Taxonomic identification methods

Prior to this study, most of the Alderton Hill insects within the museum collections had been given a preliminary identification on a label stored with the specimen (largely by Edmund Aleksander Jarzembowski in the collections of NHMUK and CHAGM), but less than 15% of the specimens have previously been taxonomically described in the literature [see 66–72]. Within this study, through direct observation and high-resolution photographs, the taxonomy of each specimen was reviewed. This review included an update of the taxonomic nomenclature, confirmation of preliminary identifications, and where possible, refinement to a finer taxonomic resolution. The Strawberry Bank insect material, held within the collections of TTNCM and originally identified by Andrew J. Ross in Williams et al. [41] was also re-examined as part of this study, and in some cases, the taxonomy of these insects was refined.

For both the Alderton Hill and Strawberry Bank collections, every fragment that could be identified as insect-derived was recorded in order to reduce bias during identification. No additional specimen preparation was undertaken. Incomplete or poorly preserved material that could not confidently be assigned to a particular insect taxon was labelled ‘indeterminate’. Following open nomenclature convention [110], where the taxonomic identification of a specimen is uncertain, a? is placed in front of the taxon name (e.g.? Dermaptera). Where possible, insects were examined with a binocular stereomicroscope at the different institutes to assess both the taxonomy and individual taphonomic grade. Following standard entomological nomenclature [111 and references therein], taxonomic identifications were predominantly based on wings and their venation.

Accurate morphological data and measurements of the preserved length and width of specimens were recorded with digital callipers to the nearest 0.1 mm. For articulated specimens, length was measured from the anterior to the posterior preserved margin of the insect, whilst width was measured across the widest section. For isolated wings, length was measured from wing tip to base, whilst width was measured across the widest dimension. However, minor vertical compression of the material during fossilization may have resulted in limited distortion, meaning that they don’t accurately reflect those in life. The orientation (dorsoventral or lateral position) of each insect was also recorded. The feeding habits of the Alderton Hill insects were determined by direct comparison with present-day taxa [e.g. 111] where possible, whereas the feeding habits of the Strawberry Bank insects could not be inferred as most specimens could not be identified to sufficient taxonomic level.

As the collections at NHMUK contained the majority of the insect material, including many well-preserved specimens, all of the Alderton Hill and Strawberry Bank insects were photographed using a Canon EOS 5DS R with a single head light source with an adjacent reflector in order to best capture the preserved wing venation. Within the smaller collections e.g. CHAGM, BRSMG and BGS, the insect material was often poorly preserved and fragmentary, and therefore only well-preserved specimens were photographed. The Strawberry Bank insects at TTNCM were not photographed due to the poor and fragmentary preservation of the majority of the material; the best preserved examples are figured in Williams et al. [41]. Detailed photographs were also taken of associated fauna from Alderton Hill within the collections of NHMUK and CAMSM (camera model: Nikon D3200 with Nikon Micro-Nikkor 40mm F/2.8 macro lens). A specimen from each insect order, family, genus and species from Alderton Hill has been incorporated into palaeontological plates using CorelDRAW Graphics Suite X8 (Figs 3–5). In each case, we have chosen the best preserved specimen within each taxa. No plates are provided for the new Strawberry Bank specimens, due to the poor preservation of the material.

Taphonomic classification method

The completeness and articulation of each insect specimen from Alderton Hill and Strawberry Bank was individually assessed using a standard semiquantitative method of grading the preservational quality (modified from [120]; Table 3). An alphanumeric code reflecting preservational quality was assigned to each individual based on the preserved morphological features, including the presence/absence of wings and appendages and their completeness, as explained below.

The specimens were first divided into three groups based on the degree of disarticulation and indeterminability (Table 3). The first group (1), including all specimens that possessed a body with forewings and/or hindwings, was then further subdivided into three secondary taphonomic grades based on the degree of completeness; grades ranged from 1.1 through 1.3, meaning that specimens assigned to 1.1 were of the highest quality of preservation and 1.3 the poorest. The second group (2) included four subgroups: isolated bodies (2a), forewing/s (2b), hindwings (2c) and unassigned wings (2d). Each subgroup was then further divided into two taphonomic grades, designated .1 and .2, based on the degree of completeness of each insect element (Table 3). Unidentifiable insect fragments could not be used for further analysis and therefore were assigned to the third group (labelled ‘??’). Insect orders that only possessed one pair of wings e.g. Diptera were included within 2b. As many specimens possessed a part and counterpart which often displayed varying taphonomic grades, the taphonomic assessment was based on the better-preserved one. To prevent researcher bias, by which an insect could be considered better preserved because it had been previously assigned to a taxon, any associated taxonomic information was not examined until after taphonomic grades had been assigned to the specimens.

Statistical analysis

As taphonomic grades can be influenced by the anatomy of certain insects creating a preservational bias, a Chi-squared statistical test was performed in PAST 4.03 [121] to quantify and compare the association between each of the taphonomic grades and order diversity. In order to determine the degree to which each primary taphonomic grade contributed to the total Chi-squared score at order level, Pearson residuals were calculated for each entry in the Chi-squared contingency table at the order level using the statistics programme R (version 4.2.0). In order to determine how representative the fossil insect assemblages are at all taxonomic ranks at Alderton Hill and Strawberry Bank, individual-based rarefaction analyses were carried out at order-, family-, genus- and species-level using PAST 4.03 [121]. This allowed taxonomic richness at different sample sizes to be meaningfully compared [122] and assessment of whether sampling was sufficient to characterise the fossil community at each locality and taxonomic level. Standard errors (square roots of resampling variances) are converted to 95 percent confidence intervals in PAST 4.03 [121].

Alderton Hill palaeoentomofauna

Class Insecta Linnaeus, 1758 [118]

Subclass Pterygota Brauer, 1885 [123]

Order Ephemeroptera Hyatt & Arms, 1890 [124]

Assigned material.–WARMS G 8084.

Remarks.–Specimen was not available for re-examination (see materials and methods). WARMS G 8084 was originally reported as “Ephemera” by Kelly [109], however this determination is uncertain, as only two undescribed ephemeropteran wings are known from the Toarcian insect-bearing localities of Grimmen and Dobbertin, Germany.

Order Odonata Fabricius, 1793 [125]

Suborder Anisozygoptera Handlirsch, 1906 [115]

Family Heterophlebiidae Needham, 1903 [126]

Genus Heterophlebia Westwood, 1849 [127]

Heterophlebia buckmani (Brodie, 1845) [62]

(Fig 3A)

Holotype.–NHMUK PI I.11343, isolated partial forewing (33.8 mm long, 8.4 mm wide). Paratype.–NHMUK PI I.3988; holotype of Heterophlebia dislocata Westwood, 1849 [127] (missing from collections, unable to be re-examined).

Assigned material.–NHMUK PI I.11310 & NHMUK PI I.11344, holotype of Heterophlebia tillyardi Handlirsch, 1939 [128]; isolated forewing, part and counterpart (33.8 mm long, 7.3 mm wide); BRSMG Cg2370 & BRSMG Cg2371, isolated partial forewing, part and counterpart (26 mm long, 8.4 mm wide); NHMUK PI I.59404 & NHMUK PI I.10496, isolated left hindwing, part and counterpart (35.8 mm long, 8.6 mm wide); NHMUK PI I.11312, holotype of Heterophlebia angulata Tillyard, 1925 [69]; isolated left partial male hindwing (20.7 mm long, 7.7 mm wide); NHMUK PI I.3317, two wing fragments (unable to be measured).

Remarks.–All additional material is fragmentary and the venation in these wing fragments is identical to those of other specimens attributed to Heterophlebia buckmani [see 73]. See [129] for an extensive synonymy list for H. buckmani.

References.–[69, 73, 115, 128, 129].

Heterophlebia sp.

Assigned material.–CHAGM F.651, partial wing fragment (12.1 mm long, 8.1 mm wide); CHAGM F.654, wing tip fragment (12.3 mm long, 8.8 mm wide).

Family Liassogomphidae Tillyard, 1935 [130]

Genus Heterothemis Handlirsch, 1906 [115]

Heterothemis brodiei (Buckman, 1843) [60]

(Fig 3B–3C)

Holotype.–NHMUK PI I.3552, isolated left female hindwing, lectotype selected as holotype (45 mm long, 13.9 mm wide).

Assigned material.–BGS Geol.Soc.Coll. 4126, isolated forewing apex (19.6 mm long, 3.4 mm wide); NHMUK PI I.11288 & NHMUK PI I.11269, isolated partial right hindwing, part and counterpart (31.5 mm long, 11 mm wide); CHAGM F.653, isolated partial basal hindwing (21.3 mm long, 12.6 mm wide); CHAGM F.657.1 & CHAGM F.657.2, partial wing fragment, part and counterpart (27.8 mm long, 8.7 mm wide); WARMS G 8079 (unable to be measured).

Remarks.–WARMS G 8079 was not available for examination but was reported as Heterothemis brodiei by Kelly [109]. All additional material is fragmentary and the venation in these wing fragments is identical to those of other specimens attributed to Heterothemis brodiei [see 73].

References.–[58, 73, 115, 128, 131].

Heterothemis sp.

Assigned material.–CHAGM F.650, partial wing fragment (13.3 mm long, 11.7 mm wide); CHAGM F.655, basal wing fragment (20.9 mm long, 5 mm wide); CHAGM F.662, partial wing tip (13.6 mm long, 4.6 mm wide); PALE-8749, partial hindwing fragment (unable to be measured).

Remarks.–PALE-8749 was originally reported as? Liassogomphus in the database of the Zoological Collections, Museum of Comparative Zoology, Harvard University.

