Search

Eltxo grimaldii sp. nov. (Figs 1, 2) LSID: urn:lsid:zoobank.org:act: 8A273D67-2B1B-4798- A67B-5D2E8AFE245B Holotype. Specimen ES- 07-40, female, housed at the Institutional Collection from the El Soplao Cave (Government of Cantabria), Celis, Cantabria, Northern Spain. It is virtually complete but lacking the apices of the two antennae and distal parts of all the legs except one of the metathoracic legs, due to the limits of the amber fragment that contains it. The specimen is in a small fragment of amber, 4 × 3 × 1 mm in size, and embedded in a regular prism of epoxy resin, 23 × 9 × 1 mm in size, for optimal viewing and curation. No syninclusions were detected. Etymology. Honoring Dr David Grimaldi (AMNH), for his relevant contributions on fossil insects and amber. Diagnosis. Female. The new species differs from the only species described, Eltxo cretaceus (which was based on a unique male), by the following features: distal section of R 1 vein elongate, vein M present in a short basal part and evanescent, arculus vein contacting M very basal and CuA vein contacting wing margin distad level of basal section of Rs. Locality and horizon. Middle Albian amber outcrop of Rábago-El Soplao (near Rábago village, Cantabria, northern Spain), Peñosas Fm. (Najarro et al., 2009, 2010). Description. Female (Figs 1, 2). Body length ca. 1.1 mm. Antennae with only scape, pedicel and 4–5 flagellomeres preserved due to limits of amber fragment (Figs 1D, 2A): preservation of flagellomeres preventing identification of potential sensoria. Ocelli absent. Palpi three-segmented and elongate (Figs 1E, 2B), with palpomeres subequal, distal palpomere slightly longer than others, palpomeres 2.5 times longer than wide; other mouthpart structures obscure. Wing 1.11 mm long, 0.45 mm wide. Wing venation reduced (Figs 1F, 2C). C extending beyond Rs with a clear end. Apex of R1 at half wing lenght.Apex of Rs meeting C just beyond wing apex. Sc preserved. Basal section of Rs slightly evanescent. Vein M present in a short basal part and evanescent. Vein CuA unforked and evanescent at its apex; apex of CuA well distad level of basal section of Rs. Arculus present, with distal half evanescent, contacting M very close to separation of M and CuA. Wings with dense microtrichia and sparse macrotrichia. Wing margin with setae in costal margin and macrotrichia in posterior margin (but only very few of them still visible) (Fig. 1B, C). Distal portions of legs not preserved due to limits of the amber fragment, except to a complete metathoracic leg, which is long and setose, with five tarsomeres; tarsomere 1 longer than 2 (Figs 1H, 2D). All tibiae lacking tibial spurs. Pretarsal claws short and with a small basal prolongation with rounded apex; empodium insconspicuous (Fig. 1G). Abdomen 0.56 mm long (up to the cerci apices). Genitalia obscure, but long ovipositor (0.20 mm); contour of the cerci well visible in lateral view., Published as part of PEÑALVER, ENRIQUE, ARILLO, ANTONIO & NEL, ANDRÉ, 2022, A review of the Cretaceous genus Eltxo (Diptera: Cecidomyiidae) with description of the new species Eltxo grimaldii from El Soplao amber, pp. 461-467 in Palaeoentomology 5 (5) on page 462, DOI: 10.11646/palaeoentomology.5.5.7, http://zenodo.org/record/7333739, {"references":["Najarro, M., Penalver, E., Rosales, I., Perez-de la Fuente, R., Daviero-Gomez, V., Gomez, B. & Delclos, X. (2009) Unusual concentration of Early Albian arthropod - bearing amber in the Basque-Cantabrian Basin (El Soplao, Cantabria, northern Spain): Palaeoenvironmental and palaeobiological implications. Geologica Acta, 7, 363 - 387. https: // doi. org / 10.1344 / 105.000001443","Najarro, M., Penalver, E., Rosales, I., Soriano, C., Perez-de la Fuente, R., Menor-Salvan, C., Lopez del Valle, R., Ortega- Blanco, J. & Delclos, X. (2010) A review of the El Soplao amber outcrop, Early Cretaceous of Cantabria (Spain). Acta Geologica Sinica (English Edition), 84, 959 - 976. https: // doi. org / 10.1111 / j. 1755 - 6724.2010.00258. x"]}

Infraorder Bibionomorpha Hennig, 1948 Family Cecidomyiidae Newman, 1835 Subfamily Micromyinae Rondani, 1856 Tribe Amediini Jaschhof, 2021 Genus Eltxo Arillo & Nel, 2000 Type species. Eltxo cretaceus Arillo & Nel, 2000 Emended diagnosis. Male and female. Cecidomyiid fly with tarsomere 1 longer than 2. Antennae with 13 flagellomeres. Ocelli absent. Wing venation reduced, without vein M or with short and evanescent vein M. Vein CuA unforked. C extending beyond Rs, far apex of wing. R1 joining C at half length of wing. Rs joining C beyond wing apex., Published as part of PEÑALVER, ENRIQUE, ARILLO, ANTONIO & NEL, ANDRÉ, 2022, A review of the Cretaceous genus Eltxo (Diptera: Cecidomyiidae) with description of the new species Eltxo grimaldii from El Soplao amber, pp. 461-467 in Palaeoentomology 5 (5) on page 462, DOI: 10.11646/palaeoentomology.5.5.7, http://zenodo.org/record/7333739, {"references":["Hennig, W. (1948) Die Larvenformen der Dipteren. 1. Akademie- Verlag, Berlin, 185 pp.","Newman, E. (1835) Attempted division on British insects into natural orders. Entomological Magazine, 2, 195 - 227.","Rondani, C. (1856) Dipterologiae Italicae prodromus. Vol 1: Genera Italica ordinis dipterorum ordinatim disposita et distincta et i familias et stirpes aggregata. Parma, 228 pp. https: // doi. org / 10.5962 / bhl. title. 8160","Jaschhof, M. (2021) Areviewofgallmidges (Diptera: Cecidomyiidae) described from Mesozoic deposits. Palaeoentomology, 4 (1), 001 - 014. https: // doi. org / 10.11646 / palaeoentomology. 4.1.1","Arillo, A. & Nel, A. (2000) Two new fossil cecidomyiids flies from the Lower Cretaceous amber of Alava (Spain) (Diptera, Cecidomyiidae). Bulletin de la Societe Entomologique de France, 105 (3), 285 - 288. https: // doi. org / 10.3406 / bsef. 2000.16673"]}

Eltxo grimaldii sp. nov. (Figs 1, 2) LSID: urn:lsid:zoobank.org:act: 8A273D67-2B1B-4798- A67B-5D2E8AFE245B Holotype. Specimen ES- 07-40, female, housed at the Institutional Collection from the El Soplao Cave (Government of Cantabria), Celis, Cantabria, Northern Spain. It is virtually complete but lacking the apices of the two antennae and distal parts of all the legs except one of the metathoracic legs, due to the limits of the amber fragment that contains it. The specimen is in a small fragment of amber, 4 × 3 × 1 mm in size, and embedded in a regular prism of epoxy resin, 23 × 9 × 1 mm in size, for optimal viewing and curation. No syninclusions were detected. Etymology. Honoring Dr David Grimaldi (AMNH), for his relevant contributions on fossil insects and amber. Diagnosis. Female. The new species differs from the only species described, Eltxo cretaceus (which was based on a unique male), by the following features: distal section of R 1 vein elongate, vein M present in a short basal part and evanescent, arculus vein contacting M very basal and CuA vein contacting wing margin distad level of basal section of Rs. Locality and horizon. Middle Albian amber outcrop of Rábago-El Soplao (near Rábago village, Cantabria, northern Spain), Peñosas Fm. (Najarro et al., 2009, 2010). Description. Female (Figs 1, 2). Body length ca. 1.1 mm. Antennae with only scape, pedicel and 4–5 flagellomeres preserved due to limits of amber fragment (Figs 1D, 2A): preservation of flagellomeres preventing identification of potential sensoria. Ocelli absent. Palpi three-segmented and elongate (Figs 1E, 2B), with palpomeres subequal, distal palpomere slightly longer than others, palpomeres 2.5 times longer than wide; other mouthpart structures obscure. Wing 1.11 mm long, 0.45 mm wide. Wing venation reduced (Figs 1F, 2C). C extending beyond Rs with a clear end. Apex of R1 at half wing lenght.Apex of Rs meeting C just beyond wing apex. Sc preserved. Basal section of Rs slightly evanescent. Vein M present in a short basal part and evanescent. Vein CuA unforked and evanescent at its apex; apex of CuA well distad level of basal section of Rs. Arculus present, with distal half evanescent, contacting M very close to separation of M and CuA. Wings with dense microtrichia and sparse macrotrichia. Wing margin with setae in costal margin and macrotrichia in posterior margin (but only very few of them still visible) (Fig. 1B, C). Distal portions of legs not preserved due to limits of the amber fragment, except to a complete metathoracic leg, which is long and setose, with five tarsomeres; tarsomere 1 longer than 2 (Figs 1H, 2D). All tibiae lacking tibial spurs. Pretarsal claws short and with a small basal prolongation with rounded apex; empodium insconspicuous (Fig. 1G). Abdomen 0.56 mm long (up to the cerci apices). Genitalia obscure, but long ovipositor (0.20 mm); contour of the cerci well visible in lateral view.

