A new fossil bramoides from the eocene london clay re aligned with the modern genus gasterochisma
The London Clay Formation is a marine geological formation of Ypresian (Lower Eocene Epoch, c. 56–49 Ma) age which crops out in the southeast of England. The London Clay is well known for its fossil content. The fossils from the Lower Eocene indicate a moderately warm climate, the tropical or subtropical flora. Though sea levels changed during the deposition of the Clay, the habitat was generally a lush forest – perhaps like in Indonesia or East Africa today – bordering a warm, shallow ocean.
The London Clay is a stiff bluish clay which becomes brown when weathered. Nodular lumps of pyrite and crystals of selenite (sometimes called "waterstones") frequently exist in the clay, and large septarian concretions are also common. These have been used in the past for making cement. They were once dug for this purpose at Sheppey, near Sittingbourne, and at Harwich, and also dredged, off the Hampshire coast. The clay is still used commercially for making bricks, tiles, and coarse pottery in places such as Michelmersh in Hampshire. It is infertile for gardens and crops.
Distribution and geology
The London Clay is well developed in the London Basin, where it thins westwards from around 150 metres (492 feet) in Essex and north Kent to around 4.6 metres (15 feet) in Wiltshire. though it is not frequently exposed as it is to a great extent covered by more recent Neogene sediments and Pleistocene gravel deposits. One location of particular interest is Oxshott Heath, where the overlying sand and the London Clay layers are exposed as a sand escarpment, rising approximately 25 metres (82 feet). This supported a thriving brick industry in the area until the 1960s. The London Clay is also well developed in the Hampshire Basin, where an exposure 91 metres (299 ft) thick occurs at Whitecliff Bay on the Isle of Wight and around 101 metres (331 ft) is spread along 6 kilometres (4 miles) of foreshore at Bognor Regis, West Sussex.
The clay was deposited in a sea up to 200 metres (660 ft) deep at the eastern end. Up to five cycles of deposition (representing transgression followed by shallowing of the sea) have been found, most markedly at the shallower, western end. Each cycle begins with coarser material (sometimes including rounded flint pebbles), followed by clay which becomes increasingly sandy. The final cycle ends with the Claygate Beds.
Claygate Beds
The youngest part of the London Clay, known as the Claygate Beds or Claygate Member forms a transition between the clay and the sandier Bagshot Beds above. This is shown separately on many geological maps, and often caps hills. It is up to 15 metres (49 ft) thick at Claygate, Surrey. It is now believed to be diachronous, with the formation at Claygate for example being the same age as the end of the fourth cycle of deposition further east.
Engineering
The presence of a thick layer of London Clay underneath London itself, providing a soft yet stable environment for tunnelling, was instrumental in the early development of the London Underground, although this is also the reason why London has no true skyscraper buildings, at least to the same degree as many other cities throughout the world. Erecting tall buildings in London requires very deep, large and costly piled foundations.
London Clay is highly susceptible to volumetric changes depending upon its moisture content. During exceptionally dry periods or where the moisture is extracted by tree root activity, the clay can become desiccated and shrink in volume, and conversely swell again when the moisture content is restored. This can lead to many problems near the ground surface, including structural movement and fracturing of buildings, fractured sewers and service pipes/ducts and uneven and damaged road surfaces and pavings. Such damage is recognised to be covered by the interpretation of subsidence in buildings insurance policies, and the periods of dry weather in 1976/77 and 1988/92, in particular, led to a host of insurance claims. As a result, many insurance companies have now increased the cost of premiums for buildings located in the most susceptible areas where damage occurred, where the clay is close to the surface.
Tunnels in London Clay
The London Clay is an ideal medium for driving tunnels, which is why the London Tube railway network expanded quickly north of the Thames, but south of the Thames the stratum at tube level is water-bearing sand and gravel (not good for tunnelling) with London Clay below, which is why there are few tube tunnels there. London Clay has a stand-up time long enough to enable support to be installed without urgency. It is also almost waterproof, resulting in virtually no seepage of groundwater into the tunnel. It is over-consolidated, which means that it is under pressure, and expands upon excavation, thus gradually loading the support, i.e. it is not necessary to stress the support against the ground.