Suborder Archizygoptera Handlirsch, 1906 [115]

Family Protomyrmeleontidae Handlirsch, 1906 [115]

Genus Protomyrmeleon Geinitz, 1887 [112]

Protomyrmeleon sp.

(Fig 3D)

Assigned material.–NHMUK PI In.59065, isolated partial supposed hindwing (6.8 mm long, 2.5 mm width).

Remarks.–NHMUK PI In.59065 possess venation identical to those of other specimens attributed to Protomyrmeleon [see 73].

References.–[132, 133].

Family? Protomyrmeleontidae Handlirsch, 1906 [115]

gen. et sp. indet

Assigned material.–NHMUK PI I.11350, isolated partial wing (5.7 mm length, 2 mm width).

Family incertae sedis

gen. et sp. indet

Assigned material.–Isolated partial forewing: BRSMG Cg2359. Isolated partial wings: NHMUK PI I.3330; NHMUK PI In.59078 & NHMUK PI In.59080, part and counterpart; CHAGM F.630; CHAGM F.652; CHAGM F.681; NHMUK PI In.59075; NHMUK PI I.11347 & NHMUK PI I.11375 (part and counterpart); CAMSM TN 3193.

Remarks.–Attributed to Odonata based on the dense cellular wing venation, but further taxonomic determination to family or genus level cannot be made due to fragmentary preservation.

Order? Dermaptera De Geer, 1773 [113]

Family incertae sedis

(Fig 3E)

Assigned material.–NHMUK PI I.11403, isolated elytron (5.6 mm long, 3.5 mm wide).

Remarks.–Only tentatively assigned to this order due to poor preservation of the specimen.

Order Reculida Handlirsch, 1906 [115]

Family Geinitziidae Handlirsch, 1906 [115]

Genus Geinitzia Handlirsch, 1906 [115]

Geinitzia carpentieri Zeuner, 1937 [71]

(Fig 3F)

Holotype.–NHMUK PI In.36200, isolated partial forewing, part and counterpart (18.4 mm long, 7.2 mm wide).

Assigned material.–BRSMG Cg2364 & BRSMG Cg2365, isolated partial forewing, part and counterpart (9.8 mm length and 7.8 mm width).

Remarks.–Additional material attributed to Geinitzia carpentieri based on species characteristics described by Zeuner [71], including: MA₁ with terminal fork; CuA₂ not undulated.

References.–[45, 47, 71, 115, 128].

Superorder Dictyoptera Latreille, 1829 [134]

Order Blattodea Latreille, 1810 [114]

Family? Raphidiomimidae Vishnyakova, 1973 [135]

Genus Liadoblattina Handlirsch, 1906 [115]

Liadoblattina blakei (Scudder, 1886) [67]

(Fig 3G)

Holotype.–NHMUK PI I.3574, isolated tegmen (14.6 mm long, 5 mm wide).

Assigned material.–NHMUK PI I.11361, tegmen and pronotum (2.5 mm long, 1.5 mm wide); NHMUK PI I.3562, holotype of Mesoblattina bensoni Scudder, 1886 [67], isolated tegmen (18 mm long, 5 mm wide); NHMUK PI I.11341, isolated partial tegmen, part and counterpart (16.1 mm long, 5 mm wide).

Remarks.–Attributed to Liadoblattina blakei based on species characteristics described by Vršanský and Ansorge [75], including an elongate tegmen, which is up to three times as long (c. 20 mm) as it is wide, and a clavus that is more than twice as long as it is wide.

References.–[67, 75, 115, 128].

Family uncertain

undescribed(Fig 3H)

Assigned material.–BRSMG Cg2374 & BRSMG Cg2375, isolated tegmen, part and counterpart (12.8 mm long, 3.8 mm wide).

Remarks.–Our observations show that this specimen is noticeably dissimilar to other known Toarcian cockroaches and represents a new genus and species. The family placement of this specimen is uncertain at present.

Family incertae sedis

gen. et sp. indet

Assigned material.–CHAGM F.660, isolated partial wing (6.8 mm long, 2 mm wide).

Remarks.–Attributed to Blattodea based on the tegmen venation, which involves numerous closely spaced parallel veins, however further taxonomic determination to family or genus level cannot be made due to poor venation preservation.

Order Orthoptera Olivier, 1789 [117]

Family Locustopsidae Handlirsch, 1906 [115]

Genus Locustopsis Handlirsch, 1906 [115]

Locustopsis sp.

(Fig 3I)

Assigned material.–Isolated forewings: NHMUK PI I.11303; NHMUK PI I.11370. Isolated partial forewings: NHMUK PI I.11334; NHMUK PI I.11330; NHMUK PI I.11363; NHMUK PI I.11351. Isolated hindwing: NHMUK PI I.11340.

Remarks.–Specimens possess venation and colour patterns similar to those of other specimens attributed to the genus Locustopsis [see 136].

References.–[115, 128, 136, 137].

Family Locustopsidae Handlirsch, 1906 [115]

gen. et sp. indet

Assigned material.–BGS GSM 117360, isolated partial wing (8.7 mm long, 2.5 mm wide).

Family Elcanidae Handlirsch, 1906 [115]

Genus “Elcana” Giebel, 1856 [138]

Remarks.–Gorochov et al. [139] synonymized Elcana Giebel, 1856 [138] with Panorpidium Westwood, 1854 [65] as both generic names were based on the same specimen, the type species of P. tesselatum Westwood, 1854 [65]. Gorochov et al. [139] also divided Elcanidae in two subfamilies: Elcaninae Handlirsch 1906 [115] [= Baisselcaninae Gorochov, 1986 [140]] and Archelcaninae Gorochov, Jarzembowski & Coram, 2006 [139].

The numerous species of Elcana described by earlier authors [see 115, 141, 142] are in need of taxonomic revision [139]. Although a possible generic name for Early Jurassic elcanids is Clathrotermes Heer, 1865 [143], here we use the name “Elcana” as a placeholder until a full revision of this genus is undertaken. At species level, we follow Zessin [142] in his treatment of Early Toarcian elcanid species, who reduced more than 100 Lower Toarcian species described by Handlirsch [115, 128] and Bode [45, 47] from Dobbertin and the vicinity of Braunschweig to four: “Elcana” minima Handlirsch, 1906 [115], “Elcana” geinitzi (Heer, 1880) [116], “Elcana” media Handlirsch, 1906 [115] and “Elcana” magna Handlirsch, 1906 [115]. Recently, Heads et al. [144] described two new elcanid species from the Lower Toarcian of Luxembourg as Archelcana tina Heads et al., 2022 [144] and A. numbergerae Heads et al., 2022 [144]. However, these species may prove to be junior synonyms.

“Elcana” cf. geinitzi (Heer, 1880) [116]

(Fig 3J–3K)

Holotype.–MBI 15.1 a,b, isolated forewing. Grimmen Formation, ‘Green Series’; Early Jurassic: Lower Toarcian of Dobbertin, Mecklenburg-Vorpommern, Germany.

Assigned material.–NHMUK PI I.11427, isolated partial forewing (8.1 mm long, 2.1 mm wide); NHMUK PI I.11295, isolated partial forewing (10 mm long, 2.8 mm wide); NHMUK PI I.3349 (specimen was not available to be measured); NHMUK PI I.3560, holotype of Elcana brodiei (Handlirsch, 1906) [115], isolated partial forewing (7.2 mm long, 2.6 mm wide).

Remarks.–NHMUK PI I.3349 was reported as Archelcana geinitzi by Kelly [109].

References.–[46, 115, 116, 128, 142, 145].

“Elcana” sp.

Assigned material.–NHMUK PI I.11394, isolated partial forewing tip (5.1 mm long, 2 mm wide); NHMUK PI I.38, part and counterpart (9.2 mm long, 4.5 mm wide)

Family Protogryllidae Zeuner, 1937 [71]

Genus Protogryllus Handlirsch, 1906 [115]

Protogryllus cf. dobbertinensis (Geinitz,1880) [116]

Holotype.–FGWG 117/7, isolated forewing. Grimmen Formation, ‘Green Series’; Early Jurassic: Lower Toarcian of Dobbertin, Mecklenburg-Vorpommern, Germany.

Assigned material.–NHMUK PI I.11421, isolated partial male forewing (16.4 mm long, 5.3 mm wide).

References.–[44, 47, 71, 115, 128].

Protogryllus magnus Zeuner, 1937 [71]

(Fig 3L)

Holotype.–NHMUK PI I.11324, isolated partial male forewing (27 mm long, 5.7 mm wide).

References.–[71].

Protogryllus sp.

Assigned material.–NHMUK PI I.11377, partial body with wings (17.7 mm long, 2.7 mm wide); NHMUK PI I.11461, isolated partial wing (12.3 mm long and 3.7 wide).

Family incertae sedis

gen. et sp. indet

(Fig 3M)

Assigned material.–Almost complete individuals: NHMUK PI I.11270. Isolated exoskeletal elements: NHMUK PI I.3314; NHMUK PI I.11321; NHMUK PI I.11405; NHMUK PI I.11333; NHMUK PI I.11392. Isolated partial wings: BGS GSM 117359; BGS GSM 117361; NHMUK PI I.3305; NHMUK PI I.11301; NHMUK PI I.11290; NHMUK PI I.11396; NHMUK PI I.3304; NHMUK PI I.11264. Indeterminable: NHMUK PI I.11342; NHMUK PI I.11332; NHMUK PI I.59084.

Remarks.–Attributed to Orthoptera based on the tegminised forewings and distinctive colouration patterns. Further determination to a lower taxonomic level cannot be made due to fragmentary wing preservation.

