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The English and London Clay: two remarkable British bony fish Lagersta¨tten

MATT FRIEDMAN1*, HERMIONE T. BECKETT1, ROGER A. CLOSE1 & ZERINA JOHANSON2 1Department of Earth Sciences, University of Oxford, South Park Road, Oxford OX1 3AN, UK 2Department of Earth Sciences, The Natural History Museum, Cromwell Road, London SW7 5BD, UK *Corresponding author (e-mail: [email protected])

Abstract: The Late () Chalk and (Ypre- sian) London Clay Formation are two British marine deposits that yield globally significant assem- blages of actinopterygian (ray-finned) fishes. Materials from these units, especially the Chalk, featured prominently in the work of Arthur Smith Woodward. Here we summarize the history of study of actinopterygian from the Chalk and London Clay, review their geological and palaeoenvironmental context and provide updated faunal lists. The Chalk and London Clay are remarkable for preserving fossil fishes in three dimensions rather than as the flattened individ- uals familiar from many other famous Lagersta¨tten, as well as capturing detailed ‘snapshots’ of marine fish faunas that bracket the major taxonomic shift that took place near the Cretaceous– Palaeogene boundary.

Gold Open Access: This article is published under the terms of the CC-BY 3.0 license.

The (Cenomanian–Maastrichtian) symposium volume. Responsible for the world’s English Chalk and Eocene () London Clay finest collection of fossil fishes from these locali- are two fossiliferous British marine deposits that ties at what is now the Natural History Museum, are significant in the history of palaeontology gener- London, Smith Woodward produced numerous ac- ally, and palaeoichthyology specifically. Fossils counts of Chalk and London Clay specimens (e.g. from both units featured prominently in early contri- Woodward 1887a, b, 1888a–d, 1889a–e, 1891a, b, butions that played a major part in the develop- 1893a, b, 1894a, b, 1895a–c, 1898, 1899a–c, ment of modern geology and palaeontology (Luidii 1900a, b, 1901, 1902, 1903, 1905, 1906a, b, 1907a, (¼Lhuyd) 1699; Woodward 1729; Jacobs 1777a, b; b, 1908, 1909, 1911, 1912a, b, 1913, 1917, 1923, Mantell 1822; Agassiz 1833–44; Fig. 1; Owen & 1926, 1936; Woodward & Sherborn 1890, 1891), Bell 1849). Fossil fishes from the English Chalk naming numerous in the process (see Smith served as touchstones in early debates on the succes- 2015). A handful of major contributions added to sion of assemblages through time (Agassiz 1833–44, understanding of fishes from these deposits in the 1859) and the rates of evolutionary change (Huxley decades following Smith Woodward’s death (body 1862, 1870; Woodward 1912a: 252; summarized fossils: Patterson 1964; Casier 1966; Goody 1969; in Patterson 1981) and, more recently, provided Forey 1973, 1977; otoliths: Stinton in Casier 1966; key ancient comparators in the discovery and inter- Stinton 1975–84). However, more recent contribu- pretation of the living coelacanth Latimeria (Smith tions have been more limited in their scope, focus- 1939; Woodward 1940). The popularity of fossil- ing on specific taxa or specimens (e.g. Forey 2004). hunting in the London Clay during the mid-1800s The actinopterygians (ray-finned fishes) of the led to James Bowerbank’s foundation of the Lon- Chalk and London Clay are notable not only for don Clay Club (1836–47), which would eventually their diversity, but also for their remarkable pre- give rise to the modern Palaeontographical Society servation. Unlike most fish faunas of comparable (Elliott 1970). age, in which specimens are preserved as greatly The English Chalk and London Clay provided flattened compression fossils, these British deposits ample material for study and description by Arthur yield three-dimensionally preserved specimens of Smith Woodward, the principal subject of this either whole individuals (Chalk) or heads (London

From:Johanson, Z., Barrett, P. M., Richter,M.&Smith, M. (eds) 2016. Arthur Smith Woodward: His Life and Influence on Modern Palaeontology. Geological Society, London, Special Publications, 430, 165–200. First published online November 23, 2015, http://doi.org/10.1144/SP430.18 # 2016 The Author(s). Published by The Geological Society of London. Publishing disclaimer: www.geolsoc.org.uk/pub_ethics Downloaded from http://sp.lyellcollection.org/ by guest on September 24, 2021

166 M. FRIEDMAN ET AL.

Fig. 1. Historical representations of English Chalk and London Clay fossil fish specimens: (a) the Chalk Halec, from Horsfield & Mantell (1824), The History and Antiquities of Lewes and Its Vicinity, Volume 1; (b)‘Beryx Lewesiensis from the Chalk, Lewes’ from Mantell (1851), Petrifactions and their Teachings; (c, d) the Chalk taxa Homonotichthys and , from Dixon (1850), The Geology and Fossils of the Tertiary and Cretaceous Formations of Sussex;(e) the Chalk taxon Ctenothrissa, from Smith Woodward (1901), Catalogue of Fossil in the British Museum (Natural History). Part IV;(f) ‘Head of a Pike’ from the London Clay (probably Aulopopsis), from Douglas (1785), A Dissertation on the Antiquity of the Earth;(g) the London Clay taxon Teratichthys, from Ko¨nig (1825), Icones Fossilium Sectiles;(h) the head of the London Clay taxon , from Smith Woodward (1901), Catalogue of Fossil Fishes in the British Museum (Natural History). Part IV.

Clay). Following in Smith Woodward’s footsteps, can only be applied to some specimens from the subsequent workers have realized the significance London Clay and, in these cases, must be completed of these fossils, applying acid preparation tech- with great care. Non-destructive computed tomog- niques to extract considerable detail on the skulls raphy (CT; Sutton et al. 2014) offers an approach of fishes from the Chalk (Patterson 1964; Goody to studying the remarkable fish fossils of the Chalk 1969; Forey 1973, 2004; Forey & Patterson 2006). and London Clay without any of the drawbacks Although powerful, this approach is not without associated with traditional preparation techniques its limitations. First-generation acid preparation (Fig. 2; Beckett & Friedman 2015). We are currently yielded detailed anatomical information, but the re-studying the ray-finned fishes of the Chalk and specimens themselves were rendered very delicate London Clay using CT scanning, with the principal and difficult to examine. The loss of positional goals of better documenting their anatomy, relation- information associated with the disarticulation of ships and, through the use of biomechanical models specimens during preparation was exacerbated by developed for living fishes (e.g. Hulsey & Wain- the loss of individual bones (e.g. the jaws of Diplo- wright 2002; Collar & Wainwright 2006), ecology. mystus in Forey 2004: fig. 2). More generally, this This contribution provides context for this ongoing preparation approach is limited by the nature of research, reviewing current understanding of fishes matrix encasing fossils; although acid preparation from these two exceptional deposits. Examination is very effective with fish fossils from the Chalk, it of actinopterygians from the Chalk and London Downloaded from http://sp.lyellcollection.org/ by guest on September 24, 2021

ENGLISH CHALK AND LONDON CLAY FISHES 167

Fig. 2. Detail of fossil fish skulls from the and London Clay Formation revealed by computed tomography (CT): (a–c) Argillichthys toombsi, NHMUK PV P 42519, London Clay Formation, Sheppey, ; (d–f)‘Sardinioides’ illustrans, NHMUK PV P 3977, Chalk Group, Burham, Kent. Models show: bones (white) visible external to matrix (brown) before segmentation of CT data (panels a and d); fossil specimen with matrix completely removed (panels b and e); and model of fossil with jaws and palate removed to reveal the branchial (green) and ventral hyoid (pink) arches.

Clay has been intermittent following the major SMUC, Sedgwick Museum, University of Cam- twentieth-century monographs of Smith Woodward bridge, Cambridge, UK. (1902, 1903, 1907a, 1908, 1909, 1911, 1912a) and Casier (1966) (both of which produced major increases in the number of named fish species Geological context: Chalk Group from the British record; see Lloyd & Friedman 2013: fig. 2), meaning that important updates are High sea-levels and low continental relief limited dispersed throughout a large body of specialist liter- clastic deposition to regions deep within modern ature. Our goal is to provide an accessible summary continental interiors during the Late Cretaceous of current understanding of the Chalk and London and early Palaeogene (Mortimore et al. 2001; Mor- Clay actinopterygian faunas by reviewing a history timore 2011). Deposition in the expansive shelf of their study, basic geological context and evolu- settings of this interval was dominated by slow tionary significance. We close with a prospectus accumulation of the calcitic skeletons of plank- for future research on these deposits, which are tonic coccolithophorid algae. The resulting pelagic rightly recognized as Lagersta¨tten that represent carbonate – chalk – represents a distinctive and exceptional windows to fish diversity in the Late widespread feature of the Late Cretaceous– and Eocene (Casier 1966; Patterson Palaeogene sedimentary record, with units found 1993a, b; Lloyd & Friedman 2013). across northern and central , central Asia, and the western interior of (Mortimore 2011). Institutional abbreviations The British record is no exception to this global pattern. While much of , Ireland, Wales BMB, Booth Museum, Brighton, UK; BIRUG, and far southwestern represented exposed Lapworth Museum of Geology, University of topographic highs during the Late Cretaceous, the Birmingham, Birmingham, UK; IRSNB, Royal remainder of the British Isles was flooded by a Belgian Institute of Natural Sciences, Brussels, shallow shelf sea dominated by chalk deposition. ; MNHN, Muse´um national d’Histoire Often referred to collectively as the ‘the English naturelle, Paris, ; NHMUK, Natural History Chalk’ or more simply ‘the Chalk’, these pelagic Museum, London, UK; OUMNH, Oxford Uni- carbonates are exposed principally in SE England versity Museum of Natural History, Oxford, UK; with additional outcrops in the NE and SW and Downloaded from http://sp.lyellcollection.org/ by guest on September 24, 2021