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)."]}

Calathus elpis n. sp. (Figs. 1–3) Description. Holotype: 1 female preserved in a piece of Baltic amber (16 mm x 5 mm / diameter x width) (Eocene) deposited in the Álava Museum of Natural Sciences (Vitoria, Spain) MCNA- 13638 (the genitalia and other soft tissues are not preserved but the specimen is a female as protarsi are not swollen). Length (from head to elytra apex): 7.5 mm. Color brown-reddish (Fig. 1 b and 1 c). Disk of elytron depigmented at its posterior part (probably due to taphonomic processes) (Fig. 1 b and 1 d). Head (Fig. 1 b, 1 c, 1 d, and 3 a): Large and convex compound eyes. Long and oblique tempora. Two supraocular setae, being the posterior one at the level of posterior edge of eyes. Two setae on the clypeus. Rectangular labrum with six antero-marginal setae. Mandibles without setae on the lateral sulcus. Slender antennae with eleven antennomeres, pubescent from 4 th to 11 th (the 4 th one lacking pubescence in its basal third part). Antennomeres bearing a crown of distal setae except the first one (with only one long seta) and the second one (shorter than the rest and without setae). Mentum tooth not visible. Maxillary and labial palps typical of Calathus. Pronotum (Fig. 1 b and 1 d) quadrangular, not transverse. Anterior margin slightly concave, being its anterior angles protuding but round shaped. Basal margin straight. Lateral margins curved, with its maximum width at the level of the anterior marginal setae. Posterior angles obtuse and blunt. Lateral channel barely evident being wider at its base. Basal fossae not very deep. Prosternum (Fig. 1 c and 2 a) with acuminate linguiform intercoxal apophysis. Intercoxal apophysis of mesosternum (Fig. 1 c and 2 c) W-shaped, that of metasternum (Fig. 1 c and 2 d) subtriangular. Long metaepisternum (Fig. 2 b). All thoracic esternal pieces with smooth tegument. Elytra (Fig. 1 b and 1 d) lateral margins are subparallel, sharped towards the apex. Epipleura not crossed. Rounded shoulders, without humeral teeth. All the striae are visible, from the base to the apex. Striae from 1 st to 7 th clearly stippled, 8 th one sulciform. Lateral umbilicate serie (Fig. 1 d) is continuous, with 15 setigerous pores over the 8 th striae. Interstriae flat and with micropunctures. Discal setae (Fig. 1 d) barely visible due to the presence of a whitish veil partially covering the dorsum of the specimen; one seta visible in the third interval, adjacent to striae 2, two setae adjacent to striae 5 and one seta close to the apex, over the 7 th striae. One apical seta is preserved over the right elytrum. Both elytra are not equal in length (Fig. 1 d) so striae 1 st to 4 th make up different anastomosis on each elytrum: setae 1–4, 2 – 3 on the left elytrum and setae 1–2, 3 – 4 on the right elytrum. This kind of light malformations are quite frequent on recent carabidofauna. Wings well developed. Legs moderately long and gracile (Fig. 1 b, 1 c and 1 d). Spherical prothoracic (Fig. 1 d and 2 a), conical mesothoracic (Fig. 1 d and 2 c) and transverse metathoracic (aliform) coxae (Fig. 1 d and 2 d). Protibial cleaner organ with internal notch and two guide setae (Fig. 3 b). Mesotibial cleaner organ (Fig. 3 c) similar to comb model (sensu Ortuño 1988). Tarsi smooth, long and dorsally glabrous (Fig. 3 b, 3 c and 3 d); onychium ventrally setose with tarsal claws pectinate (Fig. 3 b, 3 c and 3 e). Etymology: Specific epithet is after Elpis (λπίς), who was in the Greek mythology the personification of hope that remained in the Pandora’s box to help people to bear the misfortunes. Elpis (Esperanza in Spanish) is after our colleague, the anthropologist Dra. Esperanza Gutiérrez Redomero. State of preservation. Although the specimen is slightly damaged, the state of preservation is quite good, so morphological and taxonomic studies are possible. Most of soft tissues are digested, being the tegument quite transparent. Dorsal side of the fossil is partially covered by a withish coat (Fig. 1 b and 1 d) but most of diagnostic characters of the genus are visible. Abdominal sternites are partially destroyed (Fig. 1 c) allowing to observe the folded wings under the elytra. Left prothoracic leg is not complete; only the coxa, the trochanter and the base of the femur are preserved (Fig. 1 c). Left mesothoracic leg has lost the tarsi and most part of the tibia (Fig. 1 b, 1 c and 1 d). Right metathoracic leg has lost the tarsi (Fig. 1 c). Left antenna is incomplete, only preserved up to the 4 th antennomere (Fig. 1 b, 1 c and 1 d).

Chimeromyina concilia Arillo & Grimaldi, sp. n. (Fig. 13) Diagnosis. As for genus. Description. HEAD: Fairly broad, width more than 1.5 x width of notum; eyes large, bare, facets not differentiated. Ocelli on small tubercle, tubercle with 2 pairs long setae. Antenna with apical arista; 3 aristomeres present. Vibrissae not present. THORAX: 0.44 mm long, bristly, each side of notum with 2 long notopleural setae and supra-alar seta; 2 pairs of dorsocentrals, row of long setulae anterior to dorsocentrals. Acrostichal setulae in one, extensive row, extended to posterior margin of notum. Scutellum with 2 pairs setae, apical pair long (ca. 2 x length of other pair), cruciate. Wing 0.93 mm long; fairly broad, width less than half the length. Vein C extended to apex of M 1, though obscured because tip of wing is folded; Sc present but very incomplete and faint; R 1 virtually straight; R 2 + 3 parallels R 1, curved costad only at very apex; fork of R 4 and R 5 widely divergent, nearly at right angle; crossveins bm-cu slightly oblique to r-m (not parallel) and proximal to r-m by distance equal to length of r-m; cell dm and crossvein dm-cu present, cell dm large; vein M 2 complete, closing cell dm with crossvein dm-cu; CuA 1 complete; CuA 2 present, complete, forming cell cu p, with incomplete ���spur��� of veins A 1 [+CuA 2?] originating from cell cu p. Anal lobe of wing well developed. Legs long, bristly, with row of stiff setae on ventral margin of hind tibia. ABDOMEN: Very folded and not visible in dorsal view, unclear in lateral view. Type. Sex? (tip of abdomen not preserved), Holotype: SPAIN: ��LAVA, near Pe��acerrada, Escucha Formation, Lower Cretaceous; MCNA 8882. Etymology. From the Latin, concilio, unite separate parts into a whole, connect; in reference to the plesiomorphic features of this genus that link Chimeromyia with other basal Eremoneura. Comments. See discussion of phylogenetic relationships, below., Published as part of Grimaldi, David A., Cumming, Jeffrey M. & Arillo, Antonio, 2009, Chimeromyiidae, a new family of Eremoneuran Diptera from the Cretaceous, pp. 34-54 in Zootaxa 2078 on pages 50-52, DOI: 10.5281/zenodo.187264

Calathus elpis n. sp. (Figs. 1���3) Description. Holotype: 1 female preserved in a piece of Baltic amber (16 mm x 5 mm / diameter x width) (Eocene) deposited in the ��lava Museum of Natural Sciences (Vitoria, Spain) MCNA- 13638 (the genitalia and other soft tissues are not preserved but the specimen is a female as protarsi are not swollen). Length (from head to elytra apex): 7.5 mm. Color brown-reddish (Fig. 1 b and 1 c). Disk of elytron depigmented at its posterior part (probably due to taphonomic processes) (Fig. 1 b and 1 d). Head (Fig. 1 b, 1 c, 1 d, and 3 a): Large and convex compound eyes. Long and oblique tempora. Two supraocular setae, being the posterior one at the level of posterior edge of eyes. Two setae on the clypeus. Rectangular labrum with six antero-marginal setae. Mandibles without setae on the lateral sulcus. Slender antennae with eleven antennomeres, pubescent from 4 th to 11 th (the 4 th one lacking pubescence in its basal third part). Antennomeres bearing a crown of distal setae except the first one (with only one long seta) and the second one (shorter than the rest and without setae). Mentum tooth not visible. Maxillary and labial palps typical of Calathus. Pronotum (Fig. 1 b and 1 d) quadrangular, not transverse. Anterior margin slightly concave, being its anterior angles protuding but round shaped. Basal margin straight. Lateral margins curved, with its maximum width at the level of the anterior marginal setae. Posterior angles obtuse and blunt. Lateral channel barely evident being wider at its base. Basal fossae not very deep. Prosternum (Fig. 1 c and 2 a) with acuminate linguiform intercoxal apophysis. Intercoxal apophysis of mesosternum (Fig. 1 c and 2 c) W-shaped, that of metasternum (Fig. 1 c and 2 d) subtriangular. Long metaepisternum (Fig. 2 b). All thoracic esternal pieces with smooth tegument. Elytra (Fig. 1 b and 1 d) lateral margins are subparallel, sharped towards the apex. Epipleura not crossed. Rounded shoulders, without humeral teeth. All the striae are visible, from the base to the apex. Striae from 1 st to 7 th clearly stippled, 8 th one sulciform. Lateral umbilicate serie (Fig. 1 d) is continuous, with 15 setigerous pores over the 8 th striae. Interstriae flat and with micropunctures. Discal setae (Fig. 1 d) barely visible due to the presence of a whitish veil partially covering the dorsum of the specimen; one seta visible in the third interval, adjacent to striae 2, two setae adjacent to striae 5 and one seta close to the apex, over the 7 th striae. One apical seta is preserved over the right elytrum. Both elytra are not equal in length (Fig. 1 d) so striae 1 st to 4 th make up different anastomosis on each elytrum: setae 1���4, 2 ��� 3 on the left elytrum and setae 1���2, 3 ��� 4 on the right elytrum. This kind of light malformations are quite frequent on recent carabidofauna. Wings well developed. Legs moderately long and gracile (Fig. 1 b, 1 c and 1 d). Spherical prothoracic (Fig. 1 d and 2 a), conical mesothoracic (Fig. 1 d and 2 c) and transverse metathoracic (aliform) coxae (Fig. 1 d and 2 d). Protibial cleaner organ with internal notch and two guide setae (Fig. 3 b). Mesotibial cleaner organ (Fig. 3 c) similar to comb model (sensu Ortu��o 1988). Tarsi smooth, long and dorsally glabrous (Fig. 3 b, 3 c and 3 d); onychium ventrally setose with tarsal claws pectinate (Fig. 3 b, 3 c and 3 e). Etymology: Specific epithet is after Elpis (������ς), who was in the Greek mythology the personification of hope that remained in the Pandora���s box to help people to bear the misfortunes. Elpis (Esperanza in Spanish) is after our colleague, the anthropologist Dra. Esperanza Guti��rrez Redomero. State of preservation. Although the specimen is slightly damaged, the state of preservation is quite good, so morphological and taxonomic studies are possible. Most of soft tissues are digested, being the tegument quite transparent. Dorsal side of the fossil is partially covered by a withish coat (Fig. 1 b and 1 d) but most of diagnostic characters of the genus are visible. Abdominal sternites are partially destroyed (Fig. 1 c) allowing to observe the folded wings under the elytra. Left prothoracic leg is not complete; only the coxa, the trochanter and the base of the femur are preserved (Fig. 1 c). Left mesothoracic leg has lost the tarsi and most part of the tibia (Fig. 1 b, 1 c and 1 d). Right metathoracic leg has lost the tarsi (Fig. 1 c). Left antenna is incomplete, only preserved up to the 4 th antennomere (Fig. 1 b, 1 c and 1 d)., Published as part of Ortu��o, Vicente M. & Arillo, Antonio, 2009, Fossil carabids from Baltic amber ��� I ��� A new species of the genus Calathus Bonelli, 1810 (Coleoptera: Carabidae: Pterostichinae), pp. 55-61 in Zootaxa 2239 on pages 56-59, DOI: 10.5281/zenodo.190516, {"references":["Ortuno, V. M. (1988) Importancia de los caracteres tibiales en la sistematica de Harpalidae (Col. Caraboidea). Actas del III Congreso Iberico de Entomologia, Granada, pp. 263 - 270."]}

