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BLATTODEA (COCKROACHES) Figure 7A Two pieces of amber, AMNH WH-12 and AMNH WH-11, contain fragmentary remains of small roaches, probably nymphs. WH-12 contains a portion of one leg: apical portion of the femur and the entire tibia and tarsus. The tibia has 18 spinelike setae (including a pair of apical tibial spurs), in roughly four longitudinal rows. WH-11 contains portions of three legs and an antenna, all very dark and surrounded by a dark reddish ���halo��� of pyritization/oxidation. One leg is preserved as a portion of a femur; the tarsi and portions of the tibia are preserved for the other two legs. Approximately 30 filiform flagellomeres are preserved; the basal ones short (lengths about twice the width), gradually increasing in length distad to about five times the width. Leg segments of WH-11 are shorter than in WH-12, and tarsomere four has a long ventral lobe. Thus, there appears to be two taxa of roaches. Though these specimens cannot be identified to family or superfamily, they are clearly Blattodea based on the antennal structure and the tibial spines, which further have minute serrations on the ventral margin. The fossil record and natural history of roaches are reviewed by Grimaldi and Engel (2005). The natural distribution of Blattodea worldwide is almost entirely tropical to warm temperate; pest species reach higher latitudes in association with human habitations. Of the 69 species of roaches in North America, 24 are introduced from other regions, and only three native species (Parcoblatta pensylvanica, P. uhleriana, and P. virginica) have distributions that extend into southernmost Ontario and Qu��bec (Vickery and McKevan, 1985; Atkinson et al., 1991). The roaches are the most obvious example in Chickaloon amber of a taxon that has retreated from high northern latitudes., Published as part of Grimaldi, David A., Sunderlin, David, Aaroe, Georgene A., Dempsky, Michelle R., Parker, Nancy E., Tillery, George Q., White, Jaclyn G., Barden, Phillip, Nascimbene, Paul C. & Williams, Christopher J., 2018, Biological Inclusions in Amber from the Paleogene Chickaloon Formation of Alaska, pp. 1-37 in American Museum Novitates 2018 (3908) on page 20, DOI: 10.1206/3908.1, http://zenodo.org/record/4598569, {"references":["Grimaldi, D. A., and M. S. Engel. 2005. Evolution of the insects. New York: Cambridge University Press.","Atkinson, T. H., P. G. Koehler, and R. S. Patterson. 1991. Catalog and atlas of the cockroaches (Dictyoptera) of North America north of Mexico. Miscellaneous Publications of the Entomological Society of America 78: 1 - 86."]}

ACARI (MITES) Figures 5H, 11A AMNH WH-6: A small piece containing most of the remains of an oribatid mite, similar in overall structure to the family Damaeidae (fig. 11A), which consists of fewer than 100 living species in 14 genera, primarily as mycophagous and algophagous inhabitants of leaf litter and subcortical microhabitats of temperate and boreal forests. Damaeidae occur in Eocene amber from the Baltic region and Rovno, Ukraine (Weitschat and Wichard, 2010; Perkovsky et al., 2010). The mite is near the corner of the amber piece, with the appendages of one side lost or completely obscured; the piece also contains dark layers from various resin flows as well as particulate plant matter. Body length (without appendages) is approximately 400 μm, with a slight constriction between the prodorsum and notogaster; legs are long and slender, length of the longest is 550 μm. The mite is dark and opaque, generally obscuring many of the setae, sensilla, and cuticular details except those visible at margins. Anterior-most appendage (pedipalps) (only one of a pair observable), with three short, stout podites, apical one pointed, with two long, fine solenidia. A pair of long, stiff solenidia occurs at the anterior end of the prodorsum. Structure of the legs is very distinctive and quite similar to that of the living family Damaeidae, in particular the “moniliform” legs (although this habitus also occurs in oribatid superfamilies closely related to Damaeoidea). Leg I has bulbous portions of the femur, genu, tibia, and at the base of the tela + basitarsus. The only bulbous portion of leg II is on what is either the genu or tibia (boundaries between podites are barely visible). The apices of tibiae in legs II and III each have a long, stiff solenidium that is nearly equal in length to that of its respective podite. Pretarsal claws are long, slender, and sickle shaped. AMNH LC-D1: A piece of amber containing a complete mite (~280 μm body length), which is moderately well preserved (fig. 5H). Cuticle of the notogaster and prodorsum is cracked and slightly disintegrated, precluding observation of most of the chaetotaxy, glands, and sensilla, though a pair of thick, plumose trichobothria/bothridial setae is visible, one at each posterolateral corner of the prodorsum. While identification of the mite in the Oribatida is certain, a more detailed identification will be challenging. The mite is rather generalized in structure, lacking specialized (e.g., plumose) setae and obvious cuticular microstructure (e.g., reticulations). The gnathosoma is well integrated and conical, laterally with a pair of projecting solenidia; legs are relatively short, pair I with an elongate solenidium dorsoapically on what appears to be the genu; all pretarsal claws are long, slender, and hooked. The Oribatida is a highly diverse, speciose group of mites comprised of some 9000 living species in 172 families, largely inhabitants of soils, leaf litter, and moss (Norton and BehanPelletier, 2009). The fossil record of the group is ancient and diverse, beginning with unambiguous cuticular remains from the Devonian that are preserved in microscopic detail (Norton et al., 1988). Oribatids have even been implicated in the processing of plant detritus from Carboniferous swamps (Labandeira et al., 1997). Their fossil record in amber from the Cretaceous and Cenozoic is excellent (Dunlop et al., 2018). This is due partly to the improved techniques in preparation and high-magnification (400−1000×) microscopy (Sidorchuk, 2013), and the discovery of major new amber deposits. Taxa described more than a century ago in Eocene Baltic amber are being redescribed in great detail (e.g., Norton, 2000; Sidorchuk and Norton, 2010, 2011), which establish a new standard for study.

