Bembridge Marls Research Articles

The first fossil chylizine fly (Diptera: Psilidae) with a discussion on the ages of related fossil Acalyptratae

Abstract The first fossil rust fly (Diptera: Psilidae) from the Insect Bed of the Bembridge Marls of the Isle of Wight is also the first known fossil record of the subfamily Chylizinae. It is late Eocene: Priabonian in age, 34.2 Ma. Chyliza colenutti Ross, Zhou, Hoffeins & Crighton, sp. nov. is compared with 30 extant species of Chyliza and differs from them in having a relatively long 2nd basal cell compared to the discal cell. Some other fossil Acalyptratae records are reviewed and their probable ages updated. The phylogenetic position of the Psilidae is currently uncertain so the family may have appeared anytime from 70–40 million years ago.

A first species of the genus Cryptophilus (Coleoptera: Erotylidae) from Eocene Baltic amber

Based on a fossil specimen from the Late Eocene Baltic amber Cryptophilus karinae sp. nov., is described. The new species from the late Eocene Baltic amber differs from reexamined holotype of Telmatophilus britannicus from Bembridge Marls in bigger length of body and elytra/pronotum length ratio.

A New Genus of Ricaniidae (Hemiptera: Fulgoromorpha) from the Late Eocene Bembridge Marls, Isle of Wight, United Kingdom

A New Genus of Ricaniidae (Hemiptera: Fulgoromorpha) from the Late Eocene Bembridge Marls, Isle of Wight, United Kingdom

Revealing Palaeogene distribution of the Ptilodactylidae (Insecta: Coleoptera): the first Ptilodactyla Illiger, 1807 records from Rovno amber of Ukraine

ABSTRACT The beetle family Ptilodactylidae is recorded for the first time from Rovno amber, Ukraine. Ptilodactyla odnosum Telnov, Perkovsky, Kundrata and Bukejs sp. nov. and a male of P. eocenica Kundrata, Bukejs and Blank, 2021 are described and illustrated using X-ray micro-computed tomography. Our discoveries of two species in Rovno amber together with recently published findings in Baltic amber and Bembridge Marls suggest that ptilodactylids were rather diverse in Eocene Europe. Identification key to Ptilodactylidae described from European Eocene ambers is provided, and relationships of the Baltic and Rovno amber coleopterofaunas are briefly discussed.

A large Eomys antiquus (Aymard, 1853) (Mammalia, Rodentia) from the early Oligocene sedimentary deposits at Bouldnor Cliff (Isle of Wight, England, UK)

A collection of approx. 150 isolated cheek teeth of the eomyid rodent Eomys antiquus which was obtained from a very thin layer in the Hamstead Member of the Bouldnor Formation at Bouldnor Cliff (Isle of Wight, England, UK) is described. The material is post-Grande Coupure, and earliest Oligocene in age. Comparison is made with and new data are given on teeth of Eomys from localities of a similar age situated elsewhere in Europe: Eomys antiquus from Hoogbutsel (Belgium), and Möhren 13, 19, and 20 (Germany), and Eomys aff. antiquus from Montalbán 1D (Spain), and Kocayarma (Turkish Thrace, Turkey). Teeth from Bouldnor Cliff and Hoogbutsel (Mammal Paleogene zone 21) are morphologically close to those from Möhren 13 (Mammal Paleogene zone 22), but tend to be larger. Size decrease in the course of time is also suggested by the smaller size of the teeth from Montalbán 1D (Mammal Paleogene zone 23). The morphological differences observed between the English, Belgian, and German material at the one side, and the Spanish and Turkish material at the other, confirm that the species from Montalbán 1D and Kocayarma is a different, although closely related form. The taxonomic history of Eomys antiquus is reviewed. The species is the commoner of the two oldest European Eomyidae known. General knowledge on the ecology of the eomyids suggests that at the time of deposition of the Eomys-containing layer in the Hampshire Basin forest was close by.