Agriculture
Due to its impermeability especially when exposed by ploughing, London clay does not make good agricultural soil. In Middlesex, it has historically been called "ploughing up poison."
Plant fossils, especially seeds and fruits, are found in abundance and have been collected from the London Clay for almost 300 years. Some 350 named species of plant have been found, making the London Clay flora one of the world's most diverse for fossil seeds and fruits. The flora includes plant types found today in tropical forests of Asia and demonstrates the much warmer climate of the Eocene epoch, with plants such as Nypa (Nipah palms) and other palms being frequently encountered. The following plants list is incomplete and is based on the research by Marjorie Chandler. and research works done by the paleobotanist Steven R. Manchester and by professor of plant palaeobiology Margaret Collinson.
Pteridophytes
Pteridaceae
Acrostichum – mangrove fern
Salviniaceae – water ferns
Azolla
Gymnosperms
Araucariaceae – monkey puzzle, bunya pine, and Norfolk pine
†Araucarites spp.
†Doliostrobus stenbergi - fossils in the past named †Araucarites are said to be of this genus, it is now placed in its own family, †Doliostrobaceae.
Pinaceae
Pinus spp.
Taxaceae
?Taxaceae indet.
Cephalotaxus bowerbanki
Cupressaceae
†Cupressinites spp.
Libocedrus adpressa
†Quasisequoia sp.
Sequoia couttsiae
Podocarpaceae
Podocarpus argillaelondinensis?
Angiosperms
Arecaceae – palms
†Caryotispermum cantiense
Livistona minima
Nypa burtini (syn. Nipa burtini)
Oncosperma anglica
†Palmospermum bracknellense
†Palmospermum cooperi
†Palmospermum davisi
†Palmospermum elegans
†Palmospermum jenkinsi
†Palmospermum minutum
†Palmospermum ornatum
†Palmospermum ovale
†Palmospermum subglobulare
Sabal grandisperma
Cyclanthaceae
Cyclanthus lakensis
Cyperaceae - sedges
†Caricoidea obovata
†Polycarpella caespitosa
Posidoniaceae - seagrass family
Posidonia parisiensis
Nymphaeaceae – water lilies
†Protobarclaya eocenica
Magnoliaceae
Liriodendron gardneri - tulip tree
Magnolia angusta
Magnolia crassa
Magnolia davisi
Magnolia lata
Magnolia lobata
Magnolia oblonga
Magnolia pygmaea
Magnolia rugosa
Magnolia subquadrangularis
Magnolia symmetrica
Myristicaceae - nutmeg family
†Myristicacarpum chandlerae
Annonaceae - the custard apple family
†Anonaspermum anonijorme - related to the genera Anonna and Polyalthia, most similar species is Annona muricata
†Anonaspermum cerebellatum - Uvaria, Melodorum and Orophea shows closest resemblance to the fossil
†Anonaspermum commune
†Anonaspermum complanatum - related to Anonna and Melodorum
†Anonaspermum corrugatum - related to the genera Anonna and Melodorum
†Anonaspermum minimum - related to Dasymaschalon clusiflorum
†Anonaspermum obscurum
†Anonaspermum ovale - related to the genera Anonna and Melodorum
†Anonaspermum pulchrum - related to Dasymaschalon clusiflorum
†Anonaspermum punctatum - related to the genera Bocagea, Orophea, Unonopsis, and Guatteria
†Anonaspermum rotundatum - distantly related to Polyauhia
†Anonaspermum rugosum
†Anonaspermum subcompressum
Lauraceae – avocado and cinnamon family
Beilschmiedia bognorensis
Beilschmiedia eocenica
Beilschmiedia gigantea
Beilschmiedia oviformis
Beilschmiedia pyriformis
Cinnamomum grande
Cinnamomum globulare
Cinnamomum oblongum
Cinnamomum ovoideum
†Crowella globosa
Endiandra spp.
