Fossil Ants (Hymenoptera: Formicidae) of the Middle Eocene Kishenehn Formation

A broad range of interesting fossil insects have been discovered recently in Kishenehn Formation shale (middle Eocene, ca. 46 myo) in northwestern Montana, among them a diversity of ant species.  Two hundred forty-nine ant fossils were examined in this study, with 152 them assignable at least to subfamily.  Here, twelve fossil ant species are formally described.  These include a new genus of Dolichoderinae (Ktunaxia, gen. nov.), and the oldest known species from two extant genera: Crematogaster (C. aurora, sp. nov.) and Pseudomyrmex (P. saxulum, sp. nov.).  The Eocene is of particular interest for understanding ant evolution because it is during this period that many present-day speciose and ecologically dominant clades of ants apparently emerged.  In order to understand the evolution of ants, and in particular their march to the terrestrial dominance observed in modern times, it is critical to understand the tempo of ant diversity during the Eocene.  The Kishenehn provides another window into Eocene ant diversity; its relevance to some of the other major Eocene ant fossil deposits is discussed

An Electrophoretic Study of Caecal Proteins of Clinostomum marginatum (Trematoda)

Whole-worm extracts of Clinostomum marginatum were subjected to polyacrylamide gel electrophoresis. Gels of early stage metacercariae, late-stage metacercariae and adult worms contained 26,24, and 27 protein bands respectively. The protein band patterns of the three stages were similar except for 1) a hemoglobin band in gels of adult extracts and 2) a large increase in the quantity of four anodic bands of late-stage gels. The proteins constituting these four bands were present in the caecal contents of late-stage metacercariae . These same proteins were also found in metacercarial cysts as part of an exuded material left behind by the fluke after excystment

A new compression fossil, Eotriadomeroides abjunctus Huber, gen. & sp. nov. (Hymenoptera, Mymaridae), in Eocene shale from the Kishenehn Formation, USA

A new fossil genus and species of fairyfly, Eotriadomeroides abjunctus Huber & Greenwalt, gen. and sp. nov. (Hymenoptera: Chalcidoidea: Mymaridae), is described and illustrated from a female preserved as a compression fossil in middle Eocene shale from the Kishenehn Formation, Montana, USA. It is compared to extant species of Neotriadomerus Huber, known only from Australia, and Triadomerus Yoshimoto, a Cretaceous amber fossil from Canada. It is suggested that these three genera, classified together in Triadomerini, likely the most ancestral lineage of Mymaridae, are evidence of the Middle or perhaps Late Jurassic origin of the family