Order Hemiptera Linnaeus, 1758 [118]

Suborder Fulgoromorpha Evans, 1946 [146]

Family Fulgoridiidae Handlirsch, 1939 [128]

Genus Fulgoridium Handlirsch, 1906 [115]

Fulgoridium sp.

(Fig 4A)

Assigned material.–Isolated skeletal elements: NHMUK PI I.11424. Isolated forewings: NHMUK PI I.3331 & NHMUK PI I.3334, part and counterpart; NHMUK PI I.11328. Isolated partial forewing: NHMUK PI I.3079. Isolated partial wings: NHMUK PI I.11275; NHMUK PI I.11293; NHMUK PI I.11315. Indeterminable: NHMUK PI I.11278; NHMUK PI I.11453.

Remarks.–Specimens NHMUK PI I.11278 and NHMUK PI I.11453 were not available for re-examination but were reported as Fulgoridium by Kelly [109]. From the Toarcian, only the genera Fulgoridulum Handlirsch, 1939 [128], Margaroptilon Handlirsch, 1906 [115], and Elasmoscelidium Martynov, 1926 [147] are currently well defined. Until a revision of all species of Fulgoridium is undertaken, it is impossible to determine the taxonomy of the Alderton Hill specimens more precisely.

References.–[44, 47, 115, 128, 147].

Genus Margaroptilon Handlirsch, 1906 [115]

Margaroptilon brodiei Handlirsch, 1906 [115]

(Fig 4B, 4C)

1906 Margaroptilon brodiei Handlirsch: 499

1906 Margaroptilon woodwardi Handlirsch: 499 syn. nov.

1906 Margaroptilon bulleni Handlirsch: 499 syn. nov.

Holotype.–NHMUK PI I.3561, spotted isolated partial forewing (6 mm long, 2.1 mm wide); NHMUK PI I.11266 (holotype of M. bulleni) & NHMUK PI I.11286 (holotype of M. woodwardi), isolated forewing, part and counterpart (6.3 mm long, 2.5 mm wide).

Assigned material.–Isolated forewings: CHAGM F.642; CHAGM F.664; NHMUK PI I.11296. Isolated partial forewings: CHAGM F.659.1 & CHAGM F.659.2 (part and counterpart); CHAGM F.661; CHAGM F.635; NHMUK PI I.11374; NHMUK PI I.11399; NHMUK PI I.11414; NHMUK PI I.11426; NHMUK PI I.11429; NHMUK PI I.15015; NHMUK PI I.3341; NHMUK PI I.11268; NHMUK PI I.11415. Indeterminable: NHMUK PI I.3309.

Remarks.–Following the suggestion in Kelly’s PhD thesis [109], here we have synonymized Margaroptilon brodiei Handlirsch, 1906 [115]; Margaroptilon bulleni Handlirsch (1906) [115]; and Margaroptilon woodwardi Handlirsch, 1908 [115] based on morphological similarity, with the latter being the senior synonym. He recognized that H. woodwardi and M. bulleni were based on part and counterpart of the same wing. We regard the small differences in spot arrangement between M. woodwardi and M. brodiei as intraspecific variation, as Kelly [109] did. However, the name M. brodiei has priority instead of M. woodwardi. NHMUK PI I.3309 was not available for re-examination but was reported as Margaroptilon woodwardi by Kelly [109]. Margaroptilon is a planthopper with faint venation typical for fulgoromorphans, although it is difficult to study as the well sclerotized wing spots resemble those of Fulgoridulum.

References.–[46, 109, 115].

Family Fulgoridiidae Handlirsch, 1939 [128]

gen. et sp. indet.

Assigned material.–CHAGM F.636, isolated partial wing tip (7 mm long, 3.1 mm wide); CAMSM TN 3195, partial spotted wing (5.6 mm long, 2.7 mm wide).

Family Archijassidae Becker-Migdisova, 1962 [148]

Genus Archijassus Handlirsch, 1906 [115]

Archijassus heeri (Geinitz, 1880) [116]

(Fig 4D)

Assigned material.–NHMUK PI I.11458, isolated forewing (specimen not available for re-examination).

References.–[44, 116].

Suborder Coleorrhyncha Myers and China, 1929 [149]

Family Progonocimicidae Handlirsch, 1906 [115]

gen. et sp. indet.

(Fig 4E)

Assigned material.–NHMUK PI I.11274, isolated partial body (3.3 mm long, 1.1 mm wide); NHMUK PI I.11465, isolated forewing (4 mm long).

Suborder Heteroptera Latreille, 1810 [114]

Infraorder Gerromorpha Popov, 1971 [150]

Genus Engynabis Bode, 1953 [46]

Engynabis tenuis Bode, 1953 [46]

(Fig 4F)

Assigned material.–NHMUK PI I.3313, elongated body with paired wings (5.3 mm long, 1.2 mm wide).

References.–[46, 151].

Family incertae sedis

gen. et sp. indet.

(Fig 4G–4I)

Assigned material.–Partial individuals: NHMUK PI I.11362; NHMUK PI I.11446; CHAGM F.643.1 & CHAGM F.643.2. Isolated bodies: NHMUK PI I.11280; NHMUK PI I.11294; NHMUK PI I.11297; NHMUK PI I.11317; NHMUK PI I.82 & NHMUK PI I.74 (part and counterpart); NHMUK PI I.11379. Isolated skeletal elements: NHMUK PI I.11454; NHMUK PI I.3052; NHMUK PI I.11285; NHMUK PI I.11456; NHMUK PI I.11452; NHMUK PI I.3324; NHMUK PI I.68; NHMUK PI I.11445; BGS GSM 117358; CHAGM F.634.1 & CHAGM F.634.2 (part and counterpart). Isolated forewing: NHMUK PI I.78. Isolated partial hindwings: NHMUK PI I.11439. Isolated partial wings: NHMUK PI I.3342; CHAGM F.671 & CHAGM F.672 (part and counterpart); CHAGM F.670.1 & CHAGM F.670.2 (part and counterpart). Indeterminate: NHMUK PI In. 59079.

Order Coleoptera Linnaeus, 1758 [118]

gen. et sp. indet.

(Fig 4J–4M)

Assigned material.–Almost complete individuals: CHAGM F.645; CHAGM F.646; NHMUK PI I.11339; NHMUK PI I.11360. Partial individuals: CHAGM F.638; CHAGM F.647.1 & CHAGM F.647.2; CHAGM F.649.1 & CHAGM F.649.2; NHMUK PI I.3307; NHMUK PI I.11345; NHMUK PI I.11383; CAMSM TN 3196 a+b. Isolated bodies: NHMUK PI I.11267; NHMUK PI I.11287; NHMUK PI I.11292. Isolated skeletal elements: NHMUK PI I.69; NHMUK PI I.3308; NHMUK PI I.11302; NHMUK PI I.11404; NHMUK PI I.81. Isolated forewings: CHAGM F.640; CHAGM F.644; CHAGM F.663; CHAGM F.667; NHMUK PI I.3052; NHMUK PI I.3077; NHMUK PI I.3078; NHMUK PI I.11318; NHMUK PI I.11331; NHMUK PI I.11353 & NHMUK PI I.11354; NHMUK PI I.11355 & NHMUK PI I.11376; NHMUK PI I.11359; NHMUK PI I.11369; NHMUK PI I.11411; NHMUK PI I.11420; NHMUK PI I.11431; NHMUK PI I.11432; NHMUK PI I.11438; NHMUK PI I.11441; NHMUK PI I.59071. Isolated partial forewings: CHAGM F.639; CHAGM F.641; NHMUK PI I.3335; NHMUK PI I.11368. Indeterminable: NHMUK PI I.2163; NHMUK PI I.3339; NHMUK PI I.3348 & NHMUK PI I.11308 (part and counterpart); NHMUK PI In. 59072; NHMUK PI In.59286; NHMUK PI In.59285; NHMUK PI In.59287; NHMUK PI In.59289; NHMUK PI In.59294; NHMUK PI I.11381.

Remarks.–Attributed to Coleoptera based on the forewings which are modified into highly sclerotised elytra. However, the majority of the Coleoptera are represented by disarticulated and fragmented elytra which are compressed, poorly preserved, and therefore difficult to identify beyond order level. Therefore, all beetles specimens from Alderton Hill remain unassigned, and no types are based on the recorded specimens. However, at least three elytra morphotypes can be identified, including smooth (NHMUK PI I.11345; Fig 4K), striated (NHMUK PI I.3077; Fig 4L) and pitted (NHMUK PI I.11355 and NHMUK PI I.11376; Fig 4M), indicating that the beetle fauna is likely to consist of several genera or species. The thoracic fragment NHMUK PI I.3052 was originally assigned to the recent genus Coccinella Linnaeus, 1758 [118] based on the associated specimen label, however this identification cannot be supported.

Order Necrotrichoptera Engel, 2022 [152]

Family Necrotauliidae Handlirsch, 1906 [115]

Genus Necrotaulius Handlirsch, 1906 [115]

Necrotaulius parvulus (Geinitz, 1884) [66]

(Fig 4N –4O)

Holotype.–FGWG 119/7, isolated wing. Grimmen Formation, ‘Green Series’; Early Jurassic: Lower Toarcian of Dobbertin, Mecklenburg-Vorpommern, Germany.

Assigned material.–NHMUK PI I.15016 & NHMUK PI I.15014 (part and counterpart) isolated forewing, holotype of Necrotaulius pygmaeus Tillyard, 1933 [70] (3.2 mm long, 0.9 mm wide); NHMUK PI I.11425, isolated forewing (2.8 mm long, 0.7 mm wide); NHMUK PI I.3336, isolated body (3.8 mm long, 1.8 mm wide); NHMUK PI I.11389 (specimen not available for re-examination).