Ortuño, Vicente M., Arillo, Antonio (2009): Fossil carabids from Baltic amber – I – A new species of the genus Calathus Bonelli, 1810 (Coleoptera: Carabidae: Pterostichinae). Zootaxa 2239: 55-61, DOI: 10.5281/zenodo.190516

Litoleptis fossilis Arillo, Pe��alver & Garc��a-Gimeno sp. nov. Figs 1���2; Pl. 1 Derivation of name: From the Latin fossilis (dug out of the earth, excavated, etc.). This is the first fossil representative of the genus. Holotype: CPT- 3344 housed in the Fundaci��n Conjunto Paleontol��gico de Teruel-Din��polis (Teruel Province, Spain). Complete, well preserved specimen (Pl. 1, fig. 1) present in a transparent, clear piece of amber of 4 x 4 x 2 mm into an epoxy resin preparation of 22 x 13 x 2 mm. Type locality and stratigraphy: Specimen from San Just outcrop, in the municipality of Utrillas, near the village of Escucha (Teruel Province, Spain). It was collected in a thin level of grey-black claystones very rich in amber, charcoal and also fusinite formed by a palaeofire, which implicates an exceptional resin production in the palaeoforests. Escucha Fm. (La Orden Member), Lower Cretaceous (Lower Albian). Diagnosis: Antennal flagellomere III prolonged medially into a long, densely pubescent, unsegmented style with its basal part bulbous. Labium as long as antennae. Presence of a small spur on metathoracic tibia. R 4 and R 5 end in the costal margin. R 4 + 5 and M 1 + 2 forked at same distance from wing base. Description: Body length 2.17 mm from head frons to terminalia apex (Fig. 1 and Pl. 1, fig. 1). Head. Eyes bare, large, 0.49 mm long, 0.40 mm aprox. wide (estimated); dorsal and ventral facets undifferentiated (Fig. 2 a and Pl. 1, fig. 2). Eyes nearly holoptic, separated only by distance less to diameter of facet. Postocciput with a dorsal line of fine setae. Ocellar triangle prominent and setulose (ocellar diameter 0.07 mm) (Figs. 1 and 2 a). Antennae 0.38 mm long, with flagellomere I ovoid; flagellomere II short, nearly spherical (0.06 x 0.06 x 0.04 mm); flagellomere III prolonged medially into a long (0.30 mm), densely Figure 1. Camera lucida drawing of Litoleptis fossilis sp. nov. (Spaniidae), female holotype (CPT- 3344), in lateral habitus, from San Just amber. Arrow indicates the metathoracic tibial spur. Thorax: Notum high and rounded (Fig. 1). Scutum with numerous setae partially arranged in rows (Pl. 1, fig. 2). Scutellum strongly acuminate in lateral view, with 4 pairs of long setae in dorsal position (Pl. 1, fig. 2). Wings elongate, 2.02 mm length, 0.73 mm width. C, R 4 +R 5 and h veins strongly sclerotized (Fig. 2 d and Pl. 1, fig. 5). R 2 + 3 and distal part of Sc slightly sclerotized. C vein is circumambient but only strongly sclerotized up to end of R 5. Stigma large, filling space between veins R 1 and R 2 + 3. (Pl. 1, fig. 5). R 2 + 3 slightly sinuous. Stem of R 2 + 3 and R 4 + 5 contacts R 1 at level of h or slightly basally. R 4 + 5 and M 1 + 2 forked at same distance from wing base and near the distal margin, forming broad, short cells. R 4 and R 5 end in the dorsal margin. Stem of M separates cells br and bm. Discal cell absent. Vein CuA 2 complete, reaching margin of wing but not meeting A 1 before wing margin (they separate distally 0.06 mm). Basal margin of anal lobe with long, fine setae. Halter bare, ca. 0.5 mm long, having elongate knob. Legs long and thin (Fig. 1 and Pl. 1, fig. 1). Protarsus 0.60 mm length (tarsomere I 0.25, II 0.10, III 0.08, IV 0.06, V 0.09). Mesothoracic leg 1.57 mm length (femur 0.48, tibia 0.59, tarsomere I 0.26, II 0.08, III 0.06, IV 0.04, V 0.06). Metathoracic leg 1.72 mm length (femur 0.63, tibia 0.59, tarsomere I 0.21, II 0.09, III 0.07, IV 0.06, V 0.07) (Fig. 1). Mesothoracic tarsomeres with at least four strong distal setae, not present in pro- and metathoracic ones. Very short tibial spur in posterior subapical position on metathoracic leg (Fig. 1). Empodium pulvilliform; pulvilli smaller than empodium (observed in left mesothoracic tarsus) (Fig. 2 b and Pl. 1, fig. 4). Abdomen broad and short (1.19 mm long including terminalia and 0.46 mm wide) with abundant setae. Genitalia with segment 1 of cercus not visible (probably short and hidden by tergum). Segment 2 of cercus elongate. Sternum 8 nearly rhombic (Fig. 2 c and Pl. 1, fig. 3). Discussion: The new species clearly belongs to the Spaniidae due to the shape of the antenna and the pulvilliform empodium [it is remarkable that James & Turner (1981) erroneusly represented the antenna of L. alaskensis as segmented]. The lack of a discal cell is typical of the genus Litoleptis but this feature has appeared independently in other taxa. The genus Hilarimorpha Schiner (Hilarimorphidae) has a very similar wing venation (Webb 1974) but the shape of the antenna and the presence of a pulvilliform empodium easily differentiates the new species. Curiously, fossil Hilarimorphidae have a well developed discal cell as well as does the extant genus Apystomyia Melander, 1950 (Grimaldi & Cumming 1999; Mostovski 1999). Among rhagionoids, loss of the discal cell is also a character present in one species described by Webb (1969) from New York, USA: Bolbomyia andiscalcella Webb, 1969. However, the antenna of B. andiscalcella has a clearly segmented antennal style. Figure 2. Camera lucida drawings of Litoleptis fossilis sp. nov. (Spaniidae), female holotype (CPT- 3344). Details of the a) head, b) distal mesotarsomere showing claws, pulvilli and the pulvilliform empodium, c) female terminalia in dorsolateral (top), lateral and ventral views, and d) wing. The new species is differentiated from the extant species of Litoleptis by having a small spur on metathoracic tibia. It is also easy to distinguish the new species from L. alaskensis and L. orientalis by having a complete CuA vein (Grimaldi & Cumming, 1999 erroneously figured a complete CuA vein in L. alaskensis). Litoleptis fossilis sp. nov. is differentiated from L. chilensis and from the two undescribed Japanese and Nepalese species (Nagatomi 1982) by the shape of the third antennal segment. In the new Plate 1: Litoleptis fossilis sp. nov. (Spaniidae), female holotype (CPT- 3344), from San Just amber. 1) habitus in dorsal, ventral and lateral (below) views, 2) thorax and head in dorsal view showing chaetotaxy in scutum and scutellum, 3) genitalia in dorso-lateral view, 4) distal left mesotarsomere showing claws, pulvilli and the pulvilliform empodium, and 5) wing. species R 4 and R 5 end in the costal margin, while in the wing apex in all extant species whose venation is known, and R 4 + 5 and M 1 + 2 forked at same distance from wing base and near the distal margin, determining broad, short cells. Also, new species is separated from extant ones by having a long labium that protrudes as much as the antennae. The female genitalia of extant species of Litoleptis are poorly known. The genitalia of L. fossilis sp. nov. seems closely related to genera Spania and Spaniopsis (Nagatomi & Iwata 1976). Palaeoecology: Litoleptis fossilis sp. nov. has a long proboscis and could have been a hematophagous fly. In fact, it has a proboscis larger than the proboscis of ceratopogonids from the same outcrop (see Arillo et al. 2008). Blood-feeding behaviour is not common among living rhagionoids, occurring only in some Spaniidae and Rhagioniinae (Lehane 2005). Palaeobiogeography: Rhagionoids are among the earliest and most diverse brachycerans in the fossil record. The finding of a Cretaceous Litoleptis proves that Spaniidae is an old lineage that was already separated from the rest of rhagionoids in the Cretaceous. However, its appearance in Spanish amber is quite surprising if we consider its currently known Asio-Nearctic disjunt distribution (Saigusa 2006). Obviously, the genus had a wider distribution during the Mesozoic, but a more precise palaeogeographic distribution can be understood only when further fossil records are available. An undescribed specimen of Litoleptis is known from the late Eocene of Florissant (Colorado, USA) (Mostovski pers. comm.)., Published as part of Arillo, Antonio, Pe��alver, Enrique & Garc��a-Gimeno, Victoria, 2009, First fossil Litoleptis (Diptera: Spaniidae) from the Lower Cretaceous amber of San Just (Teruel Province, Spain), pp. 33-39 in Zootaxa 2026 on pages 34-38, DOI: 10.5281/zenodo.274756, {"references":["James, M. T. & Turner, W. J. (1981) [Chapter] 33. Rhagionidae. Pp 483 - 488. In McAlpine, J. F., Peterson, B. V., Shewell, G. E., Teskey, H. J., Vockeroth, J. R. & Wood, D. M. (coords.). Manual of Nearctic Diptera. Volume 1. Research Branch, Agriculture Canada. Monograph No. 27. 674 pp.","Webb, D. W. (1974) A revision of the genus Hilarimorpha (Diptera: Hilarimorphidae). Journal of the Kansas Entomological Society, 47 (2), 172 - 222.","Grimaldi, D. A. & Cumming, J. (1999) Brachyceran Diptera in Cretaceous ambers and Mesozoic diversification of the Eremoneura. Bulletin of the American Museum of Natural History, 239, 1 - 124.","Mostovski, M. B. (1999) On an interesting find of a brachycerous fly (Diptera, Brachycera) in the Jurassic of Kazakhstan. Paleontological Journal, 33 (4), 70 - 72.","Webb, D. W. (1969) A new and unusual species of Bolbomyia (Diptera: Rhagionidae). Journal of the Kansas Entomological Society, 42 (3), 285 - 288.","Nagatomi, A. (1982) The genera of Rhagionidae (Diptera). Journal of Natural History, 16, 31 - 70.","Nagatomi, A. & Iwata, K. (1976) Female terminalia of lower Brachycera - I (Diptera). Beitrage zur Entomologie, 26, 5 - 47.","Arillo, A., Penalver, E. & Delclos, X. (2008) Microphorites (Diptera: Dolichopodidae) from the Lower Cretaceous amber of San Just (Spain), and the co-occurrence of two ceratopogonid species in Spanish amber deposits. Zootaxa, 1920, 29 - 40.","Lehane, M. J. (2005) The biology of blood-sucking in insects. 2 nd Edition. Cambridge University Press, Cambridge. 321 pp.","Saigusa, T. (2006) Asio-Nearctic disjunct distribution in Diptera. 6 th International Congress of Dipterology. Fukuoka, Japan. 23 - 28 September 2006. Abstracts Volume: p. 209."]}

