A 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.

This paper reviews the changing environments, developing landforms and terrestrial stratigraphy during the Early and Middle Pleistocene stages in Scotland. Cold stages after 2.7 Ma brought mountain ice caps and lowland permafrost, but larger ice sheets were short-lived. The late Early and Middle Pleistocene sedimentary record found offshore indicates more than 10 advances of ice sheets from Scotland into the North Sea but only 4–5 advances have been identified from the terrestrial stratigraphy. Two primary modes of glaciation, mountain ice cap and full ice sheet modes, can be recognised. Different zones of glacial erosion in Scotland reflect this bimodal glaciation and the spatially and temporally variable dynamics at glacier beds. Depths of glacial erosion vary from almost zero in Buchan to hundreds of metres in glens in the western Highlands and in basins both onshore and offshore. The presence of tors and blockfields indicates repeated development of patches of cold-based, non-erosive glacier ice on summits and plateaux. In lowlands, chemical weathering continued to operate during interglacials, but gruss-type saprolites are mainly of Pliocene to Early Pleistocene age. The Middle Pleistocene terrestrial stratigraphic record in Scotland, whilst fragmentary and poorly dated, provides important and accessible evidence of changing glacial, periglacial and interglacial environments over at least three stadial–interstadial–interglacial cycles. The distributions of blockfields and tors and the erratic contents of glacial sediments indicate that the configuration, thermal regime and pattern of ice flow during MIS 6 were broadly comparable to those of the last ice sheet. Improved control over the ages of Early and Middle Pleistocene sediments, soils and saprolites and on long-term rates of weathering and erosion, combined with information on palaeoenvironments, ice extent and sea level, will in future allow development and testing of new models of Pleistocene tectonics, isostasy, sea-level change and ice sheet dynamics in Scotland.

The crustacean fauna of the Insect Bed (late Eocene), Isle of Wight is reviewed. The fauna comprises the branchiopod Branchipodites vectensis Woodward, 1879, ostracod Potamocypris brodiei Jones and Sherborn, 1889, and isopod Eosphaeroma margarum (Desmarest, 1822). In addition a new clam shrimp (Crustacea: Diplostraca: Spinicaudata) is described and named Paraleptestheria mitchelli sp. nov. This is the first record of the genus outside China and the first ‘conchostracan' to be described from the European Cenozoic.

Lithostratigraphical studies coupled with the development of new dating methods has led to significant progress in understanding the Late Pleistocene terrestrial record in Scotland. Systematic analysis and re-evaluation of key localities have provided new insights into the complexity of the event stratigraphy in some regions and the timing of Late Pleistocene environmental changes, but few additional critical sites have been described in the past 25 years. The terrestrial stratigraphic record remains important for understanding the timing, sequence and patterns of glaciation and deglaciation during the last glacial/interglacial cycle. Former interpretations of ice-free areas in peripheral areas during the Last Glacial Maximum (LGM) are inconsistent with current stratigraphic and dating evidence. Significant challenges remain to determine events and patterns of glaciation during the Early and Middle Devensian, particularly in the context of offshore evidence and ice sheet modelling that indicate significant build-up of ice throughout much of the period. The terrestrial evidence broadly supports recent reconstructions of a highly dynamic and climate-sensitive British–Irish Ice Sheet (BIIS), which apparently reached its greatest thickness in Scotland between 30 and 27ka, before the global LGM. A thick (relative to topography) integrated ice sheet reaching the shelf edge with a simple ice-divide structure was replaced after the LGM by a much thinner one comprising multiple dispersion centres and a more complex flow structure.