Remarks.–Necrotaulius parvulus is distinct due to the dense coverage of the wing with hairs. Wing venation of this species is similar to Trichoptera, however it has no distally bent CuP, a synapomorphic character of Trichoptera [77, 153]. Because of this CuP character, we here exclude Austaulius Thomson, Ross and Coram, 2018 [43] from Necrotauliidae and transfer it to Trichoptera incertae familiae.

References.–[66, 77, 109, 112, 128, 153–156].

?Necrotaulius sp.

Assigned material.–CHAGM F.637, isolated partial wing (specimen unable to be measured).

Order Mecoptera Packard, 1886 [157]

Family Bittacidae Handlirsch, 1906 [115]

Genus Protobittacus Tillyard, 1933 [70]

Protobittacus handlirschi Tillyard, 1933 [70]

(Fig 4P)

Holotype.–NHMUK PI I.11406 & NHMUK PI I.11335, isolated hindwing, part and counterpart (19.4 mm long, 4.3 mm wide).

Remarks. During our investigation of the Alderton Hill insect material, we located the unknown counterpart of the holotype (NHMUK PI I.11335), which displays most of the proximal part of the wing including the unique type of A1/A2 fork (Fig 4P).

References.–[70].

Bittacidae gen. et sp. indet.

Assigned material.–CHAGM F.632, isolated wing (8.5 mm long, 2 mm wide).

Family Orthophlebiidae Handlirsch, 1906 [115]

Genus Orthophlebia Westwood, 1845 [62]

Orthophlebia brodiei Tillyard, 1933 [70]

(Fig 4Q)

Holotype.–NHMUK PI I.15019, isolated forewing (10.7 mm long, 3 mm wide).

Remarks.–Note that Tillyard 1933 [70] referred to the holotype specimen as NHMUK PI I.15017; see Tillyard [70] for the original holotype description.

References.–[44, 70, 160, 161].

Orthophlebia sp.

(Fig 4R)

Assigned material.–Isolated forewings: NHMUK PI I.11413; NHMUK PI I.11418 & NHMUK PI I.11395 (part and counterpart); NHMUK PI I.11225; NHMUK PI I.11387. Isolated partial forewing: NHMUK PI I.11271. Isolated hindwing: NHMUK PI I.15013. Isolated partial hindwing: NHMUK PI I.11366. Isolated wing: NHMUK PI I.11448. Isolated partial wing: CAMSM TN 3192.

References.–[44, 158, 159].

Family Pseudopolycentropodidae Handlirsch, 1920 [160]

Genus Pseudopolycentropus Handlirsch, 1906 [115]

Pseudopolycentropus sp.

Assigned material.–CHAGM F.656, isolated forewing (5.4 mm long, 2 mm wide).

References.–[44, 47, 115, 128, 161].

Order Neuroptera Linnaeus, 1758 [118]

Family Osmylopsychopidae Martynova, 1949 [158]

Genus Actinophlebia Handlirsch, 1906 [115]

Actinophlebia intermixta (Scudder, 1885) [119]

(Fig 5A, 5B)

Holotype.–NHMUK PI I.3577 & NHMUK PI I.11346, isolated forewing, part and counterpart (9.2 mm length, 5.5 mm width). Actinophlebia anglicana Tillyard, 1933 [70] (NHMUK PI I.11346) is a counterpart of Actinophlebia intermixta (Scudder, 1885) [119], an objective synonym of the latter.

Assigned material.–NHMUK PI I.11349, isolated forewing (6.3 mm length and 4.1 mm width); NHMUK PI I.11319, isolated partial forewing (7.3 mm length and 4.5 mm width).

Remarks.–Neuropterans from the Toarcian with a triangular wing shape are often attributed to the genus Actinophlebia [44, 68, 70, 115]. However, Ponomarenko [162] regarded similar wings with a distally fused Sc with RA as belonging to Parhemerobius Bode, 1953 [46]. In Actinophlebia, the Sc appears to run free into the costal margin. This, however, is most likely based on a misinterpretation of the course of Sc, as this featured can be observed in the Actinophlebia sp. specimen LGA 1080 from the Lower Toarcian of Grimmen, Germany [see 45]. Note that Tillyard [70] originally measured NHMUK PI I.11346 at 11.2 mmm long, while Scudder [119] measured a length of 10 mm.

References.–[44, 47, 68, 70, 115, 119, 128, 162–165].

Actinophlebia sp.

Assigned material.–NHMUK PI I.11314, isolated partial wing (6.8 mm length, 4.4 mm width).

“Familia nova A” Makarkin et al., 2012 [166]

Genus Paractinophlebia Handlirsch, 1906 [115]

Paractinophlebia curtisii (Scudder, 1886) [67]

(Fig 5C)

Holotype.–NHMUK PI I.3585, isolated partial hindwing (27.1 mm long, 7.9 mm wide); the counterpart (PALE-8751) is housed in the Entomology collection at the Museum of Comparative Zoology, Harvard University, and assigned as Neuroptera sp.

Assigned material.–NHMUK PI I.11320, isolated partial wing fragment (17.9 mm long, 9 mm wide).

Remarks.–Scudder [67] figured the counterpart (plate 48, Fig 16) at PALE. The holotype is most likely a hindwing fragment, which is strongly dissimilar to the wings of Kalligrammatidae [see 68], in particular by the presence of rare cross-veins arranged in series and by the very narrow subcostal space [167].

References.–[67, 166, 167].

Family “Prohemerobiidae”Handlirsch, 1906 [115]

Genus “Prohemerobius”Handlirsch, 1906 [115]

“Prohemerobius”aldertonensis Whalley, 1988 [68]

(Fig 5D)

Holotype.–NHMUK PI I.11304, isolated partial forewing (4 mm long, 1.5 mm wide).

Assigned material.–NHMUK PI I.11279, isolated partial wing (5.7 mm long, 2.7 mm wide); NHMUK PI I.11390, isolated partial wing (6.6 mm long, 1.8 mm wide).

Remarks.–The genus Prohemerobius Handlirsch 1906 [115] is regarded as a waste basket taxon for small Lower Jurassic neuropterans, and therefore requires a complete revision. Ponomarenko, 1995 [161] regards the English Lower Toarcian species Archeosmylus alysius Whalley, 1988 [68] and A. complexus Whalley, 1988 [68] as belonging to Prohemerobius. The only difference is whether the Sc and RA run parallel or strongly converge. We refer here the assignment of specimen to the certain taxa as provided by Whalley [68].

References.–[68, 162, 168].

“Prohemerobius” sp.

Assigned material.–NHMUK PI I.11284, isolated wing (7.4 mm length, 3.3 mm width); NHMUK PI Ig 60, isolated wing (9.1 mm length, 2.4 mm width); NHMUK PI I.11417, partial individual with wings (specimen was not available for measurement).

“Archeosmylus” alysius Whalley, 1988 [68]

(Fig 5E)

Holotype.–NHMUK PI I.3318, body (2.3 mm long, 0.5 mm wide) with attached wings (4.3 mm long, 1.7 mm wide), part and counterpart.

Paratypes.–NHMUK PI I.11416, isolated wing (5 mm long, 2.5 mm wide); NHMUK PI I.11433, partial individual with paired wings (3.5 mm long, 1.9 mm wide).

Assigned material.–NHMUK PI I.11281, isolated wing (3.7 mm long, 1.8 mm wide); NHMUK PI I.11402, isolated partial wing (6.5 mm long, 1.7 mm wide); NHMUK PI I.11347 & NHMUK PI I.11375, partial individual, part and counterpart (specimen could not be measured).

References.–[68, 162].

“Archeosmylus” complexus Whalley, 1988 [68]

(Fig 5F)

Holotype.–NHMUK PI I.11412, isolated wing (6.7 mm long, 3.7 mm wide).

Paratype.–NHMUK PI I.3315 & NHMUK PI I.3380, isolated wing, part and counterpart (9.1 mm, 2.1 mm wide).

References.–[68, 162].

Family “Prohemerobiidae”Handlirsch, 1906 [115]

gen. et sp. indet.

Assigned material.–BRSMG Cd 1397, isolated partial wing (9 mm long, 4.7 mm wide).

Family incertae sedis

gen. et sp. indet.

Assigned material.–Partial individual: NHMUK PI I.11409. Isolated wings: NHMUK PI I.66 & NHMUK PI I.66-83, part and counterpart; NHMUK PI I.11385. Isolated partial wings: NHMUK PI I.3337; NHMUK PI I.11326; NHMUK PI I.11401; NHMUK PI I.11463; NHMUK PI I.11464. Indeterminable: WARMS G 8087; NHMUK PI I.11434.

Remarks.–WARMS G 8087 was not available for re-examination but was reported as Neuroptera by Kelly [109]. See supplementary material for individual measurements and descriptions.

Order Diptera Linnaeus, 1758 [118]

Suborder Nematocera Latreille, 1825 [169]

Family Limoniidae Speiser, 1909 [170]

Genus Architipula Handlirsch, 1906 [115]

Architipula anglicana (Tillyard, 1933) [70]

(Fig 5G)

Holotype.–NHMUK PI I.11298, isolated wing (8.2 mm long, 2.2 mm wide).

Assigned material.–NHMUK PI I.11272, isolated wing (5.9 mm long, 2 mm wide); NHMUK PI I.11365, isolated wing (7.3 mm long, 2.5 mm wide); NHMUK PI I.11313, isolated partial wing (6.5 mm long, 2.6 mm wide); NHMUK PI I.11398, isolated partial wing (8.7 mm long, 1.3 mm wide).