A previously missing link in the evolution of dasytine soft‐winged flower beetles from Cretaceous Charentese amber (Coleoptera, Melyridae)

AbstractThe soft‐winged flower beetles are a diverse cosmopolitan family of cleroid beetles with an unresolved higher classification, the adults of which feed on nectar and pollen. The fossil record of Melyridae is sparse, with only two described fossils known from the Mesozoic. Here we describe the first representative of the family from the Mesozoic of Europe, from early Cenomanian Charentese (French) amber. Due to the opaque nature of French amber, X‐ray microtomography was used to document fine morphological details of the fossil. Protodasytes cretaceus gen. et sp. nov. possesses a unique combination of characters within the subfamily Dasytinae, which is shared by the tribes Chaetomalachiini, Dasytini, and Listrini, namely the presence of a small antennal club, tarsomere 4 slightly shorter than 3 but not distinctly narrower, symmetrical claws, and a distinct pronotal and elytral fringe. This suggests that Dasytinae had begun to diversify by the Late Cretaceous, possibly in concert with the radiation of angiosperms. The fossil record of soft‐winged flower beetles is critically reviewed, and a checklist of extinct species is provided. The fragmentary Cenozoic fossils Troglops punctatissima and Malachius vertummni are removed from the family Melyridae, and the record of Attalus lusitanicus from Eocene–Oligocene Bembridge Marls is regarded as unlikely. The potential role of Protodasytes gen. nov. and Cretaceous melyrids as pollinators of early angiosperms is discussed.

An etched turtle bone from the Paleogene of the Isle of Wight, UK

Many aspects of the palaeontology of the Isle of Wight are well known, but less so its ichnology. A turtle bone, Emys? sp., from the Lower Oligocene (Rupelian) Hamstead Member, Bouldnor Formation, exposed on the north coast of the island, preserves an unusual surface etching. Dictyoporus nodosus Mägdefrau is recorded from the Paleogene for the first time; hitherto, in northern Europe, it was only known from the Mesozoic. This is also its first record in bone. The trace fossil is an elongate reticulate structure which extended onto adjacent parts of the skeleton (not preserved). Morphologically, identical borings in hard substrates should be included in the same ichnogenus; the nature of the substrate – wood, limestone, bone – is a poor ichnotaxobase. Thus, the clavate bone boring, Karethraichnus Zonneveld et al. is synonymized with Gastrochaenolites Leymerie, more common in limestone. The rarity of D. nodosus may be due to collection failure, in part; perhaps collectors are not looking for surface etchings.

<p><strong>A new species of <em>Sympherobius</em> Banks (Neuroptera: Hemerobiidae) from the late Eocene Rovno amber</strong></p>

Sympherobius irinae sp. nov. (Neuroptera: Hemerobiidae: Sympherobiinae) is described from the late Eocene Rovno amber, Ukraine. It differs from other species of the genus by small details in the forewing venation: (1) crossvein 3rp1-rp2 is not strongly shifted distally relative to neighboring crossveins in the third gradate series, and (2) the humeral veinlet has only two branches. All four late Eocene species form a distinctive species group, which is named here the completus species group, distinguished from other species of the genus by the presence of the crossvein 3rp2-rp3 in the forewing in distal position, connecting RP2 and RP3. The neuropteran assemblages of the Bembridge Marls and succinites (particularly Baltic and Rovno ambers) are briefly compared. The assemblages of some families are relatively similar, but there are obvious differences in others. It is impossible to draw conclusions about the age of Baltic or Rovno ambers based on this weak similarity.

Morphometric analysis of fossil bumble bees (Hymenoptera, Apidae, Bombini) reveals their taxonomic affinities.