168 M. FRIEDMAN ET AL. more minor ones in (Mortimore The Southern Province comprises most of the et al. 2001; Gale & Kennedy 2002). Extending Chalk Group exposed in southeastern England, from the Cenomanian to the early Maastrichtian, extends to continental Europe in northern France, the Chalk spans much of the Late Cretaceous. It and has Tethyan affinities. The two realms are overlies Early Cretaceous () deposits, which connected in East Anglia by sediments of the Tran- vary locally from the Upper to the sitional Province, in which lithologies and fos- Clay. The thickest successions of the Chalk sils characteristic of the two realms interdigitate. preserve roughly 500 m of strata, with thicker pack- Although the same subgroups are recognized in ages present in the . Although there is var- the Northern and Southern provinces, separate litho- iability between different geographical provinces stratigraphic divisions have been developed for (see below), some major compositional trends are these regions at the formation level. Apart from bio- apparent across the Chalk. The most prominent of stratigraphy (see below), marker beds like glauco- these is a decrease in clay content in younger sec- nitic marl seams, flint bands and layers of nodular tions, which is reflected in the principal division of chalk represent important tools for correlation the formal Chalk Group into the marly sediments within the Chalk (Mortimore 2011). of the Grey Chalk Subgroup and the nearly pure Global correlation of the Chalk succession suf- carbonates of the White Chalk Subgroup. Marine fers from a lack of ammonites in post-Cenomanian transgression throughout the Late Cretaceous, sections (Jenkyns et al. 1994; Gale & Kennedy which further removed sources of clastic input from 2002; but see Gale et al. 2005), probably owing to areas of deposition, seems to be responsible for early diagenetic dissolution of aragnonite (Jefferies this pattern (Mortimore et al. 2001). 1962; Hudson 1967). In the absence of ammonites, and with planktonic foraminiferans rare, assem- Stratigraphy blage zones based on calcite-mineralizing benthic invertebrates dominate historical biostratigraphic The English Chalk has been divided under a variety schemes for the Chalk (note that correlations with of lithostratigraphic and biostratigraphic schemes standard ammonite zones in Fig. 3 are only approx- over the past two centuries (Fig. 3). Because older imate). Tracing their ancestry to studies in the Paris accounts of fossils from the Chalk apply these his- Basin (He´bert 1863), these classifications were torical schemes, they are briefly summarized here. refined in a series of studies (e.g. Barrois 1876) Early work resulted in initial subdivisions based that culminated with Rowe’s (1900, 1901, 1903, upon clay content and the presence or absence of 1904, 1908) influential set of divisions (reviewed flints. Townsend (1813) gave an initial description in Gale & Cleevely 1989). This classification repre- of the chalk in , followed by Mantell sented the state of the art at the time of publication of (1818, 1822) for Sussex and the , and The Fossil Fishes of the English Chalk, and was the Phillips (1821) for Kent. All of these authorities scheme applied by Smith Woodward (1902, 1903, adopted a threefold division of the Chalk: a ‘Grey 1907a, 1908, 1909, 1911, 1912a) to describe the Chalk’ or ‘Chalk Marl’ as the lowest unit, overlain stratigraphic distribution of Chalk fishes. Rowe’s by the ‘Lower Chalk’, and the ‘Upper Chalk’. zonation remained little modified until the late Following the synthesis of Jukes-Browne & Hill twentieth century (Rawson et al. 1978), and it was (1903, 1904) the of the English not until this time that lithostratigraphic and bio- Chalk was subdivided into ‘Lower’, ‘Middle’ and stratigraphic divisions of the Chalk began to be ‘Upper’ Chalk and remained that way for almost separated (Wood & Smith 1978). Although persis- a century (Mortimore et al. 2001). Detailed but tent, zonal schemes based on macrofossils are defi- accessible historical overviews of Chalk stratigra- cient by modern biostratigraphic standards: they are phy are provided by Mortimore et al. (2001), Gale coarse in resolution, inexplicitly defined and do & Kennedy (2002) and Mortimore (2011). not allow for application over broad geographical Modern classifications recognize the Chalk of scales (Gale & Kennedy 2002). Despite these limi- historical usage as the Chalk Group, which in turn tations, macrofossil zones give the narrowest readily is divided into the Grey Chalk and White Chalk available constraints on the age of many old fish subgroups (Fig. 3). This subdivision reflects some specimens from the Chalk, and remain widely components of older classifications, with the Grey used for historical reasons (Fig. 3). Chalk and White Chalk subgroups equivalent to the Lower Chalk and Middle plus Upper Chalk, Key exposures respectively. The Chalk Group shows regional var- iation in sediment and fossil content leading to the Fossil fishes are found throughout the Chalk, but recognition of two distinct provinces. The North- most of the articulated bony fish material described ern Province extends from to NE England, by Smith Woodward and others derives from the and shows biological links with the Boreal Realm. ‘Lower’ and ‘Middle’ Chalk of historical usage, Downloaded from http://sp.lyellcollection.org/ by guest on September 24, 2021

ENGLISH CHALK AND LONDON CLAY FISHES 169

Fig. 3. Chronological, lithostratigraphic and biostratigraphic divisions of the English Chalk and underlying formations. (a) Upper Cretaceous zones from Gale & Kennedy (2002), Lower Cretaceous zones from Gale & Owen (2010) and Dineley & Metcalf (1999). International Biostratigraphic Zones and chronological divisions are taken from Gradstein et al. (2012). Local divisions, initially proposed by Rowe (1901) and used in Smith Woodward (1902, 1903, 1907a, 1908, 1909, 1911, 1912a) to define fossil age ranges are taken from Mortimore et al. (2001) and Gale & Kennedy (2002) (correlations between local stratigraphy and zones, and global ammonite biozonation, are only approximate). Approximate stratigraphic extents of select localities given on the right based on Dineley & Metcalf (1999), Mortimore et al. (2001), and zones reported by Smith Woodward (1902, 1911, 1912a; Cherry Hinton and South Ferriby). (b) Geographical extent of Chalk outcrop, after Mortimore et al. (2001). Downloaded from http://sp.lyellcollection.org/ by guest on September 24, 2021

170 M. FRIEDMAN ET AL. and thus the Grey Chalk Subgroup and base of the Chalk fauna (Woodward 1902, 1903, 1907a, 1908, White Chalk Subgroup of the contemporary litho- 1909, 1911, 1912a). Significant as rare representa- stratigraphic classification. Remains are present in tives of relatively complete fish remains from the younger parts of the Chalk succession, but are Northen Province of the Chalk, these fossils were not as abundant as they are in older sections, and destroyed when the Hull Municipal Museum was tend to consist of disarticulated remains or teeth hit by an incendiary bomb in 1943. Images of (Table 1). This uneven stratigraphic distribution is these now lost fossils are given in Figure 4. matched by geographical heterogeneity, with most Of these localities, two are particularly signifi- fossil fishes collected from rocks belonging to the cant from a historical and palaeontological perspec- Southern Province in SE England (Table 1). tive: (GCR 2906) and Southerham Although numerous individual sites yield re- (GCR 217). specimens from Blue Bell Hill mains of bony fishes, these are largely fragmentary derive from two chalk pits. The Lower Culand Pit in nature. Articulated and associated materials over- exposes a section of the Grey Chalk Group and whelmingly derive from a handful of key localities base of the White Chalk Group roughly 40 m (Fig. 3), and were collected at a time when chalk thick. The Upper Culand Pit exposes sections of pits were worked by hand rather than excavated the White Chalk Group roughly 60 m. Fossils with machinery (Woodward 1902, 1903, 1907a, from these pits are generally labelled as from ‘Bur- 1908, 1909, 1911, 1912a; Patterson 1964; Longbot- ham’ or ‘Blue Bell Hill’, but information on collec- tom & Patterson 2002). These materials formed tion horizons is highly variable. It is clear, however, the basis of key historical treatments of Chalk that multiple levels within these exposures yield fishes: Mantell (1822), Agassiz (1833–45), Dixon articulated fish fossils. Based on the account given (1850), Newton (in Dixon 1878) and Woodward in Smith Woodward (1902, 1903, 1907a, 1908, (1902, 1903, 1907a, 1908, 1909, 1911, 1912a). 1909, 1911, 1912a), Dineley & Metcalf (1999) con- Owing to the rarity of articulated remains, additions clude that the two Blue Bell Hill localities collec- to the Chalk fauna based on field collection of com- tively yield over 20 genera of bony fishes. Several plete bony fish specimens are unlikely. Discoveries chalk pits in the area of Southerham, near Lewes, based on reinterpretation of old museum specimens Sussex have yielded important fish material, includ- are more probable (e.g. Forey 2004; Friedman et al. ing specimens first reported by Mantell (1822) 2010), while bulk sampling and acid preparation and Dixon (1850; Woodward 1902, 1903, 1907a, of Chalk continue to yield abundant microvertebrate 1908, 1909, 1911, 1912a). Dineley & Metcalf remains, especially chondrichthyan teeth (Dineley (1999) suggest that most of these specimens derive & Metcalf 1999). from the Machine Bottom Pit, which exposes Dineley & Metcalf (1999) provide a detailed over 40 m of the Grey Chalk Subgroup (Zig Zag overview of important fossil fish-bearing sites in Chalk formation of Cenomanian age) and White the Chalk. Localities yielding articulated bony fish Chalk Subgroup (Holywell Nodular Chalk Forma- specimens that figure most prominently in Smith tion of Cenomanian and earliest age). Fos- Woodward’s (1902, 1903, 1907a, 1908, 1909, 1911, sils with collection localities given as ‘Lewes’ or 1912a) research include Amberley Station Quarry ‘Southerham’ probably derive from this site, but (north of Arundel, Sussex; Cenomanian–Turonian), as with Blue Bell Hill, controls on stratigraphic hori- Clayton Limeworks and Railway Tunnel (south zon are often poor. Together the deposits around of Clayton, Sussex; Cenomanian), Glynde Station Lewes yield nearly 30 bony fish genera (Woodward Quarry (SE of Lewes, Sussex; Cenomanian– 1902, 1903, 1907a, 1908, 1909, 1911, 1912a; Dine- Turonian); Houghton (SE of Amberley, Sussex; ley & Metcalf 1999). Turonian), Southerham (SE of Lewes, Sussex; Cenomanian–Turonian), Folkestone (Folkestone, Palaeoenvironment Kent; Cenomanian), Dover (Dover, Kent; Turo- nian–), Halling (Halling, Kent; Cenoma- The Late Cretaceous coincides with the breakup nian–Turonian), Blue Bell Hill (Burham, Kent; of the supercontinents of Laurasia and Gondwana- Cenomanian–Turonian) and Cherry Hinton (Cherry land (Mortimore et al. 2001). Sea-levels reached Hinton, Cambridgeshire; Cenomanian). With the their Phanerozoic peak of roughly 100 m above exception of Cherry Hinton, which is located within those of the present day (Miller et al. 2005), associ- the Transitional Province, these localities sample ated with elevated levels of atmospheric carbon the Southern Province of the Chalk. Also notewor- dioxide and clear evidence for elevated global tem- thy is South Ferriby (Lincolnshire; Cenomanian). peratures (Bice et al. 2005). Vertebrate assemblages Smith Woodward described material from this site and plants from Arctic deposits comparable in age principally through short publications in The Natu- to the Chalk indicate warm, ice-free poles (Tarduno ralist (Woodward 1907b, 1912b , 1923), but men- et al. 2001; Friedman et al. 2003; Vandermark tioned some specimens in his overview of the et al. 2009), while a variety of proxies for sea Table 1. Actinopterygian fauna of the Chalk Group Downloaded from Taxon Type/relevant specimen Specimens Province Cenom. Turon. Con. San. Ca.

abcdefghijkl

Chondrostei? Chondrostei incertae sedis Pholidurus disjectus Woodward, 1889a NHMUK PV OR 33221-4 Single Southern x

Holostei http://sp.lyellcollection.org/

‘Lepidotes’ pustulatus Woodward, 1895a NHMUK PV P 6201a-c Multiple Southern X FISHES CLAY LONDON AND CHALK ENGLISH Halecomorphi *Lophiostomus dixoni Egerton, 1852 NHMUK PV OR 23023 Multiple Southern X X X *Neorhombolepis excelsus Woodward, 1888a NHMUK PV OR 43077 Single Southern X Neorhombolepis? punctatus Woodward, 1895a NHMUK PV P 4705 Multiple Southern, Northern X *Tomognathus mordax Dixon, 1850 BMB 007225 Multiple Southern x X Teleostei Anomoeodus angustus Agassiz, 1844 NHMUK PV P 1616 Multiple Southern XXXXXX Anomoeodus pauciseriale Kriwet, 2002 BMB 011147 Single Southern Anomoeodus willetti Woodward, 1893b BMB 007270 Single Southern X ‘’ parallelus (Dixon, 1850) BMB 007276a-b Single Southern x x x byguestonSeptember24,2021 ‘Coelodus’ fimbriatus Woodward, 1893b NHMUK PV OR 43090 Single Southern X ‘’ cretaceus Agassiz, 1844 Lost Multiple Southern XXX ‘Pycnodus’ scrobiculatus Reuss, 1844 ‡Unknown Multiple Southern X ‘Phacodus’ punctatus Dixon, 1850 NHMUK PV OR 25829 Multiple Southern x x x Acrotemnus faba Agassiz, 1844 NHMUK PV OR 4584 Single Southern x x * ferox Leidy, 1857 NHMUK PV OR 4088-9, 4096, 4135 Multiple Southern XXXX Protosphyraena compressirostris Woodward, 1895d NHMUK PV P 5631 Single Southern x x x Protosphyraena minor Agassiz, 1844 NHMUK PV OR 4078 Single Southern X Rhinconichthys taylori Friedman et al. (2010) NHMUK PV OR 33219 Single Southern cinctus Agassiz, 1844 NHMUK PV OR 4266 Multiple Southern XXXX basalis Dixon (1850) NHMUK PV OR 49014 Single Southern XXXXXX Pachyrhizodus dibleyi Woodward, 1901 NHMUK PV P 9115 Multiple Southern X *Pachyrhizodus subulidens (Owen, 1842) SMUC B.9097 Multiple Southern, Transitional X X *Pachyrhizodus magnus (Woodward, 1901) NHMUK PV OR 37981 Multiple Southern x *Pachyrhizodus sp. Municipal Museum, Hull (destroyed) Single Northern X 171 (Continued) 172 Downloaded from