DERMESTIDAE (CARPET BEETLES) Figures 8 A���C; 14F AMNH LC-II-B4: A partial larva that is missing the head and legs, but has seven abdominal segments largely to entirely preserved (portions of the anterior segments are lost at the amber surface on the right side) (fig. 8A). The dorsum of the abdomen is covered with a dense vestiture of long setae having short, thick plumosity; presence/absence of bare patches on tergites is not observable. The apical abdominal segments have tu��s of peculiar spear-shaped setae, which are very well preserved. These specialized setae have a bullet-shaped head that is hollow, with an asymmetrical, sharp basal rim; the setal sha�� has evenly spaced nodes, each node with a crenulated collar of small spines or tubercles (fig. 14F). Such setae, called hastisetae, allowed identification of the partial larva to the Dermestidae, and in fact hastisetae of this structure are confined to the subfamily Megatominae (Kiselyova and McHugh, 2006), most similar to the genus Cryptorhopalum. The hastisetae in extant dermestids are defensive, being dehiscent and snagging together when the larva is attacked, entangling the attacker (Nutting and Spangler, 1969). There are 1300 living species of Dermestidae in 53 genera, well-known for their larval diet of dried animal remains (including carrion, and shed feathers, hairs, and skin in nests). The genus Anthrenus (also a megatomine) is the notorious museum pest that decimates unprotected collections of skins and pinned insects. The oldest putative dermestid is in Jurassic shale (Deng et al., 2017), with definitive larvae and adults in Early Cretaceous amber from Lebanon (Kirejtshuk et al., 2009), and the oldest Attageninae from the mid-Cretaceous of Myanmar (Cai et al., 2017) and Late Cretaceous of New Jersey (Peris and H��va, 2016). Hastisetae of megatomine dermestids are preserved in Upper Albian���aged amber from Spain, snagged in the legs and body of ticks (Pe��alver et al., 2017). The ticks most likely acquired the hastisetae in the arboreal nest of a vertebrate host (Pe��alver et al., 2017). Diverse modern genera of dermestids occur in Eocene Baltic amber (e.g., H��va et al., 2008) and Miocene Dominican amber. The Chickaloon amber specimen is the most northerly fossil record of the Dermestidae, the prior ones being in Baltic amber., Published as part of Grimaldi, David A., Sunderlin, David, Aaroe, Georgene A., Dempsky, Michelle R., Parker, Nancy E., Tillery, George Q., White, Jaclyn G., Barden, Phillip, Nascimbene, Paul C. & Williams, Christopher J., 2018, Biological Inclusions in Amber from the Paleogene Chickaloon Formation of Alaska, pp. 1-37 in American Museum Novitates 2018 (3908) on page 25, DOI: 10.1206/3908.1, http://zenodo.org/record/4598569, {"references":["Kiselyova, T., and J. V. McHugh. 2006. A phylogenetic study of Dermestidae (Coleoptera) based on larval morphology. Systematic Entomology 31: 469 - 507.","Nutting, W. L., and H. G. Spangler. 1969. The hastate setae of certain dermestid larvae: an entangling defense mechanism. Annals of the Entomological Society of America 62: 763 - 769.","Deng, C., A. Slipinski, D. Ren, and H. Pang. 2017. The oldest dermestid beetle from the Middle Jurassic of China (Coleoptera: Dermestidae). Annales Zoologici 67: 109 - 112.","Kirejtshuk, A. G., D. Azar, P. Tafforeau, R. Boistel, and V. Fernandez. 2009. New beetles of Polyphaga (Coleoptera, Polyphaga) from Lower Cretaceous Lebanese amber. Denisia 26: 119 - 130.","Cai, C., J. Hava, and D. Huang. 2017. The earliest Attagenus species (Coleoptera: Dermestidae: Attageninae) from Upper Cretaceous Burmese amber. Cretaceous Research 72: 95 - 99.","Peris, D., and J. Hava. 2016. New species from Late Cretaceous New Jersey amber and stasis in subfamily Attageninae (Insecta: Coleoptera: Dermestidae). Journal of Paleontology 90: 491 - 498.","Penalver, E., et al. 2017. Ticks parasitised feathered dinosaurs as revealed by Cretaceous amber assemblages. Nature Communications 8: 1924. [doi: 10.1038 / s 41467 - 017 - 01550]","Hava, J., J. Prokop, and A. Herrmann. 2008. New fossil dermestid beetles (Coleoptera: Dermestidae) from the Baltic amber-III. Acta Societatis Zoologicae Bohemicae 17 (2007): 151 - 157."]}

FORMICIDAE (ANTS) Figures 9 A���C AMNH WH-1: A single worker ant specimen, approximately 1.7 mm total body length (excluding antennae), attributable to the subfamily Formicinae. The acidopore, a primary diagnostic feature of formicine ants, appears faintly visible as a circular opening at the terminus of the abdomen. The petiole���a waistlike segment separating the trunk and gaster���is conspicuously scale shaped, with its dorsal margin reaching the same approximate height as the propodeum, a syndrome found in multiple extant and fossil formicine genera. Specimen WH-1 is heavily desiccated and partially disarticulated, making precise placement difficult. Nevertheless, the specimen clearly does not fit into any currently described genus of formicine, in particular due to its widely spaced mandibular dentition and shortened antennal scape (figs. 9B, C). The isolated antenna of another ant specimen, in piece GC-A4 (fig. 14E), has different segmental proportions than the formicine above, indicating the existence of another ant species in this amber. Very little can be determined taxonomically on the basis of an antenna. The subfamily Formicinae is presently distributed worldwide, and fossils are similarly