†Laurocalyx bowerbanki
†Laurocalyx dubius
†Laurocalyx fibrotorulosus
†Laurocalyx globularis
†Laurocalyx magnus
†Laurocarpum crassum
†Laurocarpum cupuliferum
†Laurocarpum davisi
†Laurocarpum inornatum
†Laurocarpum minimum
†Laurocarpum minutissimum
†Laurocarpum ovoideum
†Laurocarpum paradoxum
†Laurocarpum proteum
†Laurocarpum pyrocarpum
†Laurocarpum sheppeyense
Litsea pyriformis
†Protoaltingia europaea
†Protoravensara sheppeyensis
Dilleniaceae
Hibbertia bognorensis
Tetracera ?cantiensis
Tetracera crofti
Tetracera eocenica
Platanaceae – sycamore or plane tree
†Plataninium decipiens
Proteaceae
Dryandra acutiloba
Betulaceae – birch, alder and hornbeams
Alnus richardsoni (syn. Petrophiloides richardsoni) – an alder
Hamamelidaceae – witch-hazel family
Corylopsis? bognorensis
Corylopsis? latisperma
Corylopsis venablesi
†Protoaltingia europaea
Myricaceae - wax-myrtle, bayberry family
Myrica boveyana
Juglandaceae – walnut, hickory and pecan family
†Juglandicarya bognorensis
†Juglandicarya cooperi
†Juglandicarya depressa - the most common Juglandacarya species in the London Clay
†Juglandicarya lubbocki
†Juglandicarya minuta
†Petrophiloides richardsoni - among the two most abundant Juglandaceae species in the London Clay
†Pterocaryopsis bognorensis - related to the genus Pterocarya
†Menispermoxylon - close to the extant genus Tinomiscium
†Microtinomiscium foveolatum
†Palaeococculus lakensis
†Palaeosinomenium pulchrum
†Tinomiscoidea scaphiformis
Tinospora excavata
†Wardenia davisi
Aristolochiaceae - birthwort family
Aristolochia sp.
Torricelliaceae
Toricellia sp. - very similar to Torricellia bonesii from the Clarno Formation of Oregon
Altingiaceae
Liquidambar palaeocenica - sweetgum
Oleaceae
Fraxinus sp.
Styracaceae
Styrax sp.
Symplocaceae – sapphire-berry, sweet leaf.
Symplocos curvata
Symplocos quadrilocularis
Symplocos trilocularis
Symplocos bognorensis
Ericaceae
Leucopogon quadrilocularis
Pentaphylacaceae
Sapotaceae
†Sapoticarpum rotundatum
†Sapoticarpum latum
†Sapoticarpum duhium
†Sapotispermum sheppeyense - allied to Chrysophyllum and Sideroxylon
Myrtaceae
†Hightea elliptica
†Hightea turgida
†Myrtospermum variabile
†Palaeorhodomyrtus subangulata - allied to Rhodomyrtus
Fabaceae
Dalbergia sp.