Chimeromyia reducta Grimaldi & Cumming 1999 C. reducta Grimaldi & Cumming 1999: 74. Diagnosis. Immediately distinguished from all other species in the genus by the terminal arista, short posterior ocellar setae; long R 2 + 3 vein gradually sloping towards C; base of M and vein M 2 lost. In Lebanese amber.

Chimeromyia alava Arillo & Grimaldi, sp. n. (Figs. 11, 12) Diagnosis. Similar to C. pilitibia in having crossvein bm-cu slightly proximal to r-m and the male terminalia with a lobed surstylus, but differs by having the surstylus shorter and being trilobed instead of bilobed, R 2 + 3 is shorter and curved more abruptly costad, and the male hind tibia without a brush of long, fine setae. Description. Body length 1.0– 1.45 mm. HEAD: Eyes large, bare, facets not differentiated. Two pairs long ocellar setae present; ocelli on small raised tubercle. Arista situated dorsally on flagellomere one; arista with three articles, basal two very small. Vibrissae present. THORAX: Bristly, with notopleural and supra-alar setae present. Acrostichals in short, single row; one pair of dorsocentals. Scutellum with 2 pairs setae, apical pair longer (this pair cruciate in MCNA 8883). Legs bristly, but without brushes on tibiae or tarsi. Wing: Sc vestigial; C extended to slightly past apex of vein M 1; R 1 meeting C at slightly less than length half length of wing; R 2 + 3 strongly curved toward and meeting C, distances between apices of R 1 and R 2 + 3 only ca. 1 / 2 length of vein R 1 from where it forks with R 2 + 3; vein R 4 + 5 forked with R 4 and R 5 widely divergent, at nearly a right angle; crossveins bm-cu slightly proximal to r-m, separated by distance ca. 0.3 x length of r-m; M 1 straight, nearly parallel to R 5; base of vein M 2 absent, M 2 not reaching wing margin; CuA 1 straight, complete apically. Anal lobe reduced. ABDOMEN: Male terminalia: Fused cerci with pointed apex; epandrium with row of stiff setae on dorsoapical margin; surstylus with three lobes (two lateral ones, one longer, thin mesal one), setulae at apex of only lateral lobes; phallus apically scoop-shaped, slightly S-shaped; postgonites with apex having minute denticles on ventral surface; hypandrium obscured. Female genitalia (MCNA 9318) with cercus large and ovoid in lateral view. Types. All Specimens from SPAIN: ÁLAVA: Peñacerrada, Escucha Formation, Early Cretaceous. Holotype, male, MCNA 9238 (body length 1.45 mm, wing 1.05 mm), with genitalia well preserved. Paratypes: female, MCNA 9318, genitalia well preserved but head and thorax badly preserved (body length 1.25 mm, wing 1.1 mm); male/female, MCNA 8883, a very small specimen (1.00 mm body length), but with venation identical to others in type series; MCNA 8743.2 female, the venation identical to other specimens of C. alava; female, MCNA 8886, head is very distorted and gives an appearance of the arista not being dorsal, venation also identical to others in type series (body length 1.45 mm). All deposited in Álava Museum of Natural Sciences, Vitoria-Gasteiz, Spain. Etymology. Taken directly from Álava, Basque Country of Spain.

Chimeromyia acuta Grimaldi & Cumming 1999 C. acuta Grimaldi & Cumming 1999: 74. Diagnosis. Base of M complete; vein M 2 incomplete, anal lobe and alula lost; readily distinguished on basis of relatively large flagellomere 1, which has an acute ventral margin. Arista dorsal. Sursylus simple (not multilobed), digitiform; postgonite long, apex slightly hooked; apical margin of hypandrium with pair of long thin lobes. In Lebanese amber., Published as part of Grimaldi, David A., Cumming, Jeffrey M. & Arillo, Antonio, 2009, Chimeromyiidae, a new family of Eremoneuran Diptera from the Cretaceous, pp. 34-54 in Zootaxa 2078 on pages 39-40, DOI: 10.5281/zenodo.187264, {"references":["Grimaldi, D. & Cumming, J. (1999) Brachyceran Diptera in Cretaceous ambers and Mesozoic diversification of the Eremoneura. Bulletin of the American Museum of Natural History, 239, 1 - 124."]}

Chimeromyia intriguea Grimaldi & Cumming 1999 (Fig. 4) C. intriguea Grimaldi & Cumming 1999: 71. Diagnosis (revised). Arista dorsal. Base of vein M (proximal to crossveins r-m and bm-cu) complete, but very faint; crossvein r-m either connected to M or with a slight break near connection; vein M 2 incomplete at proximal end (either evanescent or abruptly), distal end either incomplete or connected to wing margin; CuA 1 complete apically, sometimes faint; anal lobe and alula lost; small spur of vein (CuA 2 + CuP) at base of wing; wing very broad, width approximately 0.5 x the length. Female terminalia simple, with small pair of cerci broadly connected to abdomen. In Lebanese amber. Comments. The diagnosis is slightly revised based on a new AMNH female specimen (Fig. 4).

Chimeromyina Arillo & Grimaldi, gen. nov. Diagnosis. At present, defined only on the basis of characters that are symplesiomorphic with respect to Chimeromyia: Arista terminal, with 3 aristomeres; wing with crossveins r-m and bm-cu separated by distance approximately equal to length of r-m; cell dm present, large, with crossvein dm-cu; vein CuA 2, cell cu p and A 1 [+ CuA 2?] present; anal lobe and alula present. Type Species. Chimeromyina concilia, n.sp. Etymology. Derived from the name of the sister genus, Chimeromyia., Published as part of Grimaldi, David A., Cumming, Jeffrey M. & Arillo, Antonio, 2009, Chimeromyiidae, a new family of Eremoneuran Diptera from the Cretaceous, pp. 34-54 in Zootaxa 2078 on page 48, DOI: 10.5281/zenodo.187264