cosmopolitan. In total, 196 fossil formicine species are described, comprising 43 genera from 44 localities across North America, Europe, Asia, and New Zealand (Barden, 2017). Potentially a result of high sampling bias (nearly 13,000 ant inclusions were utilized in a recent analysis of ant species richness [Penney and Preziosi, 2014]), nearly 40 formicine species are described from Baltic amber, the greatest of any deposit. The Chickaloon amber species has not yet been formally described, however, it may represent a stem relative of the primarily Palearctic and Nearctic tribe Formicini or the cosmopolitan tribe Lasiini, based on current understanding of relationships (e.g., Lapolla et al., 2010; Ward et al., 2016)���both represented in the fossil record, including in Baltic amber. This preliminary diagnosis is based on the large circular propodeal spiracle, gradually sloping mesonotum, 5:4 palpomere formula, and mandibular shape. Interestingly, this new specimen is contemporaneous with formicine ants described from Fushun amber (Hong, 2002), making it, along with its counterparts in Asia, the oldest formicines known following Kyromyrma neffi in Turonian-aged New Jersey amber (Grimaldi and Agosti, 2000). This is a valuable window into ant evolution as the Fushun amber holotypes have since been lost. Slightly younger ants (Dolichoderinae, Myrmecinae, Formicinae, and Myrmeciinae) are reported in mid to Late Eocene shales and Hat Creek amber from British Columbia (Archibald et al., 2018). It should be noted that the identification of Technomyrmex (Dolichoderinae) in Poinar et al. (1999) was changed to Formicinae incertae sedis in Archibald et al. (2018)., Published as part of Grimaldi, David A., Sunderlin, David, Aaroe, Georgene A., Dempsky, Michelle R., Parker, Nancy E., Tillery, George Q., White, Jaclyn G., Barden, Phillip, Nascimbene, Paul C. & Williams, Christopher J., 2018, Biological Inclusions in Amber from the Paleogene Chickaloon Formation of Alaska, pp. 1-37 in American Museum Novitates 2018 (3908) on page 28, DOI: 10.1206/3908.1, http://zenodo.org/record/4598569, {"references":["Barden, P. 2017. Fossil ants (Hymenoptera: Formicidae): ancient diversity and the rise of modern lineages. Myrmecological News 24: 1 - 30.","Penney, D., and R. F. Preziosi. 2014. Estimating fossil ant species richness in Eocene Baltic amber. Acta Palaeontologica Polonica 59: 927 - 929.","Lapolla, J. S., S. G. Brady, and S. O. Shattuck. 2010. Phylogeny and taxonomy of the Prenolepis genus-group of ants (Hymenoptera: Formicidae). Systematic Entomology 35: 118 - 131.","Ward, P. S., B. B. Blaimer, and B. B. and B. L. Fisher. 2016. A revised phylogenetic classification of the ant subfamily Formicinae (Hymenoptera: Formicidae), with resurrection of the genera Colobopsis and Dinomyrmex. Zootaxa 4072: 343 - 357.","Hong, Y. 2002. Amber insects of China. Beijing: Beijing Scientific and Technological Publishing House.","Grimaldi, D., and D. Agosti, D. 2000. A formicine in New Jersey Cretaceous amber (Hymenoptera: Formicidae) and early evolution of the ants. Proceedings of the National Academy of Sciences of the United States of America 97: 13678 - 13683.","Poinar, G., B. Archibald, and A. Brown. 1999. New amber deposit provides evidence of early Paleogene extinctions, paleoclimates, and past distributions. Canadian Entomologist 131: 171 - 177."]}

CHIRONOMIDAE (MIDGES) Figures 7E, 14 A���D AMNH WH-3: A complete male nonbiting midge (Chironomidae) (body length 1.60 mm), preserved adjacent to a small juvenile spider (fig. 7E). Eyes are bare; pedicel large, subspherical; antenna with long plumosity, apparently having 11 flagellar articles, apical article longest; maxillary palp with four palpomeres, lengths 4> 2 = 3> 1 (fig. 14C). Legs: mesotibia having two bladelike apical spurs (one with fine pectination), apical comb of 11���12 thick, sclerotized, slightly clavate setae (fig. 14D); pretarsal claws simple (untoothed), pulvilli either minute or lost. Wings are very faint, obscuring the venation; no macrotrichia occur on the wing membrane. Male genitalia well preserved: tergite 9 (epandrium) large, shieldlike; gonocoxite large; gonostylus articulating with (not fused to) gonocoxite, bare, flattened and hatchetlike, without discernable apical peg/tooth; pair of inner lobes present, bare; anal point absent (fig. 14A, B). Chironomidae have a rich fossil record, partly because the larvae are aquatic and semiaquatic and both adults and larvae are readily fossilized in lacustrine sediments. The oldest Chironomidae are Triassic, and they are frequently among the most abundant and diverse winged insects in many deposits of amber, such as Eocene Baltic amber and Late Cretaceous ambers from western Canada, New Jersey, and Siberia. The fossil record has been reviewed by Evenhuis (1994). Critical study relies on various microscopic features, and most of the described fossils, done decades to a century ago, require re-description based on modern standards. The male genitalia of the Chickaloon fossil appear most similar to those in the large, widespread subfamily Tanypodinae., Published as part of Grimaldi, David A., Sunderlin, David, Aaroe, Georgene A., Dempsky, Michelle R., Parker, Nancy E., Tillery, George Q., White, Jaclyn G., Barden, Phillip, Nascimbene, Paul C. & Williams, Christopher J., 2018, Biological Inclusions in Amber from the Paleogene Chickaloon Formation of Alaska, pp. 1-37 in American Museum Novitates 2018 (3908) on page 27, DOI: 10.1206/3908.1, http://zenodo.org/record/4598569, {"references":["Evenhuis, N. L. 1994. Catalogue of the fossil flies of the world (Insecta: Diptera). Leiden: Backhuys Publishers."]}