Remarks.–Note that NHMUK PI I.11298 was formerly described within Liassotipula Tillyard, 1933 [70], which was then synonymized with Architipula by Kopeć et al. [76]. Venation of additional material is identical to other specimens attributed to Architipula anglicana as described by Kopeć et al. [76].

References.–[70, 76, 115].

Architipula sp.

Assigned material.–CHAGM F.633, isolated wing (6.2 mm long, 1.7 mm wide).

Genus Grimmenia Krzemiński and Zessin, 1990 [171]

Grimmenia tillyardi Kopeć et al., 2017b [76]

(Fig 5H)

Holotype.–NHMUK PI I.3328, isolated wing (3.8 mm long, 0.9 mm wide).

References.–[76].

Genus Mesotipula Handlirsch, 1920 [160]

Mesotipula slatteri Kopeć et al., 2018 [172]

Holotype.–NHMUK PI I.11316 & NHMUK PI I.11311, isolated female imago, part and counterpart (5.7 mm long, 1.9 mm wide).

Assigned material.–NHMUK PI I.3320 & NHMUK PI I.3323, isolated partial wing, part and counterpart (6.4 mm long, 1.7 mm wide).

Remarks.–Venation of additional material is identical to other specimens attributed to Mesotipula slatteri as described by Kopeć et al. [172].

References.–[172].

Family Limoniidae Speiser 1909 [170]

gen. et sp. indet

Assigned material.–Partial individual: NHMUK PI I.11329. Isolated skeletal elements: NHMUK PI I.71; NHMUK PI I.11328; NHMUK PI I.11348; NHMUK PI I.11380; NHMUK PI I.11391; NHMUK PI I.3319; NHMUK PI I.3345; NHMUK PI I.3350. Isolated partial wings: NHMUK PI In.59069; NHMUK PI In.59074; NHMUK PI In.59076.

Family incertae sedis

gen. et sp. indet

Assigned material.–NHMUK PI I.11276, isolated segmented body (5.3 mm long, 0.7 mm wide). Isolated partial wings: NHMUK PI I.73 (3.7 mm long, 1 mm wide); NHMUK PI I.11315 (7.9 mm long, 2.2 mm wide); NHMUK PI I.11358 (7.8 mm long, 0.5 mm wide).

Strawberry bank palaeoentomofauna

Class Insecta Linnaeus, 1758 [118]

Subclass Pterygota Brauer, 1885 [123]

Order Odonata Fabricius, 1793 [125]

Suborder Anisozygoptera

Family Liassogomphidae Tillyard, 1935 [130]

gen. et sp. indet.

Assigned material.–TTNCM 39/2011/0364, isolated wing fragment (10.7 mm long, 8 mm wide); TTNCM 39/2011/0509, isolated wing fragment (26.9 mm long, 7.9 mm wide).

Remarks.–TTNCM 39/2011/0509 is figured in Williams et al. [41] (Fig 3C).

References.–[41, 73].

Family incertae sedis

gen. et sp. indet.

Assigned material.–TTNCM 39/2011/0362a-b; TTNCM 39/2011/0403a-b; TTNCM 39/2011/0408; TTNCM 39/2011/0420a-b; TTNCM 39/2011/0503 & TTNCM 39/2011/0504 (part and counterpart); TTNCM 39/2011/0505; TTNCM 39/2011/0506; TTNCM 39/2011/0507a-b; TTNCM 39/2011/0511; TTNCM 39/2011/0512; TTNCM 39/2011/0514; TTNCM 39/2011/0515; TTNCM 39/2011/0516; TTNCM 39/2011/0517; TTNCM 39/2011/0528; TTNCM 39/2011/0545; TTNCM 39/2011/1013; TTNCM 39/2011/1014.

Superorder Dictyoptera Latreille, 1829 [133]

Order Blattodea Latreille, 1810 [114]

Family Caloblattinidae Vršanský and Ansorge, 2000 [173]

Genus Rhipidoblattina Handlirsch, 1906 [115]

Rhipidoblattina sp.

Assigned material.–TTNCM 39/2011/0537 & TTNCM 39/2011/0538, isolated forewing, part and counterpart (8.4 mm long, 2.8 mm wide).

References.–[173].

Family incertae sedis

gen. et sp. indet

Assigned material.–Isolated partial forewing: TTNCM 39/2011/1008. Isolated partial wings: TTNCM 39/2011/0391; TTNCM 39/2011/0392 & TTNCM 39/2011/0543 (part and counterpart); TTNCM 39/2011/0393 & TTNCM 39/2011/0395 (part and counterpart); TTNCM 39/2011/0396; TTNCM 39/2011/0398; TTNCM 39/2011/0405; TTNCM 39/2011/0410; plus an additional specimen with no accession number.

Order Dermaptera De Geer, 1773 [113]

Family Dermapteridae Vishnyakova, 1980 [174]

Genus Trivenapteron Kelly, Ross and Jarzembowski,2016 [42]

Trivenapteron moorei Kelly et al., 2016 [42]

Holotype.–TTNCM 39/2011/0489, isolated sub-quadrate, truncate tegmen, pointed apically and without ornamentation.

References.–[41, 43].

Order Orthoptera Olivier, 1789 [117]

Family Locustopsidae Handlirsch, 1906 [115]

Genus Locustopsis Handlirsch, 1906 [115]

Locustopsis sp.

Assigned material.–NHMUK PI I.11485, isolated hindwing fragment (14 mm long, 2.4 mm wide).

References.–[115, 128, 138].

Family Locustopsidae Handlirsch, 1906 [115]

gen. et sp. indet.

Assigned material.–TTNCM 39/2011/0520 & TTNCM 39/2011/0522, isolated wing fragment, part and counterpart fragment (16 mm long, 3.8 mm wide); TTNCM 39/2011/0527, isolated forewing fragment (16.5 mm long, 4.2 mm wide); TTNCM 39/2011/0731, isolated wing (13 mm long, 2.6 mm wide); CAMSM TN 3200, partial wing (7.2 mm long, 2.8 mm wide).

Family Elcanidae Handlirsch, 1906 [115]

gen. et sp. indet.

Assigned material.–Isolated forewing: TTNCM 39/2011/0523. Isolated partial forewing: TTNCM 39/2011/0424a-b; TTNCM 39/2011/0525. Isolated wing: TTNCM 39/2011/0544. Isolated partial wing: TTNCM 39/2011/0732; TTNCM 39/2011/0411; TTNCM 39/2011/0397; TTNCM 39/2011/0419; TTNCM 39/2011/0434; TTNCM 39/2011/0415. Indeterminable: TTNCM 39/2011/0530; TTNCM 39/2011/0533; TTNCM 39/2011/0486a-b.

Remarks.–See supplementary material for individual measurements and descriptions. TTNCM 39/2011/0523 is figured in Williams et al. [41] (Fig 3D).

References.–[41, 115, 128, 142].

Family Protogryllidae Zeuner, 1937 [71]

gen. et sp. indet.

Assigned material.–TTNCM 39/2011/1006

Remarks.–Specimen was not available to be re-examined. TTNCM 39/2011/1006 was originally reported as belonging to Protogryllidae by Andrew J. Ross in the database he originally compiled on the specimens from TTNCM.

Family incertae sedis

gen. et sp. indet.

Assigned material.–NHMUK PI I.11490; TTNCM 39/2011/0389; TTNCM 39/2011/0401; TTNCM 39/2011/0409; TTNCM 39/2011/0421; TTNCM 39/2011/0521; TTNCM 39/2011/0524; TTNCM 39/2011/0526; TTNCM 39/2011/0529; TTNCM 39/2011/0531; TTNCM 39/2011/0532a-b; TTNCM 39/2011/0534; TTNCM 39/2011/0552; TTNCM 39/2011/1007.

Remarks.–See supplementary material for individual measurements and descriptions.

Order Hemiptera Linnaeus, 1758 [118]

Suborder Fulgoromorpha Evans, 1946 [146]

Family Fulgoridiidae Handlirsch, 1939 [128]

Genus Margaroptilon Handlirsch, 1906 [115]

Margaroptilon brodiei Handlirsch, 1906 [115]

Holotype.–NHMUK PI I.3561, isolated forewing. Whitby Mudstone Formation: Dumbleton Member; Early Jurassic: Lower Toarcian of Alderton Hill, Gloucestershire, UK.

Assigned material.–TTNCM 39/2011/0593, isolated forewing (5 mm long, 2.4 mm wide); NHMUK PI I.11492, isolated forewing (unable to be measured).

Remarks.–See Alderton Hill remarks under Margaroptilon for discussion on M. brodiei.

References.–[46, 109, 115].

Margaroptilon sp.

Assigned material.–Isolated spotted wings: TTNCM 39/2011/0535 (8 mm long, 2.2 mm wide); TTNCM 39/2011/0592 (6.6 mm long, 2 mm wide); TTNCM 39/2011/0594 (5 mm long, 2 mm wide); TTNCM 39/2011/0595 (7.8 mm long, 2.2 mm wide); Isolated spotted wing fragment: TTNCM 39/2011/0995.

Remarks.–Williams et al. [41] identified and figured TTNCM 39/2011/0594 as Fulgoridulum sp. (Fig 3A), however here we attribute this specimen to Margaroptilon sp.