Bumble bees (Bombus spp.) are a widespread corbiculate lineage (Apinae: Corbiculata: Bombini), mostly found among temperate and alpine ecosystems. Approximately 260 species have been recognized and grouped recently into a simplified system of 15 subgenera. Most of the species are nest-building and primitively eusocial. Species of Bombus have been more intensely studied than any other lineages of bees with the exception of the honey bees. However, most bumble bee fossils are poorly described and documented, making their placement relative to other Bombus uncertain. A large portion of the known and presumed bumble bee fossils were re-examined in an attempt to better understand their affinities with extant Bombini. The taxonomic affinities of fossil specimens were re-assessed based on morphological features and previous descriptions, and for 13 specimens based on geometric morphometrics of forewing shape. None of the specimens coming from Eocene and Oligocene deposits were assigned within the contemporary shape space of any subgenus of Bombus. It is shown that Calyptapis florissantensis Cockerell, 1906 (Eocene-Oligocene boundary, Florissant shale, Colorado, USA) and Oligobombus cuspidatus Antropov, 2014 (Late Eocene, Bembridge Marls) likely belong to stem-group Bombini. Bombus anacolus Zhang, 1994, B. dilectus Zhang, 1994, B. luianus Zhang, 1990 (Middle Miocene, Shanwang Formation), as well as B. vetustus Rasnitsyn & Michener, 1991 (Miocene, Botchi Formation) are considered as species inquirenda. In the Miocene, affinities of fossils with derived subgenera of Bombus s. l. increased, and some are included in the shape space of contemporary subgenera: Cullumanobombus (i.e., B. pristinus Unger, 1867, B. randeckensis Wappler & Engel, 2012, and B. trophonius Prokop, Dehon, Michez & Engel, 2017), Melanobombus (i.e., B. cerdanyensis Dehon, De Meulemeester & Engel, 2014), and Mendacibombus (i.e., B. beskonakensis (Nel & Petrulevičius, 2003), new combination), agreeing with previous estimates of diversification.

True flies (Insecta: Diptera) from the late Eocene insect limestone (Bembridge Marls) of the Isle of Wight, England, UK

ABSTRACTThe Diptera fauna from the late Eocene of the Isle of Wight (Bembridge Marls) is studied including redescriptions of formerly described material. The fauna includes the following taxa: Anisopodidae – one species; Bibionidae – 11 species; Ceratopogonidae – one described and two unidentified species; Chironomidae – undetermined species of three subfamilies; Culicidae – four species; Cylindrotomidae – one species; Dixidae – one species; Keroplatidae – one described and four unidentified species; Limoniidae – 31 species; Mycetophilidae – 14 species; Psychodidae – two species; Scatopsidae – two species; Sciaridae – three species; Simuliidae – an unnamed species; Tipulidae – nine species. Diptera Brachycera: Agromyzidae – unidentified species; Anthomyzidae – two species; Asilidae – two species; Bombyliidae – one species; Chloropidae – one species; Dolichopodidae – unidentified species; Empididae – unidentified species; Ephydridae – two species; Heleomyzidae – unidentified species; Lauxaniidae – unidentified species; Otitidae – one species; Phoridae – unidentified species; Rhagionidae – two species; Stratiomyidae – two species; Syrphidae – unidentified species; Tabanidae – one species. Following new taxa are described:Leptotarsus(Longurio)fragmentatusKrzemiński, n. sp.,Leptotarsus(Longurio)wegierekiKrzemiński, n. sp.,Tipula(s. lato)anglicanaKrzemiński, n. sp. (Tipulidae);CyttaromyiarossiKrzemiński, n. sp. (Cylindrotomidae);Gurnardia coramiKrzemiński, n. gen, n. sp.,Pilaria hooleyiKrzemiński, n. sp.,Pilaria volodiiKrzemiński, n. sp.,Cheilotrichia (Cheilotrichia) duplicataKrzemiński, n. sp.,Cheilotrichia (Empeda) szwedoiKrzemiński, n. sp.,Symplecta (Psiloconopa) gurnetensisKrzemiński, n. sp.,Orimarga (Orimarga) lenaeKrzemiński, n. sp.,Dicranomyia (Dicranomyia) azariKrzemiński, n. sp.,Dicranomyia (Dicranomyia) fasciataKrzemiński, n. sp.,Helius (Helius) edmundiKrzemiński, n. sp.,Helius (Helius) popoviKrzemiński, n. sp.,Dicranoptycha staryiKrzemiński, n. sp. (Limoniidae);WightipsychodaAzar, n. gen. (Psychodidae);Dilophus andrewrossiNel, Colomb & Waller, n. sp.,Bibiodes massiliensisNel, Colomb & Waller, n. sp. (Bibionidae);Azana cockerelliBlagoderov, n. sp.,Aglaomyia vectisBlagoderov, n. sp.,Dziedzickia oligocenicaBlagoderov, n. sp.,Palaeoempalia saxeaBlagoderov, n. sp.,MycomyahooleiBlagoderov, n. sp.,Leia gurnardensisBlagoderov, n. sp. (Mycetophilidae);RegmocleminahaenniiKrzemiński, n. sp. (Scatopsidae);Sylvicola problematicaKrzemiński, n. sp. (Anisopodidae);Palaeoberidops barkeriNicholson,n. gen., n. sp. (Stratiomyidae). The Diptera fauna and palaeohabitat of the Insect Bed is discussed.