†Leguminocarpon nervosum
†Mimosites browniana
Myrsinaceae
Ardisia eocenica
Rhizophoraceae - mangrove
†Palaeobruguiera elongata
†Palaeobruguiera lata
Malpighiaceae
Salicaceae
Oncoba variabilis
† Saxifragispermum spinosissimum
Linaceae
†Decaplatyspermum bowerbanki
†Wetherellia variahilis - related to Hugonia
Nyctaginaceae - the four o'clock family
Pisonia sp. - the birdcatcher tree
Olacaceae
Erythropalum europaeum - Only one extant species of this genus which is a scandent shrubs or liana
Erythropalum jenkinsi
Erythropalum turbinatum
Olax depressa
Boraginaceae
Ehretia clausentia
Ehretia ehretioides
Solanaceae
†Cantisolanum daturoides
Apocynaceae - the dogbane family
†Ochrosella ovalis
†Ochrosoidea sheppeyensis
Burseraceae - the incense tree family
†Bursericarpum aldwickense
†Bursericarpum bognorense
†Bursericarpum ovale
†Bursericarpum venablesi
†Palaeobursera bognorensis
†Protocommiphora europaea
Anacardiaceae
Choerospondias sheppeyensis
Dracontomelon minimum
Dracontomelon subglobosum
Lannea europaea
Lannea jenkinsi
†Lobaticarpum variabile
Odina europaea
Odina jenkinsi
Odina subreniformis
†Pseudosclerocarya subalata
†Spondiaecarpon operculatum - according to several botanists, the pyritized specimens, originally described as Spondiaecarpon operculatum, represent locule casts of Torricellia sp.
Onagraceae
†Palaeeucharidium cellulare - allied to Eucharidium
Lythraceae
†Cranmeria trilocularis
†Minsterocarpum alatum - closely related to the crape myrtle or crepe myrtle genus Lagerstroemia
†Pachyspermum quinqueloculare
†Tamesicarpum polyspermum
Malvaceae
†Cantitilia polysperma
Elaeocarpaceae
†Echinocarpus sheppeyensis
Moraceae – mulberry and fig family
?Morus sp.
Urticaceae
†Urticicarpum scutellum
Euphorbiaceae - the spurge family
†Euphorbiospermum bognorense
†Euphorbiospermum cooperi
†Euphorbiospermum eocenicum
†Euphorbiospermum obliquum
†Euphorbiospermum subglobulare
†Euphorbiospermum subovoideum
†Euphorbiospermum venablesi
†Euphorbiotheca minima
†Euphorbiotheca sheppeyensis
†Lagenoidea bilocularis
†Lagenoidea trilocularis
†Wetherellia variabilis
Cucurbitaceae - the gourd family
†Cucurbitospermum cooperi
†Cucurbitospermum equiaelaterale
†Cucurbitospermum sheppeyense
†Cucurbitospermum triangulare
Vitaceae
Tetrastigma corrugata
Tetrastigma davisi
Tetrastigma elliotti
Tetrastigma sheppeyensis
†Vitacexoylon sp. - close to the extant genus Rhoicissus
Vitis bilobata
Vitis bognorensis
Vitis bracknellensis
Vitis elegans
Vitis longisulcata
Vitis magnisperma
Vitis obovoidea
Vitis platyformis
Vitis pygmaea
Vitis rectisulcata
Vitis semenlabruscoides
Vitis subglobosa
Vitis venablesi
Sapindaceae - soapberry
†Cupanoides grandis
†Cupanoides tumidus
†Palaeallophylus minimus
†Palaeallophyllus ovoideus
†Palaeallophylus rotundatus
†Palaealectryon spirale
†Sapindospermum cooperi
†Sapindospermum davisi
†Sapindospermum grande
†Sapindospermum jenkinsi
†Sapindospermum ovoideum
†Sapindospermum revolutum
†Sapindospermum subovatum
Meliaceae - the mahogany family
Cedrela sp.