Suborder Heteroptera Latreille, 1810 [114]

Family Archegocimicidae Handlirsch, 1906 [115]

Assigned material.–TTNCM 39/2011/0707a-b, isolated partial forewing (3.2 mm long, 2.2 mm wide); TTNCM 39/2011/0394, isolated wing (3.1 mm long, 1 mm wide); TTNCM 39/2011/0549, isolated wing (3.6 mm long, 1.4 mm wide).

Remarks.–TTNCM 39/2011/0707a-b is figured in Williams et al. [41] (Fig 3E).

Family incertae sedis

gen. et sp. indet.

Assigned material.–Almost complete individuals: TTNCM 39/2011/0692; TTNCM 39/2011/0694; TTNCM 39/2011/0689a-b. Partial individual: TTNCM 39/2011/0665; TTNCM 39/2011/0693; TTNCM 39/2011/0696; NHMUK PI I.11493. Isolated bodies: TTNCM 39/2011/0646; TTNCM 39/2011/0648. Isolated skeletal elements: TTNCM 39/2011/0440; TTNCM 39/2011/0563a-b; TTNCM 39/2011/0570; TTNCM 39/2011/0647. Isolated partial forewings: TTNCM 39/2011/0700a-b; NHMUK PI I.11489. Isolated wing: TTNCM 39/2011/0416; TTNCM 39/2011/0429; TTNCM 39/2011/0480a-b; TTNCM 39/2011/0547; TTNCM 39/2011/0572; TTNCM 39/2011/0540a-b; TTNCM 39/2011/0562a-b. Isolated partial wing: TTNCM 39/2011/0399; TTNCM 39/2011/0431; TTNCM 39/2011/0541; TTNCM 39/2011/0542; NHMUK PI I.11488. Indeterminable: TTNCM 39/2011/0452; TTNCM 39/2011/0585; TTNCM 39/2011/0635; TTNCM 39/2011/1004.

Order Coleoptera Linnaeus, 1758 [118]

Family Cupedidae Laporte, 1836 [175]

gen. et sp. indet

Assigned material.–TTNCM 39/2011/0624, isolated forewing (6.3 mm long, 1.7 mm wide); TTNCM 39/2011/0712, isolated partial forewing (11.5 mm long, 2.1 mm wide).

Family incertae sedis

gen. et sp. indet

Assigned material.–Almost complete individuals: TTNCM 39/2011/0365; TTNCM 39/2011/0377; TTNCM 39/2011/0640; TTNCM 39/2011/0643; TTNCM 39/2011/0644; TTNCM 39/2011/0645; TTNCM 39/2011/0666; TTNCM 39/2011/0667; TTNCM 39/2011/0697 & TTNCM 39/2011/0698 (part and counterpart); TTNCM 39/2011/0709; CAMSM TN 3199. Partial individuals: TTNCM 39/2011/0366; TTNCM 39/2011/0374; TTNCM 39/2011/0375; TTNCM 39/2011/0376; TTNCM 39/2011/0378; TTNCM 39/2011/0379; TTNCM 39/2011/0380; TTNCM 39/2011/0381; TTNCM 39/2011/0382; TTNCM 39/2011/0597; TTNCM 39/2011/0622; TTNCM 39/2011/0663 & TTNCM 39/2011/0664 (part and counterpart); TTNCM 39/2011/0669; TTNCM 39/2011/0670; TTNCM 39/2011/0671; TTNCM 39/2011/0672; TTNCM 39/2011/0673; TTNCM 39/2011/0708; TTNCM 39/2011/0710; TTNCM 39/2011/0711; NHMUK PI I.11486; NHMUK PI I.11487; NHMUK PI I.11497; NHMUK PI I.11500; CAMSM TN 3202. Isolated bodies: TTNCM 39/2011/0726; TTNCM 39/2011/0725. Isolated skeletal elements: TTNCM 39/2011/0441; TTNCM 39/2011/0443; TTNCM 39/2011/0471; TTNCM 39/2011/0496; TTNCM 39/2011/0662; TTNCM 39/2011/0695. Isolated forewings: TTNCM 39/2011/0363; TTNCM 39/2011/0367; TTNCM 39/2011/0386; TTNCM 39/2011/0387; TTNCM 39/2011/0388; TTNCM 39/2011/0406; TTNCM 39/2011/0471; TTNCM 39/2011/0596; TTNCM 39/2011/0598; TTNCM 39/2011/0599; TTNCM 39/2011/0600; TTNCM 39/2011/0601; TTNCM 39/2011/0602; TTNCM 39/2011/0603; TTNCM 39/2011/0604; TTNCM 39/2011/0605; TTNCM 39/2011/0606; TTNCM 39/2011/0607; TTNCM 39/2011/0608; TTNCM 39/2011/0609; TTNCM 39/2011/0610; TTNCM 39/2011/0613; TTNCM 39/2011/0614; TTNCM 39/2011/0615; TTNCM 39/2011/0616; TTNCM 39/2011/0617; TTNCM 39/2011/0620; TTNCM 39/2011/0621; TTNCM 39/2011/0627; TTNCM 39/2011/0630; TTNCM 39/2011/0631; TTNCM 39/2011/0668; TTNCM 39/2011/0713; TTNCM 39/2011/0714; TTNCM 39/2011/0715; TTNCM 39/2011/0717; TTNCM 39/2011/0718; TTNCM 39/2011/0719; TTNCM 39/2011/0720; TTNCM 39/2011/0721; TTNCM 39/2011/0722; TTNCM 39/2011/0723; TTNCM 39/2011/0724; TTNCM 39/2011/0727; TTNCM 39/2011/0728; TTNCM 39/2011/0760a-b; TTNCM 39/2011/0998; TTNCM 39/2011/1002; NHMUK PI I.11484; NHMUK PI I.11491; NHMUK PI; I.11495; NHMUK PI I.11496 & NHMUK PI I.11494 (part and counterpart); NHMUK PI I.11498; NHMUK PI I.11502; CAMSM TN 3203; CAMSM 3204. Isolated partial forewings: TTNCM 39/2011/0373; TTNCM 39/2011/0407; TTNCM 39/2011/0493; TTNCM 39/2011/0498; TTNCM 39/2011/0611; TTNCM 39/2011/0612; TTNCM 39/2011/0618; TTNCM 39/2011/0619; TTNCM 39/2011/0623; TTNCM 39/2011/0625; TTNCM 39/2011/0628 & TTNCM 39/2011/0629 (part and counterpart); TTNCM 39/2011/0716; TTNCM 39/2011/0949. Indeterminable: TTNCM 39/2011/0447; TTNCM 39/2011/0448; TTNCM 39/2011/0449; TTNCM 39/2011/0450; TTNCM 39/2011/0451; TTNCM 39/2011/0452; TTNCM 39/2011/0453; TTNCM 39/2011/0636; TTNCM 39/2011/0641a-b; TTNCM 39/2011/0657; TTNCM 39/2011/0658; TTNCM 39/2011/0659; TTNCM 39/2011/0660; TTNCM 39/2011/0661; TTNCM 39/2011/0674; TTNCM 39/2011/0675; TTNCM 39/2011/0676; TTNCM 39/2011/0677; TTNCM 39/2011/0678; TTNCM 39/2011/0679; TTNCM 39/2011/0680 & TTNCM 39/2011/0681 (part and counterpart); TTNCM 39/2011/0682; TTNCM 39/2011/0683; TTNCM 39/2011/0684a-b; TTNCM 39/2011/0685; TTNCM 39/2011/0686; TTNCM 39/2011/0687; NHMUK PI I.11501.

Remarks.–TTNCM 39/2011/0640 is figured in Williams et al. [41] (Fig 3B). At least two elytra morphotypes can be identified, including smooth (TTNCM 39/2011/0727) and striated elytra (TTNCM 39/2011/0614), indicating that the beetle fauna of Strawberry Bank was likely to consist of several species. Other beetle elements consist of well-preserved partial bodies (TTNCM 39/2011/0644).

Superorder Amphiesmenoptera Kiriakoff, 1948 [176]

Order incertae sedis

Assigned material.–TTNCM 39/2011/0996a-b, isolated partial wing (9 mm long, 2.2 mm wide); TTNCM 39/2011/1009, isolated partial wing (6.5 mm long, 3 mm wide); TTNCM 39/2011/1010, isolated partial wing (5.2 mm long, 3 mm wide).

Order Necrotrichoptera Engel, 2022 [152]

Family Necrotauliidae Handlirsch, 1906 [115]

Genus Necrotaulius Handlirsch, 1906 [115]

Necrotaulius parvulus Geinitz, 1884 [66]

Holotype.–FGWG 119/7, isolated wing. Grimmen Formation, ‘Green Series’; Early Jurassic: Lower Toarcian of Dobbertin, Mecklenburg-Vorpommern, Germany.

Assigned material.–TTNCM 39/2011/0733, isolated wing (7.3 mm long, 3 mm wide).

Remarks.–See above for the remarks and discussion of N. parvulus for the Alderton Hill specimens.

References.–[43, 66, 77].

Order Mecoptera Packard, 1886 [157]

Family Orthophlebiidae Handlirsch, 1906 [115]

Assigned material.–TTNCM 39/2011/0400, isolated wing (9.9 mm long, 2.2 mm wide).

Family incertae sedis

gen. et sp. indet.

Assigned material.–TTNCM 39/2011/0546, isolated wing (6.1 mm long, 3.4 mm wide).

Order Diptera Linnaeus, 1758 [118]

Family Limoniidae Speiser, 1909 [170]

Genus Architipula Handlirsch, 1906 [115]

Architipula sp.

Assigned material.–TTNCM 39/2011/0402, isolated wing (7.1 mm long, 1.9 mm wide).

References.–[41, 76].