Chimeromyia burmitica Grimaldi & Cumming, sp. n. (Figs. 1���3) Diagnosis. Immediately distinguished from all other species in the family by the loss of vein R 2 + 3; vein M very faint, anal lobe of wing highly reduced, CuA 2 + CuP veins highly reduced to small spur at base of wing; male terminalia with glabrous, sclerotized epandrium; surstyli simple, base enlarged, apical half slender and digitiform. In Burmese amber. Description. A minute fly, body length 1.1 mm. Body apparently yellowish, no apparent color patterns. HEAD: Eyes reddish, without interfacetal setulae, occupying most of lateral surface of head, no differentiation of facets. Two pairs ocellar setae, posterior pair slightly behind posterior ocelli, anterior ocellars slightly posterior to anterior ocellus; ocellar triangle not particularly tuberculate. No setae on frons. One pair of long inner vertical setae present, plus smaller postoculars. Antenna with pedicel cup-shaped, no visible setae; basal flagellomere setulose, lobe-like, with depression and possibly with inserted condyle on mesal surface; rest of flagellum aristate, having two small basal articles and a long, fine, setulose terminal article. Clypeus large, protrudent, apex with pair of thick, stiff setae. Palpus and labellum small; details not visible. THORAX: Mesoscutum with one row of acrostichal setulae; two long rows dorsocentrals flanking acrostichals (slightly larger than acrostichals); each postpronotal lobe with erect seta. Mesoscutellum with single pair of setae (apicals), slightly erect. Wing: Broadly rounded on apex, base narrow, with anal lobe lost. Sc incomplete, apically evanescent; C ending at apex of R 5; R 1 short, apex reaching to midline of C; R 2 + 3 lost; vein R 4 + 5 forked with R 4 widely divergent from R 5, approximately 80 �� divergent from R 5. Vein M very faint, basally and apically evanescent; M 2 present, barely connected to M 1 in short basal fork; crossveins r-m and bm-cu nearly in line, with gap where M intersects. CuA 1 basally sclerotized, portion apical to bm-cu is faint. CuA 2 + CuP highly reduced to small, spur-like vein at base of wing. Legs: Simple, with no distinctive setation or spines, save for row of ca. 6 longer, thin, stiff setae on ventral surface of fore femur. ABDOMEN: fairly long and narrow, tapered apicad, with eight tergites fully visible. Male terminalia: Epandrium short, seemingly heavily sclerotized (darker than rest of sclerites), glabrous save for row of 4 stiff setulae on posterior surface; posteroventral margin with slight notch. Cerci fused medially, long and narrow in lateral view, extended slightly beyond apex of surstyli. Surstyli apparently articulated with epandrium, with basal half broad and apical half slender and digitiform; apparently bare of setulae. Only apex of phallus visible, extended to three-quarters the length of surstyli. Postgonites present, with small dorsal lobe bearing apical seta, several other stiff setae. Hypandrium setulose, simple and lobe-like. Female terminalia (best seen in AMNH Bu 298, where abdomen is cleared, but spermathecal capsules not visible): Cerci lost or not apparent, sternite VIII subtriangular, minute triangular sclerite on ventral surface of apex; no acanthophorite spines. Types. All types in amber from northern Myanmar, Kachin state, near the village of Tanai (ca. 100 km NW of the Mytchkina), coll. 1999 by Leeward Capital Corp. Holotype, male, AMNH Bu 352; Paratypes, two females, AMNH Bu 298, AMNH Bu 504. The holotype is particularly well preserved, though the wings are folded (reconstructed in Fig. 2 B). Etymology. Taken directly from the former name of the Republic of Myanmar., Published as part of Grimaldi, David A., Cumming, Jeffrey M. & Arillo, Antonio, 2009, Chimeromyiidae, a new family of Eremoneuran Diptera from the Cretaceous, pp. 34-54 in Zootaxa 2078 on pages 36-39, DOI: 10.5281/zenodo.187264

Chimeromyia reducta Grimaldi & Cumming 1999 C. reducta Grimaldi & Cumming 1999: 74. Diagnosis. Immediately distinguished from all other species in the genus by the terminal arista, short posterior ocellar setae; long R 2 + 3 vein gradually sloping towards C; base of M and vein M 2 lost. In Lebanese amber., Published as part of Grimaldi, David A., Cumming, Jeffrey M. & Arillo, Antonio, 2009, Chimeromyiidae, a new family of Eremoneuran Diptera from the Cretaceous, pp. 34-54 in Zootaxa 2078 on page 40, DOI: 10.5281/zenodo.187264, {"references":["Grimaldi, D. & Cumming, J. (1999) Brachyceran Diptera in Cretaceous ambers and Mesozoic diversification of the Eremoneura. Bulletin of the American Museum of Natural History, 239, 1 - 124."]}

Chimeromyina Arillo & Grimaldi, gen. nov. Diagnosis. At present, defined only on the basis of characters that are symplesiomorphic with respect to Chimeromyia: Arista terminal, with 3 aristomeres; wing with crossveins r-m and bm-cu separated by distance approximately equal to length of r-m; cell dm present, large, with crossvein dm-cu; vein CuA 2, cell cu p and A 1 [+ CuA 2?] present; anal lobe and alula present. Type Species. Chimeromyina concilia, n.sp. Etymology. Derived from the name of the sister genus, Chimeromyia.

Chimeromyia alava Arillo & Grimaldi, sp. n. (Figs. 11, 12) Diagnosis. Similar to C. pilitibia in having crossvein bm-cu slightly proximal to r-m and the male terminalia with a lobed surstylus, but differs by having the surstylus shorter and being trilobed instead of bilobed, R 2 + 3 is shorter and curved more abruptly costad, and the male hind tibia without a brush of long, fine setae. Description. Body length 1.0��� 1.45 mm. HEAD: Eyes large, bare, facets not differentiated. Two pairs long ocellar setae present; ocelli on small raised tubercle. Arista situated dorsally on flagellomere one; arista with three articles, basal two very small. Vibrissae present. THORAX: Bristly, with notopleural and supra-alar setae present. Acrostichals in short, single row; one pair of dorsocentals. Scutellum with 2 pairs setae, apical pair longer (this pair cruciate in MCNA 8883). Legs bristly, but without brushes on tibiae or tarsi. Wing: Sc vestigial; C extended to slightly past apex of vein M 1; R 1 meeting C at slightly less than length half length of wing; R 2 + 3 strongly curved toward and meeting C, distances between apices of R 1 and R 2 + 3 only ca. 1 / 2 length of vein R 1 from where it forks with R 2 + 3; vein R 4 + 5 forked with R 4 and R 5 widely divergent, at nearly a right angle; crossveins bm-cu slightly proximal to r-m, separated by distance ca. 0.3 x length of r-m; M 1 straight, nearly parallel to R 5; base of vein M 2 absent, M 2 not reaching wing margin; CuA 1 straight, complete apically. Anal lobe reduced. ABDOMEN: Male terminalia: Fused cerci with pointed apex; epandrium with row of stiff setae on dorsoapical margin; surstylus with three lobes (two lateral ones, one longer, thin mesal one), setulae at apex of only lateral lobes; phallus apically scoop-shaped, slightly S-shaped; postgonites with apex having minute denticles on ventral surface; hypandrium obscured. Female genitalia (MCNA 9318) with cercus large and ovoid in lateral view. Types. All Specimens from SPAIN: ��LAVA: Pe��acerrada, Escucha Formation, Early Cretaceous. Holotype, male, MCNA 9238 (body length 1.45 mm, wing 1.05 mm), with genitalia well preserved. Paratypes: female, MCNA 9318, genitalia well preserved but head and thorax badly preserved (body length 1.25 mm, wing 1.1 mm); male/female, MCNA 8883, a very small specimen (1.00 mm body length), but with venation identical to others in type series; MCNA 8743.2 female, the venation identical to other specimens of C. alava; female, MCNA 8886, head is very distorted and gives an appearance of the arista not being dorsal, venation also identical to others in type series (body length 1.45 mm). All deposited in ��lava Museum of Natural Sciences, Vitoria-Gasteiz, Spain. Etymology. Taken directly from ��lava, Basque Country of Spain., Published as part of Grimaldi, David A., Cumming, Jeffrey M. & Arillo, Antonio, 2009, Chimeromyiidae, a new family of Eremoneuran Diptera from the Cretaceous, pp. 34-54 in Zootaxa 2078 on page 48, DOI: 10.5281/zenodo.187264