The last Scottish Ice Sheet (SIS) expanded from a pre-existing ice cap after ∼35 ka. Highland ice dominated, with subsequent build-up of a Southern Uplands ice mass. The Outer Hebrides, Skye, Mull, the Cairngorms and Shetland supported persistent independent ice centres. Expansion was accompanied by ice-divide migration and switching flow directions. Ice nourished in Scotland reached the Atlantic Shelf break in some sectors but only mid-shelf in others, was confluent with the Fennoscandian Ice Sheet (FIS) in the North Sea Basin, extended into northern England, and fed the Irish Sea Ice Stream and a lobe that reached East Anglia. The timing of maximum extent was diachronous, from ∼30–27 ka on the Atlantic Shelf to ∼22–21 ka in Yorkshire. The SIS buried all mountains, but experienced periods of thickening alternating with drawdown driven by ice streams such as the Minch, the Hebrides and the Moray Firth Ice Streams. Submarine moraine banks indicate oscillating retreat and progressive decoupling of Highland ice from Orkney–Shetland ice. The pattern and timing of separation of the SIS and FIS in the North Sea Basin remain uncertain. Available evidence suggests that by ∼17 ka, much of the Sea of the Hebrides, the Outer Hebrides, Caithness and the coasts of E Scotland were deglaciated. By ∼16 ka, the Solway lowlands, Orkney and Shetland were deglaciated, the SIS and Irish Ice Sheet had separated, the ice margin lay along the western seaboard, nunataks had emerged in Wester Ross, the ice margin lay N of the Cairngorms and the sea had invaded the Tay and Forth estuaries. By ∼15 ka, most of the Southern Uplands, the Firth of Clyde, the Midland Valley and the upper Spey valley were deglaciated, and in NW Scotland ice was retreating from fjords and valleys. By the onset of rapid warming at ∼14.7 ka, much of the remnant SIS was confined within the limits of Younger Dryas glaciation. The SIS, therefore, lost most of its mass during the Dimlington Stade. It is uncertain whether fragments of the SIS persisted on high ground throughout the Lateglacial Interstade.

The fossil cockroaches (Blattodea), praying mantises (Mantodea) and earwigs (Demaptera) are described from the Insect Limestone (Priabonian) of the Isle of Wight, southern England. Three new species of cockroach are described in the family Ectobiidae – Phyllodromica protosardea sp. nov., Balta protosimilis sp. nov. and Malaccina? wightensis sp. nov. – and a further nine indeterminate species are presented (based on ten specimens). The only known specimen of praying mantis is described as Protohierodula crabbi gen. et sp. nov. in the family Manteidae, which constitutes the first record of Mantodea from the UK. The only known specimen of earwig is an incomplete juvenile belonging to the superfamily Forficuloidea.

During Lateglacial cold periods, permafrost developed throughout Scotland, sediment-mantled slopes were extensively modified by solifluction and other forms of periglacial mass movement, large-scale sorted patterned ground formed on plateaux, and enhanced rockfall resulted in talus accumulation below cliffs. Most rock-slope failures occurred within five millennia following ice-sheet deglaciation, with many probably triggered by uplift-induced earthquakes; numerous debris-free scarps represent sites where Lateglacial rockslide debris was excavated by glaciers during the Younger Dryas Stade (∼12.9–11.7 ka). Sandar and outwash fans deposited by glacial rivers during ice-sheet retreat were incised to form high-level terraces. Under the cool temperate but relatively stable climate of the Holocene, solifluction and patterned ground formation continued to operate on high ground, though accumulation of high-level aeolian deposits on most mountains was terminated by erosion during the Little Ice Age. Drift-mantled slopes and talus slopes have been extensively eroded by translational failures and debris flows, the latter depositing debris cones on upland valley floors. The incidence of Holocene rockslides has been much lower than during the Lateglacial period. Dating of alluvial deposits and low Holocene terraces suggests no consistent pattern of Holocene floodplain evolution: incision has apparently dominated in the Highlands, aggradation in the lowlands, and floodplains in the Southern Uplands have asynchronous histories of incision and aggradation. Studies of floodplain behaviour over the past 200–300 years suggest that though major floods rework the floodplains of braided and piedmont rivers, there is no tendency towards net floodplain aggradation or incision. Most valley-side alluvial fans accumulated episodically in the last 4000 years, many in response to lowering of hillslope stability by woodland clearance. For many postglacial landsystems, disentangling the effects of declining paraglacial sediment supply, climate change and local influences (extreme rainstorm events or anthropogenic impacts) remains challenging.

The late Eocene mantis genus and species Protohierodula crabbi can be attributed to the extant clade Artimantodea incertae sedis sit. nov., but its original placement in the Manteidae is not supported. It remains the oldest reliable described Artimantodea.