Order Neuroptera Linnaeus, 1758 [118]

Family incertae sedis

gen. et sp. indet.

Assigned material.–TTNCM 39/2011/0423; TTNCM 39/2011/0432; TTNCM 39/2011/0433; TTNCM 39/2011/0553; TTNCM 39/2011/1011; TTNCM 39/2011/1012.

Superorder Panorpida Hinton, 1958 [177]

Family incertae sedis

gen. et sp. indet.

Assigned material.–Isolated wing: TTNCM 39/2011/0539. Isolated partial wings: TTNCM 39/2011/0368; TTNCM 39/2011/0372; TTNCM 39/2011/0390; TTNCM 39/2011/0404; TTNCM 39/2011/0412; TTNCM 39/2011/0418; TTNCM 39/2011/0425; TTNCM 39/2011/0428; TTNCM 39/2011/0430; TTNCM 39/2011/0436; TTNCM 39/2011/0437; TTNCM 39/2011/0548.

Remarks.–See supplementary material for individual measurements and descriptions.

Alderton Hill

From the 370 individual insects documented and examined from Alderton Hill, 29.2% (n = 108; Table 4) of the total assemblage was considered indeterminate due to poor preservation and therefore could not be confidently assigned to any taxonomic group; these specimens were not included within further taxonomic analysis. The remaining 70.8% (n = 262) could be taxonomically identified to at least the level of order (Table 4). Of these individuals, 37.6% (n = 139) could be further assigned to family level, whilst 31.9% (n = 118) and 20.3% (n = 75) could be further assigned to genera and species respectively (Table 4). The 12 insect orders identified from the 262 identifiable individuals collected at Alderton Hill are Hemiptera (true bugs), Coleoptera (beetles), Orthoptera (grasshoppers), Neuroptera (lacewings), Odonata (dragonflies), Diptera (flies), Mecoptera (scorpionflies), Blattodea (cockroaches), Necrotrichoptera (caddisflies), Reculida (‘winged insects’),? Dermaptera (earwigs) and Ephemeroptera (mayflies); and their relative abundance is shown in Fig 7A.

The Hemiptera of Alderton Hill are the most diverse and abundant order within the fossil insect assemblage, comprising 22.1% (n = 58; Fig 7A) of all the identifiable insect specimens collected from this locality. This order is represented by four families (Table 5). The majority of the hemipteran material, in particular those attributed to Fulgoridiidae, are comprised of isolated wings. The genus Margaroptilon is the most common taxon within this order, and is represented by well-preserved forewings with spotted pigmentation (i.e. NHMUK PI I.11266; Fig 4C). Other hemipteran elements also include partial bodies (Figs 3I–4G), which cannot be identified further due to their poor preservation. Coleoptera is the second most common insect order of the Alderton Hill fossil assemblage (20.6%; n = 54). This order is mostly represented by isolated elytra, but also paired elytra and partial individuals, and the poor preservation prevents further identification (Table 5). However, at least three elytra morphotypes can be identified (Fig 4K–Fig 4M), indicating that the beetle fauna is likely to consist of several genera or species. Orthoptera (13.4%, n = 35; Fig 7A) are attributed to three families (Table 5) and are primarily represented by fragmented, yet well-preserved tegmina. Some tegmina possess the original striped and spotted pigmentation (Fig 3I–3L), which aids their taxonomic assignment. Isolated skeletal elements including saltatorial hind legs are also present (Fig 3M). Whilst Neuroptera are relatively abundant from Alderton Hill (12.2%, n = 32; Fig 7A), they are mostly represented by poorly preserved isolated wings. However, several well-preserved examples of Neuroptera are present (Fig 5A–5F), representing three families (Table 5).

Odonata is represented by three families (Table 5) and accounts for 11.5% (n = 30) of the total Alderton Hill palaeoentomofauna (Fig 7A). The order is mainly represented by partial wing fragments that possess clear venation (Fig 3C and 3D) aiding identification, however complete, well-preserved forewings (Fig 3A) and hindwings (Fig 3B) are also present. Diptera comprises 9.5% (n = 25) of the fossil assemblage (Fig 7A), in which all specimens identified to family level are exclusively assigned to the family Limoniidae (craneflies) and primarily composed of isolated bodies and wings. Well-preserved isolated wings belonging to the order Mecoptera comprises 5% (n = 13) of the total insect assemblage (Figs 3R and 4Q; Fig 6A), and there are three families present. Blattodea comprise 2.3% (n = 6) (Fig 7A) and are represented by rather well-preserved, isolated tegmina with the clavus attached (Fig 3G–3H). One specimen, BRSMG Cg2374 & BRSMG Cg2375, is noticeably dissimilar to all other Toarcian cockroaches and therefore represents a new genus and species. Necrotrichoptera accounts for only 1.9% (n = 5) of the Alderton Hill palaeoentomofauna (Fig 7A), in which all specimens belong to the family Necrotauliidae (Fig 4N and 4O). Similarly, Reculida is also uncommon (0.8%; n = 2) (Fig 7A) and is only represented by partial, yet well preserved forewing fragments of Geinitzia carpentieri (Fig 3F).? Dermaptera (NHMUK PI I.11403; Fig 3E) and Ephemeroptera (WARMS G 8084) are rare orders within the deposit, each representing 0.4% (n = 1) of the total insect assemblage (Fig 7A); these orders are all represented by poorly preserved fragmentary remains.

Strawberry bank

Of the 799 individual insects from Strawberry Bank, 30% (n = 240; Table 4) of the insect assemblage are considered indeterminate due to poor preservation and therefore cannot be confidently assigned to any taxonomic group. Of the total insect assemblage, 70% (n = 559) can be taxonomically identified to at least order level (Table 4). Of these individuals, only 4.8% (n = 38) of the Strawberry Bank insect assemblage can be assigned to family level, whilst 1.5% (n = 12) and 0.5% (n = 4) can be assigned to genera and species (Table 4). Nine insect orders and two superorders are identified among 559 identifiable individuals from Strawberry Bank: Hemiptera, Coleoptera, Orthoptera, Neuroptera, Odonata, Diptera, Mecoptera, Blattodea, Dermaptera, Amphiesmenoptera and Panorpida; and their relative abundance is shown in Fig 7B.

Within the Strawberry Bank fossil insect assemblage, Coleoptera is the most abundant order comprising 74.2% (n = 415; Fig 7B) of all the identifiable insect specimens collected from this locality. The majority of the coleopteran material is represented by disarticulated and fragmented elytra that are compressed and rather badly preserved, and not possible to identify beyond order level (Table 6). Similar to Alderton Hill, at least two elytra morphotypes can be identified, indicating different species. Hemiptera is the second most common insect order of the Strawberry Bank palaeoentomofauna (8.9%, n = 50) (Fig 7B), and can be assigned to two families (Table 6). This order is primarily represented by partial individuals (NHMUK PI I.11493) and isolated wings; wings attributed to Margaroptilon (Table 6) in particular possess spotted pigmentation (TTNCM 39/2011/0594). Orthoptera, the third most abundant insect order (6.6%, n = 37) (Fig 7B), is represented by three families (Table 6) and comprised of isolated, fragmented tegmina, some of which show original striped and spotted pigmentation (TTNCM 39/2011/0523). Odonata accounts for 3.6% (n = 20) of the Strawberry Bank palaeoentomofauna (Fig 7B) and can be attributed to only one family (Table 6). This order is mainly represented by partial wing fragments that possess some venation (e.g. TTNCM 39/2011/0516). Although disarticulated, segmented abdominal elements (TTNCM 39/2011/0362a-b) are also documented. Panorpida (2.3%, n = 13; Fig 7B) is represented exclusively by isolated and often poorly preserved wing fragments (e.g. TTNCM 39/2011/0436), preventing taxonomic refinement below superorder level (Table 6).

Blattodea accounts for 1.8% (n = 10) of the palaeoentomofauna (Fig 7B), and is represented by only one family (Table 6). Similar to the rest of the Strawberry Bank insect assemblage, the material is preserved as disarticulated tegmina, with no hindwings present. Neuroptera comprises 1.1% (n = 6) of the Strawberry Bank fossil insect assemblage (Fig 7B) and is mostly represented by poorly preserved, isolated wings that are difficult to identify beyond order level (Table 6); however, some better preserved examples are present (TTNCM 39/2011/0423). Amphiesmenoptera comprises 0.7% (n = 4) of the palaeoentomofauna (Fig 7B), with one family identified (Table 6). Whilst the rest of these individuals cannot be taxonomically refined any further than superorder level, they most likely belong to either Lepidoptera or Necrotrichoptera. This material is represented exclusively by isolated, preserved wing and wing fragments (TTNCM 39/2011/1009). Whilst Dermaptera is uncommon within the Strawberry Bank palaeoentomofauna (0.2%, n = 1) (Fig 7B), a single tegmen was described by Kelly et al. [42] as a new species—Trivenapteron moorei (TTNCM 39/2011/0489), which was the first record of an earwig from the Toarcian of the UK [41]. Finally, Mecoptera and Diptera are rare within the Strawberry Bank deposit, and represent 0.4% (n = 2) and 0.2% (n = 1) of the total insect assemblage respectively (Fig 7B).

In summary, approximately 70% of the insect assemblages at both Alderton Hill and Strawberry Bank can be identified to order level, and whilst the Coleoptera, Hemiptera and Orthoptera are the three most common orders at both localities, their relative abundances differ (Fig 7). Alderton Hill is significantly more diverse at all taxonomic ranks recognised in the fossil assemblage (Table 3), as more specimens can be assigned to family (AH = 37.6% cf. SB = 4.8%), genera (AH = 31. 9% cf. SB = 1.5%) and species (AH = 20.3% cf. SB = 0.5% (Table 3).

