Dineley, D. & Metcalf, S. GCR Editor: D. Palmer. 1999. Fossil Fishes of Great Britain. Geological Conservation Review Series No. 16. JNCC, Peterborough, ISBN 1 86107 470 0. The original source material for these web pages has been made available by the JNCC under the Open Government Licence 3.0. Full details in the JNCC Open Data Policy
Southerham (Machine Bottom Pit)
Highlights
The Machine Bottom Pit at Southerham in Sussex is a site of special historic interest, being the locality from which most of the large 19th century collections of Chalk fossil fishes were made by famous gentleman scholars, such as Gideon Mantell and Frank Dixon. The collections include 28 type specimens (many more or less complete specimens) and still form the basis for much Upper Cretaceous fish taxonomy.
Introduction
The chalk pits around the East Sussex town of Lewes
Most of the complete fish specimens described from the English Chalk were recovered in the 19th century and nowhere was fossil collecting more fashionable than in the Cretaceous succession of the South Downs. Both Gideon Mantell and Frank Dixon made extensive collections in the Chalk of Sussex, and many of their fish specimens are from the Lower and Middle Chalk around Lewes (Mantell, 1822; Dixon, 1850; Jukes-Browne and Hill, 1903). The Lewes fish collections have been the subject of much scientific description, including the work of Agassiz in the early 19th century and Smith Woodward's monographic work on Chalk fishes (1902–1912). They are the source of at least 28 type specimens.
The Lower and Middle Chalk succession in the Machine Bottom Pit was described by Barrois (1876), Jukes-Browne and Hill (1903), White (1926), Gastcr (1951), Kennedy (1969) and Lake et al. (1987). The pit is now disused and largely overgrown, and the higher beds of the section (Plenus Marls and Melbourn Rock), exposed high up in the north-east face, are inaccessible (Lake et al., 1987). Yet re-excavation of the lower beds is possible and could yield more specimens.
Description
The confusion over the naming of the Machine Bottom Pit at Southerham arose because prior to White's recording of the section there in 1926, the pit was known as the 'Grey Pit'. When White (1926) made his observations the pit was disused. Later the pit was reopened and greatly enlarged, until final closure in the 1970s and was given its new name (Lake et al., 1987). The new Southerham Grey Pit (q.v.) is situated about 200 m to the west of the older pit.
The Machine Bottom Pit exposes approximately 30 m of Lower Chalk and the lowest beds of the Middle Chalk
Thickness (m) | |
Middle Chalk | |
Ranscombe Griotte Chalk Member Melbourn Rock: creamish white, hard, nodular chalk with marl partings; some of the nodular beds are markedly iron-stained | 0.9 |
Lower Chalk | |
Plenus Marls: pale grey, very marly chalkwith slight greenish tinge; two well-marked hard beds up to 0.3 m thick; sharp erosional base | 4.8–5.7 |
Grey Chalk: greyish white chalk, massive, thickly bedded, with thin marly partings | 29.2 |
Chalk Marl: rhythms of up to 0.9 m thick, consisting of medium grey marly chalk grading up into pale fawn hard chalk commonly with large uncrushed ammonites; the hard beds generally have sharp tops, commonly burrowed; scattered pyrite nodules | 9.1 |
The Lewes district is particularly important for Chalk lithostratigraphy, providing stratotypes for many members, beds and marker horizons (Mortimore, 1983). The relationship between the formal lithostratigraphical nomenclature, informal terminology and the biostratigraphy of the Lewes Chalk succession is given in
The Chalk Marl (M. dixoni Zone) at the Machine Bottom Pit is not as clearly bedded as the same unit in other sections around Southerham (e.g. Grey Pit (q.v.; Lake et al., 1987). Locally, the harder limestone beds are extremely rich in invertebrate fossils, with the ammonite Acanthoceras particularly common. The overlying Grey Chalk is thickly bedded, although again individual beds are not easily defined in the section
The rich collections made at the Southerham chalk pits in the 19th century are preserved in the BMB, NHM and CAMSM. However, the earlier workers generally labelled their material as from the 'Chalk: Southerham', 'Chalk: Lewes' or in some cases 'Chalk: Sussex'. Similar labelling is found in the large collections made by Mantell and Dixon, which were described by Agassiz and include some of the type specimens described below. Mantell's list (1822) contains a few specimens which evidently came from the Chalk Marl (M. dixoni Zone) and also from upper whiter part of the Grey Chalk (M. geslinianum Zone) of Southerham, which has led most subsequent authors (e.g. Jukes-Browne and Hill, 1903; Lake et al., 1987) to record Southerham and Lewes fish specimens as from the older S. varians and H. subglobosus Zones of the Lower Chalk. Mantell's and Dixon's specimens may have also come from the Middle Chalk, as suggested by Woodward (1902–1912). This is borne out by the listing of species recorded by the Geological Survey for the Middle Chalk (mainly T. lata Zone) of the Lewes region (Jukes-Browne and Hill, 1903).