Vertebrate remains from the Insect Limestone (latest Eocene), Isle of Wight, UK

ABSTRACTA small fauna of vertebrates is recorded from the Insect Limestone, Bembridge Marls Member, Bouldnor Formation, late Priabonian, latest Eocene, of the Isle of Wight, UK. The taxa represented are unidentified teleost fishes, lizards including a scincoid, unidentified birds and the theridomyid rodent Isoptychus. The scincoid represents the youngest record of the group in the UK. Of particular note is the taphonomic interpretation based on the preservation of anatomical parts of land-based tetrapods that would have been most likely transported to the site of deposition by wind, namely bird feathers and pieces of shed lizard skin. These comprise the majority of the specimens and suggest that the dominant transport mechanism was wind.

First Record of the Late Eocene Ichneumon Fly Rasnitsynites tarsalis Kasparyan (Ichneumonidae, Townesitinae) in Ukraine Confirms Correlation of the Upper Eocene Lagerstätten

Rasnitsynites tarsalis Kasparyan (Ichneumonidae, Townesitinae) described from the Baltic amber is recorded for the first time in the Rovno amber (Gulyanka, Zhitomir Region); therefore, this is the southernmost and easternmost find of Townesitinae. The presence of numerous extinct taxa of Townesitinae and tribes of Hybrizoninae Blanchard 1845 = Paxylommatinae Forster, 1862 = Hybrizontinae Marshall, 1872 confirms close relation of the Bembridge Marls fauna (Upper Eocene of England) and that of the succinites (Baltic, Rovno, and other Late Eocene European ambers); they are common in the succinites and in the Bembridge Marls and are only known from these localities.

Tertiary bark lice (Insecta: Psocodea) from the Insect Limestone (Bembridge Marls, Late Eocene) of the Isle of Wight, UK

ABSTRACTLate Eocene fossil Psocodea from the Isle of Wight are studied and classified within three new genera and seven species (Wightipsocus acourti (Cockerell 1921) gen. nov. et n. comb., Wightipsocus intasi gen. et sp. nov., Wightipsocus rossi gen et sp. nov., Awightipsocus jarzembowskii gen. et sp. nov., Awightipsocus megaloi gen. et sp. nov. Awightipsocus minimus gen. et sp. nov. and Wightimicropsocus inexpectatus gen. et sp. nov.) without however attributing them to any family. A key for fossil Psocodea from the Isle of Wight is given.

Thrips (Insecta: Thysanoptera) from the Insect Limestone (Bembridge Marls, Late Eocene) of the Isle of Wight, UK

ABSTRACTThirty-two thrips fossils were studied from the Bembridge Marls, late Eocene of the Isle of Wight, S. England. Two extant families are recorded, Aeolothripidae with two species and Thripidae with three species. Described as new areSinaeolothripsShmakov gen. n. (Aeolothrips brodieiCockerell, 1917, type species),Aeolothrips jarzembowskiiShmakov sp. nov (Aeolothripidae) andCoccothrips hoffeinsorumShmakov, sp.nov.(Thripidae: Panchaetothripinae). The genusThripsis tentatively recorded as fossil for the first time. Taxonomic composition of thrips is close to the Recent ones.