Table 1. Continued

Taxon Type/relevant specimen Specimens Province Cenom. Turon. Con. San. Ca.

abcdefghijkl

*Elopopsis crassus (Dixon, 1850) BMB 007207 Multiple Southern, Transitional, xXx Northern http://sp.lyellcollection.org/ *Thrissopater megalops Woodward, 1901 NHMUK PV OR 49826 Single Southern x *Apsopelix anglicus (Dixon, 1850) NHMUK PV P 611 Multiple Southern XXXX sp. (Ichthyodectes minor [Egerton, 1850] NHMUK PV OR 28894 Multiple Southern XXXX

nomen dubium) FRIEDMAN M. Prosaurodon sp. (Ichthyodectes elegans NHMUK PV OR 41687 Multiple Southern X Newton, 1877) *Ichthyodectes tenuidens Woodward, 1901 NHMUK PV OR 49054 Single Southern X sp. (including X. mantelli [Newton, 1877], NHMUK PV OR 4066-7 (X. mantelli), Multiple Southern, Transitional X X X x x Xiphactinus daviesi [Newton, 1877]) NHMUK PV OR 28388 (X. daviesi) *‘’ intermedius (Newton, 1878) NHMUK PV OR 47250 Single Southern x TAL. ET

Saurocephalidae indet. NHMUK PV P 42017 Single Southern x byguestonSeptember24,2021 *Dinelops ornatus Woodward, 1907a NHMUK PV OR 39432 Multiple Southern X Protelops anglicus Woodward, 1901 NHMUK PV OR 49906 Multiple Southern X ‘Albuliformes’ *Osmeroides lewesiensis (Mantell, 1822) NHMUK PV OR 4294 Multiple Southern XXX *Osmeroides levis Woodward, 1901 NHMUK PV P 5681 Multiple Southern X *Osmeroides latifrons Woodward, 1907a NHMUK PV P 10465 Multiple Southern XX *Enchelurus anglicus Woodward, 1901 NHMUK PV P 4249 Multiple Southern X Anguilliformes *Urenchelys anglicus Woodward, 1900a BMB 007246 Multiple Southern x x Anogmius ornatus Woodward, 1923 Municipal Museum, Hull (destroyed) Single Northern X Plethodus expansus Dixon (1850) BMB 008254 Multiple Southern XXXX Pentanogmius pentagon Woodward, 1901 NHMUK PV OR 41716a Multiple Southern X *Dixonanogmius oblongus (Dixon, 1850) BMB 007300 Multiple Southern X ‘Protosphyraena’ stebbingi Woodward, 1909 NHMUK PV P 11216 Multiple Southern, Northern X Ellimichthyiformes *Diplomystus sp. NHMUK PV P 5695, BIRUG 1580 Multiple Southern x Downloaded from Aulopiformes *‘Sardinioides’ illustrans Woodward, 1901 NHMUK PV P 3977 Single Southern x x x x *Acrognathus boops Agassiz, 1844 NHMUK PV OR 4304 Multiple Southern x x x *Apateodus striatus (Dixon, 1850) NHMUK PV OR 49821 Multiple Southern, Transitional X X X X Apateodus lanceolatus Woodward, 1901 NHMUK PV OR 39080 Multiple Southern X Prionolepis angustus Egerton, 1850 NHMUK PV P 9253 Multiple Southern, Transitional X X *Cimolichthys lewesiensis Leidy, 1857 NHMUK PV OR 4039 Multiple Southern X X X X X *Halec eurypterygius (Dixon, 1850) BMB 007183 Multiple Southern X X X X X *Enchodus lewesiensis (Mantell, 1822) NHMUK PV OR 4004, 4049, 4157, Multiple Southern X X X X X http://sp.lyellcollection.org/ 4160, 4180-1, 4183-4, 4198 *Eurypholis pulchellus (Woodward, 1901) NHMUK PV P 1703 Multiple Southern X X X X FISHES CLAY LONDON AND CHALK ENGLISH ‘Dercetis’ latiscutatus Woodward, 1903 BMB 007264 Multiple Southern x x x *‘Dercetis’ maximus Woodward, 1903 NHMUK PV OR 31075-82 Single Southern X *Leptotrachelus elongatus (Agassiz, 1844) NHMUK PV OR 4132-3 Multiple Southern, Transitional x x x X Ctenothrissiformes *Ctenothrissa radians (Agassiz, 1835) NHMUK PV OR 4029 Multiple Southern XXXX *Ctenothrissa microcephala (Agassiz, 1835) NHMUK PV OR 4034 Multiple Southern XXXX *Aulolepis typus (Agassiz, 1844) NHMUK PV OR 4033 Multiple Southern X x x x Lampridiformes *‘Aipichthys’ nuchalis (Dixon, 1850) NHMUK PV OR 25770 Single Southern X byguestonSeptember24,2021 Polymixiiformes *Berycopsis elegans Dixon, 1850 BMB 007204 Multiple Southern XXXX *Homonotichthys dorsalis (Dixon, 1850) BMB 007210 Multiple Southern X x x *Homonotichthys rotundus (Woodward, 1902) NHMUK PV P 315 Multiple Southern X *Homonotichthys pulchellus (Dixon, 1850) NHMUK PV OR 25886 Multiple Southern X *Caproberyx superbus (Dixon, 1850) NHMUK PV OR 25959 Multiple Southern XX *Hoplopteryx lewesiensis (Mantell, 1822) NHMUK PV OR 4014-5 Multiple Southern XXXXX *Hoplopteryx macracanthus Patterson, 1964 NHMUK PV P 33230 Multiple Southern XXX *Hoplopteryx gephyrognathus Patterson, 1964 NHMUK PV OR 41104 Single Southern X *Hoploptyerx simus Woodward, 1902 NHMUK PV OR 49073 Multiple Southern X X *Trachichthyoides ornatus Woodward, 1902 NHMUK PV OR 39076 Single Southern x

Stratigraphic distributions largely from Smith Woodward (1902, 1903, 1907a, 1908, 1909, 1911, 1912a). ‘Relevant’ specimens are non-types identified at generic or higher level. Occurrences marked by ‘X’ are definitive, while those indicated by ‘x’ are less certain. Taxa marked with ‘*’ are known from relatively intact skulls. ‘‡’ preceding holotype specimen information indicates that type not from the Chalk Group. Dates of publication in Agassiz (1833–44) follow Brown (in Woodward & Sherborn 1890). Traditional zones are indicated by characters as follows: a, Schloenbachia varians;b,Holoaster subglo- bosus;c, plenus;d,Rhynchonella cuvieri;e,Terebratulina gracilis;f,Holaster planus;g, cortestudinarium;h,Micraster coranguinum;i,Uintacrinus socialis;j, testu- dinarius;k,Offaster pilula;l,Gonioteuthis quadrata. Correspondences between these zones and -level divisions of the international timescale shown here are inexact (see Fig. 3). Cenom., Cenomanian; Turon., Turonian; Con., ; San., Santonian; Ca., . 173 Downloaded from http://sp.lyellcollection.org/ by guest on September 24, 2021

174 M. FRIEDMAN ET AL.

Fig. 4. Specimens of bony fishes from the Northern Province of the Chalk Group at South Ferriby, Lincolnshire, destroyed during aerial bombing of Hull in 1943. All specimens are reported as coming from the zone of Holaster subglobusus.(a) Skull of Pachyrhizodus sp. in left lateral view (from Anonymous 1912). Scale not indicated. (b) Rostrum of ‘Protosphyraena’ stebbingi in multiple views, with anterior to right (from Woodward 1912b). Scale bar represents 50 mm. (c) Anogmius ornatus in right-lateral view (from Woodward 1923). Scale bar represents 50 mm. surface temperatures indicate warm conditions in Forster et al. 2007). The early Late Cretaceous is both polar and equatorial regions (Pearson et al. marked by a major ocean anoxic event at the Ceno- 2001; Schouten et al. 2003; Jenkyns et al. 2004; manian–Turonian Boundary: the Bonarelli Event Downloaded from http://sp.lyellcollection.org/ by guest on September 24, 2021

ENGLISH CHALK AND LONDON CLAY FISHES 175 or Ocean Anoxic Event (OAE) 2 (Schlanger & Jen- all – or even most – benthic species known from kyns, 1976; Jenkyns 1980; Mortimore et al. 2001). the Chalk. Instead, the weight of palaeontological This well-defined OAE at the Cenomanian–Turo- evidence suggests that the seafloor during depo- nian boundary is marked by a +3‰ excursion of sition of the Chalk was sufficiently firm to allow d13C of bulk organic matter and appears to have vagile benthic organisms to easily move across been a global, rather than regional, event (Turgeon it, and for infaunal organisms to construct stable & Creaser 2008). burrows within it (Gale & Smith 1982; Gale & lay at a palaeolatitude of Kennedy 2002). 408 N during the deposition of the Chalk Group, and was covered by a shallow shelf sea. This geo- Taphonomy and graphical placement coincides with a transition between the Boreal and Tethyan faunal realms, Although major taphonomic features of the Chalk reflected in the Transitional Zone within the Chalk Group are well known (e.g. a clear preservational of East Anglia. The water depth during the deposi- bias against aragonite-biomineralizing taxa; sum- tion of the Chalk has been greatly debated in the marized in Gale & Kennedy 2002 and elsewhere), past, but current estimates favour values between remarkably little is known about the conditions 100 and 300 m but no deeper than 500 m (Bell that led to the exceptional preservation of articu- et al. 1999). d18O values for the Chalk Group sug- lated fossil fishes in this deposit. Formal studies of gest early Cenomanian temperatures of 20–228C Chalk fish taphonomy are restricted to a brief note rising to a peak of 26–288C at the Cenomanian– on a specimen of the aulopiform Dercetis preserved Turonian boundary, and gradually declining through within a flint, an uncommon mode of preservation the remainder of the Cretaceous to between 16 and among Chalk fishes limited to a handful of examples 188C by the early Maastrichtian (Jenkyns et al. (Davis & Hitchin 1996). 1994, 2004). OAE 2 punctuates the deposition of In the absence of formal study, a few observa- the Chalk Group, with the Plenus Marl (Southern tions on Chalk fishes seem relevant in the context Province) and Black Band (Northern Province) of available studies of fish taphonomy (Scha¨fer members corresponding to this event (Mortimore 1972; Elder & Smith 1984, 1988; Ferber & Wells et al. 2001). The biotic effects of OAE 2 are widely 1995). First, some articulated specimens show debated. Although many attribute faunal turn- strong curvature of the spine (e.g. Osmeroides, over at this interval to genuine of shelf Woodward 1907a: pl. 23, figs 1, 9), gaping mouths fauna owing to anoxia (Jarvis et al. 1988; Harries (e.g. Berycopsis, Woodward 1902: pl. 1, figs 1 & 2; & Little 1999), others suggest these patterns are an Hoplopteryx, Woodward 1902: pl. 3, fig. 1; Apsope- artefact generated by a combination of migration– lix, Woodward 1902: pl. 10, fig. 1; Tomognathus, immigration and a biased stratigraphic record Woodward 1908: pl. 29, figs 5 & 13; Lophiostomus, (Gale et al. 2000; Smith et al. 2001). Woodward 1909: pl. 33, fig. 4a) or a combination Although the ultimate cause of faunal changes of the two (e.g. Caproberyx (misidentified as Hop- within the Chalk Group remains debated, the com- lopteryx), Woodward 1902: pl. 6; Fig. 5a). Such position of fossil assemblages and the anatomies postures are referred to as tetany, and often taken of individual taxa provide clues to the palaeo- as evidence of death by anoxia, heat shock or alka- ecology of chalk seas. Sediments of the Chalk Group linity shock by comparison with modern examples are often bioturbated to the degree that the origi- (Elder & Smith 1984, 1988; Fig. 5). Second, many nal sedimentary structure is obliterated (Mortimore fishes show minimal disruption of individual skel- et al. 2001; Gale & Kennedy 2002). Apart from etal elements. This suggests a lack of scavenging providing clues about organisms dwelling on and that would otherwise scatter bones, implying rapid within the substrate, Chalk ichnology provides a burial or an absence of potential scavengers. Both means to assess the degree of sediment compaction, mechanisms are difficult to reconcile with con- as well as distinguishing between depositional ventional views of deposition and palaeoecology and diagenetic features visible in outcrop (Gale & of the Chalk. Mean sedimentation rates are esti- Kennedy 2002). A large body of literature focuses mated at 25 mm ka21 and there is abundant evi- on the geological and environmental implications dence (in the form of body and trace fossils) of of trace fossils in chalk settings (e.g. Bromley 1967, both benthic organisms that would have scavenged 1984, 1986; Kennedy 1967; Ekdale & Bromley fish carcasses on the seafloor and burrowers that 1984). Historical interpretations of the Chalk sea would disrupt shallowly buried remains. Classic proposed an ooze-like benthos, with anatomical views of chalk formation entail a gentle pelagic structures of some invertebrates (e.g. bivalves) rain, but it is now clear that deposition of the interpreted as adaptations to prevent individuals Chalk and similar deposits elsewhere was dynamic from sinking into the soupy seafloor (Carter 1968, (Bromley & Ekdale 1987; Buchbinder et al. 1988; 1972). Such features, however, are not found in Esmerode et al. 2008). Although diagnostic features Downloaded from http://sp.lyellcollection.org/ by guest on September 24, 2021