THYSANOPTERA (THRIPS) Figures 7C, D; 12A, B AMNH LC-B6: A piece of amber containing a complete but minute (0.60 mm body length), wingless thrips, presumably a nymph (fig. 7C, D). The body is slightly compressed and distorted, the opaque body contents preventing observation of cuticular microsculpture, chaetotaxy, and sutures. The head is not well preserved, being partially collapsed, but the postocular region does not appear long (as in Phlaeothripidae); mouthparts are not visible. Antennae are preserved well enough to reveal segment proportions, cuticular microstructure, and sensilla (fig. 12B), their total lengths 440���460 ��m. There are seven antennomeres, plus a minute terminal stylus, with relative lengths: 7> 5 = 3> 4> 6 = 1> 2. Most antennomeres have very fine transverse wrinkles and minute setigerous pimples; article 3 is the broadest and bears sensorial plaques, several of which also appear to occur on article 2; sensory cones appear to be absent, as do any thickened, blunt, setiform sensilla (fig. 12B). Femora are moderately thick (approximately twice the thickness of the tibiae); tarsi appear to be 1-segmented (best seen ventrally on the midlegs). Abdomen is somewhat fusiform in shape, with a small lateral lobe on most tergites (6 are visible) bearing a pair of recurved setae (fig. 12A). Presumably there is a pair of these lobes on each of the segments. Apex of the abdomen appears laterally compressed, the tip rounded and bearing a pair of long setae. In lieu of wings and other structures it is difficult to definitively place the thrips to family, though the antennal structure provides good information. Seven antennomeres are found in the Uzelothripidae and in most Thripidae and Phlaeothripidae (which also have 1-segmented tarsi); thrips plesiomorphically have nine antennomeres. Uzelothripidae is highly doubtful, since the fossil does not have antennomeres 3 + 4 fused or an annulate apical segment (also, the family is monotypic in the Recent fauna, and has just one fossil species, in Early Eocene Oise amber from France [Nel et al., 2011]). Phlaeothripidae further have sensoria on segments three and four, but given that the postocular region of the head in LC-B6 does not appear long (perhaps preservational), and the terminal abdominal segment is not tubular, placement of the fossil in this family is uncertain. Phlaeothripidae and Thripidae comprise some 85% of the approximately 6000 living species of thrips, and they have diverse diets, which include phytophagy, pollenivory, and mycophagy. The oldest Thysanoptera are Late Triassic (Grimaldi et al., 2004); the oldest Thripidae and Phlaeothripidae occur in Early Cretaceous amber and are diverse in Cenozoic ambers (Nel et al., 2010). This is the northernmost fossil occurrence of Thysanoptera., Published as part of Grimaldi, David A., Sunderlin, David, Aaroe, Georgene A., Dempsky, Michelle R., Parker, Nancy E., Tillery, George Q., White, Jaclyn G., Barden, Phillip, Nascimbene, Paul C. & Williams, Christopher J., 2018, Biological Inclusions in Amber from the Paleogene Chickaloon Formation of Alaska, pp. 1-37 in American Museum Novitates 2018 (3908) on pages 20-22, DOI: 10.1206/3908.1, http://zenodo.org/record/4598569, {"references":["Nel, P., A. R. Schmidt, C. Bassler, and A. Nel. 2011. Fossil thrips of the family Uzelothripidae suggest 53 million years of morphological and ecological stability. Acta Palaeontologica Polonica 58 (3): 609 - 614.","Grimaldi, D., A. Shmakov, and N. Fraser. 2004. Mesozoic thrips and early evolution of the order Thysanoptera (Insecta). Journal of Paleontology 78: 941 - 952.","Nel, P., E. Penalver, D. Azar, G. Hodebert, and A. Nel. 2010. Modern thrips families Thripidae and Phlaeothripidae in Early Cretaceous amber (Insecta: Thysanoptera). Annales de la Societe Entomologique de France 46: 154 - 163."]}

Deinocrotonidae Peñalver, Arillo, Anderson and Pérez-de la Fuente fam. nov. Type genus. Deinocroton gen. nov. Monotypic. Etymology. From Greek deinos, “ terrible ”, and krotÓn, “ tick ”. Gender: neutral. Diagnosis (both sexes). Integument with closely spaced, deep pits, and mound-like elevations between pits; integument not convoluted, lacking microsculpture. Pseudoscutum distinct (abbreviated in females), pitted but without elevations. Eyes absent. Hypostome subterminal. Basis capituli not bordered by coxae I. Palpi elongated, gracile; palpomere II distally thickened and bent in ventral direction, palpomeres III and IV elongated, tubular, fully mobile. Genital aperture transverse, close to the capitulum in males and slightly posteriad in females. Presence of a conspicuous anteroventral depressed area, post-genital in position. Spiracles smooth, medium sized, located at the level of coxae IV. Genital groove distinct, medially divided in two sections and extending posteriorly. Anal pore terminal. Preanal groove prolonged posteriorly, with sides closing. Legs ruffled. All coxae with short spurs in rows. Leg joints not of the ball and socket type but notch-like processes present. Haller ’ s organ proximal capsule completely open. Festoons absent. Deinocroton draculi Peñalver, Arillo, Anderson and Pérez- de la Fuente gen. et sp. nov. Etymology. Patronym