Feeding traits

The diet of 74.3% (n = 278) of the Alderton Hill fossil insect assemblage is classified as indeterminable. Specific diets for the majority of taxa cannot be inferred because of the low taxonomic resolution (e.g. Coleoptera). At present-day, carnivorous, herbivorous and parasitic examples are present in almost all insect orders; demonstrating that generalisations cannot be made. For taxa where their specific diets can be determined (n = 92), 58.7% (n = 54) are herbivorous (e.g. Hemiptera: Fulgoromorpha and Coleorrhyncha; Orthoptera: Elcanidae, Locustopsidae and Protogryllidae) and 41.3% (n = 38) are carnivorous (e.g. Blattodea: Raphidiomimidae; Mecoptera: Bittacidae; and Odonata). However, the low numbers in each feeding trait mean that analysis is not very representative. As the majority of the Strawberry Bank palaeoentomofauna is comprised of Coleoptera, which are largely unidentifiable beyond order level, further insights in the trophic feeding of this insect community and specific diets cannot be inferred.

Alderton hill

Based on the matrix of the insect specimens examined, all of the insect material collected from Alderton Hill to date appears to have come from the limestone nodules within the Fish Bed (Bed 10 of Richardson [79]). The insect material of Alderton Hill is mostly preserved as disarticulated, fragmented compression fossils. Some Coleoptera are preserved in three-dimensions, but show minor compression (NHMUK PI I.11318; NHMUK PI I.11369). As the insects are preserved in early diagenetic nodules formed within fine-grained sedimentary deposits, the intricate detail of the insects is well preserved, including wing venation and evidence of colour pigmentation and patterns.

From the 370 individual insects documented and examined, 20.5% (n = 76) could not be taphonomically assigned because of poor preservation or, in a few cases, they were not available for examination. These specimens were assigned the taphonomic grade ‘??’. Of the taxonomically identifiable specimens, 56.1% (n = 147) of the total insect assemblage were comprised of heavily sclerotised taxa i.e. Hemiptera, Coleoptera and Orthoptera. No aquatic larval stages were observed within the collections, and thus all of the specimens examined were represented by terrestrial adult life stages, including flying terrestrial and semiaquatic taxa.

The taphonomic grades from 1 – 2d (Table 2) were assigned to 294 individuals (Fig 8A). Whilst a wide range of preservation was observed among the Alderton Hill insects (Table 3), the material was mostly represented by isolated wings (n = 198, 67.3%; Fig 8A). The most common primary taphonomic grade within the deposit was isolated forewing/s (grade 2b) with 101 (34.4%) individuals; 50 (17%) and 51 (17.3%) of these forewings could be assigned to the secondary taphonomic grades 2b.1 and 2b.2 respectively (Fig 8A). Eighty-six individuals (29.3%) were isolated unassigned wing/s (grade 2d), with 12 (4.1%) being almost complete (grade 2d.1) and 74 (25.2%) wing fragments (grade 2d.2) (Fig 8A); the latter was the most common secondary taphonomic grade recorded from the insect assemblage. Seventy-one individuals (24.1%) were isolated bodies (grade 2a); 12 (4.1%) and 59 (20.1%) were almost complete (grade 2a.1) and skeletal body elements (grade 2a.2) (Fig 8A). Twenty-five individuals (8.5%) were bodies with forewings and/or hindwings (grade 1); 7 (2.4%) insects were assigned to the secondary taphonomic grade 1.2, and 18 (6.1%) individuals to 1.3 (Fig 8A). No complete, fully articulated individuals (grade 1.1) have been collected from this locality to date (Fig 8A). One of the most complete insect collected from Alderton Hill was the paratype of Heterophlebia buckmani (NHMUK PI I.3988), comprised of a head, thorax and fore- and hindwings, with only the distal half of abdomen absent [63]; however, this type specimen was not located within the NHMUK collection during this study and is presumed lost. Whilst this specimen could not be included within this research as it could not be re-examined, it demonstrates that more complete individuals were preserved at Alderton Hill.

The rarest primary taphonomic grade within the deposit was isolated hindwing/s (grade 2c) with 11 individuals (3.7%); 4 (1.4%) of these hindwings were almost complete (grade 2c.1) and 7 (2.4%) were fragments (grade 2c.2) (Fig 8A). The level of disarticulation is high in all insect orders; with only 8.5% (n = 25) of the total assemblage being considered either partially articulated or fully articulated (e.g. abdomen associated with elytra and thorax); therefore, much of the Alderton Hill collection consisted of fully disarticulated insect elements (n = 269).

In total, 230 (78.2%) of the individuals assigned a taphonomic grade could be taxonomically assigned to at least order level (Table 3). A Chi-squared test indicated that there was a statistically significant association between the taxonomic rank and the taphonomic grade at order level (x² = 234.58, df = 99, p = < 0.001). The excellent preservation of bodies with wings (grade 1), isolated complete bodies (grade 2a.1) and wings >75% complete (i.e. grades 2b.1 and 2c.1) in the majority of the specimens allowed for a high level of taxonomic identification. Where the wings were less than 75% complete (i.e. grades 2b.2, 2c.2 and 2d.2) or there were only isolated skeletal elements (grade 2a.2), the taxonomic identification was poorer. In order to determine the degree to which each primary taphonomic grade contributed to the total Chi-squared score at order level, Pearson residuals were calculated for each entry in the Chi-squared contingency table and displayed graphically (Fig 9). Within the individual orders, the largest range of taphonomic grades were observed within the Hemiptera, with 30% (n = 15) of this material being assigned to 2b.2. Coleoptera were primarily represented by isolated or paired elytra, as 45.5% (n = 20) of the coleopteran material was assigned to 2b.1 (Fig 9).a

The Alderton Hill insect fossils had a minimum preserved length of 0.3 mm and a maximum of 45 mm, with a median preserved length of 6.4 mm and a mean preserved length of 7.98 mm (SD = 6.10 mm; n = 318) (Fig 10A). Preserved specimen widths ranged from 0.1 mm to 13.9 mm, with a median preserved width of 2.3 mm and a mean preserved width of 2.84 mm (SD = 2.12 mm; n = 318) (Fig 10C); only 5% of the insect material was greater than 20 mm long and 11.9% was greater than 5 mm wide.

Strawberry bank

Similar to Alderton Hill, the levels of disarticulation were high in all insect orders at Strawberry Bank; only 9.3% (n = 49) of the total insect assemblage was considered either partially articulated or fully articulated, therefore much of the deposit consisted of fully disarticulated insect elements (n = 477) (Fig 8B).

As a high proportion (74.2%) of the specimens from Strawberry Bank are taphonomically robust Coleoptera, most of the material is disarticulated but preserved in three-dimensions, with minor amounts of compression. The remainder of the material is preserved as highly disarticulated, and fragmented compression fossils, similar to Alderton Hill. From the 799 individual insects documented and examined from the Strawberry Bank insect assemblage, 34.2% (n = 273) could not be taphonomically assigned and were given the taphonomic grade ‘??’. Of the taxonomically identifiable specimens, 89.8% (n = 502) of the total insect assemblage of Strawberry Bank was comprised of heavily sclerotised taxa i.e. Coleoptera, Hemiptera and Orthoptera. The taphonomic grades from 1–2d (Table 2) were assigned to 526 individuals (Fig 8B).

Similar to Alderton Hill, the insect material of Strawberry Bank was mostly represented by isolated wings (n = 436, 82.9%; Fig 8B). The commonest primary taphonomic grade within the deposit was isolated forewing/s (grade 2b) with 337 (64.1%) individuals. From this, 308 (58.6%) of these forewings were almost complete (grade 2b.1) and 29 (5.5%) were fragments (grade 2b.2) (Fig 8B). The former was the most common secondary taphonomic grade recorded from the Strawberry Bank insect assemblage.

Ninety-seven individuals (18.4%) were isolated unassigned wing/s (grade 2d), with 21 (4%) and 76 (14.4%) of these wings belonging to the secondary taphonomic grades 2d.1 and 2d.2 respectively (Fig 8B). Forty-nine individuals (9.3%) were bodies with forewings and/or hindwings (grade 1); 15 (2.9%) insects were assigned to the secondary taphonomic grade 1.2, and 34 (6.5%) individuals to 1.3 (Fig 8B). Forty-one individuals (7.8%) were isolated bodies (grade 2a); 5 (1%) and 36 (6.8%) were given the secondary taphonomic grades 2a.1 and 2a.2 respectively (Fig 8B). No insects could be assigned to taphonomic grade 1.1 (Fig 8B), meaning no complete, fully articulated individuals have been recovered from Strawberry Bank to date. The rarest primary taphonomic grade within this deposit was isolated hindwing/s (grade 2c) with only 2 individuals (0.4%) assigned to taphonomic grade 2c.1 (Fig 8B).

The Strawberry Bank insects had a minimum and maximum preserved length of 0.8 mm and 51.2 mm respectively, with a median preserved length of 5 mm and a mean preserved length of 6.46 mm (SD = 5.44 mm; n = 275; Fig 10B). Preserved width ranged from 0.3 mm to 18.4 mm, with a median preserved width of 2.2 mm and a mean preserved width of 2.54 mm (SD = 1.82 mm; n = 275; Fig 10D). The distribution of the length and width shows a greater frequency of small sized elements at Strawberry Bank. Further statistical comparison of the size data from Strawberry Bank and Alderton Hill was not undertaken, as the taphonomic and taxonomic bias at Strawberry Bank render any results from this location meaningless.