176 M. FRIEDMAN ET AL.

Fig. 5. Examples of tetany (gaping mouths or spinal curvature induced during death by anoxia, heat shock or alkalinity shock) in fossil fishes of the Chalk. (a) Detail of slab documenting a mass-mortality event of Caproberyx superbus (NHMUK PV P 9153), from the Lower Chalk. Most individuals exhibit strong spinal curvature, and the two individuals shown here demonstrate the gaping mouths characteristic of tetany in fishes. (b) An example of Osmeroides lewesiensis (NHMUK PV OR 4207) that exhibits extremely pronounced gaping, including flaring of the branchiostegals. (c) Osmeroides levis (NHMUK PV OR 49903). (d) Lophiostomus dixoni (NHMUK PV OR 23023) specimen preserving a terebratulid in its mouth, suggesting energetic sediment/water flows and rapid burial. Left image is transparent rendering of specimen derived from CT data, with brachiopod shown in white and marked with an asterisk. Scale bars represent 10 mm. Downloaded from http://sp.lyellcollection.org/ by guest on September 24, 2021

ENGLISH CHALK AND LONDON CLAY FISHES 177 of mass sediment transport in chalk deposits have European (Friedman 2012; Bien´kowska- been identified, these cannot easily be detected in Wasiluk et al. 2015). museum specimens. Despite these limitations, there is some circumstantial evidence for rapid bur- Geological context: London Clay ial of fishes. Such a mechanism seems to be the only plausible origin of remarkable Chalk fossils like Formation NHMUK PV P 9153, which includes several three- The London Clay Formation comprises a 90– dimensionally preserved individuals of Caproberyx 130 m-thick sequence of monotonous, fine-grained strewn throughout a slab of Chalk (Fig. 5a). Com- marine sediments of early Eocene (Ypresian, c. puted tomography study of Chalk fishes (see 54–50 Ma) age. The formation underlies most of below) reveals more subtle evidence in the form Greater London, but crops out over a broad area in of a complete terebratulid brachiopod within the southeastern England. Equivalent strata also occur oral cavity of the holotype of Lophiostomus in Europe as part of the Anglo- (Davis (Fig. 5d). The brachiopod is an unlikely prey item & Elliott 1957; Curry 1965), which extends from the (Lophiostomus has piercing, rather than crushing, southeastern coast of the UK to the western coast of dentition), and as an epifaunal organism, would mainland Europe. Today, this single basin has been not have burrowed into this position. It seems divided into a number of smaller sub-basins. In more probable that both were entrained in sediment southern England, London Clay Formation occurs and rapidly buried. Anecdotal evidence like this within the and London basins; offshore, aside, there is a lack of geological context for many these connect to the Dieppe and North Sea basins, Chalk fishes for testing taphonomic hypotheses. while on mainland Europe, contiguous deposits However, if slumps were genuinely a significant occur in Holland (known from boreholes only), contributor to the preservation of fishes in the NW and (mostly known from Chalk, then there might be striking differences in borings; Davis & Elliot 1957), France (small expo- the mode of preservation of fishes as a function sures from the Paris Basin possibly present in north- of their ecology. Sediment mass-transport would ern France; Davis & Elliot 1957) and Belgium, principally entrain fishes living in proximity to the where the 140 m-thick Argile d’Ypres is well docu- seafloor rather than those dwelling high in the mented (Davis & Elliot 1957; Steurbaut 2006). The water column, and it is these taxa that should be London Clay Formation and the underlying Har- preserved most frequently as articulated specimens. wich Formation (defined by Ellison et al. 1994; Taxa inhabiting surface waters would be less likely formerly known as the Blackheath or Oldhaven to become engulfed in sediment flows than those Beds) together form the Thames Group (King 1981). living at or near the seafloor. Examples of these Most of the London Clay Formation consists of species might be expected to represent individuals heavily bioturbated but homogeneous argillaceous that sank to the bottom after death rather than or slightly clay, with intervals of silty those buried alive. Such taxa might be characterized to sandy clays. The upper part of the sequence is by high degrees of scattering and disarticulation sandier than the lower, and the sequence terminates by benthic scavengers, to the degree that small- to at the base of the ‘Bagshot Sands’ (Whitaker 1866; medium-sized forms might be very rare. Anecdot- King 1981). Unweathered London Clay is blue- ally, pelagic taxa like clupeomorphs are relatively grey, but rapidly oxidizes to a reddish-brown colour. common at penecontemporaneous sites in Lebanon Well-defined horizons of calcareous (Forey et al. 2003), but are represented by only or nodules occur throughout the sequence. The two specimens from the entirety of the Chalk western regions of the London Clay Formation succession (Table 1). Examination of large collec- were closer to the Eocene shoreline and are coarser- tions of Chalk fossils might be able to determine grained than the fine-grained muds of the east, whether there is such a disparity in preservational which were deposited in deeper waters (King 1981). mode between benthic and pelagic fishes consis- tent with this taphonomic model. However, caution Stratigraphy must be applied to any such inference owing to the non-systematic collection of nearly all Chalk The most commonly used (though not the most up- specimens. to-date) subdivision of the London Clay Formation Although articulated fishes are rare in the Chalk, is that of King (1981). King’s informal Divisions invertebrate burrows lined with fish debris are A–E reflect five major transgressive–regressive more common. Mantell (1822) and Agassiz (1833– cycles, defined based on lithological and biostrati- 44) mistook these as articulated body fossils of elon- graphic criteria. Divisions are separated by marker gate fishes, which were informally known among horizons, and each is marked by a coarsening- quarrymen as ‘fossil ’ (Davies 1879, p. 145). upwards depositional sequence (King 1984). Occur- Such burrows are common in Late Cretaceous rences of layers and laminations also Downloaded from http://sp.lyellcollection.org/ by guest on September 24, 2021

178 M. FRIEDMAN ET AL. aid the delineation of divisions (King 1981). Ellison outcrops in many locations across southeastern et al. (2004) revised King’s (1981) stratigraphy England and is best accessible along the coast, par- based on information from boreholes, dividing the ticularly in Kent (e.g. Isle of Sheppey, Seasalter, London Clay Formation into informal Units A–D; Medway, Tankerton, Whitstable, Herne Bay and although the two schemes are similar, unit and divi- Swalecliffe), Essex (e.g. Maylandsea, Burnham- sion boundaries do not correspond exactly. Correla- on-Crouch, Aveley, Harwich, Steeple Bay, Stans- tions between King’s divisions and Ellison et al.’s gate Abbey Farm, Mundon Stone Point, Iltlney, units, together with their thicknesses and litholo- Wrabness, Ockendon, Ongar and Walton-on-the- gies, are given in Figure 6. Naze), Sussex (e.g. Bognor Regis), Bournemouth Only the Claygate Member (¼‘Claygate Beds’), and Studland, and the . Many ephem- located at the top of the London Clay Formation eral exposures have also been created by road cut- (contemporary with Division E), has been for- tings and pits or quarries. Fossil fish material is mally mapped (Aldiss 2012). However, the British widespread within the London Clay, with a list Geological Survey plans to replace King’s (1981) of localities and their associated faunas given by and Ellison et al.’s (2004) informal divisions Hooker & Ward (1980; see also Hooker et al. 1980). with official members (Aldiss 2012). In addition Here we single out the four exceptional localities to the Claygate Member, these will comprise the (Bognor Regis, the Isle of Sheppey, Maylandsea ‘Sheppey Member’, which approximately corre- and Burnham-on-Crouch) selected by Dineley & sponds to Ellison et al.’s (2004) Unit D or to the Metcalf (1999) for the Geological Conservation upper part of King’s (1981) Division C, Division Review for more detailed summary. D and most of Division E; the ‘Aveley Member’, A 90 m sequence of London Clay from the east- which corresponds to Unit C/Division B and the ern edge of the is accessible lower part of Division C; the ‘Ockendon Member’, along the foreshore of Bognor Regis in Sussex, which corresponds to Unit B/Division A3; and representing Divisions A2 through to C; most verte- the Walton Member, which corresponds to Unit brate material derives from the Aldwick Beds A/Division A2 (King, unpublished manuscript; (Division B; Venables 1962; King 1981). Abundant cited by Aldiss 2012). insect and beetle fossils (Venables 1962) together An early Ypresian age (c. 52–49 Ma) for the with plant fossils (Chandler 1964; Collinson 1983) London Clay Formation is supported by the identi- and nautiloids (Hewitt 1988) suggest a near-shore fication of calcareous nannoplankton species corre- marine environment with a water depth of less sponding to zones NP11 (Division B) and NP 12 than 70 m. Fishes, and are also (Divisions C–E; Martini 1971; King 1981; Aubry represented. The rich, well-preserved fish fauna of 1985), while the existence of NP10 (Division A2 the Aldwick Beds mostly comprises teeth, scales, or the ‘Walton Member’) has been inferred (Ellison otoliths and other disarticulated material collected et al. 1994; Ali & Jolley 1996). Biostratigraphic by bulk-processing from accumulations of zonation microfossils that assist correlations within along the foreshore or beach sand, although semi- the London Clay include dinoflagellates (Zones articulated fishes can be found in phosphatic nod- D5–D8; Costa & Downie 1976; Powell et al. ules (Venables 1962). Although diverse, 1996), (Keen 1977; King 1981), benthic form a more minor component of the fauna than at and planktonic foraminiferans (King 1981), Sheppey (perhaps because large concretions are and radiolarians (Ellison et al. 2004). Although less less prevalent; Venables 1962); the assemblage is prevalent, some diagnostic macrofossils (molluscs: dominated by predatory elasmobranchs, suggesting nautiloids, bivalves, gastropods and scaphopods; a highly productive marine environment (Dineley : Isselicrinus subbasaltiformis, I. basalti- & Metcalf 1999). formis and Cainocrinus; crustaceans: crabs and The exposures of the London Clay along the lobsters) are also recognized. The chronology of northern and northeastern shores of the Isle of the lower Palaeogene sediments of the London and Sheppey, Kent, are among the best known and most Hampshire basins has also been correlated with productive for fossil collecting (King 1981, 1984). related sections in NW Europe via magnetostratig- Although the full 165 m sequence is accessible by raphy (Aubry et al. 1986). Sequence stratigraphical drill-coring, only the upper 53.5 m crops out, repre- treatments of the London Clay in the context of the senting Divisions D–E (King 1981, 1984). Biostra- English early Palaeogene are given by Knox tigraphic relationships between Sheppey and rest (1996a) and Neal (1996). of the London Clay Formation have been establi- shed via , dinoflagellate zones, although Key exposures no diagnostic calcareous nannoplankton species have been identified (King 1981, 1984). Bands The type area is Greater London and Essex (Aldiss of concretionary nodules are common within the 2012). However, the London Clay Formation monotonous clays, with some sandier layers (Davis Downloaded from http://sp.lyellcollection.org/ by guest on September 24, 2021

ENGLISH CHALK AND LONDON CLAY FISHES 179

Fig. 6. Lithostratigraphic and biostratigraphic subdivisions, and geographical extent of the London Clay. (a) Lithostratigraphic and biostratigraphic subdivisions of the London Clay, showing the relationship between King’s (1981) informal Divisions A–E, and Ellison et al.’s (2004) Units A–D. NP Zones, nannoplankton zones given by Gradstein et al. (2012). Stratigraphic extents of key fish localities are given on the right. (b) Geographical extent of the London Clay outcrop, after King (1984).