The Flora of the Insect Limestone (latest Eocene) from the Isle of Wight, southern England

ABSTRACTLatest Eocene fossil plant remains occur in concentrations within blue-grey micrite known as Insect Limestone near the base of the Bembridge Marls Member (Bouldnor Formation, Solent Group), Isle of Wight, southern England. Some of the previously reported taxa (collections in the Natural History Museum, London) are not preserved within the Insect Limestone. These (e.g., all Arecaceae (palms)) are excluded from the floral list. New non-destructive techniques have yielded additional taxonomic information. Leaves previously assigned to Ficus and Fagus are now incertae sedis. Wetland elements are abundant, particularly Typha, but also Acrostichum, Azolla, Potamogeton, Sparganium and others. Non-wetland elements are rare. Trees and shrubs included representatives of Betulaceae, Caprifoliaceae, Juglandaceae, Lauraceae, Rhamnaceae (the sclerophyllous Zizyphus), other flowering plants and several genera of conifers. There are rare specimens of possible herbaceous plants and propagules with plumes or awns, the latter possibly an early fossil record of Clematis. The common plant remains were probably derived from vegetation near a freshwater body, sometimes with slight brackish influence, whilst rarer elements were probably blown in from a greater distance. There is little evidence of plant–insect interaction; one leaf with small galls, a stem containing an insect larva and a possible association between stratiomyid flies and Typha.

Mollusca from the Insect Limestone (Bembridge Marls Member: Bouldnor Formation: Solent Group), Palaeogene, Isle of Wight, southern England

ABSTRACTMolluscs from the Insect Limestone of the Bembridge Marls Member are described. The gastropodsLymnaea(Galba)longiscata(Brongniart 1810),Planorbarius discus(Edwards 1852),Gyraulus similisFérussac, 1814,Hippeutis headonensisJodot, 1942, Viviparuslentus(Solander 1766) andViviparus angulosus(J. Sowerby 1817) indicate freshwater alkaline conditions. The cerithioid gastropodTarebia acuta(J. Sowerby 1822) probably represent more brackish conditions.Polymesoda(Pseudocyrena)obovata(J. Sowerby 1817) is the only bivalve present. All were potentially washed into the depocentre. Despite a wide outcrop of the Insect Limestone on the Isle of Wight's northwest and northeast coasts, there are no records of molluscs other than from the Gurnard/Thorness Bay area on the northwest coast of the island.

Caddis-flies (Insecta: Trichoptera) from the Insect Limestone (Bembridge Marls, Late Eocene) of the Isle of Wight, UK

ABSTRACTThe trichopteran assemblage of the Late Eocene Bembridge Marls is revised. six species in five genera and four families are identified, with four species and two genera described as new: (Beraeidae: Beraeodes vectensis Cockerell, 1921 and B. anglicus Cockerell, 1921; Phryganeidae: Bembridgea insularia gen. sp. nov.; Polycentropodidae: ?Plectrocnemia incompleta Sukatsheva, sp. nov.; Philopotamidae: Wormaldia longaeva Sukatsheva, sp. nov.; incertae familiae: Paleodicella anomala Sukatsheva, gen. sp. nov.). The assemblage is unique in being absolutely dominated by the generally uncommon family Beraeidae, and particularly by the single species Beraeodes vectensis. Except for a representative of ubiquitous Phryganaeidae, all the caddis-flies recorded apparently had oxy- and psychrophilous larvae confined to springs and cold streams (or else mosses near water), thus indicating the absence of fresh lakes or slow running waters near the depositional water body.