This paper reviews the evidence for environmental change during the Lateglacial period (c.14.7–11.7 ka), perhaps the most intensively studied episode in the Quaternary history of Scotland. It considers first the stratigraphic subdivision and nomenclature of the Lateglacial, before proceeding to a discussion of the various lines of proxy evidence that have been used to reconstruct the spatial and temporal patterns of environmental change during this time period. These include pollen and plant macrofossil data; coleopteran and chironomid records; diatom data; stable isotope and geochemical records; and evidence for human activity. The paper then considers the principal methods that have been employed to date and correlate Lateglacial events: radiocarbon dating; surface exposure dating; varve chronology; and tephrochronology. This is followed by an examination of the constraints imposed on environmental reconstructions, an account of the ways in which the evidence can be employed in the development of an event stratigraphy for the Lateglacial in Scotland, and a proposal for a provisional Lateglacial type sequence (stratotype) at Whitrig Bog in SE Scotland. Emphasis is placed throughout on the potential linkages between the Scottish records and the isotopic signal in the Greenland ice cores, which forms the stratigraphic template for the N Atlantic region. The paper concludes with a discussion of the strategies and approaches that should underpin future research programmes on Lateglacial environmental change in Scotland.

I accept Schubnel & Nel's (2019) opinion that Protohierodula belongs to the clade Artimantodea, that it cannot be reliably assigned to the family Manteidae and should be regarded as family incertae sedis.

All taxa of Orthoptera known from the latest Eocene deposits of the Isle of Wight are redescribed or discussed, based on the study of their type material. The following new taxa are described: Ontogryllus gen. nov., O. rossi sp. nov. (type species), Achetomorpha gen. nov., A. irregularis sp. nov. (type species), and Proeneopterotrypus gen. nov. The latter genus is here proposed for Pteroplistes danicus Rust, described from the Palaeocene/Eocene of Denmark, as this find is important for understanding the systematic position of the genus Eneopterotrypus Zeuner. Remarks about the taxonomy of the taxa studied, age of these deposits, and ecology are given.

Palaeoecology has been prominent in studies of environmental change during the Holocene epoch in Scotland. These studies have been dominated by palynology (pollen, spore and related bio-and litho-stratigraphic analyses) as a key approach to multi- and inter-disciplinary investigations of topics such as vegetation, climate and landscape change. This paper highlights some key dimensions of the pollen- and vegetation-based archive, with a focus upon woodland dynamics, blanket peat, human impacts, biodiversity and conservation. Following a brief discussion of chronological, climatic, faunal and landscape contexts, the migration, survival and nature of the woodland cover through time is assessed, emphasising its time-transgressiveness and altitudinal variation. While agriculture led to the demise of woodland in lowland areas of the south and east, the spread of blanket peat was especially a phenomenon of the north and west, including the Western and Northern Isles. Almost a quarter of Scotland is covered by blanket peat and the cause(s) of its spread continue(s) to evoke recourse to climatic, topographic, pedogenic, hydrological, biotic or anthropogenic influences, while we remain insufficiently knowledgeable about the timing of the formation processes. Humans have been implicated in vegetational change throughout the Holocene, with prehistoric woodland removal, woodland management, agricultural impacts arising from arable and pastoral activities, potential heathland development and afforestation. The viability of many current vegetation communities remains a concern, in that Scottish data show reductions in plant diversity over the last 400 years, which recent conservation efforts have yet to reverse. Palaeoecological evidence can be used to test whether conservation baselines and restoration targets are appropriate to longer-term ecosystem variability and can help identify when modern conditions have no past analogues.