for the main character of the gothic horror novel by Irish writer Abraham “ Bram ” Stoker, which is a fictionalised account of Vlad III, or Vlad Dracula (ca. 1429 – 1476). Holotype. Adult male (AMNH Bu-SA 5 a), ca. 3.9 mm long from posterior margin to apex of hypostome (Figs. 3 a, e – g, j, k, 4 a, f – h, 5 a, c, d, f, g; Supplementary Fig. 2 c, e). Additional material. Allotype: female (CM 63007) (Fig. 4 b, c; Supplementary Figs. 2 d, 3). Paratypes: male (AMNH Bu- SA 5 b) (Figs. 3 a, d, i, l, 4 d, e, 5 b, e; Supplementary Fig. 2 b) and engorged female (CM 63001) (Figs. 3 b, c, h, m, 5 h, i). All adults (see Supplementary Note 1 for more details). Locality and horizon. Southwest of Tanai (close to Maingkhwan village) in the Hukawng Basin, Kachin State area (northern Myanmar), likely from the Noije bum opencast system of mines; earliest Cenomanian7. Diagnosis for genus and species. As for the family. Description. See Supplementary Note 2 for body measurements. Male: Body outline subcircular. Integument surface with closely spaced, deep pits and with single, mound-like elevations between pits (Figs. 3 d, k, 5 a – c, e), as in females (Fig. 4 c). Integument not convoluted (cf. Nuttalliella), lacking microsculpture (e.g., granulations). Body without conspicuous setal vestiture, except setae present on palpi, legs and anal valves, and very sparse setae present on dorsal and ventral integument. Integumentary pits lacking any associated setae. Dorsum. Pseudoscutum distinct (not highly chitinised as in Ixodidae, with integument resembling the rest of body), occupying most part of dorsum, reaching anterior margin of dorsum (Fig. 3 d), with anterolateral margin broadened posteriorly (Fig. 5 b). Cervical grooves present, relatively shallow (Fig. 5 a, b). Pseudoscutum integument with closely spaced, deep pits, but without mound-like elevations as in the rest of body, rendering a surface with smooth appearance in which pits are very apparent (Fig. 3 g). Pits separated by a length equal to their diameter or less. Festoons absent. Eyes absent. Venter. Capitulum partially visible in dorsal view. Hypostome subterminal (sensu Mans et al. 16) (Figs. 3 a, d, f, 5 b), well developed, reaching apex of palpomere II. Hypostome ultrastructure obscure, dental formula indeterminate. Chelicerae only partially visible in the paratype male. Palpi elongated, gracile (around two times the length of hypostome), fully mobile (Fig. 4 a; Supplementary Fig. 2 b – d), as in females (Fig. 4 b). Palpomere I short. Palpomere II the longest, distally thickened in width and height, bent distally in ventral direction (creating a ventral concavity, with surface of articulation with palpomere III facing that direction). Palpomeres III and IV elongated, tubular, tapering basally. Palpomere III about two times as long as wide, with surface slightly ruffled. Palpomere IV in terminal position, about four times as long as wide. Palpi without spurs but bearing abundant, fine setae. Basis capituli not bordered by coxae I, with anterior margin rimmed and surface smooth (Fig. 3 f; Supplementary Fig. 2 b); auriculae, cornua and porose areas absent. Genital aperture a transverse slit in an oval area between anterior half of coxae II (Fig. 3 f, i), close to capitulum. Presence of a conspicuous anteroventral depressed area (Fig. 5 g) that is quadrangular in shape and post-genital, laterally limited by anterior section of genital groove. Genital groove well developed and extending posteriorly; medially divided (immediately after coxae IV) into two sections (Fig. 5 g). Anterior genital groove section extending from coxae II to IV, briefly bordering coxae IV distally (i.e., diverging towards body margin). Posterior genital groove section the longest, grooves progressively diverging posteriorly, slightly bordering anal plate. Spiracle well developed and very close to body margin at level of coxae IV, smaller than in Ixodidae (and in a different position) and larger than in Nuttalliella namaqua 17. Spiracle plate structure sub-triangular in shape and consisting of a small macula and a smooth triangular plate, not fenestrated but bearing two small concavities (Fig. 5 e, f), as in females (Figs. 3 h, 5 h); macula projecting towards ostium to form a lip; entire plate arising from a depressed cuticular area. Preanal groove prolonged posteriorly, with sides closing, delimiting a guitar pick-shaped anal plate (Fig. 3 l), as in females (Fig. 3 m). Anal pore close to posterior margin of body. Anal valves with a few long and fine setae. Legs. Long and strongly flattened laterally from trochanters to tarsi; arising within anterior two-fifths of total body length. Leg joints not of ball and socket type as in Nuttalliella, but leg articles with paired, notch-like ventrodistal processes (without forming sockets for the articulation, balls not distinct), more apparent in basal articulations (Figs. 4 f, h, 5 c, d). Slight separation between coxae, except coxa I contiguous with II. Coxae armed with rows of small, shallow spurs (i.e., rounded tubercles, such as in some ixodids and Nuttalliella) (Figs. 3 f, 4 d, 5 f): one spur on coxa I — in medioposterior position — and three on each coxa II, III, and IV. Three coxal spurs forming a row in coxa II, with two of them in a medial, posterior position while third one in a distal, anterior position. Three coxal spurs aligned in medial position in coxae III and IV (two close