†Melicarya variabili
Toona sulcata
Rutaceae - the rue family
†Canticarya gracilis
†Canticarya ovalis
†Canticarya sheppeyensis
†Canticarya ventricosa
†Caxtonia elongata
†Caxtonia glandulosa
†Caxtonia rutacaeformis
†Citrispermum sheppeyense
†Clausenispermum dubium
†Eozanthoxylon glandulosum
Euodia costata
†Rutaspermum bognorense
†Rutaspermum minimum
†Shrubsolea jenkinsi
†Zanthoxylon compression
†Zanthoxylon bognorense
Celastraceae - the staff vine or bittersweet family
†Canticarpum celastroides
†Cathispermum pulchrum
†Celastrinoxylon ramunculiformis
Sterculiaceae
†Sphinxia ovalis
Cornaceae
†Beckettia mastixioides
Cornus ettingshausenii
†Dunstania ettinghauseni
†Dunstania multilocularis
†Langtonia bisulcata
†Lanfrancia subglobosa
Mastixia cantiensis
Mastixia grandis
Mastixia parva
†Portnallia bognorensis
†Portnallia sheppeyensis
Nyssaceae - the tupelo family
Nyssa bilocularis
Nyssa cooperi
†Palaeonyssa multilocularis
Curtisiaceae
Curtisia quadrilocularis
Alangiaceae
Alangium jenkinsi
Icacinaceae
†Faboidea crassicutis
†Icacinicarya amygadaloidea
†Icacinicarya bartonensis
†Icacinicarya becktonensis
†Icacinicarya bognorensis
†Icacinicarya echinata
†Icacinicarya elegans
†Icacinicarya emarginata
†Icacinicarya forbesii
†Icacinicarya foveolata
†Icacinicarya glabra
†Icacinicarya inornata
†Icacinicarya jenkinsi
†Icacinicarya minima
†Icacinicarya mucronata
†Icacinicarya nodulifera
†Icacinicarya ovalis
†Icacinicarya ovoidea
†Icacinicarya platycarpa
†Icacinicarya pygmaea
†Icacinicarya reticulata
†Icacinicarya rotundata
†Icacinicarya transversalis
Iodes acutiform
Iodes bilinica
Iodes corniculata
Iodes davisii
Iodes eocenica
Iodes multireticulata
Natsiatum eocenicum
†Palaeophytocrene ambigua
†Palaeophytocrene foveolata
†Sphaeriodes ventricosa
†Stizocarya communis
Animals
Animal fossils include bivalves, gastropods, nautilus, worm tubes, brittle stars and starfish, crabs, lobsters, fish (including shark and ray teeth), reptiles (particularly turtles), and a large diversity of birds. A few mammal remains have also been recorded. Preservation varies; articulated skeletons are generally rare. Of fish, isolated teeth are very frequent. Bird bones are not infrequently encountered compared to other lagerstätten, but usually occur as single bones and are often broken.
The following fauna species list follows Clouter (2007).
Vertebrates
Mammals
Argillotherium
Coryphodon eocaenus – a pantodont
Hyracotherium – the famous horse ancestor
Oxyaena – a creodont
Platychoerops richardsoni - from Herne Bay, a primate belonging to the order Plesiadapiformes
Birds
Anatalavis oxfordi – a waterbird possibly related to the magpie-goose of Australia
Argillipes – perhaps a landfowl
Dasornis, Odontopteryx and Pseudodontornis – pseudo-tooth birds
Eocolius – a coliiform
Eostrix – an owl
Gastornis - from the Isle of Grain, a very large flightless bird
Halcyornis – a parrot or roller relative
Lithornis and Promusophaga – paleognaths
Parvigyps – perhaps a diurnal raptor
Pediorallus – a paleognath or landfowl
Percolinus – perhaps another landfowl
"Precursor" – apparently a chimera of Charadriiformes and Psittaciformes (and possibly other) bones
Primapus – a swift-like bird
Primodroma – a tubenose, possibly a storm-petrel
Prophaeton – a tropicbirds relative
Proherodius – another waterbird
Proplegardis – a stork or ibis
Pulchrapollia – a parrot relative