1936; King 1981, 1984). Sheppey represents a low- through the succession; the lithology becomes energy, well-oxygenated middle-shelf environment coarser towards the top, reflecting progressively with a depth fluctuating between c. 20 and c. 100 m shallower sea-levels (King 1984; Islam 1984). Downloaded from http://sp.lyellcollection.org/ by guest on September 24, 2021

180 M. FRIEDMAN ET AL.

Sheppey is particularly noted for the tremen- In addition to these sites, a nearly complete dous variety of exceptionally preserved, three- sequence of London Clay strata could formerly be dimensional fossils, including many articulated fish found in a quarry at Aveley in Essex; unfortunately, specimens (mostly skulls, although some preserve the site has been infilled and is no longer acces- postcranial elements). Most fossils derive from sible. An account of the geology and palaeontology Division D, in an interval 9.5–16 m below the of this site is given by Williams (2002). base of Division E (key fossil localities occur at Minster, Royal Oak, Bugsby’s Hole, East Church Palaeoenvironment Gap, Barrow Brook and Warden Point; Davis 1936; King 1981). Non-vertebrate macrofossils comprise In the early Eocene, the southern UK lay at a palae- a wide diversity of shallow-marine (mostly epifau- olatitude of approximately 428 N (Smith et al. nal or infaunal) forms along with forms washed 1994). Much of the southeastern portion of England in from terrestrial or near-shore habitats, including was submerged, with the shoreline probably extend- crustaceans, molluscs, , , ing NE–SW, from the Wash to near the Isle of bryozoans, coelenterates, , annelids and Wight (Wills 1951). The Isle of Sheppey there- plants (Dineley & Metcalf 1999). The most common fore represents a depositional environment at least vertebrate fossils are fish (mostly represented by 80 km from the shoreline (Collinson 1983). teeth and vertebrae), but reptiles, and Although the London Clay deposits are exclu- birds are also well known. are exceptionally sively marine, water depth and distance from the diverse, and Dineley & Metcalf (1999) speculate shoreline vary somewhat owing to the series of that ray-finned fishes may be less well represented transgressive–regressive cycles recorded through- for taphonomic reasons. However, King (1981, out the sequence – five major cycles, overprinted pp. 126–128) suggests that by 19 more minor cycles (Neal 1996). Conditions were predominantly deeper-water and low-energy, the relative abundance of fish remains at localities coinciding with the elevated sea-levels of the early such as the Isle of Sheppey ... and Bognor Regis ... Eocene (King 2006). In contrast to the shallow- is more apparent than real, and is dependent on their marine conditions represented by the underlying concentration by wave-action on a clay foreshore Harwich Formation, the base of the London Clay with an extensive outcrop. At other localities with London Clay Formation exposures but no such con- Formation marks a substantial increase in depth centration [resulting from wave-action] (such as Alum associated with the sea-level highstand phase and Bay and ), records [of fish remains] establishment of a true oceanic environment, as are few. muds were deposited over most of the southern UK (as indicated by the influx of planktonic forami- Another remarkably productive locality is that of nifera and gastropods at the level of the ‘planktonic Maylandsea, Essex, where Division C of the London datum’; King 1981; Knox 1996b). This low-energy Clay outcrops intermittently along the low banks of deeper marine environment (either an open shelf Lawling Creek and in the intertidal zone of the or a more restricted lagoon/embayment; Ellison Blackwater Estuary. These facies are thought to rep- et al. 2004) was generally deeper in the London resent the deepest-water environments recorded in Basin than in the Hampshire Basin (Murray & the London Clay, and this environmental difference Wright 1970; Collinson 1983 (forams); Keen 1977 is reflected in the fish species present (known mostly (ostracods); Walder 1964; and Gilkes 1968 (miner- from teeth and otoliths (Stinton in Casier 1966; alogy)). Some of the deepest-water facies are seen at Stinton 1975–84), although one skull has been Maylandsea (the deeper-water faunal assemblage found). includes species not seen at other sites; Dineley & Exposures along the tidal-river cliffs and fore- Metcalf 1999). Sandier intervals scattered through- shore at Burnham-on-Crouch, Essex, have yielded out the London Clay succession reflect subtidal, an extraordinarily rich macrofauna (Dineley & Met- stormy conditions (Huggett & Gale 1998); at the calf 1999), although the fish assemblage is heavily top of the London Clay Formation, the Claygate dominated by elasmobranchs. The exposures repre- Member represents tidal conditions (Ellison et al. sent 2–3 m of the topmost interval of Division D, 2004), while the overlying Bagshot Beds represent 26 m below the Claygate Member. The palaeo- a fall in sea-level and return to sublittoral sedimen- environment and fish fauna are similar to those tation (Knox 1996b). Marker horizons associated recorded on Sheppey, but remains are limited to with fluctuations in sea-level were one line of evi- disarticulated teeth and bones extracted by bulk dence used by King (1981) to delineate his Divi- sampling techniques. Although concretions are sions A–E (the start of each division is thought common along the foreshore, few have been found to coincide with a transgressive event; see Daley to contain fossils. However, preservation is better & Balson 1999, fig. 2.6). Although sediments com- than at Sheppey (Dineley & Metcalf 1999). prising the London Clay could have originated Downloaded from http://sp.lyellcollection.org/ by guest on September 24, 2021

ENGLISH CHALK AND LONDON CLAY FISHES 181 from any land surrounding the North Sea Basin (insects), and terrestrial transported (Collinson 1983), they were at least partially locally into deep waters (Casier 1966; Ward 1979; Hooker derived from the Chalk, West Country granites & Insole 1980; Hooker & Ward 1980; Walker 1980; (Gilkes 1968), and pyroclastic ashes (Knox & Benton & Spencer 1995). Most of the vertebrate Harland 1979). material is preserved in diagenetic nodules or con- The deposition of the London Clay occurred cretions, primarily those with phosphatic or calcitic either during or just after the –Eocene chemistries (Allison 1988). Thermal Maximum; the period of slow cooling The or ‘septarian’ nodules are that followed the Paleocene–Eocene Thermal Max- lenticular and tabular concretions, ranging from imum began later, in the mid-Eocene (Collinson 300 mm to over 1 m in diameter, that contain radial 1983; King 2006). There can be no doubt that the cracks infilled by yellow-brown ferroan calcite or climate was warm and frost-free, although floristic other (e.g. iron pyrite, baryte or vivianite; evidence is not entirely consistent with purely trop- Huggett 1994; Huggett & Gale 1995; Williams ical conditions (Collinson 2000). The fossilized 2002) and are generally concentrated in discrete flora of the London Clay is presented in detail by bands within the strata (King 1981, 1984); many Collinson (1983; early work was done by Reid & nucleate around fragments of fossil wood (Huggett Chandler 1933). The assemblage is one of the 1994; Huggett & Gale 1995) and some preserve col- world’s most diverse, with over 500 species pre- onies of teredinid bivalves (Huggett et al. 2000). served, of which over 350 are named (Collinson The ubiquitous bioturbation within the London 1983); the presence of such a diverse and abundant Clay extends to the interiors of concretions. The flora in a deep-marine environment is probably presence of uncompacted burrows within these explained by masses of plant debris rafting down structures indicates that they formed during early rivers and floating out to sea (Collinson 1983). Well- diagenesis (Huggett 1994). preserved seeds, fruits, twigs, wood fragments and The phosphatic nodules (composed of francolite, other plant debris from a range of flowering plant or carbonate fluorapatite; Hewitt 1982) are gener- (magnolias, palms, vines, lianas, dogwoods, laurels ally smaller than those composed of calcium car- and bays) and gymnosperm lineages are indicative bonate, and typically have softer, buff-coloured of a range of different climatic zones. Most are exteriors with harder, darker brown cores; fossils from groups that are, today, exclusively tropical are more commonly found in the smaller, ovoid (131 species). However, 86 subtropical species are nodules, although larger, lenticular nodules and also present, along with temperate (60 species) cylindrical, vertically oriented nodules (perhaps and even north temperate (17 species) groups, indi- representing infilled burrows) have also been docu- cating some seasonality (Collinson 1983). Nipa mented (King 1984; Williams 2002). Phosphatic palm seeds, along with Ceriops and other rhizophor- nodules are found on the Isle of Sheppey, and have acean taxa (and additionally by the preservation been recovered at the now-inaccessible quarry at of teredinid bivalve genera associated with man- Aveley, Essex (King 1984; Williams 2002; Young groves; Huggett et al. 2000), indicate that man- & Williams 2008). groves fringed the nearby coast, while further Preservation of hard parts almost exclusively inland, dense forests and reed marshes grew along occurs within early-diagenetic pyritic, calcareous rivers; grasses appear to have been rare or absent. or phosphatic concretions or nodules. Minerali- Collinson (1983, p. 24; see also Daley 1972) zation of skeletal material is strongly correlated draws parallels with modern ‘para-tropical’ Indo- with its original composition (Allison 1988): phos- Malaysian environments, but cautions that ‘the phatic tissues such as vertebrate bone are usually London Clay flora, like other Tertiary floras, cannot preserved as brown-black calcium phosphate within be precisely equated with any vegetation or climatic phosphate nodules, while skeletons mineralized regime which exists today’ in a world where temper- with calcium are usually found in calcium car- ate conditions reached as far as the poles. bonate concretions (Balson 1980; Hewitt 1982; Allison 1988; Huggett 1994; Dineley & Metcalf Taphonomy and diagenesis 1999). Most of the best-preserved vertebrate fos- sils are found within phosphatic nodules. This pres- Most and plant fossils from the London ervation bias can be explained by the sequence of Clay Formation are exceptionally preserved in mineralization events (first apatite, then calcite three dimensions, often with very little taphonomic and pyrite), as the process of decay is halted only distortion, and it has been argued that the deposits by early diagenetic mineralization (Allison 1988; represent a Konservat Lagersta¨tten (Allison 1988). Huggett 1994). In addition to actinopterygian and chondrichthyan Preservation of soft parts is exceedingly rare, but fishes, the London Clay biota includes invertebrates, not unknown (e.g. a fossil insect larva, Rowell & both marine (crustaceans, molluscs) and terrestrial Rundle 1967; and a lophophore of a terebratulid Downloaded from http://sp.lyellcollection.org/ by guest on September 24, 2021