Representatives of the Hemiptera: Sternorrhyncha, Fulgoromorpha, Cicadomorpha and Heteroptera from the Late Eocene of the Isle of Wight, UK, are analysed and discussed. Psylloidea were reviewed and a key to the described taxa is given. Aphidoidea were studied, previously described taxa revised and new taxa described. New taxa of Aphidoidea are Hormaphididae: Hormaphis? longistigma Wegierek sp. nov.; Eriosomatidae: Eriosoma gratshevi Wegierek sp. nov. and Colopha? incognita Wegierek sp. nov.; Drepanosiphidae: Panfossilis anglicus Wegierek gen. et sp. nov. and Betulaphis kozlovi Wegierek sp. nov. Previously described Fulgoromorpha were revised and new taxa are described. The homonym Hastites Cockerell, 1922 (Cixiidae) preoccupied by Hastites Mayer-Eymar, 1883 is replaced by Catulliastites Szwedo nom. nov. for Catulliastites muiri (Cockerell) comb. nov. New taxa described are Cixiidae: Klugga gnawa Szwedo gen. et sp. nov., Klugga regoa Szwedo sp. nov., Liwakka gelloa Szwedo gen et sp. nov., Delwa morikwa Szwedo gen. et sp. nov., Kommanosyne wrikkua Szwedo gen. et sp. nov., Kernastiridius nephlajeus Szwedo gen. et sp. nov., Margaxius angosus Szwedo gen. et sp. nov., Dweivera reikea Szwedo gen. et sp. nov., Samaliverus bikkanus Szwedo gen. et sp. nov., Komsitija tuberculata Szwedo gen. et sp. nov., Langsmaniko marous Szwedo gen. et sp. nov., Komnixta jarzembowskii Szwedo gen. et sp. nov. and Worodbera nimakka gen. et sp. nov.; Tropiduchidae: Reteotissus hooleyi Szwedo gen. et sp. nov., Phatanako gen. nov. for Phatanako wilmattae (Cockerell) comb. nov., Senogaetulia kwalea Szwedo gen. et sp. nov., Dakrutulia mikhailkozlovi Szwedo gen. et sp. nov., Keriophettus atibenus Szwedo gen. et sp. nov. and Sognotela emeljanovi Szwedo gen. et sp. nov.; Issidae: Krundia korba Szwedo gen. et sp. nov., Breukoscelis vadimgratshevi Szwedo gen. et sp. nov., Breukoscelis phrikkosus Szwedo sp. nov. and Uphodato garwoterus Szwedo gen. et sp. nov.; Nogodinidae: Ambitaktoinae Szwedo subfam. nov., Ambitaktoa stoumma Szwedo gen. et sp. nov., Phariberea gurdonika Szwedo gen. et sp. nov., Wixskimoa torxsea Szwedo gen. et sp. nov., and Nadrimini trib. nov. with Niadrima yulei Szwedo gen. et sp. nov.; Lophopidae: Ankomwarius brodiei Szwedo gen. et sp. nov.; Ricaniidae: Ankwlanno bluga Szwedo gen. et sp. nov. Previously described Cicadomorpha were revised and new taxa are described: Cicadidae: Kintusamo boulardi Szwedo gen. et sp. nov; Aphrophoridae: Blenniphora Szwedo gen. nov. for Blenniphora woodwardi (Cockerell) comb. nov., Blenniphora skaka Szwedo sp. nov. and Blenniphora bikkanoa Szwedo sp. nov.; Luisphantyelus briwus Szwedo gen. et sp. nov., Natajephora lijanka Szwedo gen. et sp. nov.; Cercopidae: Berro enissuextaensis Szwedo gen. et sp. nov.; Cicadellidae: Mileewinae: Teniwitta andrewrossi Szwedo gen. et sp. nov. Formerly described true bugs (Heteroptera) are revised and several new taxa are described – Nepomorpha: Corixidae: Diacorixites szwedoi Popov gen. et sp. nov.; Cimicomorpha: Tingidae: Parasinalda wappleri Popov sp. nov., Viktorgolubia Popov gen. nov. for Viktorgolubia seposita (Cockerell) comb. nov.; Miridae: Gurnardinia herczeki Popov gen. et sp. nov.; Pentatomomorpha: Lygaeidae: Gurnardobayini Popov trib. nov., Gurnardobaya rossi Popov gen. et sp. nov.; Cydnidae: Eocenocydnus lisi Popov gen. et sp. nov.; Pentatomidae: Podopinites coloratus Popov gen. et sp. nov. and Podopinites acourti (Cockerell) comb. nov. An overview of the Late Eocene fauna of the Hemiptera is presented. Ecological and biogeographical patterns of the Hemiptera from the Isle of Wight deposits are discussed.