Stintonornis – probably a hawk relative
Reptiles
Crocodylians
Diplocynodon – an alligatoroid
Kentisuchus spenceri – a crocodylid
Snakes
Palaeophis toliapicus and P. typhaeus
Turtles and tortoises
Allaeochelys – a pig-nosed turtle
Argillochelys, Eochelone, Puppigerus and "Thalassochelys" sp. – true sea-turtles
Chrysemys bicarinata and C. testudiniformis – pond turtles
Eosphargis – a leatherback sea-turtle
Homopus comptoni – a tortoise
Lytoloma crassicostatum and L. planimentum – prehistoric sea-turtles
Palaeaspis – an African sideneck turtle
Podocnemis bowerbanki – an American sideneck turtle
Trionyx pustulatus and T. sp. – softshell turtles
Dacochelys and Pseudotrionyx – incertae sedis
Bony fish
Acestrus elongatus, A. ornatus, Aglyptorhynchus sulcatus, A. venablasi, Xiphiorhynchus parvus and X priscus – swordfish relatives
Acipenser toliapicus – a true sturgeon
Albula oweni – a bonefish
Ampheristus toliapicus – a scorpionfish
Ardiodus marriotti – incertae sedis
Argillichthys toombsi – a lizardfish relative
Aulopopsis depressifrons, A. egertoni and Labrophagus esocinus – flagfins
Beerichthys ingens and B. sp. – Two species of luvar or luvar-like fish
Bramoides brieni and Goniocranion arambourgi – pomfrets
Brychaetus muelleri – an arowana
Bucklandium diluvii – a naked catfish
Cylindracanthus rectus and Hemirhabdorhynchus elliotti – Blochiidae
Cymbium proosti, Eocoelopoma colei, E. curvatum, E. gigas, E. hopwoodi, Eothynnus salmoneus, Scombramphodon crassidens, S. sheppeyensis, Scombrinus macropomus, S. nuchalis, Sphyraenodus priscus, Tamesichthys decipiens, Wetherellus brevior, W. cristatus, W. longior and Woodwardella patellifrons – mackerel and tuna relatives
Diodon sp. – a porcupinefish
Egertonia isodonta and Phyllodus toliapicus – Phyllodontidae
Elops sp., Esocelops cavifrons, Megalops oblongus, M. priscus, Promegalops sheppeyensis and P. signeuxae – ladyfish
Enniskillenus radiatus – acanthomorph
Eutrichurides winkleri – a cutlassfish
Halecopsis insignis – Halecopsidae
Laparon alticeps and Whitephippus tamesis – spadefish
Lehmanamia sheppeyensis – a bowfin
Myripristis toliapicus, Naupygus bucklandi and Paraberyx bowerbanki – soldierfish
Percostoma angustum, Plesioserranus cf. wemmeliensi and Serranopsis londinensis – groupers
Podocephalus curryi, P. nitidus, Sciaenuropsis turneri and Sciaenurus bowerbanki – porgies
Progempylus edwardsi – a snake mackerel
Pseudosphaerodon antiquus and P. navicularis – wrasses?
Pycnodus bowerbanki and P. toliapicus – Pychnodontidae
Rhinocephalus planiceps and Trichurides sagittidens – hakes
Rhynchorhinus branchialis and R. major – Eccelidae
Tetratichthys antiquitatis – a jack mackerel
Cartilaginous fish
Abdounia beaugi, Carcharhinus sp. and Physogaleus secundus – requiem sharks
Aetobatis irregularis, Burnhamia daviesi, Myliobatis dixoni, M. latidens, M. raouxi and M. toliapicus – eagle rays
Aporrhais sowerbii and Eotibia lucida – true conchs
Bathytoma granata, B. turbida, Clavatula conica, Cochlespira gyrata, Conolithus concinnus, Endiatoma cerithiformis, Fusiturris selysi, F. simillima, F. wetherelli, Gemmula koninckii, Hemipleurotoma fasciolata, H. prestwichi, Pseudotoma topleyi, Surculites errans, S. velatus, Turricula crassa, T. helix, T. latimarginata, T. nanodis, T. symmetrica and T. teretrium – Conoidea
Bonellitia clathratum and B. laeviuscula – nutmeg shells