182 M. FRIEDMAN ET AL. brachiopod, Rundle & Cooper 1971). Allison (1988) 1901) and The Fossil Fishes of the English Chalk concluded that soft tissues had decayed prior (1902, 1903, 1907a, 1908, 1909, 1911, 1912a). The to phosphatization, a process that usually causes most important actinopterygian specimens had an animal’s remains to collapse and be flattened; been collected from the Chalk by Smith Wood- routine three-dimensional preservation at localities ward’s time, with a rarity of subsequent discoveries such as Sheppey and Aveley quarry, particularly in the field reflecting a decline in collectors com- the preservation of fish scales in situ, implies bined with a shift to mechanical mining of chalk ‘extremely rapid rates of sedimentation’ (Allison pits in the late nineteenth and early twentieth centu- 1988, p. 1081) that would have enabled sedi- ries (Patterson 1964). ments to infill the body cavity during decomposi- There has been little study of bony fishes from tion. Calcite-infilled septarian nodules, by contrast, the English Chalk in the more than a century since are thought to have formed during periods of slow Smith Woodward’s (1902, 1903, 1907a, 1908, burial (Huggett 1994), perhaps partially account- 1909, 1911, 1912a) synoptic overview. A handful ing for the paucity of vertebrate fossils contained of monographs in the second half of the twentieth within. Furthermore, dysaerobic or anaerobic bot- century provided more detail on some groups: tom-waters are indicated by the preservation of Patterson (1964) on ctenothrissiforms and acan- phosphate-rich tissues as apatite (Smith 1987). thomorphs, Goody (1969) on aulopiforms, Forey Hard parts in both phosphatic and calcareous (1973) on elopomorphs and on pachyrhizodontids concretions tend to be pyritized to some extent, (Forey 1977). In reviewing modern groups, Regan particularly if exposed to the elements for a substan- (1911; acanthomorphs) and Rosen (1973; aulopi- tial period of time before collection. While moder- forms, ctenothrissiforms) made detailed compari- ate, uniform pyritization can improve the contrast sons with Chalk taxa in an attempt to place fossils obtained from CT scans (contrast between fos- relative to living groups. More recent contributions sil and matrix may be low when pyritization has are limited in taxonomic scope and include: partial not occurred), excessive pyritization obscures the redescriptions of the elopomorphs Urenchelys and boundaries between fossil and matrix and can sub- Enchelurus (Belouze 2002, pp. 180–182, 227– stantially hinder the processing of CT data. Small 229); revision of tselfatiiforms erecting two new pyritic nodules also occur in isolation and tend to genera (Petanogmius and Dixonanogmius; Taverne preserve plant materials, insects, stems, 2000a; Taverne & Gayet 2005) and rejection molluscs, wood fragments, algal tubes and burrows of Smith Woodward’s (1908; 1936) stomiiform (Allison 1988). Williams (2002) reports that pyrite interpretation of Tomognathus in favour of an amii- nodules are found in association with at least half form one (Forey & Patterson 2006); and reports of a of the fish skulls discovered at Aveley. new taxa or occurrences – a new species of the pyc- nodont Anomoeodus (Kriwet 2002), a specimen identified by Smith Woodward (1902) as Hoplop- A history of study of Chalk and London teryx but reinterpreted as the clupeomorph Dip- Clay fishes lomystus (Forey 2004), and the recognition of the new suspension-feeding pachycormid Rhin- Modern study of fossil fishes from the English Chalk conichthys (Friedman et al. 2010). These additions can be traced back to the early nineteenth century to the Chalk fish fauna stem from restudy of his- when Mantell described his collection in Fossils torical material, all of which was collected prior of the South Downs (1822), followed by the more to Smith Woodward’s (1902, 1903, 1907a, 1908, exhaustive descriptions by Agassiz (1833–43), 1909, 1911, 1912a ) monograph. Given the rarity who recognized a diverse actinopterygian fauna of new discoveries of articulated material in the of roughly 30 species from the Chalk. Specimens Chalk, future additions to the Chalk fauna will gathered together by Dixon, Egerton, Enniskillen most likely arise from examination of existing col- and other private collectors were later acquired by lections. For example, the elongate rostra named the British Museum and now form the core of the by Smith Woodward (1909, 1912b; Fig. 4b) as comprehensive collection of English Chalk fish Protosphyraena stebbingi bear midline grooves fossils housed at the Natural History Museum and paired canals indicating that they probably (Woodward 1902, 1903, 1907a, 1908, 1909, 1911, constitute paired bones rather than the median ros- 1912a). Isolated papers from the late nineteenth trodermethmoid that contributes to the snout of century gave descriptions of some of these fossils Protosphyraena and other pachycormids. Elongate (e.g. Newton 1877) before more extensive treatment rostra composed of paired premaxillae are known by Smith Woodward in his Catalogue of Fossil from some tselfatiiforms (e.g. Martinichthys; Tav- Fishes (elasmobranchs: 1889c; holocephalans and erne 2000b), which are similar to fossils of ‘P.’steb- coelacanths: 1891a; ‘lower’ teleosts and other non- bingi in being comparatively flat in cross section actinopterygians: 1895a; ‘higher’ teleosts: with one surface covered with a dense network of Downloaded from http://sp.lyellcollection.org/ by guest on September 24, 2021

ENGLISH CHALK AND LONDON CLAY FISHES 183 denticles; we advocate a tselfatiiform interpretation substantial revision of scombroids and xiphioids of this taxon here (Table 1). (Monsch 2000, 2005). Detailed study of fishes from the London Clay, Several authors have questioned Casier’s (1966) like those of the Chalk, commenced in the early higher-level attribution of fishes from the London part of the nineteenth century. Ko¨nig (1825) named Clay without any substantial reanalysis or rede- and figured skulls belonging to three genera still scription of relevant material (Table 2). Patterson recognized today: Bucklandium, Ampheristius and doubted the amiid affinities of Lehmanamia (Patter- Teratichthys. Agassiz recognized over 20 actino- son 1973, p. 277; reiterated by Lambers 1994; pterygian species from the London Clay by the Grande & Bemis 1998), as well as the interpretation time he completed his study of fossil fishes, but of Bramoides and Goniocranion as bramids (Patter- many of these were published solely as names and son 1993b, p. 644), but did not offer alternative were formally described only by later authorities. placements in any of these cases. Patterson also Owen (1840–45), Cocchi (1866), Egerton (1877) reassigned Beerichthys, regarded by Casier as an and Noetling (1885) made minor contributions to aulopiform, to Luvaridae (p. 649), an assessment the fauna in the decades following Agassiz. It was borne out by our ongoing CT study of this material. Smith Woodward who made the greatest contribu- It has been suggested that Whitephippus and tion to the London Clay fauna near the turn of the Goniocranion – interpreted by Casier as an ephip- nineteenth and twentieth centuries, principally pid and a bramid, respectively – are lampridiforms through the fourth volume of his Catalogue (1901) (attributed to Bonde 1995 by Carnevale 2004, but but also through a series of more minor publica- not appearing in that publication), while Bannikov tions (Woodward 1889a, b, d, 1893a). Smith Wood- (1979, 1985) reassigned Eothynnus from Scombri- ward named numerous new species and formally dae to . In contrast to the Chalk, new dis- described taxa that had been nomina nuda since coveries of articulated fish remains continue to be Agassiz (1833–43). Although he undertook mono- made in the London Clay, thanks to a combination graphic descriptions of fishes from the Chalk of active seaside erosion and a dedicated community (Woodward 1902, 1903, 1907a, 1908, 1909, 1911, of collectors. Some of the most spectacular remains 1912a) and Wealden & Purbeck (1916–19), there have been featured in popular books (e.g. Clouter is no evidence that Smith Woodward made plans et al. 2000; Rayner et al. 2009). to do the same for the London Clay fauna. Errol White, who succeeded Smith Woodward as the principal palaeoichthyologist at the British Museum Significance of the Chalk and London (Natural History), intended to review Eocene British Clay faunas vertebrates – including the fishes of the London Clay – but succeeded in producing only a single The significance of the bony fish faunas of the Chalk contribution covering the Thanet Sands and Base- Group and London Clay Formation stems princi- ment Bed of the London Clay (White 1931). As pally from two key features, both fully appreciated White’s research interests shifted, the task of deliv- by Smith Woodward (1912a, 1942): the unique ering the monograph on London Clay fishes was preservation within these units and their age relative passed to Edgar Casier of the Institut royal des Sci- to major events in actinopterygian evolution. ences naturelles de Belgique, who was familiar with Ypresian fishes from his own work in lateral equiv- Remarkable preservation complements to alents of the London Clay in Belgium (e.g. Casier other fossil Lagersta¨tten 1946). Casier’s (1966) volume, with a supplement later joined by a monograph on British Eocene The Chalk and London Clay are notable for their otoliths by Frederick Stinton (1975–84), stands as three-dimensional preservation of fish specimens, the principal reference on fossil fishes from this particularly skulls. This quality has long been recog- deposit. Subsequent revisions to the actinopterygian nized, with Mantell (1844, p. 663) writing that body fossil record have been limited. The most Chalk fishes ‘are exceedingly beautiful Ichthyolites, substantive accounts based on historical material and are almost invariably found with body but little available to Smith Woodward and Casier include: compressed; the fish, in many examples, is as round redescriptions of the osteoglossomorph Brychaetus and perfect as when living’. Smith Woodward (Roelling 1974; Taverne 1974, 1979), accompanied (1902, p. 1) noted that Chalk fishes ‘exhibit the by its temporary synonymization with Phaerodus essential details of their osteology as well as modern (Li 1994; Li et al. 1997) followed by reinstatement skeletons’ and represent an essential complement to (Taverne 2009); revised anatomical accounts of elo- ‘whole fishes [from other coeval localities] in a piform and ‘albuliform’ elopomorphs (Forey 1973); crushed state, displaying the , redescription of the sparid Sciaenurus and place- fins, and scales, in undisturbed position, but show- ment relative to modern forms (Day 2003); and a ing almost nothing of the cranial osteology’. Casier Table 2. Actinopterygian fauna of the London Clay Formation 184

Taxon Type/relevant specimen Specimens Locality London Clay Divisions Downloaded from

ABCDE

Chondrostei Acipenseriformes Acipenser toliapicus Woodward, 1889a NHMUK PV P 3372, P 3372a, P Multiple Sheppey X 529, P 37762, P 24619, P 44093

Acipenser sp. Private collection Multiple Sheppey X X x X x http://sp.lyellcollection.org/ Halecomorphi *Lehmanamia sheppeyensis Casier, 1966 MNHN PTE 11 Single Sheppey x Teleostei Pycnodontiformes Pycnodus toliapicus Agassiz, 1839 OUMNH P 3759 Multiple Sheppey x FRIEDMAN M. Pycnodus bowerbanki Egerton, 1877 NHMUK PV OR 38824 Single Sheppey x Pycnodus sp. NHMUK PV P 3783 Multiple Sheppey, Swanwick x Elopiformes *Protarpon priscus (Woodward, 1901) NHMUK PV OR 36070 Multiple Sheppey x Protarpon cf. priscus NHMUK PV P 4153 Single Sheppey x TAL. ET Protarpon oblongus (Woodward, 1901) NHMUK PV P 634 Single Sheppey x byguestonSeptember24,2021 *Promegalops signeuxae Casier, 1966 MNHN PTE 9 Single Sheppey x Promegalops sheppeyensis Casier, 1966 NHMUK PV P 9192 Single Sheppey x X x *Esocelops cavifrons Woodward, 1901 NHMUK PV P 1749 Multiple Sheppey X x X x *Elops sp. NHMUK PV OR 39443, P 1762 Multiple Sheppey x ‘Albuliformes’ *Albula oweni (Owen, 1845) NHMUK PV P 9158 Multiple Sheppey, Bognor Regis x x X x Anguilliformes *Echelus branchialis (Woodward, 1901) NHMUK PV P 633 Multiple Sheppey X X x X x Osteoglossiformes *Brychaetus muelleri Woodward, 1901 NHMUK PV P 3893 Multiple Sheppey, Chalk Hill Tunnel x x x X x cf. Heterosteoglossum sp. NHMUK PV P 65206 Single Aveley Quarry X Osteoglossidae indet. NHMUK PV P 66354 Single Sheppey x x x x Clupeiformes indet. NHMUK PV P 47286, P 61527 Multiple Sheppey, Burnham-on-Crouch x X X x Gonorynchiformes *Halecopsis insignis (Delvaux & Ortlieb, 1888) ‡IRSNB P 00143-5 Multiple Primrose Hill, Highgate XXxXx Archway, Sheppey, Southend-on-Sea, Walton-on-the-Naze Siluriformes