Fauna
The following list is from Woodward (1902–1912) and includes specimens recorded as from 'Chalk: Lewes' and 'Chalk: Southerham'. The collections are considered to have been made from the Machine Bottom Pit at Southerham (Jukes-Browne and Hill, 1903; Lake et al., 1987; D. Ward, pers. comm., 1994).
Chondrichthyes: Elasmobranchii: Euselachii: Hybodontoidea
Polyacrodus (Acrodus) illingworthi (Dixon, 1850)
Ptychodus decurrens Agassiz, 1835–1839
P. latissimus Agassiz, 1835–1843
P. mammillaris Agassiz, 1835–1839
P. polygyrus Agassiz, 1835–1839
P. polygyrus var. marginalis Agassiz, 1835–1839
Chondrichthyes: Elasmobranchii: Neoselachii: Squalomorphii
Hexanchus (Notidanus) microdon (Agassiz, 1843)
Chondrichthyes: Elasmobranchii: Neoselachii: Galeomorphii
Cretolamna (Lamna) appendiculata (Agassiz, 1843)
C. (Isurus) mantelli (Agassiz, 1843)
C. Woodwardi (Herman, 1975)
'Lamna'spp.
Plicatolamna (Oxyrhina) crassidens (Dixon, 1850)
P. (Lamna) sulcata (Gcinitz, 1843)
Scapanorhynchus rhaphiodon (Agassiz, 1844)
S. subulatus (Agassiz, 1843)
Squalicorax (Corax) falcatus (Agassiz, 1843)
S. (C) kaupi (Agassiz, 1843)
Chondrichthyes: Holocephali: Chimaeriformes
Edaphodon (Ischyodus) agassizi (Buckland, 1835–1836)
E. (I.) mantelli (Buckland, 1835–1836)
E. (Psittacodon) sedgwicki (Agassiz, 1843)
Elasmodectes willetti Newton, 1878
Osteichthyes: Actinopterygii: Neopterygii: Halecostomi
Acrotemnus faba Agassiz, 1837–1844
Anomoedus (Gyrodus) angustus (Agassiz, 1837–1844)
Coelodus (Pycnodus) parallelus (Dixon, 1850)
Phacodus punctatus Dixon, 1850
Polygyrodus (Gyrodus) cretaceus (Agassiz, 1839–1844)
Polygyrodus bennetti White, 1927
Osteichthyes: Actinopterygii: Neopterygii: Halecomorphi
Lophiostomus dixoni Egerton, 1852
Tomognathus mordax Dixon, 1850
Osteichthyes: Actinopterygii: Neopterygii: Teleostei
Belonostomus cinctus Agassiz, 1837–1844
Protosphyraena ferox Leidy, 1857
P. minor (Agassiz, 1837–1844)
Ichthyodectes (Hypsodon) minor (Egerton, 1850)
Osmeroides lewesiensis (Mantell, 1822)
O. levis Woodward, 1901
Pachyrhizodus (Acrodontosaurus) gardneri (Mason, 1869) ( = P. basalis Forey, 1977)
P. (Thrissopater) megalops (Woodward, 1901)
Plethodus expansus Dixon, 1850
P. pentagon Woodward, 1902–1911
Protelops anglicus Woodward, 1888
Xiphactinus (Portheus) mantelli (Newton, 1877)
Osteichthyes: Actinopterygii: Neopterygii: Euteleostei
Acrognathus hoops Agassiz, 1844
Aulolepis typus Agassiz, 1837–1844
Apateodus striatus Woodward, 1901
Platycormus (Berycopsis) elegans Dixon, 1850
Cimolichthys levesiensis Leidy, 1857
Ctenothrissa (Beryx) microcephala (Agassiz, 1838)
C. (B) radians (Agassiz, 1835–1838)
Dercetis elongatus Agassiz, 1835–1844
Enchodus (Esox) lewesiensis (Mantell, 1822)
Eurypholis pulchellus Woodward, 1901
Halec (Pomognathus) eupterygius (Dixon, 1850)
Hoplopteryx (Beryx) lewesiensis (Mantell, 1822)
H. (B.) superbus (Dixon, 1850)
H. simus Woodward, 1902–1911
Osteichthyes: Sarcopterygii: Actinistia
Macropoma mantelli Agassiz, 1835–1844
Macropoma sp. (coprolites)
Interpretation
