bed; CHM, Castle Hill Marls; SM, Solent Marls 1,2,3; CF, Charmandean Flint; WWF, Whitecliff Wispy Marl; VF, Vectis Flint;

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ACCEPTED MANUSCRIPT Warren Farm Marls 1,2,3; WF, Whitecliff Flint; PM, Portsdown Marl; SM, Scratchell’s Marls. Photo taken by Prof. Ian Jarvis. Compare with Appendix Fig. 8.

Appendix Fig. 10. Section exposed in Whitecliff Bay, Isle of Wight. Data in lower part of section taken from Gale et al. 2013, upper part from Gale et al. 2015. See

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Table for details of localities. Some microcrinoid zonal boundaries are marked.

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5

North Lancing Sussex Halewick Lane, Lancing, Sussex Lambley’s Lane Sompting, Worthing, Sussex

6

Cote Bottom, Worthing, Sussex

Lat-long

Sea cliff

Mortimore 1986a,b Mortimore 1986a,b Wood and Mortimore 1988 Wood and Mortimore 1988 Gaster 1924

TV488982

50o45’51.40”N 0o06’31.94W 50o46’52.50N 0o02’49.66W 50o47’07.59”N 0o01’10.79”W

Sea cliff Steps down sea cliff Path down sea cliff Disused quarry, pit no. 2 cutting Disused quarry, pit no. 7 Disused quarry, pit no. 10 Disused quarry, no. 17

Disused quarry, pit no. 19 Warningcamp, Disused Sussex, quarry, pit no. 26 Warningcamp,Sussex Disused quarry, pit no. 24 Cliff Dale Gardens Disused Cosham Hampshire quarry Candy’s Pit, Disused Cosham,Hampshire quarries Paulsgrove Pit, Disused Hampshire quarry

8

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Clapham, Sussex

7

9 9 10

Gaster 1924 Mortimore 1986a,b Gaster 1924 Mortimore 1986a,b Gaster 1924 Mortimore 1986a,b Gaster 1924

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Charmandean Lane Worthing Sussex

Grid ref.

TQ447982 TQ424004

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Black Rock, Brighton Sussex

Key refs

TQ344032

50o48’44.14”N 0o05’33.93”W

TQ184063

50o50’36.95”N 0o19’12.54”W

TQ176066

50o50’49.11”N 0o19’51.01”W 50o50’37.32”N 0o21’28.66”W

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4

type

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No. locality in Fig. 2 1 Seaford Head Sussex 2 Newhaven Sussex 3 Friar’s Bay Steps, Peacehaven Sussex

Gaster 1924 Mortimore 1986a,b Gaster 1924 Mortimore 1986a,b Brydone 1912 Brydone 1912 Brydone 1912

TQ156058

TQ148057

50o50’23.06”N 0o22’13.98”W

TQ116065

50o50’48.44”N 0o24’59.43”W

TQ094066

50o50’56.19”N 0o26’50.19W

TQ037074

50o51’29.73”N 0o31’42.41”W

TQ045077

50o51’24.64”N 0o31’42.41”W

50o51’08.23”N 1o30’30.70”W SU666063 50o51’11.08”N 1o30’17.16W SU664063- 50o51’21.94”N SU639066 1o06’10.54”W SU664063

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13 14 15 16 17 18

Sea cliff Disused quarry Disused quarry Disused quarry Active quarry

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19

Sea cliff

White 1913 White 1913 Gale et al. 2015 Gale et al. 2013,2015 Hopson et al. 2014 Brydone 1914 Bristow et al. 1995 Mortimore et al. 2001 Mortimore 1986a,b Brydone 1914; Mortimore 1986a,b

SU618065 SU618066 SU604069 SZ619855 SZ026847

50o51’16.70”N 1o07’24.00”W 50o51’18.34” 1o07’28.03”W 50o51’28.28”N 1o08’30.21”W 50o40’00.96”N 1o49’07.10”W 50o39’42.72”N 1o34’56.72”W 50o37’28.65”N 2o17’59.14”W 50o51’34.55”N 2o04’58.96”W 51o03’28.36”N 1o49’07.10W 51o02’33.36”N 1o40’38.04”W 51o03’41.92”N 1o31’41.92”W

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12

Disused quarry Disused quarry Disused quarry Sea cliff

SY790805

TR942066 SU127287

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11

Portchester east pit, Hampshire Portchester west pit, Hampshire Warren Farm Hampshire Whitecliff Bay, Isle of Wight Scratchell’s Bay, Isle of Wight Middle Bottom, Dorset Tarrant Rowston, Dorset West Harnham, Wiltshire East Grimstead Wiltshire Mottisfont Hampshire

SU227271

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