Chimeromyia mediobscura Grimaldi & Cumming, sp. n. (Figs. 5���8) Diagnosis. Base of M (proximal to crossveins r-m and m-cu) virtually lost, fold-like; R 2 + 3 gradually tapered toward C (not abruptly turned); crossveins r-m and bm-cu contiguous or nearly so, but interrupted where M intersects; anal lobe barely evident, no alula, a small spur of a vein at base of wing (CuA 2 + CuP); two pairs of scutellar setae (posterior pair cruciate); arista dorsal; male: surstylus simple and digitiform, postgonites spatula-shaped, hypandrium cleft with apices of lobes pointed, stVIII with row of large setae on apical margin; female terminalia: simple, tapered to narrow apex, cerci present. In Lebanese amber. Description. A small, bristly species, slightly more robust than other Chimeromyia. HEAD: Wider than long; eyes large, bare, facets not differentiated. Ocellar triangle on small, slightly raised tubercle, with 2 long pairs of ocellar setae; posterior ocellars 0.7 x length of anteriors. Pair of long vertical setae, nearly erect; 4���5 postoccipital setae very close to posterior margin of eye. Antenna with flagellomere I setulose, rounded; arista situated near dorsal margin; 3 aristal articles, basal two aristomeres minute, approximately equal in length. Clypeus protuberant; with pair of facial setae present, projecting anteriad, length 0.5 x that of anterior ocellar setae. Palpus and labellum small; labrum visible in one specimen (Az 285), length c. 3 x the width, with pointed apex. Epipharyngeal teeth, if present, not visible. THORAX: Fairly broad, setose. Scutum with one row 6���8 acrostichal setulae; pair of large dorsocentral setae between supra-alars, ca. 5 small dorsocentrals in each row; two supra-alar setae present, two notopleural setae; one erect postpronotal seta; scutellum with apical pair of long, cruciate setae, subapically with smaller, finer pair, 0.7 x length of apicals. Legs long, femora stout (particularly hind femora). Femora with ventral row of ca. 4���6 stiff, fine setae, lengths about equal to width of femur. Legs largely yellowish, apical two tarsomeres dark brown to blackish, very dark in fore tarsi. Hind tibia of male without plumosity, but with dorsal and ventral row of short, stiff setae (Fig. 5 b). Wing with vein C extended to apex of M 1; Sc incomplete, apex evanescent, crossvein h not present; R 1 meeting C near middle of wing; R 2 + 3 nearly straight, sloped gradually toward C; vein R 4 + 5 forked with R 4 and R 5 widely divergent, length of fork significantly less than base of fork; base of M (proximal to r-m and m-cu) fold-like, virtually absent; M 1 parallel to base of fork R 4 + 5; crossveins r-m and bm-cu in line, sometimes with slight break where they would meet M; vein M 2 thin, almost connected distally to wing margin, proximal end evanescent; CuA 1 tapered, small spur of a vein (CuA 2 + CuP) present at base of vein; although anal lobe present but highly reduced. ABDOMEN: Slender, with male genitalia projected posteriad, not dorsoflexed. Male terminalia: Cerci fused at base, narrow and slightly cleft apically, formed hood-like over phallus and surstyli. Surstyli long, simple lobes with 4���5 small setulae at apex; lengths approximately equal to that of phallus. Phallus narrow, apically scoop-shaped; postgonites laterally flat and spatula-shaped, with stiff apical setae on ventral margin, length slightly less than that of phallus. Hypandrium largely hidden by large sternite VIII; apex of hypandrium cleft, with apex of both lobes pointed. Sternite VII with row of ca. 10 large, stiff setae near apical margin. Female terminalia: simple, tapered to narrow apex with cerci present. Types. All types in Lebanese amber. Holotype, male, Az 649; Paratypes: male, Az 440; male, 274; female, Az 285. All in MHNP. Etymology. In reference to the median vein and its faint structure. Chimeromyia pilitibia Grimaldi & Cumming , sp. n. (Figs. 9, 10) Diagnosis. Antennal arista dorsal, not apical; crossvein bm-cu slightly proximal to r-m, not directly in line; base of vein M sclerotized (not evanescent), bisecting cells br and bm; R 2 + 3 abruptly upturned toward C; two pairs scutellar setae, posterior pair cruciate; hind tibia plumose (probably in males only), with 3���4 longitudinal rows of long, fine setae; male terminalia with bilobed (vs. simple) surstylus. Description: Body length 1.40 mm, wing length 1.0 mm. HEAD: Wider than long; eyes large, bare, facets not differentiated. Ocellar triangle on small, slightly raised tubercle, with 2 long pairs of ocellar setae (setal lengths ca. same lengths as dorsocentral setae). 4���5 fronto-orbital setae very close to margin of eye. Antenna with flagellomere I setulose; arista situated dorsally; 3 aristal articles, basal two minute. Face with clypeus protuberant, pair of large, projecting setae. THORAX: Fairly broad, setose. Mesoscutum with row of acrostichal setulae; large pair of posterior dorsocentral setae, smaller dorsocentrals anteriad, several supra-alar and notopleural setae; scutellum with apical pair of long, cruciate setae, subapically with smaller pair (ca. 0.3 x length of apicals). Legs long and slender; most distinctive feature is plumose hind tibia (probably a male character only), with 4 longitudinal rows of long, fine setae. Wing with vein C extended to apex of M 1; Sc not observable; R 1 meeting C at slightly less than basal third of wing; R 2 + 3 curved upward toward C, distance between its apex and apex of R 1 1 / 2 the length of cell r 1; vein R 4 + 5 forked with R 4 and R 5 widely divergent, length of fork slightly less than base of fork; M 1 parallel to base of fork R 4 + 5; crossvein r-m slightly distal to bm-cu, separated by segment less than 1 / 3 length of either crossvein; vein M 2 very diffuse, fold-like, incomplete at both ends; CuA 1 tapered and incomplete apically; veins CuA 2 + CuP not apparent, although anal lobe well developed. ABDOMEN: Slender, with male genitalia dorsoflexed. Male terminalia: Cerci fused, formed hood-like over phallus and surstyli; surstylus with two long lobes, dorsal one slender and digitiform, both lobes with apical setulae (vental lobe with 2���3 setulae on ventral margin). Phallus apically scoop-shaped; postgonite with apical setae, length approximately equal to that of ventral lobe of surstylus. Hypandrium large, convex. Female terminalia simple, apical segment a long and slender cone; cerci small. Type. Holotype, male, LEBANON: ���Ambre de Hammana/Mdeiru, Aptian inf��rieur, Collection Dany Azar, no. 230 ��� on the label. Specimen is in a tiny (4 x 4 x 1 mm) clear yellow chip of amber, mounted in balsam on a microscope slide. The tiny chip includes the pyritized remains of two other complete Chimeromyia (both females) and the head of a fourth individual. In MNHP. Etymology. From the Greek, pilos (hair), in reference to the hind tibia having a feathering of dense, long, fine setae. Comments. The remains of four of these flies in such a minute piece of amber indicate they were either congregating at a resin flux or were swarming. Swarms in empidoids and other Diptera are usually comprised of males, into which females fly to become mated. The eyes of male Eremoneura that swarm are typically holoptic, but those of Chimeromyiidae are not. This species appears to be one of the most primitive species of Chimeromyia, by plesiomorphically possessing a fairly well developed anal lobe at the base of the wing. In those empidoids that have crossveins bm-cu and r-m nearly contiguous (e.g., some extant tachydromiines), bm-cu is usually distal to r-m, unlike C. pilitibia where bm-cu is slightly proximal to r-m. The primitive new genus Chimeromyina, however, shares the same configuration of crossveins with Chimeromyia., Published as part of Grimaldi, David A., Cumming, Jeffrey M. & Arillo, Antonio, 2009, Chimeromyiidae, a new family of Eremoneuran Diptera from the Cretaceous, pp. 34-54 in Zootaxa 2078 on pages 40-48, DOI: 10.5281/zenodo.187264

Chimeromyia intriguea Grimaldi & Cumming 1999 (Fig. 4) C. intriguea Grimaldi & Cumming 1999: 71. Diagnosis (revised). Arista dorsal. Base of vein M (proximal to crossveins r-m and bm-cu) complete, but very faint; crossvein r-m either connected to M or with a slight break near connection; vein M 2 incomplete at proximal end (either evanescent or abruptly), distal end either incomplete or connected to wing margin; CuA 1 complete apically, sometimes faint; anal lobe and alula lost; small spur of vein (CuA 2 + CuP) at base of wing; wing very broad, width approximately 0.5 x the length. Female terminalia simple, with small pair of cerci broadly connected to abdomen. In Lebanese amber. Comments. The diagnosis is slightly revised based on a new AMNH female specimen (Fig. 4)., Published as part of Grimaldi, David A., Cumming, Jeffrey M. & Arillo, Antonio, 2009, Chimeromyiidae, a new family of Eremoneuran Diptera from the Cretaceous, pp. 34-54 in Zootaxa 2078 on page 39, DOI: 10.5281/zenodo.187264, {"references":["Grimaldi, D. & Cumming, J. (1999) Brachyceran Diptera in Cretaceous ambers and Mesozoic diversification of the Eremoneura. Bulletin of the American Museum of Natural History, 239, 1 - 124."]}

Genus Litoleptis Chillcott, 1963 Litoleptis Chillcott, 1963. Type species: Litoleptis alaskensis Chillcott, 1963, by original designation., Published as part of Arillo, Antonio, Pe��alver, Enrique & Garc��a-Gimeno, Victoria, 2009, First fossil Litoleptis (Diptera: Spaniidae) from the Lower Cretaceous amber of San Just (Teruel Province, Spain), pp. 33-39 in Zootaxa 2026 on page 34, DOI: 10.5281/zenodo.274756, {"references":["Chillcott, J. G. (1963) A new genus of Rhagionidae with notes and descriptions of Bolbomyia Loew. The Canadian Entomologist, 95, 1185 - 1190."]}

Key to the identification of Cretaceous Microphorites 1. Eyes occupying most of the head. Basal flagellomere abruptily tapered, with base bulbous...................... ................................................................................................................ M. oculeus Grimaldi & Cumming –. Eyes normal sized. Basal flagellomere shorter............................................................................................ 2 2. Wing vein C ending at R 4 + 5 ....................................................................... M. similis Grimaldi & Cumming –. Wing vein C ending beyond R 4 + 5................................................................................................................. 3 3. Antennal arista short, as long as basal flagellomere. Vein R 2 + 3 clearly shorter than R 4 + 5.............................. ..................................................................................................................... M. utrillensis Peñalver sp. nov. –. Antennal arista long, around twice basal flagellomere. Vein R 2 + 3 barely shorter than R 4 + 5......................... 4 4. Four strong setae on the scutellum ............................................................................... M. extinctus Hennig –. Two strong setae on the scutellum .............................................................................. M. deploegi Nel et al.