together in a slightly basal, posterior position and third one in anterior position at middle of coxa). Trochanter without spurs. Femur, genu, and tibia bearing a sculptured surface of transverse ridges (ruffles), especially marked in genu (Figs. 4 e, 5 d). Trochanters I and II with very shallow ruffles, almost indistinct. First pair of legs with deeper ruffles. Femora I and II positioned very high and strongly flattened laterally. Femur III flattened laterally and high only basally. Femur IV tubular. Haller ’ s organ conspicuous; although only observed in right tarsus I of holotype (Figs. 3 e, 4 g; Supplementary Fig. 2 e) due to preservation of remaining specimens, situated on a dorsal elevation of tarsus I and composed of two parts, a completely open (without a transverse slit) proximal capsule having long setae and a distal pit followed by more long, distinct setae, capsule larger than pit. Basitarsus as long as tarsus in legs II – IV. Pretarsi with two curved pretarsal claws and abundant, long setae. Pretarsal claws large. Pulvilli poorly developed (Fig. 3 j). Female: As in male with the following exceptions: Integument, including that of pseudoscutum, with pits not as well defined as in males. Pseudoscutum abbreviated (Figs. 3 c, 4 c; Supplementary Fig. 3), occupying the anteriormost part of dorsum. Genital aperture in a more posterior position than in males, between coxae II and III, and apparently showing a smooth surface (Supplementary Fig. 3). Marginal groove absent. Remarks. A suite of unique, presumably derived characters defines Deinocrotonidae: the integument structure, the palp morphology, and the shape of the preanal groove. Likewise, the discontinuous genital groove is unique among ticks. The subterminal hypostome and the presence of a pseudoscutum suggest a close relationship between Deinocrotonidae and Nuttalliellidae. Pending a phylogenetic analysis when more material is available (see Supplementary Note 3), we propose here that both families are sister to (Ixodida + Argasidae). So far, a few more deinocrotonids have been found in Burmese amber, and one additional undescribed immature specimen from 105 Ma old Spanish amber most likely belongs to this new family. Apart from the unique characters among ticks, the new family differs from Nuttalliellidae in the following features (see Supplementary Tables 1 and 2): (1) pseudoscutum pitted (vs. mesh-like), (2) pseudoscutum reaching the anterior margin of the dorsum in males, (3) cervical grooves present, (4) capitulum not bordered laterally by coxae I, (5) basis capituli simple and with smooth surface, (6) cornua absent, (7) genital area smooth (vs. irregularly striated), (8) anteroventral depressed area in post-genital position (vs. in pre-genital position), (9) all coxae armed and spurs forming rows, (10) leg joints not of the ball and socket type, at least as in Nuttalliella, (11) proximal capsule of Haller ’ s organ completely open, (12) different morphology and size of the spiracle, and (13) preanal groove different in microscopic detail (smooth vs. posterior and anterior margins with dentate integumental projections). The pseudoscutum in Deinocrotonidae occupies most of the dorsum in males and is abbreviated in females, as occurs in ticks with a scutum/pseudoscutum. The special shape of palpomere II, distally thickened and bending distally in a ventral direction (Fig. 4 a, b; Supplementary Fig. 2 b – d), appears to be an adaptation to protect the distal part of the gnathosoma dorsally and anteriorly, especially the delicate teeth of the hypostome and the chelicerae. Such expansion of the distal part of the palpomere II is present in all ixodids (namely their upper inner margin, creating an inner groove), although palpomere III is also expanded, taking part in the protection of the gnathosoma, and both palpomeres are straight, directed forwards18, 19. In Deinocroton, palpomere III is elongated and tubular, directed ventrally due to the surface of articulation between palpomeres II and III facing that direction and due to the shape of the palpomere II. In Nuttalliella, palpomere II is massive, expanded laterally and provides most of the gnathosomal protection; palpomere III is smaller, triangular in shape and slightly laterally expanded ventrally, whereas both palpomeres are straight, directed forwards as in ixodids20. Argasids lack any palpomere expansion for gnathosomal protection due to the ventral position of their capitulum in adults. On the other hand, the Haller ’ s organ in deinocrotonids has a generalised morphology, with a proximal capsule and a distal small pit, but fine details are obscure under optical microscopy and they have remained unresolved using CT-scanning. Nevertheless, the proximal capsule is fully open (lacking a transverse slit) as in Ixodes, and unlike in other ixodids, argasids and Nuttalliella 21 – 23. Furthermore, CT-scanning revealed the spiracular morphology and position in detail, which are very similar to those of Argasidae 24. Although the spiracle position in Deinocroton is coincident with that of Nuttalliella, the latter has a minute spiracle with a cribose spiracular plate20. Also, the spiracle of the new family is quite different from that in Ixodidae (i.e., bigger and in a posterior position, not hidden by coxae IV 19, 25). Lastly, the ventroposterior grooves that are posterior to coxae IV and diverge towards the posterior body margin have been