The Southerham Chalk Pits reveal a flourishing assemblage of both chondrichthyans and osteichthyes, almost all of which appear to have been predatory or durophagous. The numbers of fossils and taxa suggest relatively large populations of these vertebrates near the top of broad trophic pyramids in the Chalk sea. The environment was that of a relatively shallow warm to tropical water, of high organic productivity and with little clastic input. Terrigenous sedimentation was for the most part negligible, except in Chalk Marl time. Coccolith production was high throughout. These conditions promoted a generous regime of producer organisms upon which the vertebrate populations were founded. The chondrichthyes included both benthonic and free-swimming nektonic kinds, both large and small. The feeding activities of Ptychodus are thought to be responsible for the widespread and thorough fragmentation of the ubiquitous inoceramid (bivalve) shells (Kennedy, in McKerrow, 1978). Other shark-like forms were adapted for the crushing of nektonic or planktonic hard-shelled invertebrates, which were abundant in numbers and kinds. Chimaeroids were common but the hybodont sharks were in decline. The latter are said to be more common in freshwaters during the Cretaceous.
Hybodont sharks are well represented in the Lewes collections by abundant isolated teeth, from which five type specimens were described in the early 19th century. Polyacrodus illingworthi (Dixon, 1850) is known only from the large, low-crowned teeth originally described by Dixon (1850) as from the Jurassic hybodont Acrodus, and have also been assigned to Hybodus (Longbottom and Patterson, 1987) and Synechodus (Woodward, 1902–1912).
The large crushing teeth of the Cretaceous hybodont Ptychodus are fairly common in the Southerham collections (
Neoselachian sharks are also well represented in the Lewes fauna and include both squalomorphs (Hexanchus microdon (Agassiz)) and abundant galeomorphs (Woodward, 1902–1912). WO species of lamniform shark, the cretoxyrhinid Cretolamna appendiculata (Agassiz, 1843), and the anacoracid Squalicorax falcatus (Agassiz, 1843) were named from detached teeth recovered from the Lower Chalk at Southerham. The medium-sized (15–25 mm) piercing teeth of Cretolamna appendiculata are fairly common Chalk fossils (Longbottom and Patterson, 1987). In the past most lamnid teeth found within the Chalk were referred to 'Lamna' appendiculata, but many now have been reassigned to several other species, including C. woodwardi (Herman). Squalicorax is an extinct Cretaceous anacoracid or thresher shark, notable for possessing teeth that bear a distinctly serrated cutting edge (Longbottom and Patterson, 1987). The small teeth (12–15 mm) of S. falcatus are extremely common in the Cenomanian to Campanian Chalk of southern and eastern England (Longbottom and Patterson, 1987).
Other lamnids common in the Southerham galeomorph assemblage are a second species of Squalicorax, S. (C.) kaupi (Agassiz), and several cretoxyrhinids, including Cretoxyrhina mantelli (Agassiz), Plicatolamna crassidens (Dixon) (Cappetta, 1975) and P. sulcata (Geinitz), in association with the mitsukurinids Scapanorhynchus rhapiodon (Agassiz) and S. subulatus (Agassiz) (Woodward, 1902–1912).