Protoculicoides skalskii Szadziewski & Arillo, 1998 Fig. 4 & Pl. 1, figs. d���e. Material: the specimen CPT- 970 (Figs. 4 a���d & Pl. 1, figs. d���e) in a preparation of 12 x 12 x 2 mm, in origin together with some syninclusions (four hymenopterans of the family Scelionidae, CPT- 966 to CPT- 969, and one thysanopteran, CPT- 971), and the specimen CPT- 3343 (Fig. 4 e) in an amber piece that was divided for the study of its biological inclusions (syninclusions include one Homoptera Perforissidae, 5 Hymenoptera, and 3 Diptera Keroplatidae). For future comparisons, the number of the holotype of Protoculicoides skalskii Szadziewski & Arillo, 1998 was erroneously published in the original description, actually it is MCNA- 8645. Emended and completed description of the species: Largely based on original description by Szadziewski & Arillo (1998) and the two new specimens from San Just (Fig. 4). Total length: 1.3���1.4 mm. Eyes well separated (Figs. 4 b and 4 e). Flagellum with 13 flagellomeres being 2���8 subglobular and 9���13 more elongated (Fig. 4 e). Antennal ratio around 0.93. Proboscis very long, twice as long as the eye height. Total length of head with proboscis half the length of thorax plus abdomen. Palpus very slender, 5 -segmented. Third palpal segment cylindrical, 0.10 mm long, sensilla capitata not visible. Fourth and fifth palpal segments equal in length. Scutellum rounded with four marginal setae. Legs slender, unmodified. Tibial spur of fore leg distinct. Hind tibial spur and tibial comb not visible. Fourth tarsomere almost cylindrical. Claws short, simple, equal, similar on all legs. Tarsal ratio of fore leg 1.7, of mid leg 2.1 and hind leg 2.5 (tarsal ratio of hind leg emended from original description as ratio of holotype was erroneous due to body angle). Wing length 0.8���0.9 mm. Wing membrane without macrotrichia; microtrichia distinct. C and R veins with long hairs on dorsal margin (Fig. 4 d). Media petiolate. Both radial cells well developed, second one somewhat longer than first. Costa ending at tip of vein R 3, costal ratio 0.73. Halter normal shaped (Fig. 4 a). Abdomen not modified, cerci short., Published as part of Arillo, Antonio, Pe��alver, Enrique & Delcl��s, Xavier, 2008, Microphorites (Diptera: Dolichopodidae) from the Lower Cretaceous amber of San Just (Spain), and the co-occurrence of two ceratopogonid species in Spanish amber deposits, pp. 29-40 in Zootaxa 1920 on pages 35-37, DOI: 10.5281/zenodo.184689, {"references":["Szadziewski, R. & Arillo, A. (1998) Biting midges (Diptera: Ceratopogonidae) from the Lower Cretaceous amber from Alava, Spain. Polskie Pismo Entomologiczne, 67, 291 - 298."]}

Genus Protoculicoides Boesel Protoculicoides Boesel, 1937: 50. Type species: Protoculicoides depressus Boesel, 1937, Published as part of Arillo, Antonio, Pe��alver, Enrique & Delcl��s, Xavier, 2008, Microphorites (Diptera: Dolichopodidae) from the Lower Cretaceous amber of San Just (Spain), and the co-occurrence of two ceratopogonid species in Spanish amber deposits, pp. 29-40 in Zootaxa 1920 on page 35, DOI: 10.5281/zenodo.184689

Genus Leptoconops Skuse Leptoconops Skuse, 1889: 288. Type species: Leptoconops stygius Skuse, 1889, Published as part of Arillo, Antonio, Pe��alver, Enrique & Delcl��s, Xavier, 2008, Microphorites (Diptera: Dolichopodidae) from the Lower Cretaceous amber of San Just (Spain), and the co-occurrence of two ceratopogonid species in Spanish amber deposits, pp. 29-40 in Zootaxa 1920 on page 37, DOI: 10.5281/zenodo.184689

Microphorites utrillensis Pe��alver, nov. sp. Figs. 1���3 & Pl. 1, figs. a���c. 2006 Microphorites sp.; Pe��alver et al.; fig. 1. 2007 Microphorites sp.; Delcl��s et al. 2007 Microphorites sp.; Pe��alver el al.; figs. 6 D & 7. 2007 Dipteran Microphorites sp.; Delcl��s & Soriano: fig. 4 H. Holotype: Adult female. Specimen CPT- 963, housed in the Fundaci��n Conjunto Paleontol��gico de Teruel- Din��polis (Teruel Province, Spain). Complete specimen present in a transparent, clear piece of amber of 9 x 5 mm into an epoxy resin preparation of 15 x 9 x 2 mm. The specimen, along with other insects, is trapped within a spider orb web, which is the oldest web in the fossil record (Pe��alver et al., 2006). Type locality: Specimen collected in San Just outcrop, in the municipality of Utrillas, near the village of Escucha (Teruel Province, Spain). Escucha Fm. (La Orden Member), Lower Cretaceous (Lower-Middle Albian). See Pe��alver et al. (2007) and Delcl��s et al. (2007). Derivation of name: After Utrillas, the name of the municipality where the type locality is located. Diagnosis: Distinghished from other female and male Microphorites by the following combination of characters: very short arista (shorter than basal flagellomere), scutellum with two pairs of strong apical setae, costal vein ends at apex of M 1, R 1 ends at a basal position with respect to m-cu crossvein, R 2 + 3 ends in a medial position between the ends of R 1 and R 4 + 5, crossvein r-m at 2 / 5 of the wing length, dm-cu not arched and 3.5 times longer than m-cu, abdomen elongate (not short and compact). Description: Body length 1.70 mm (from head to the evaginated distal part of the abdomen, see Fig. 1 & Pl. 1, fig. a). Head 0.25 mm long, 0.32 mm greatest width. Eye dichoptic (slightly separate in dorsal position), bare, not emarginated near antennae, diameter 0.20 mm, occupying entire lateral surface of head. Ommatidia uniform (Fig. 2 & Pl. 1, fig. b). Postocular setae apparently absent. Three large ocelli in a narrow triangle without ocellar setae. Frons broad and face broader than frons. Clypeus bare. Proboscis relatively long, with abundant fine, long hairs apically. Palp short and one-segmented, flat, slightly longer than broad (Fig. 2), 0.07 mm long and 0.04 mm wide, with short, fine hairs. Strong setae around mouthparts. Aristate antenna 0.26 mm long with fine, dense microtrichia. Scape reduced. Pedicel 0.04 mm long, 0.05 mm wide, with a ring of strong setae apically. Basal flagellomere subtriangular, bilaterally symmetrical, laterally compressed, elongate, basally swollen, 0.12 mm long, 0.04 mm greatest width, 0.05 mm greatest height. Arista apical and short, 0.10 mm long, and 2 -articulated, basal article very small, slightly longer than width (0.01 mm long). Distal article 0.09 mm long. Thorax 0.45 mm long. Proepisternum and prosternum bare. Few strong thoracic setae; scutum with two long, strong setae in a dorsal position and scutellum with two pairs of strong apical setae. Wings 1.18 mm long, 0.44 mm wide, without a costal bristle, and only very slightly pointed at apex. Apart from the diagnostic venational characters the wing venation (Fig. 3 a) shows a C with one row of spinules, a Sc parallel with R 1 except for a short curved stem which apparently touches it apically (the connection with C is not visible; possibly distal part is evanescent, i.e., Sc vein incomplete), all longitudinal veins complete, an R 4 + 5 unbranched and reaching very close to the wing apex, an M 1 straight beyond crossvein dm-cu and flexed apically, a crossvein r-m short and near wing base, an m-cu vein tubular and inclined towards wing apex, a bm-cu crossvein complete, and a CuA 2 curved. The veins are not setose, except C, and the crossvein h is obscured (not drawn in Figs. 1 & 3 a). Anal lobe not well developed. Legs not modified and moderately bristly. Fore leg 1.00 mm long (femur 0.33 mm; tibia 0.35 mm; ta 1 0.12 mm, ta 2 0.06 mm, ta 3 0.05, ta 4 0.03 mm and ta 5 0.06 mm). Mid legs difficult to measure (tarsus approximately 0.36 mm). Hind leg 1.02 mm long (femur 0.35 mm; tibia 0.32 mm; ta 1 0.16 mm, ta 2 0.06 mm, ta 3 0.04 mm, ta 4 0.03 mm and ta 5 0.06 mm); tibia not clavate and first tarsomere of normal shape. Empodium apparently small or setiform, and pulvilli well-developed (Fig. 3 b). Abdomen telescoped, elongate, 1.14 mm long, 0.30 mm wide, with the terminalia extruded (Fig. 3 c & Pl. 1, fig. c) (abdomen partially destroyed and impregnated with resin). Tergite 10 short (0.03 mm long and 0.03 mm wide), with sclerite apparently undivided, and without acanthophorite apical spines. Apical segments of the abdomen with long setulae (segment 10 with setulae only in the cerci). Cerci sclerotized (apparently heavily sclerotized), short (0.03 mm long), held horizontally, lateral in position, with at least four strong bristles each (up to 0.05 mm) and few setulae. Discussion: This female specimen found in San Just Amber clearly belongs to the genus Microphorites Hennig, 1971 (Dolichopodidae: Microphorinae). It shows all the characters indicated by Grimaldi & Cumming (1999) in the translated and summarized diagnosis of Hennig (1971). It corresponds to a new microphorine species which differentiates from the four species found in the Early Cretaceous ambers from Lebanon [M. extinctus, female (Hennig, 1971), M. similis, female, M. oculeus, male (Grimaldi & Cumming, 1999)] and France [M. deploegi males and females (Nel et al., 2004)]. It is closest to the last one mentioned. Microphorites deploegi from Charente-Maritime (France) was described from male and female specimens, but the female was not illustrated. (Nel et al., 2004). The well preserved new female specimen of Microphorites discovered in San Just has been compared in the MNHN of Paris to two paratype females of M. deploegi (ARC 118 -R and ARC 198.4 -R). Female Microphorites utrillensis differs from female M. deploegi in having body that is graceful and smaller (1.7 mm long vs. ca. 2.0 mm), arista approximately as long as the flagellomere I (vs. approximately twice as long in M. deploegi), wing approximately half long (1.2 mm vs. 2.2 mm in French specimen ARC 118 -R), costal vein ending at the end of M 1, abdomen not globular (1.1 x 0.3 mm vs. 0.8 x 0.4 mm), and fewer and finer setae on head, thorax and legs, with two pairs (vs. one pair) of strong apical setae on the scutellum. Interestingly, Microphorites utrillensis has cerci with very long apical setae as in extant genera Heteropsilopus and Microphor, among other dolichopodids (see Sinclair & Cumming, 2006, p. 46), an unlike M. deploegi which has short setulae. The male holotype of M. deploegi has the crossvein r-m spectral and m-cu evanescent (similarly in the female ARC 118 -R), both of which are clearly tubular in the new species (like in the female ARC 198.4 -R of M. deploegi), and its Sc vein is different as well (curved back towards R 1, closely parallel with R 1 but not touching it, connecting C far from the end of R 1 in M. deploegi). In contrast with the new species, a complete Sc connecting C is also present in fossil M. similis and M. oculeus, and in the extant Schistostoma (see Grimaldi & Cumming, 1999, figs. 37���38 and Sinclair & Cumming, 2006, p. 35). In M. extinctus Sc connects C but it is evanescent apically (see Hennig, 1971, fig. 21). Nel et al. (2004) did not indicate why males and females were put in the same M. deploegi species; apparently the specimens were found in different amber masses, but the co-occurrence in the same outcrop of several specimens which exhibit similar morphological characteristics suggests they are of the same species. The French males and females have the same antennal morphology and it is in agreement with the sexually dimorphic variation seen in the Microphorinae, which affects to other characters as was indicated by Grimaldi & Cumming (1999: 61). Nel et al. (2004) added to the diagnosis of the genus that the female tergite 10 bears no ���acanthophorite��� apical spines and that the female cercus is heavily sclerotized, held horizontally, bearing short setulae. The new specimen from San Just is the best known preserved female Microphorites and it is in a very clear amber mass, showing the terminalia extruded. This Spanish specimen confirms all the characters for the genus Microphorites added to the diagnosis by Nel et al. (2004), with the single exception that the cercus has long setulae. The wing venation of the new species is very similar to those of Microphorites extinctus described by Hennig (1971) from Lebanese Amber, but differs mainly in having a different Sc vein, an R 1 that ends at a basal position respect to m-cu crossvein, an R 2 + 3 that ends more basally (in a medial position between the ends of R 1 and R 4 + 5), an m-cu vein inclined towards wing apex, a dm-cu not arched, and an elongate abdomen. Microphorites utrillensis nov. sp. differs from M. similis, described by Grimaldi & Cumming (1999) on the basis of a female specimen from Lebanon Amber, mainly in having a different morphology of the basal flagellomere and a distal one that is very short in comparison, a shorter proboscis, a costal vein ending at apex of M 1, a complete Sc, an R 1 ending in a basal position respect to m-cu crossvein, an m-cu vein inclined towards wing apex, a dm-cu 3.5 times longer than m-cu, and an abdomen elongated with distinct terminalia. Microphorites oculeus was described on the basis of a male specimen from Lebanon Amber by Grimaldi & Cumming (1999). This species is easily distinguished from all other known Microphorites, including the new species, by its longer and very abruptly tapered basal flagellomere with a strongly bulbous base and eyes occupying most of the head. Also differs from the new species in having clearly longer articles of the arista, a complete Sc and a costal vein ending at apex of R 4 + 5. Several male and female specimens of Microphorites are known from ��lava Amber probably representing at least four different forms not yet described. Thus an important goal for future researches will be to elucidate if the species of San Just Amber is already know in the ��lava deposit. Nel et al. (2004) provided an extensive discussion about the relationships of the family Dolichopodidae and the ¨ Microphorinae ¨ and ¨ Parathalassiinae ¨ groups, indicating the relevance of the genus Microphorites for the cladistic analysis. Recently, an extensive cladistic analysis of the Empidoidea and basal lineages of the Cyclorrhapha based on morphological characters has been published by Sinclair & Cumming (2006). That analysis has confirmed the monophyly of both groups and the authors included into Empidoidea the five families Empididae, Hybotidae, Atelestidae, Brachystomatidae and Dolichopodidae s.lat. Last, the family Microphoridae was not recognized, and the Microphorinae and Parathalassiinae were assigned to the Dolichopodidae s.lat. by the authors cited., Published as part of Arillo, Antonio, Pe��alver, Enrique & Delcl��s, Xavier, 2008, Microphorites (Diptera: Dolichopodidae) from the Lower Cretaceous amber of San Just (Spain), and the co-occurrence of two ceratopogonid species in Spanish amber deposits, pp. 29-40 in Zootaxa 1920 on pages 31-35, DOI: 10.5281/zenodo.184689, {"references":["Penalver, E., Grimaldi, D. A. & Delclos, X. (2006) Early Cretaceous Spider Web with Its Prey. Science, 312, 1761.","Hennig, W. (1971) Insektenfossilien aus der unteren Kreide. III. Empidiformia (\" Microphoridae \") aus der unteren Kreide und aus dem Baltischen Bernstein; ein Vertreter der Cyclorrhapha aus der unteren Kreide. Stuttgarter Beitrage zur Naturkunde, 232, 1 - 28.","Grimaldi, D. A. & Cumming, J. (1999) Brachyceran Diptera in Cretaceous ambers and Mesozoic diversification of the Eremoneura. Bulletin of the American Museum of Natural History, 239, 124 pp.","Nel, A., Perrichot, V., Daugeron, Ch. & Neraudeau, D. (2004) A new Microphorites in the Lower Cretaceous amber of the Southwest of France (Diptera: Dolichopodidae, \" Microphorinae \"). Annales de la Societe Entomologique de France (n. s.), 40 (1), 23 - 29.","Sinclair, B. J. & Cumming, J. M. (2006) The morphology, higher-level phylogeny and classification of the Empidoidea (Diptera). Zootaxa, 1180, 1 - 172."]}