named herein “ posterior genital groove sections ”, despite not being connected to the longitudinal grooves that arise from the genital area. Although the origin of these posterior grooves is unclear, the set of the anterior and posterior sections is very similar in position and extension to the genital grooves of some ixodids. Other ixodids, such as Boophilus, have posterior grooves due to the presence of adanal shields; however, since Deinocroton lacks any structure resembling this shield, the posterior section of the genital groove in the new family appears to be unique among ticks. The holotype and paratype male Deinocroton, preserved together, have at least seven spear-headed, multi-segmented setae of exogenous origin attached to their bodies (Fig. 6; Supplementary Fig. 4). The longer setae remains are 311 µm (Fig. 6 a; Supplementary Fig. 4 b) and 286 µm (Fig. 6 f; Supplementary Fig. 4 e) in length as preserved and contain 27 segments plus its spear-head and 23 segments, respectively. The spear-head is 27 µm long, 5 µm wide (11 µm in the base), more sclerotised than the rest of the seta and with six basal knobs arranged in circle. The basalmost segments are long (23 µm long the longest preserved) and quickly decrease in length towards the apex of the seta. The distal setal section shows short segments of similar length (ca. 9 µm long in the 20 distal segments), with the distalmost segment (that in connection with the spear-head) not differing in shape and size from the immediately preceding ones. Despite the dilated body of the engorged specimen (paratype female), it belongs to Deinocroton draculi based on the virtually identical size and morphology of the capitulum (including the basis capituli), pseudoscutum, legs (including the relative length of leg segments), two sections of the genital groove, spiracle and anal plate. The morphoanatomical changes in the engorged specimen when compared to the three unengorged ones (attributed to engorgement) are as follows (Figs. 3 b, c, h, m, 5 h, i, 7; Supplementary Table 3): (1) the body increased ca. 1.7 times its length, ca. 1.4 times its greatest width, and ca. 3.6 times its greatest height — this corresponds to an approximate volume change from 15.0 to 126.0 mm3 (i.e., a volume increase of ca. 8.5 times); (2) the dorso-ventrally planar body became inflated (more pronouncedly so medially along the longitudinal axis) and its subcircular outline became elongated (bean-shaped), particularly in the transverse medial portion of the body or area that separates the anterior and posterior sections of the genital groove; (3) the body integument became smooth, without evidence of the original pits; (4) the post-genital depressed, quadrangular area disappeared; (5) coxae became strongly separated from one another, particularly coxae II from III and III from IV; (6) the genital aperture became deformed to a plate with a globular extruded protrusion; (7) the spiracle was displaced to a posterior position regarding coxae IV, but without changes in its morphology and size; and (8) the anal plate became dilated (its greatest width increased by one-and-half times) but the anal valves remained unchanged in morphology and size. It is noteworthy that the pseudoscutum preserved its size and pits in the engorged specimen, without signs of dilation, as in the allotype. The pseudoscutum does not change its morphology with engorgement in Nuttalliella either5. The engorged Deinocroton represents the third engorged tick known in the fossil record; the other records have been found in Cretaceous Burmese amber7 and Miocene Dominican amber26., Published as part of Enrique Peñalver, Antonio Arillo, Xavier Delclòs, David Peris, David A. Grimaldi, Scott R. Anderson, Paul C. Nascimbene & Ricardo Pérez-de la Fuente, 2017, parasitised feathered dinosaurs as Cretaceous amber assemblages revealed, pp. 1-13 in Nature Communications 8 (1924) on pages 2-9, DOI: 10.1038/s41467-017-01550-z, http://zenodo.org/record/1116358, {"references":["5. Mans, B. J., de Klerk, D., Pienaar, R. & Latif, A. A. Nuttalliella namaqua: a living fossil and closest relative to the ancestral tick lineage: implications for the evolution of blood-feeding in ticks. PLoS ONE 6, e 23675 (2011).","3. Poinar, G. O. First fossil soft ticks, Ornithodoros antiquus n. sp. (Acari: Argasidae) in Dominican amber with evidence of their mammalian host. Experientia 51, 384 - 387 (1995).","4. Lewis, R. E. & Grimaldi, D. A pulicid flea in Miocene amber from the Dominican Republic (Insecta: Siphonaptera: Pulicidae). Am. Mus. Novit. 3205, 1 - 9 (1997).","2. Voigt, E. Ein Haareinschluss mit Phthirapteren-Eiern im Bernstein. Mitt. Geol. Staatsinst. Hamburg 21, 59 - 74 (1952).","1. Wappler, T., Smith, V. S. & Dalgleish, R. C. Scratching an ancient itch: an Eocene bird louse fossil. Proc. Biol. Sci. 271, S 255 - S 258 (2004).","16. Mans, B. J. et al. Ancestral reconstruction of tick lineages. Ticks Tick Borne Dis. 7, 509 - 535 (2016).","6. Mans, B. J., de Klerk, D. G., Pienaar, R., de Castro, M. H. & Latif, A. A. Nextgeneration sequencing as means to retrieve tick systematic markers, with the focus on Nuttalliella namaqua (Ixodoidea: Nuttalliellidae). Ticks Tick Borne Dis. 6, 450 - 462 (2015).","7. Shi, G. et al. Age constraint on Burmese amber based on UePb dating of zircons. Cret. Res. 37, 155 - 163 (2012).","8. Lamarck, J. B. Systeme des animaux sans vertebres, ou Tableau general des