Bucklandium diluvii Ko¨nig, 1825 NHMUK PV P 9230 Single Sheppey x Downloaded from Aulopiformes *Aulopopsis egertoni Casier, 1966 NHMUK PV P 9410 Single Sheppey x *Aulopopsis depressifrons Casier, 1966 NHMUK PV P 26712 Multiple Sheppey, Xxx Southend-on-Sea, Aveley Quarry (B2) *Labrophagus esocinus Casier, 1966 NHMUK PV P 621 Multiple Sheppey x *Argillichthys toombsi Casier, 1966 NHMUK PV P 42519 Single ?Sheppey x

Lampridiformes http://sp.lyellcollection.org/ *Whitephippus tamesis Casier, 1966 NHMUK PV P 6479 Multiple Sheppey x Whitephippus sp. NHMUK PV OR 35057 Single Sheppey x FISHES CLAY LONDON AND CHALK ENGLISH *Rhinocephalus planiceps Casier, 1966 NHMUK PV OR 47985 Multiple Sheppey, Herne Bay X x X x Trichiurides sagittidens Winkler, 1874 ‡IRSNB P 00264 Multiple Sheppey, Bognor Regis X x X x Beryciformes *Argilloberyx prestwichae Casier, 1966 NHMUK PV P 12944 Single Southend-on-Sea x *Naupygus bucklandi Casier, 1966 NHMUK PV P 1764 Multiple Sheppey x *Paraberyx bowerbanki Casier, 1966 NHMUK PV OR 38912 Single Sheppey x *Sciaenuropsis turneri Casier, 1966 NHMUK PV P 644a Multiple Sheppey x

(including ‘Myripristis toliapicus’) byguestonSeptember24,2021 Ophidiiformes *Ampheristus toliapicus Ko¨nig, 1825 NHMUK PV P 9432 Multiple Sheppey x x X x Syngnathiformes cf. Ramphosus sp. NHMUK PV P 60896 Single Sheppey (Warden Point) X Scombroidei *Eutrichiurides winkleri Casier, 1946 ‡IRSNB P 00320, P 00324-9 Multiple Highgate Archway, XXxXx Sheppey, Herne Bay, Hampton, Bognor Regis *Progempylus edwardsi Casier, 1966 NHMUK PV OR 32388 Single Sheppey x ?Gempylidae NHMUK PV OR 41318 Single Sheppey x *Scombrinus nuchalis Woodward, 1901 NHMUK PV P 4148 Multiple Sheppey, XXxXx Primrose Hill Gymnosarda prisca Monsch, 2000 NHMUK PV P 6485 Single Sheppey x Gymnosarda sp. NHMUK PV P 1773b Single Sheppey x cf. Gymnosarda sp. NHMUK PV P 4546 Single Sheppey x 185 (Continued) Table 2. Continued 186

Taxon Type/relevant specimen Specimens Locality London Clay Divisions Downloaded from

ABCDE

Sardini incertae sedis NHMUK PV P 9459 Single Sheppey x Sardini incertae sedis NHMUK PV P 45150 Single Sheppey x Thunnini incertae sedis NHMUK PV P 4300 Single Sheppey x aff. Acanthocybium NHMUK PV P 27010 Single Sheppey x Palaeocybium proosti (Storms, 1897) ‡IRSNB P 00396 Single Sheppey x http://sp.lyellcollection.org/ Acanthocybiinae incertae sedis NHMUK PV OR 241686c, Multiple Sheppey x P 38883 *Eocoelopoma colei Woodward, 1901 NHMUK PV P 623a Multiple Sheppey x *Eocoelopoma curvatum Woodward, 1901 NHMUK PV OR 44877a Multiple Sheppey, XXxXx Hampstead Tunnel *Eocoelopoma gigas Woodward, 1901 NHMUK PV OR 39221 Multiple Sheppey x FRIEDMAN M. Eocoelopoma sp. NHMUK PV P 29983, P 26706 Multiple Sheppey, xx Bognor Regis *Tamesichthys decipiens Casier, 1966 NHMUK PV OR 41319 Single Sheppey x *Duplexdens macropomus (Agassiz, 1844) MNHN PTE 10 Multiple Sheppey x x X x ?Duplexdens sp. NHMUK PV OR 38903 Single Sheppey x TAL. ET

*Micrornatus hopwoodi (Casier, 1966) NHMUK PV OR 36136 Single Sheppey x byguestonSeptember24,2021 *Scombramphodon crassidens NHMUK PV OR 1779 Multiple Sheppey x X x X x Woodward, 1901 *Sphyraenodus priscus Agassiz, 1844 NHMUK PV P 3957 Multiple Sheppey x *Wetherellus cristatus Casier, 1966 NHMUK PV OR 28498 Multiple Sheppey x *Woodwardella patellifrons Casier, 1966 NHMUK PV P 26903 Single Sheppey x Sphyraenidae Sphyraena bognorensis Casier, 1966 NHMUK PV P 29984-5 Multiple Bognor Regis X (lower fish tooth bed) Carangoidei *Teratichthys antiquitatis Ko¨nig, 1825 NHMUK PV OR 28760 Single Sheppey x *Eothynnus salmoneus Woodward, 1901 NHMUK PV P 623 Multiple Sheppey x X x Xiphioidei *Xiphiorhynchus priscus (Agassiz, 1844) MNHN PTE 747 Multiple Sheppey, XXxXx Herne Bay ‘Makaira’ sp. NHMUK PV OR 30798 Single Sheppey x Xiphioidei incertae sedis NHMUK PV P 21306 Single Sheppey x Aglyptorhynchus sulcatus (Casier, 1946) ‡IRSNB P 00315 Multiple Sheppey, xx Bognor Regis Aglyptorhynchus venablesi Casier, 1966 NHMUK PV P 26157 Multiple Sheppey, Bognor Regis XxXx

(lower fish tooth bed) Downloaded from *Enniskillenus radiatus Casier, 1966 NHMUK PV P 646 Multiple Sheppey X X x X x Rotundorhynchus brittanicus Monsch, 2005 NHMUK PV P 1765 Multiple Sheppey x Acestrus ornatus Woodward, 1901 NHMUK PV P 627 Multiple Sheppey x x x X x ‘Serranidae’ Plesioserranus cf. wemmeliensis NHMUK PV OR 35771 Single Sheppey x Ephippidae *Laparon alticeps Casier, 1966 NHMUK PV P 620 Single Sheppey x http://sp.lyellcollection.org/ ‘Sparidae’

*Sciaenurus bowerbanki Agassiz, 1845 NHMUK PV P 3975 Multiple Sheppey, XXxXx FISHES CLAY LONDON AND CHALK ENGLISH Bognor Regis Podocephalus nitidus Casier, 1966 NHMUK PV OR 37760 Multiple Sheppey x *Podocephalus curryi Casier, 1966 NHMUK PV OR 35697 Single Sheppey X x X x Acanthuroidei *Beerichthys ingens Casier, 1966 NHMUK PV OR 43105 Multiple Sheppey X x X x Beerichthys sp. NHMUK PV P 26799 Single Sheppey x Tetraodontiformes *Triodon antiquus Leriche, 1905 ‡IRSNB P 00590-00606, P 00644 Multiple Sheppey x *Tetraodontiformes n. gen. et sp. NHMUK PV P 63336 Single Aveley Quarry X byguestonSeptember24,2021 incertae sedis *Percostoma angustum Casier, 1966 NHMUK PV P 639 Single Sheppey x Pseudosphaerodon antiquus Casier, 1966 NHMUK PV P 164 Multiple Sheppey x X x Egertonia isodonta Cocchi, 1866 NHMUK PV OR 38814 Multiple Sheppey x toliapicus Agassiz, 1839 OUMNH lost Multiple Sheppey, xXxXx Beltinge, Frinton-on-Sea Bramoides brieni Casier, 1966 NHMUK PV P 6481 Multiple Sheppey x Goniocranion arambourgi Casier, 1966 NHMUK PV P 9478a Multiple Sheppey x *Serranopsis londinensis Casier, 1966 NHMUK PV OR 36135 Single Sheppey x incertae sedis Congorhynchus? elliotti (Casier, 1966) NHMUK PV P 21304 Multiple Sheppey x X x Cylindracanthus rectus (Dixon, 1850) NHMUK PV OR 25859 Multiple Sheppey, x Highgate

Stratigraphic distributions largely from Casier (1966) and Rayner et al. (2009). Occurrences marked by ‘X’ are definitive, while those indicated by ‘x’ are less certain. Horizon occurrences as marked represent those taken cumulatively across all geographical localities, rather than individual sites. Taxa marked with ‘*’ are known from relatively intact skulls. ‘‡’ preceding holotype specimen information indicates that type not from the London Clay Formation. Dates of publication in Agassiz (1833–44) follow Brown (in Woodward & Sherborn 1890). 187 Downloaded from http://sp.lyellcollection.org/ by guest on September 24, 2021

188 M. FRIEDMAN ET AL.

(1966, p. 1) was similarly impressed by the preser- are not soluble in acid and, perhaps most signifi- vation of material from the London Clay Formation. cantly, preserves positional information about There are marine faunas with three-dimensional bones not firmly sutured to other parts of the skele- preservation of similar age to both the Chalk (e.g. ton. This is particularly relevant for interpreting Greenhorn Formation and Turonian Car- anatomy of the gill skeleton and associated bones, lile Shale of , USA: Teller-Marshall & Bar- a key source of morphological characters for infer- dack 1978; Fielitz & Shimada 1999; Turonian ring relationships among extant fishes (e.g. McAllis- Teplice Formation of the Czech Republic: Ekrt ter 1968; Nelson 1969; Rosen 1973; Wiley & et al. 2008; Turonian of Goulmima, Morocco: Johnson 2010). Bones of the branchial arches are Cavin et al. 2010; Campanian Ripley Formation of rarely studied in detail or even observed in fossil tel- , USA: Schein et al. 2013; Maastrichtian eosts (but see Taverne & Nolf 2010), significantly Ciply–Malogne Phosphatic Chalk Formation of limiting the ability of palaeontologists to place fos- Belgium and Formation of the Nether- sils relative to modern species using existing sys- lands; Friedman 2012) and London Clay (e.g. tematic frameworks. –Ypresian of NW Peru: Friedman & Three-dimensional specimens provide more than Johnson 2005; Ypresian Flanders Clay of Belgium: just additional systematic characters unavailable in Casier 1946; ‘cementstone’ layers within the Ypre- flattened fossils. Uncrushed skulls permit applica- sian Fur Formation of Denmark: Bonde et al. 2008; tion of a broader range of ecomorphological mea- Ypresian–Lutetian Lillebælt Clay of Denmark: surements developed for modern teleost fishes Bonde et al. 2008; Lutetian of Mali; uncatalogued (e.g. suction index: Wainwright et al. 2007). This material at NHMUK; Lutetian Lede and Wemmel expands upon the range of functionally relevant sands of Belgium: Storms 1896; Taverne & Nolf metrics that can be explored in fossil fishes, which 2010; late Eocene of Florida, USA: Swift & Ell- have largely been limited to simple linear measure- wood 1972), but none of these localities have yet ments taken from flattened specimens (Bellwood yielded comparable taxonomic richness to their 2003; Bellwood & Hoey 2004; Friedman 2009; British counterparts. In contrast, the Chalk Group Goatley et al. 2010; Bellwood et al. 2014). An and London Clay Formation yield a diversity of acti- expanded repertoire of ecological measurements nopterygian species on a par with all but the most for extinct fishes provides new avenues for explor- spectacular coeval Lagersta¨tten that bear compres- ing ideas about major patterns of functional diver- sion fossils (Fig. 7). sification and change during the and Three-dimensionally preserved specimens from (Vermeij 1977; Friedman 2010; Friedman these British sites provide an important complement & Sallan 2012; see below). to flattened material from these contemporary sites. Many fishes from the Chalk or London Clay are con- Documenting a key evolutionary interval generic with, or otherwise closely related to, taxa represented elsewhere by complete – but highly A major shift in the composition of actinopterygian compressed – specimens. These contrasting tapho- faunas during the Cretaceous and early Palaeogene nomic styles yield complementary morphological has been apparent since Agassiz (1843–44), and information, permitting better constraints on anat- attracted the attention of Darwin (1859, p. 305) in omy for those groups represented in deposits with his treatment of perceived inadequacies of the fossil different preservational modes. Flattened – but record. Smith Woodward (1901, 1902, 1912a, 1942) essentially complete – fossils deliver information reviewed this transition, with a particular emphasis about the postcranial skeleton. When transfer- on the origin of many modern teleost lineages in prepared, such specimens yield details of external the Cretaceous and the appearance of numerous anatomy in three dimensions, but are unlikely to anatomically and ecologically divergent acantho- yield many details about systematically relevant morph clades by the Eocene. The contrast between structures like the gill skeleton and braincase that the ‘very little specialised’ (Woodward 1901, p. v) are largely obscured by other skull bones. Fine acanthomorphs of the Late Cretaceous and the strik- external detail of the head is readily apparent in ing modernity of Palaeogene species, which show many specimens from the English Chalk and Lon- differences ‘of quite a minor character’ (Woodward don Clay, with internal details historically obtained 1901, p. v) relative to extant forms, was obvious to for the former through acid preparation (e.g. Patter- Smith Woodward. He lamented the lack of infor- son 1964; Goody 1969; Forey 2004). Computed mation provided by fossils on this transition and tomography offers a virtual means of obtaining drew parallels to the roughly contemporary radiation these internal details from three-dimensional mate- of placental mammals in terrestrial settings: ‘Palae- rial (e.g. Beckett & Friedman 2015). This approach ontology has, indeed, hitherto revealed as little con- is non-destructive, can also be applied to specimens cerning the origin of the dominant Tertiary fishes as from the London Clay encased in concretions that of the Tertiary mammals’ (Woodward 1901, p. v). Downloaded from http://sp.lyellcollection.org/ by guest on September 24, 2021