This paper summarises developments in understanding sea level change during the Quaternary in Scotland since the publication of the Quaternary of Scotland Geological Conservation Review volume in 1993. We present a review of progress in methodology, particularly in the study of sediments in isolation basins and estuaries as well as in techniques in the field and laboratory, which have together disclosed greater detail in the record of relative sea level (RSL) change than was available in 1993. However, progress in determining the record of RSL change varies in different areas. Studies of sediments and stratigraphy offshore on the continental shelf have increased greatly, but the record of RSL change there remains patchy. Studies onshore have resulted in improvements in the knowledge of rock shorelines, including the processes by which they are formed, but much remains to be understood. Studies of Late Devensian and Holocene RSLs around present coasts have improved knowledge of both the extent and age range of the evidence. The record of RSL change on the W and NW coasts has disclosed a much longer dated RSL record than was available before 1993, possibly with evidence of Meltwater Pulse 1A, while studies in estuaries on the E and SW coasts have disclosed widespread and consistent fluctuations in Holocene RSLs. Evidence for the meltwater pulse associated with the Early Holocene discharge of Lakes Agassiz–Ojibway in N America has been found on both E and W coasts. The effects of the impact of storminess, in particular in cliff-top storm deposits, have been widely identified. Further information on the Holocene Storegga Slide tsunami has enabled a better understanding of the event, but evidence for other tsunami events on Scottish coasts remains uncertain. Methodological developments have led to new reconstructions of RSL change for the last 2000 years, utilising state-of-the-art GIA models and alongside coastal biostratigraphy to determine trends to compare with modern tide gauge and documentary evidence. Developments in GIA modelling have provided valuable information on patterns of land uplift during and following deglaciation. The studies undertaken raise a number of research questions which will require addressing in future work.

New fossils of the families Nemopteridae and Chrysopidae are recorded and a new species of Hemerobiidae, Sympherobius yulei sp. nov., is described from the late Eocene Insect Bed of the Isle of Wight. The new nemopterid belongs to the tribe Stenonemiini and is very similar to a specimen already recorded from the Early Oligocene of Alsace (France), suggesting that these warm-climate insects were not affected by the ‘Oi-1 Glaciation' cooling.

Quaternary deposits and landforms are an integral component of Scotland's geodiversity and natural heritage with intrinsic, scientific, educational, cultural, aesthetic and ecological values. Their conservation is founded on the assessment and safeguard of key protected areas principally for their scientific values. The evaluation of site networks for Quaternary deposits and landforms (including glacial, fluvial, coastal, mass movement, karst and cave features) has evolved since the late 1940s, culminating in the Great Britain Geological Conservation Review (GCR) site assessments undertaken principally between 1977 and the early 1990s. Significant scientific progress since then has arisen, for example, from re-investigation of existing sites and discoveries of new sites, developments in geochronology and the formulation and application of new concepts and models. Both the GCR site lists and the supporting site documentation now require updating in the light of this progress. Today there is greater emphasis on the wider, non-scientific values of geoconservation including, for example, on ecosystem services, links with biodiversity and cultural heritage, geotourism and the benefits for human health and wellbeing through improved understanding of dynamic landscapes, climate change and natural hazards. Involvement of wider public support beyond the geoscience community and fostering better integration of geoheritage within the developing nature conservation agenda, including a land systems approach, protected area planning and management, natural capital and connecting people and nature, will help further to protect our Quaternary geoheritage.