classes, des ordres et des genres de ces animaux. Chez L ' auteur, au Museum d ' Hist. Naturelle, Paris, 1 - 472 (1801).","9. Reuter, E. R. Zur Morphologie und Ontogenie der Acariden mit besonderer Berucksichtigung von Pediculopsis graminum (E. Reut.). Acta Soc. Sci. Fenn. 36, 1 - 288 (1909).","10. Leach, W. E. A tabular view of the external characters of four classes of animals, which Linne arranged under Insecta; with the distribution of the genera composing three of these classes into orders, & c. and descriptions of several new genera and species. Trans. Linn. Soc. Lond. 11, 306 - 400 (1815).","11. Duges, A. L. Recherches sur l ' ordre des Acariens en general et de la famille Trombididies en particulier. Ann. Sci. Nat. Zool. 2, 5 - 46 (1834).","12. Poinar, G. O. & Brown, A. E. A new genus of hard ticks in Cretaceous Burmese amber (Acari: Ixodida: Ixodidae). Syst. Parasitol. 54, 199 - 205 (2003).","13. Poinar, G. O. & Buckley, R. Compluriscutula vetulum (Acari: Ixodida: Ixodidae), a new genus and species of hard tick from lower Cretaceous Burmese amber. Proc. Entomol. Soc. Wash. 110, 445 - 450 (2008).","19. Furman, D. P. & Loomis, E. C. The ticks of California (Acari: Ixodida). Bull. Calif. Insect Surv. 25, 1 - 239 (1984).","25. Roshdy, M. A., Hoogstraal, H., Banaja, A. A. & El Shoura, S. M. Nuttalliella namaqua (Ixodoidea: Nuttalliellidae): spiracle structure and surface morphology. Z. Parasitenkd. 69, 817 - 821 (1983).","23. Keirans, J. E., Clifford, C. M., Hoogstraal, H. & Easton, E. R. Discovery of Nuttalliella namaqua Bedford (Acarina: Ixodoidea: Nuttalliellidae) in Tanzania and redescription of the female based on scanning electron microcopy. Ann. Entomol. Soc. Am. 69, 926 - 932 (1976).","20. Latif, A. A., Putterill, J. F., de Klerk, G., Pienaar, R. & Mans, B. J. Nuttalliella namaqua (Ixodoidea: Nuttalliellidae): first description of the male, immature stages and re-description of the female. PLoS ONE 7, e 41651 (2012)."]}

Cambay amber originates from the warmest period of the Eocene, which is also well known for the appearance of early angiosperm-dominated megathermal forests. The humid climate of these forests may have triggered the evolution of epiphytic lineages of bryophytes; however, early Eocene fossils of bryophytes are rare. Here, we present evidence for lejeuneoid liverworts and pleurocarpous mosses in Cambay amber. The preserved morphology of the moss fossil is inconclusive for a detailed taxonomic treatment. The liverwort fossil is, however, distinctive; its zig-zagged stems, suberect complicate-bilobed leaves, large leaf lobules, and small, deeply bifid underleaves suggest a member of Lejeuneaceae subtribe Lejeuneinae (Harpalejeunea, Lejeunea, Microlejeunea). We tested alternative classification possibilities by conducting divergence time estimates based on DNA sequence variation of Lejeuneinae using the age of the fossil for corresponding age constraints. Consideration of the fossil as a stem group member of Microlejeunea or Lejeunea resulted in an Eocene to Late Cretaceous age of the Lejeuneinae crown group. This reconstruction is in good accordance with published divergence time estimates generated without the newly presented fossil evidence. Balancing available evidence, we describe the liverwort fossil as the extinct species Microlejeunea nyiahae, representing the oldest crown group fossil of Lejeuneaceae. Open-Access-Publikationsfonds 2016 peerReviewed

Spiloconis glaesaria Meinander Figures 1A, B; 2B; 4A Spiloconis glaesaria Meinander, 1998: 33. Engel and Grimaldi, 2007: 19. DIAGNOSIS: Oral margin in frontal view broad (width 0.7× the distance between compound eyes), much of frontal portion of head membranous/lightly sclerotized, most of it covered with fine setulae; dorsal margin of head barely protruding (cf. Spiloconis oediloma Engel and Grimaldi); wing with r-rs proximal to fork of R 2+3 –R 4+5 (i.e., r-rs connected to Rs); L/W basal discal cell 3.2; antenna with 21–22 flagellomeres (23–24 antennomeres), size of basal flagellomere nearly equal to that of other flagellomeres. TYPE AND OTHER MATERIAL: Known only from Dominican amber. Holotype, AMNH DR14-1094, in clear yellow amber 9 × 4 × 5 mm in size, embedded in EpoTek 301 resin, 12 × 3 × 5 mm. The amber contains many bubbles, and the holotype is best observed ventrally. Meinander (1988) stated that the body of the specimen was covered with “wax” (a fine layer of which indeed coats most modern dustywings), but the fossil is actually covered with a milky froth, a preservational artifact common among inclusions in amber. Another specimen was reported (AMNH DR14-1094) with a photograph, in Engel and Grimaldi (2007). It is a wellpreserved female in a small, clear yellow square of amber, 4 × 4 × 1.5 mm., Published as part of Grimaldi, David, Engel, Michael S., Nascimbene, Paul c. & Singh, Hukam, 2013, Coniopterygidae (Neuroptera: Aleuropteryginae) in amber from the Eocene of India and the Miocene of Hispaniola, pp. 1-20 in American Museum Novitates 2013 (3770) on page 4, DOI: 10.1206/3770.2, http://zenodo.org/record/5361898, {"references":["Meinander, M. 1998. Coniopterygidae (Neuroptera) in amber from the Dominican Republic. Journal of Neuropterology 1: 33 - 36.","Engel, M. S., and D. A. Grimaldi. 2007. The neuropterid fauna of Dominican and Mexican amber (Neuropterida: Megaloptera, Neuroptera). American Museum Novitates 3587: 1 - 58."]}

Open-Access-Publikationsfonds 2016