Chimaeroids are also common in the fish assemblages from the Machine Bottom Pit and include three type specimens (Newton, 1878; Woodward, 1902–1912). Edaphodon is a typical Cretaceous genus which survived into the Miocene, and is known from isolated teeth and tooth plates in the Chalk at Southerham
The list of bony fishes from Southerham Machine Bottom Pit is as lengthy as that from Burham in Kent, but importantly contains a pycnodont component not known from the Blue Bell Hill faunas. Most of the five species of pyc-nodont from the Chalk are based upon the arrangement of teeth within fragmentary jaws, as the isolated button-like crushing teeth are not taxonomically diagnostic (Longbottom and Patterson, 1987). The best known Cretaceous pycnodonts are Anomoeodus angustus (Agassiz), of which articulated teeth and jaws have been recovered from rocks ranging through the Lower to Upper Chalk successions, and Coelodus parallelus (Dixon) which was described from a splenial dentition in the Southerham collection at BMB (Woodward, 1902–1912). Several Coelodus species are based upon more complete robust skeletal material from the Late Cretaceous of Italy and Austria (Woodward, 1902–1912). Other Chalk pycn-odonts are much less precisely known: the sple-nial dentition of Acrotemnus faba Agassiz found at Lewes by Mantell (Woodward, 1908) is the only specimen of this genus that has bean-shaped smooth teeth with keel-like coronal tips. Phacodus punctatus Dixon is only known from a few Turonian localities in the English Chalk and has ovoid or irregularly rounded teeth with a smooth crown (Woodward, 1908); and Polygyrodus cretaceus (Agassiz), also known only from the Turonian Chalk, is characterized by dentition made up entirely of small circular and conically crowned teeth, with roughly enamelled surfaces (Longbottom and Patterson, 1987). The Southerham pycnodont species P. bennetti White, 1927 was described from an imperfect vomerine dentition.
The halecomorph neopterygian component of the Southerham bony fish fauna is much the same as that from the Blue Bell Hill pits (q.v.) at Burham, and includes the type specimen of the large aspidorhynchid Belonostomus cinctus Agassiz, 1837–1844, rostrum fragments the two species of the swordfish-like pachycormid Protosphyraena, and remains of the rare Tomognathus mordax Dixon (Woodward, 1908;
The teleosts were conspicuous members of the osteichthyan fauna; some were also very large, active predatory fishes. Deep-bodied and slender fusiform types existed, and the eel-like Dercetis, with its wide gape and formidable array of small teeth, indicate a range of lifestyles comparable to that in many Recent shallow tropical seas. There were also numerous small fishes such as Acrognathus, and deep-bodied larger forms like Berycopsis that may have swum in schools.
Nine species of teleosts are present in the Lewes Chalk assemblage, and Southerham is the type locality for six (Woodward, 1902–1912). The fauna includes representatives of the Elopiformes, the Icththyodectiformes, an anguilliform and the enigmatic plethodonts.
Elopiforms are represented by the type specimen of the small albuloid Osmeroides lewesiensis (Mantell). Another species, O. levis Woodward, is also found in the Southerham fauna and both were described in detail by Forey (1973a). The third 'pseudotarpon' elopiform Protelops anglicus Woodward is known only from fragmentary jaws lined with long, slender teeth arranged in two or more series on the border of the mouth or fused to the palate (Woodward, 1908, 1902–1912).
The large predaceous pachyrhizodontid Pachyrhizodus, with its characteristic dentition (Woodward, 1902–1912; Longbottom and Patterson, 1987; see Blue Bell Hill report), has also been recovered from the Lower and Middle Chalk succession in the Machine Bottom Pit. Two species, P. gardneri (Mason) and the type specimen of P. megalops (Woodward) (an imperfect head), are recorded in the collections (Forey, 1977). It is now known that Thrissopater is a junior synonym of Pachyrhizodus (Frickhinger, 1991;
The large carnivorous Ichthyodectiformes (Nelson, 1973) are represented by the type specimens of two species, Ichthyodectes minor (Egerton) and Xiphactinus mantelli (Newton), in the Southerham fauna (Woodward, 1902–1912). Fragmentary jaws of I. minor are known from several Thronian Chalk localities and are characterized by the sharp hollow teeth set into deep sockets (Woodward, 1902–1912; Bardack, 1965; Nelson, 1973; see Blue Bell Hill report). Better-preserved specimens of the genus were recovered from the Upper Cretaceous of Kansas and named Portheus by Cope (1872) and Xiphactinus by Leidy (1857). Newton (1877) was able to reassign much of the British material to X. mantelli, a large form probably attaining lengths of up to 4–5 m (Woodward, 1902–1912). The fossil material includes fragmentary jaws and large isolated bones, such as fin rays and vertebrae.