A review of the faunistic composition of the coleopterous taphocenoses from Bembridge Marls is given. Only two families (Cupedidae and Carabidae) have been recorded from this site before. A total of 31 families have been revealed and determined in the course of the recent study, and 42 species have been described: Agabus latissimus Ponomarenko, sp. nov. and Ilybius gratshevi Ponomarenko, sp. nov. from Dytiscidae; Neothanes europaeus Ponomarenko, sp. nov. from Carabidae; Spercheus punctatus Ponomarenko, sp. nov., Spercheus wightensis Ponomarenko, sp. nov. from Spercheidae, Hydrochara woodwardi Ponomarenko & Soriano, sp. nov. and Berosus barclayi Ponomarenko & Soriano, sp. nov. from Hydrophilidae; Ochthebius rossi Kirejtshuk, sp. nov., Eolimnebius fossilis Kirejtshuk, sp. nov., Hydraenites gracilimmus Kirejtshuk, sp. nov., Metacoxites ventritalis Kirejtshuk, sp. nov., Davidraenites gratshevi Kirejtshuk, sp. nov., D. interruptus Kirejtshuk, sp. nov. and D. spurcus Kirejtshuk, sp. nov. from Hydraenidae; Aphodius vectis Krell, sp. nov. and Pentodon dorcus Krell, sp. nov. from Scarabaeidae; Scirtes calcariferens Kirejtshuk & Ponomarenko, sp. nov., Scirtes khnzoryani Kirejtshuk & Ponomarenko, sp. nov., Scirtes metepisternalis Kirejtshuk & Ponomarenko, sp. nov., Scirtes wightensis Kirejtshuk & Ponomarenko, sp. nov., Contacyphon insularis Kirejtshuk & Ponomarenko, sp. nov., Contacyphon involutus Kirejtshuk & Ponomarenko, sp. nov. and Contacyphon kozlovi Kirejtshuk & Ponomarenko, sp. nov. from Scirtidae; Eucinetes nikolaevae Kirejtshuk & Ponomarenko, sp. nov. from Eucinetidae; Macropunctum rossi Alexeev, sp. nov. from Elateridae; Byrrhites bembridgensis Kirejtshuk, sp. nov. from Byrrhidae; Paralichas striatopunctatus Kirejtshuk, sp. nov. from Ptilodactylidae; Trixagus barclayi Kirejtshuk, sp. nov. from Throscidae; Themus (?Telephorops) polyaki Kirejtshuk, sp. nov. from Cantharidae; Attalus flexus Kirejtshuk, sp. nov. from Malachiidae; Epuraea (Epuraea) kozlovi Kirejtshuk & Kurochkin, sp. nov., Phenolia (Lasiodites) vanescens Kirejtshuk & Kurochkin, sp. nov., Prometopia europaea Kirejtshuk & Kurochkin, sp. nov., Cyllodes argutus Kirejtshuk & Kurochkin, sp. nov. and Coxollodes palaeogenicus Kirejtshuk & Kurochkin, sp. nov. from Nitidulidae; Telmatophilus britannicus Kirejtshuk & Kurochkin, sp. nov. from Cryptophagidae; Corticariites kozlovi Kirejtshuk, sp. nov. from Latridiidae; Orthoperites antiquus Kirejtshuk & Kurochkin, sp. nov. from Corylophidae; Octotemnites sepultus Kirejtshuk, sp. nov. from Ciidae; Cyclodinus efficax Kirejtshuk, sp. nov. from Anthicidae; and Plateumaris robustus Kurochkin & Kirejtshuk, sp. nov., Plateumaris rubiconis Kurochkin & Kirejtshuk, sp. nov. and Plateumaris wightensis Kurochkin & Kirejtshuk, sp. nov. from Chrysomelidae. Byrrhites Kirejtshuk, gen. nov., Corticariites Kirejtshuk, gen. nov., Davidraenites Kirejtshuk, gen. nov., Eolimnebius Kirejtshuk, gen. nov., Hydraenites Kirejtshuk, gen. nov., Metacoxites Kirejtshuk, gen. nov., Octotemnites Kirejtshuk, gen. nov. and Orthoperites Kirejtshuk & Kurochkin, gen. nov. are proposed as taxa, partly as formal ones with generic rank and include species described herein. For Plateumaris rubiconis sp. nov., a new subgenus Necrodexis Kurochkin & Kirejtshuk, subgen. nov. is proposed. A brief review of the published fossil records for the groups considered in the paper is made. The probable ecological circumstances of the lives of the groups and species considered are discussed in the paper, and comparison with other Palaeogene sites and some conclusions on probable climatic circumstances have been elaborated. The taxonomic interpretation of three fossil species from the Caenozoic is reconsidered. It is shown that the genus Miocyphon Wickham, 1914 can be scarcely regarded as a close relative of representatives of either Dascillidae or Scirtidae. ‘Phenolia' incapax Scudder, 1890 and Lithomacratria Wickham, 1914 are regarded here without family attribution, the first as a member of Cucujiformia (i.e., out of Nitidulidae) and the latter as a member of the superfamily Tenebrionoidea (i.e., out of both Anthicidae and Pyrochroidae).