ENGLISH CHALK AND LONDON CLAY FISHES 189

Fig. 7. The phylogenetic, geological, and palaeogeographical context of the bony fish faunas of the Chalk Group and London Clay Formation. (a) Phylogenetic distribution of acanthomorph lineages known from body fossils from the Chalk Group and London Clay Formation. Phylogeny adapted from Near et al. (2013). Coloured circles indicate approximate duration of these deposits. (b) Richness and composition of the Chalk and London clay bony fish faunas in comparison to other coeval Lagersta¨tten. Pie charts indicate taxonomic breakdown of sites at the species level, with the area of discs scaled proportional to richness. Faunas are: 1, Hakel, Lebanon; 2, Hadjula, Lebanon; 3, Namoura, Lebanon; 4, Komen, Slovenia; 5, , USA; 6, Sahel Alma, Lebanon; 7, Westphalia, Germany; 8, Nardo`, Italy; 9, Danatinsk Suite, Turkmenistan; 10, Fur Formation, Denmark; 11, Bolca (Pesciara and Monte Postale), Italy. Coloured bands indicate duration of the Chalk Group and London Clay Formation. Data on faunal composition compiled from: Siegfried (1954), Bannikov & Parin (1996), Bonde (1997), Forey et al. (2003), Shimada & Fielitz (2006), Belmonte (2014), Carnevale et al. (2014). Palaeogeographical reconstructions by Ron Blakey, Colorado Plateau Geosystems, Arizona USA. Downloaded from http://sp.lyellcollection.org/ by guest on September 24, 2021

190 M. FRIEDMAN ET AL.

Smith Woodward was uncertain as to the mechanism study of material from the English Chalk and Lon- responsible for this apparently sudden increase in don Clay has considerable potential to contribute diversity, and argued for both elevated rates of evolu- to both. With respect to palaeontological studies, tionary change (1901, p. xii) and, like Darwin before the three-dimensional preservation of specimens him, imperfections in the geological record, suggest- from these sites could aid in better constraining ing that ‘many unknown Acanthopterygii ...became functional inferences drawn from fishes of Late Cre- rapidly differentiated into various families in seas of taceous and early Palaeogene age, and thus provid- which the sediments still remain undiscovered or ing a critical test of hypotheses that suggest that unexplored’ (1901, p. xii). ecological release following the end-Cretaceous The picture provided of the evolutionary history extinction might have been an important factor in of acanthomorphs is much the same today, despite contributing to diversification in at least some acan- more than a century of further collection and study: thomorph groups (Cavin 2001; Friedman 2009, the oldest acanthomorphs are still – as in Smith 2010; Friedman & Sallan 2012; Miya et al. 2013; Woodward’s or Agassiz’s days – from the earliest Sibert & Norris 2015). Fossils from the Chalk and Late Cretaceous deposits, including the Chalk especially London Clay are also likely to contribute Group; a handful of percomorph taxa are known to molecular studies in the form of palaeontological from the close of the Late Cretaceous; and units of calibrations for estimating evolutionary timescales. early Eocene ages such as the London Clay Forma- As a source of fossil calibrations for acanthomorph tion yield acanthomorph assemblages of a strikingly phylogeny, Bolca remains a critical resource owing modern character. As bookends to this diversifi- to a combination of its age, rich fauna and active cation event, the Chalk Group and London Clay For- study that has placed many extinct species precisely mation represent key resources for investigating relative to their living relatives. In fact, fossils from the evolutionary origin of a substantial fraction of Bolca represent nearly a quarter (nine of 37) of the modern vertebrate diversity. Compositionally, fau- calibrations applied by Near et al. (2013) in their nas are broadly similar to ones of comparable age study of acanthomorph divergence times and diver- that preserve fishes as highly compressed speci- sification. The London Clay fauna is older than that mens. The bony fish fauna of the Chalk is dominated of Bolca, but the placement of many of its acantho- by non-acanthomorphs, and includes late-surviving morphs is poorly constrained despite excellent pres- marine holosteans and representatives of numerous ervation. Further study of London Clay material is lineages of stem teleosts (e.g. Pycnodontiformes, likely to result in more confident systematic inter- Pachycormiformes, Aspidorynchiformes, Cross- pretations, which in turn would yield new calibra- ognathiformes, Ichthyodectiformes), along with a tions for acanthomorph phylogeny. great diversity of pelagic Aulopiformes. To extend Patterson’s (1993a, p. 33) mammalian analogy, the spiny-rayed component of the Chalk fauna is Conclusions and a prospectus for dominated by acanthomorph ‘monotremes’: deep, future research relatively species-poor and often ecologically mar- ginalized branches within the modern radiation The significance of the fish faunas of the Chalk and (e.g. non-percomorph groups like Polymixiiformes, London Clay has been apparent to palaeoichthyo- Lampridiformes, Beryciformes; Fig. 7). The compo- logists since the days of Smith Woodward and sitional shift apparent in early Palaeogene deposits even earlier. Despite their significance in capturing like the London Clay reflects the decimation exceptionally preserved ‘snapshots’ of Late Creta- of stem teleost groups at the end of the Cretaceous ceous and early Palaeogene marine assemblages, associated with the appearance of numerous per- they have received comparatively little attention in comorph lineages (Cavin 2001; Friedman 2009; recent years. Numerous aspects of the Chalk and Friedman & Sallan 2012; Guinot & Cavin 2015). London Clay actinopterygian faunas require further In contrast to the Chalk and other Cretaceous investigation, but we identify a few key topics here. assemblages, the London Clay fauna is dominated First, better stratigraphic constraints are required by percomorphs – the ‘placentals’ of acantho- for specimens, particularly for those from the long- morphs – and includes some of the earliest represen- ranging Chalk Group. Because future finds in the tatives of a host of trophically and structurally Chalk are unlikely, the most practical approach diverse modern groups, ranging from tunas to puffer- would entail sampling of matrix associated with fishes to swordfishes. individual specimens in museum collections in The diversification of acanthomorphs during this order to provide microfossils for biostratigraphic interval has been investigated from palaeontological analysis. (Patterson 1993a, b; Friedman 2010; Friedman & Second, future work should strive to provide Sallan 2012) and, increasingly, molecular perspec- more robust placements of fossil specimens relative tives (Near et al. 2013; Price et al. 2014). Renewed to modern lineages. This is most pressing in the case Downloaded from http://sp.lyellcollection.org/ by guest on September 24, 2021

ENGLISH CHALK AND LONDON CLAY FISHES 191 of the London Clay, which yields taxa that probably two archives are largely treated independently of nest within groups with considerable modern diver- one another (Nolf 1985, 1995; Patterson 1993b; sity. Explicit phylogenetic interpretations of fos- but see Storms 1896; Schwarzhans 2007, 2014; Tav- sils will yield not only an improved picture of the erne & Nolf 2010). As Patterson (1977) has noted, London Clay fauna itself, but also new potential cal- otoliths lend a density to the teleost fossil record ibrations for molecular clock analysis. This is par- afforded to that of mammals by their teeth. There ticularly well illustrated by the diverse scombroids will obviously be limits to any such effort, not of the London Clay (Casier 1966; Monsch 2005), least of which is the considerable imbalance none of which have ever been formally placed between research effort on Mesozoic and Cenozoic within a published phylogeny. Further palaeonto- otoliths. The study of mammalian evolution has logical constraints would particularly benefit study benefited enormously from the synthesis of records of this clade, with current estimates of the evolution- of body fossils and isolated teeth; three-dimensional ary timescale for tunas, mackerels and their kin dif- teleost skulls might permit greater integration of fering by tens of millions of years (Miya et al. 2013; currently separate palaeontological records of this Near et al. 2013; Santini et al. 2013). Although the major vertebrate radiation. London Clay probably offers the greatest scope for We thank Mike Smith for his energy in organizing this the reinterpretation of existing fossils, restudy of tribute to Arthur Smith Woodward and his work; Emma specific material from the Chalk might provide Bernard (NHMUK), John Cooper (BMB), Annelise Folie critical new morphological data bearing on regions (IRSNB) and Paula Gentil (Hull and East Riding Museum) of the teleost tree where patterns of relationships for access to specimens or information on material in remain unstable despite continued investigation their care; and Dan Sykes (NHMUK), and Farah Ahmed using molecular and morphological data from living (NHMUK) and Rebecca Summerfield (NHMUK) for assis- species (e.g. non-anguilliform and non-elopiform tance with CT scanning. Our studies on London Clay fishes Elopomorphs; Dornburg et al. 2015). benefited from interactions with Fred Clouter, Tony Mitch- Third, the London Clay might present a unique ell, David Rayner and Martin Rayner. Peter Forey and Lio- nel Cavin provided comments that substantially improved system for beginning to reconcile the otolith and an earlier version of this contribution. This research was body fossil records of teleosts. While the preserva- supported by a Leverhulme Research Project Grant tional bias against aragonite means that even less (RPG-2012-658) and Leverhulme Prize (PLP-2012-130), stable vaterite otoliths are unlikely to be preserved both to MF, and a NERC studentship to HB through the within fishes from the Chalk, otoliths are known to University of Oxford Environmental Research Doctoral be found intact within skulls from the London Training Partnership (NE/L0021612/1). Clay (Casier 1966, p. 327; Stinton in Casier 1966, p. 456), as well as fish remains from other argilla- ceous deposits of comparable age (Lillebælt Clay; References Schwarzhans 2007). Such a work will not be without Agassiz, J. L. R. 1833–44. Recherches sur les Poissons its challenges. The striking discrepancy between the Fossiles, 5 vols. Petitpierre, Neuchaˆtel. London Clay fauna as recorded by body fossils and Agassiz, J. L. R. 1859. Essay on Classification. Longman, otoliths has long been apparent (White in Casier Brown, Green, Longmans, & Roberts, London. 1966, p. vi). Despite clear issues concerning the Agassiz, L. 1845. Report on the fossil fishes of the London interpretation of some London Clay fishes discussed Clay. Report of the British Association, London, 1844, above, the discrepancy is sufficiently large that 279–310. errors of identification are unlikely to be the sole Aldiss, D. T. 2012. The Stratigraphical Framework for cause. 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