The plethodont Plethodus is the only genus of this enigmatic group recorded from the Chalk (Longbottom and Patterson, 1987; see Blue Bell Hill report). The typical crushing dentition is clearly seen in the type species, P. expansus Dixon, described from detached dental plates from the Turonian Chalk of Lewes. These plates are up to 0.1 m in length, and the species may have reached 1 m or so in length. The pentagonal dental plates of P. pentagon Woodward have also been recorded in the Machine Bottom Pit.
The higher teleost or euteleost fauna
Among the neoteleosts represented at Southerham are the type specimens of three extinct Cretaceous ctenothrissoid fishes, Aulolepis typus Agassiz, Ctenothrissa microcephala (Agassiz) and C. radians (Agassiz). These genera are also found at Blue Bell Hill (q.v.). Both species of Ctenothrissa were named from fairly well-preserved fish from the Mantel Collection of Lower-Middle Chalk fishes from Southerham (Woodward, 1902–1912). Aulolepis is a monospecific genus restricted to the English Chalk, which differs from Ctenothrissa in the morphology of the head and jaws, and in possessing non-pectinate scales (Patterson, 1964, 1968;
The common Chalk euteleost predatory fishes, the 'enchodonts' (aulopiforms; Goody, 1969; Rosen, 1973; Longbottom and Patterson, 1987), are represented in the Southerham fauna by six species (Woodward, 1902–1912), and like the aulopiforms at Blue Bell Hill (q.v), include representatives of three groups: the enchodon-toids, the cimolichthyids and halecoids. The eel-like dercetid Dercetis is also found at Southerham (Woodward, 1902–1912). The enchodont Apateodus striatus Woodward, 1901 (
The sardinoidid myctophiform Acrognathus boops Agassiz is a small fish, attaining a size of a little over 0.1 m, with thick uniform cycloid scales (Rosen, 1973). Acrognathus possessed an elongated trunk, and a large head with huge orbits. The small delicate jaws were lined with minute teeth (Woodward, 1902–1912). Although A. boops is the type species of the genus, the material recovered from the Chalk at Lewes and pits in Surrey is fragmentary (Woodward, 1902–1912). The genus is better known from a second species, A. libanicus Woodward, from the Upper Cretaceous of Lebanon (Rosen, 1973).
Acanthomorphs in the Chalk fauna of Lewes include the polymixiid Berycopsis elegans (Dixon), the trachichthyoids Hoplopteryx lewesiensis (Mantell), H. superbus (Dixon) and H. simus Woodward, 1902, and the holocentroid Caproberyx. Hoplopteryx was a deep-bodied fish that is characterized by expanded sensory canals upon the frontal region of the skull, an upturned mouth lined with tiny teeth and large eye (Woodward, 1902–1912;
The Southerham fauna includes a nearly complete type specimen of the Cretaceous coelacanth Macropoma mantelli Agassiz (
Fossil reptiles recovered from the Lower and Middle Chalk succession of the Machine Bottom Pit (Jukes-Brown and Hill, 1903, pp. 46–58, 404) include turtles, plesiosaurs and mosasaurs, as well as the piscivorous pterosaurs. Predatory tetrapod communities, dependent upon fish in the Late Cretaceous seas, are known from many parts of the world. Those in southern England would have apparently had an abundant food supply, to judge from the record of fishes in the Chalk here. Although the quarry is not an SSSI for these marine reptiles, the fauna is listed in the review of fossil reptile sites (Benton and Spencer, 1995). However, the authors mistakenly ascribe the finds to the Grey Pit at Southerham (q.v.), because of the confusion over names.
Comparison with other localities
The faunas are broadly similar in composition and preservation to those described from Burham in Kent (q.v.). However, in detail there are minor differences, such as in the absence of pycnodonts in the Burham assemblages, but overall the two faunas complement one another.
Conclusion
The Machine Bottom Pit at Southerham has yielded one of the most complete Lower and Middle Chalk fish faunas, including 27 type specimens, which provide its conservation value. Eleven of these are amongst the earliest neoteleosts, the largest living group of bony fishes, and the Southerham specimens are important in defining basal characteristics of the group.