The wasps, bees and ants (Insecta: Vespida=Hymenoptera) from the Insect Limestone (Late Eocene) of the Isle of Wight, UK

ABSTRACTThe types and undescribed material of the hymenopteran fossils of the Insect Bed of the Bembridge Marls from the Isle of Wight (UK) are critically revised and studied. A total of 1460 fossils are recorded and attributed to 20 families: Gasteruptiidae s.l. (1); Proctotrupidae (3); Diapriidae (24); Cynipidae (7); Figitidae (6); Pteromalidae (1); Agaonidae (3); Scelionidae (12); Platygastridae (2); Ichneumonidae (32); Braconidae (75); Bethylidae (3); Crabronidae (2); Sphecidae (1); Apidae (2); Scoliidae (1); Tiphiidae (2); Vespidae (4); and Formicidae (1220). Described as new are 51 species, 13 genera, two tribes and two subfamilies. Minimum number of species recorded (either as described species or representing higher taxa with no described species in the assemblage) is 118. The composition of the hymenopteran assemblage is most similar to that of Baltic amber and indicative of a well forested territory, as well as of a humid, equable (aseasonal but not very hot) climate, more typically equable than in the Baltic amber source area, judging from the absence of Aphidiinae and scarcity of aphids.

A radiometric airborne geophysical survey of the Isle of Wight

A high resolution airborne geophysical survey across the Isle of Wight and Lymington area conducted in 2008 provided the first modern radiometric survey across the geological formations that characterise much of southern England. The basic radiometric data are presented and it is evident that bedrock geology exerts a controlling influence on the broad response characteristics of the naturally occurring radioelements. A GIS-based geological classification of the data provides a quantitative assessment and reveals that a relatively high percentage of the variability of the data is explained by the Cretaceous bedrock geology but this is much reduced in the Palaeogene. The three traditional Chalk units (Lower, Middle and Upper Chalk depicted on the currently available Geological Map) provide the lowest and most distinct behaviour within the Cretaceous sequence. Mineral content within the Chalk appears to increase with increasing age. A new method of representing the baseline radiometric information from the survey in terms of the mean values of the geological classification is presented. The variation of radioelement geochemistry within individual formations is examined in two case studies from the Cretaceous Lower Greensand Group and the Palaeogene Hamstead Member (Bouldnor Formation). The Cretaceous sequences provide the higher levels of discrimination of localised variations in radioelement distributions. A more detailed case study examines the potential influences from the degree of water saturation in the soil and superficial deposits.

Anguimorph lizards (Squamata, Anguimorpha) from the Middle and Late Eocene of the Hampshire Basin of southern England

The Hampshire Basin in southern England is the only sequence virtually spanning the Eocene-Oligocene boundary in western Europe. It has provided an opportunity to observe changes in populations across the Grande Coupure event, as an overturn in the squamate assemblage of continental Europe has also been noted. Five Tertiary formations have yielded lizard material: Creechbarrow Limestone, Barton Clay, Headon Hill, Bembridge Limestone and Bouldnor Formation. These formations span the period from the late Middle Eocene to the earliest Oligocene. Localities in the Hampshire Basin represent a depositional environment with an increasing tendency towards lacustrine and riparian influences in later horizons. The anguimorph fauna includes one new anguine, Headonhillia parva gen. et sp. nov., diagnosed by: (1) small size, (2) large parietal area levis, and (3) short parietal crista postfovealis. Another anguine, Ophisaurus sp., is identified on the basis of the morphology of the parietal. Three further morphotypes of anguines have been distinguished based on dentary, vertebrae and osteoderm specimens. Two types of indeterminate Anguidae are distinguished on the basis of maxillae and dentaries and three morphotypes of indeterminate Anguidae on the basis of vertebrae. Glyptosaurines have been identified in three members on the basis of osteoderms. ‘Necrosaurs’ have also been identified from the Hampshire Basin: two types, distinguished on the basis of dentary material and osteoderms, are identified as indeterminate ‘necrosaurs’. One type of indeterminate Varanoidea is distinguished on the basis of dentaries. The presence of anguids in the Hampshire Basin is well documented, and they appear to form a consistently diverse assemblage throughout the sequence. Anguine osteoderms are present in both pre- and post-Grande Coupure material. Pre-Grande Coupure vertebrae and osteoderms are similar in size to those of adult Recent anguine Pseudopus apodus, although many of the Hampshire Basin anguines, including Headonhillia parva, represent the smallest anguines known to date.