Genus Microphorites Hennig Microphorites Hennig 1971: 16. Type species: Microphorites extinctus Hennig, 1971, Published as part of Arillo, Antonio, Pe��alver, Enrique & Delcl��s, Xavier, 2008, Microphorites (Diptera: Dolichopodidae) from the Lower Cretaceous amber of San Just (Spain), and the co-occurrence of two ceratopogonid species in Spanish amber deposits, pp. 29-40 in Zootaxa 1920 on page 30, DOI: 10.5281/zenodo.184689, {"references":["Hennig, W. (1971) Insektenfossilien aus der unteren Kreide. III. Empidiformia (\" Microphoridae \") aus der unteren Kreide und aus dem Baltischen Bernstein; ein Vertreter der Cyclorrhapha aus der unteren Kreide. Stuttgarter Beitrage zur Naturkunde, 232, 1 - 28."]}

Leptoconops zherikhini Szadziewski & Arillo, 2003 Fig. 5 & Pl. 1, fig. f. Material: One female CPT- 3345 in a preparation of 4 x 5 x 2 mm. Description: Body length about 1 mm. Eyes widely separated. Antenna with 12 flagellomeres, being 2��� 11 more or less spherical and 12 th cylindrical (Fig. 5 c). Proboscis relatively short with 4 -segmented palpus. Wing length 0.6���0.7 mm, with costal vein not prolonged beyond vein R 3. Vein R 4 + 5 well developed to wing apex (Fig. 5 a). Legs slender, claws simple (Fig. 5 b), and tarsal ratio of hind leg about 1.4. Female cerci elongate, slender, 3.1 times longer than broad (Fig. 5 d). Discussion: The new ceratopogonid specimens from San Just Amber clearly belong to the two species described from ��lava Amber, despite some minor differences in the specimens of P. s k a l s k i i that fit within species variability. These two species possess a long proboscis, indicating haematophagous habits. According to Grimaldi & Engel (2005), vertebrate feeding occurred in the ancestral ceratopogonid, and this habit would be inherited from the ancestor of probably all culicomorphans. Adults in extant species of Leptoconops feed on blood of mammals, birds and reptiles., Published as part of Arillo, Antonio, Pe��alver, Enrique & Delcl��s, Xavier, 2008, Microphorites (Diptera: Dolichopodidae) from the Lower Cretaceous amber of San Just (Spain), and the co-occurrence of two ceratopogonid species in Spanish amber deposits, pp. 29-40 in Zootaxa 1920 on page 37, DOI: 10.5281/zenodo.184689, {"references":["Szadziewski, R. & Arillo, A. (2003) The oldest fossil record of the extant subgenus Leptoconops (Leptoconops) (Diptera: Ceratopogonidae). Acta Zoologica Cracoviensia, 46 (suppl. - Fossil Insects), 271 - 275.","Grimaldi, D. A. & Engel, M. S. (2005) Evolution of the Insects. Cambridge University Press, New York, 755 pp."]}

Key to the identification of Cretaceous Microphorites 1. Eyes occupying most of the head. Basal flagellomere abruptily tapered, with base bulbous...................... ................................................................................................................ M. oculeus Grimaldi & Cumming ���. Eyes normal sized. Basal flagellomere shorter............................................................................................ 2 2. Wing vein C ending at R 4 + 5 ....................................................................... M. similis Grimaldi & Cumming ���. Wing vein C ending beyond R 4 + 5................................................................................................................. 3 3. Antennal arista short, as long as basal flagellomere. Vein R 2 + 3 clearly shorter than R 4 + 5.............................. ..................................................................................................................... M. utrillensis Pe��alver sp. nov. ���. Antennal arista long, around twice basal flagellomere. Vein R 2 + 3 barely shorter than R 4 + 5......................... 4 4. Four strong setae on the scutellum ............................................................................... M. extinctus Hennig ���. Two strong setae on the scutellum .............................................................................. M. deploegi Nel et al., Published as part of Arillo, Antonio, Pe��alver, Enrique & Delcl��s, Xavier, 2008, Microphorites (Diptera: Dolichopodidae) from the Lower Cretaceous amber of San Just (Spain), and the co-occurrence of two ceratopogonid species in Spanish amber deposits, pp. 29-40 in Zootaxa 1920 on page 35, DOI: 10.5281/zenodo.184689

Arillo, Antonio, Ortuño, Vicente M. (2008): Did dinosaurs have any relation with dung-beetles? (The origin of coprophagy). Journal of Natural History 42 (19-20): 1405-1408, DOI: 10.1080/00222930802105130, URL: http://dx.doi.org/10.1080/00222930802105130