Eolestes syntheticus Cockerell 1940

<i>Eolestes syntheticus</i> Cockerell, 1940 <p>(Figures 1, 2, 3, 4, 5 A, 5B, 6A, 6B)</p> <p>Synonomy.</p> <p> v. 1940 <i>Eolestes synthetica</i> Cockerell 1940; p. 105, figs. 1, 2. v. 1974 <i>Eolestes syntheticus</i> Fisher 1974; p. 218.</p> <p> <b>Range.</b> Eocene of Northwestern United States (Green River Formation near DeBeque, Colorado, and Kishenehn Formation, Pinnacle, Montana).</p> <p> <b>Holotype</b> (UCM 19170) (Figures 1, 2, 3, 4, 5 A, 6A)</p> <p> <b>Type locality and stratum.</b> 39.5° N, 108.4, Roan Creek near de Begue, Garfield County, Colorado, USA. Green River Formation, Early Eocene, 53.5–48.5 mya (Smith <i>et al.</i> 2003).</p> <p> <b>Re-description.</b> Part and counterpart of a fossil damselfly with all or most of four wings, five legs, the dorsal aspect of the thorax and a poorly preserved head (Fig. 1); pterothorax square, about as long as high (lateral aspect), approximately 4.8 mm long at the mid-dorsal stripe of the mesanepisternum and 3.0 mm wide (between the humeral stripes) (Fig. 2); pro-, meso- and metafemora 3.10, 4.15 and 5.47 mm in length; pro- and mesotibiae 3.36 and 4.17 mm in length; tarsi 3-segmented with pro- and mesotarsi 1.37 and 1.98 mm in length respectively; pro- and mesofemora and pro- and mesotibiae with spines, mesotibial spines gradually tapering distally with a maximum length of 0.6 mm; pro- and mesotibial apical spurs 0.32 and 0.35 mm respectively; protarsal claw 0.36 mm in length.</p> <p>Forewing: Intact, hyaline, 30.62 mm long and 5.34 mm and 7.01 mm wide at nodus and widest point respectively; pterostigma four and a half cells and 2.52 mm long, 0.70 mm wide; pterostigmal brace oblique; distance between wing apex and pterostigma, pterostigma and nodus, nodus and arculus and arculus and base 3.03, 14.17, 5.74 and 5.23 mm respectively; the post-pterostigmal cell area does not appear to contain a supplementary sector; two antenodal and two antesubnodal crossveins with Ax1 basal of the separation of AA’ & AA’’ and separated from Ax2 by 1.61 mm (Fig. 3); Ax2 distal of anterior arcular crossvein; supplementary antenodal crossveins absent; posterior arcular crossvein separating from discoidal cell 0.39 mm below RA; RP separates from RP +MA just below RA+ RP so that RP +MA very short; nodal and subnodal crossveins oblique; dorsal subnodal bracket thickened (Fig. 4); 13 postnodal crossveins most of which are aligned with postsubnodal crossveins; ‘‘lestine’’ oblique vein ‘‘O’’ present 3.5 cells distal of base of RP 2 between RP 2 and IR2; base of IR1 four cells and 3.28 mm distal of RP 2 origin; IR1 underlies a single row of cells that, proximal to pterostigma, are higher than they are wide and overlies a row of seven single cells that transition through three double cells and, distally, an area of reticulated cells. Apical fifth of left forewing torn/split, causing gross misalignment of veins/cells; base of RP 2 three cells and 2.41 mm distal of subnodus and at least six cells and 6.56 mm distal of origin of IR2; IR2 and PR3/ 4 originate much closer to arculus than nodus. IR2 arched abruptly toward RP at base, one half cell and 0.86 mm from origin of RP 3/4; IR2 underlies a row of single cells basally, after which a supplementary longitudinal sector originates proximal to level of pterostigma and branches to five supplementary sectors near wing margin; RP 3/4 originates 1.33 mm from arculus and underlies a single row of single cells that extends to level of pterostigma, after which it is no longer visible; MA of right forewing slightly zigzagged only distal to subnodus; MP underlies a single row of cells that is obscured between subnodus and pterostigma; CuA a prominent vein that leaves subdiscoidal cell 0.33 mm below MP and transitions to a slightly zigzagged pattern at about level of origin of subnodus. It underlies a broad cubital field 1.81 mm in height that starts with a single cell below base of MP but transitions to contain three supplementary longitudinal veins with four rows of cells; anal field consists of a single row of cells that are either square or higher than wide; terminus of CuA at a point 2/3 of the way between nodus and pterostigma; cubito-anal field 2.64 mm in height; CuP distal of separation of AA’ & AA’’ and halfway between Ax1 and Ax2; petiole well defined, 3.69 mm long (from base to separation of AA’ and AA’’) and 12.05% of wing length; Petiole length relative to distance from petiole to nodus = 0.56; discoidal cell closed basally, 1.63 mm long (end of posterior arcular vein to origin of MP) and 0.81 mm wide (origin of arcular vein to origin of MAb) with an acute posterior internal angle of 25 degrees; distal side of discoidal cell (MAb) nearly perpendicular to RA (12 degrees from vertical relative to RA), and 1.01 mm in length; anterior, posterior and basal (posterior arculus) sides of discoidal cell 0.70 mm, 1.55 mm and 0.45 mm long; ratio of lengths of anterior and posterior sides of discoidal cell = 0.45; subdiscoidal cell elongate with no fusion of CuP & AA’ to posterior wing margin distal of CuP; MP slightly arched as it leaves discoidal cell at an angle of 93 degrees between MP and MAb.</p> <p>Hind wing: intact, hyaline, 28.84 mm long and 5.17 mm and 6.68 mm wide at nodus and widest point respectively (Fig. 5 A); pterostigma three and a half cells and 2.62 mm long, 0.72 mm wide; pterostigmal brace oblique; distance between wing apex and pterostigma, pterostigma and nodus, nodus and arculus and arculus and base 2.48, 13.06, 5.36 and 5.43 mm respectively; post-pterostigmal cell area (five cells preserved only) does not appear to contain a supplementary longitudinal sector; two antenodal and two antesubnodal crossveins with Ax2 opposite arcular crossvein and separated from Ax1 by 1.65 mm; supplementary antenodal crossveins absent; subnodal crossvein oblique, nodal crossvein slightly less so; dorsal subnodal bracket slightly thickened; 11 postnodal crossveins exactly aligned with postsubnodal crossveins with exception of last two; ‘‘lestine’’ oblique vein ‘‘O’’ present four cells distal of base of RP 2; base of IR1 four cells and 3.04 mm distal of RP 2 origin; IR1 zigzagged to level of pterostigma and underlies a single row of cells that, proximal to the pterostigma, are higher than wide and overlies a row of seven single cells that expand distally through four double cells, four triple cells via two and then three supplementary longitudinal sectors—distal edge of wing not preserved. Base of RP 2 two and a half cells and 2.23 mm distal of subnodus; IR2 and PR3/4 originate closer to arculus than nodus. IR2 underlies a row of single cells basally, followed by a gradual increase in the number of supplementary longitudinal sectors until there are three supplementary longitudinal sectors distal of pterostigma; RP 3/4 underlies a row of single cells that transitions from cells longer than high to higher than long proximal to stigma; MA zigzagged from a point two cells proximal of subnodus—basal of that point, MA not preserved; MA underlies a row of cells that transitions to double cells at level of IR1 origin and then gradually to 16 small cells at wing margin; MP, not zigzagged, underlies a single row of cells that gradually transitions from square-shaped to cells higher than wide; CuA leaves subdiscoidal cell 0.29 mm below MP, becomes zigzagged as it approaches subnodus, and underlies a broad cubital field, 1.92 mm in height, that starts with a single cell below base of MP and quickly transitions through one and then two supplementary longitudinal sectors and then back to a single row of cells at wing margin (Fig. 6 A); cubito-anal field (between CuA and hind margin) with a single row of cells higher than wide with the exception of the first and a terminus at a point three or four cells from terminus of MP; height of cubito-anal field 2.51 mm; CuP origin distal of separation of AA’ & AA’’ approximately halfway between Ax1 and Ax2; petiole well defined, 3.65 mm long (from base to separation of AA’ and AA’’) and 12.65 % of wing length; Petiole length relative to distance from petiole to nodus = 0.59; discoidal cell poorly preserved; distal side of discoidal cell (MAb) nearly perpendicular to RA and 1.2 mm in length; MP slightly arched as it leaves the discoidal cell.</p> <p>Both part and counterpart also contain a small fossil tipulid dipteran.</p> <p> <b>New specimen</b> (USNM 559049) (Figures 5 B, 6B)</p> <p> <b>Deposition.</b> USNM 559049, National Museum of Natural History, Washington, D.C.</p> <p> <b>Locality and stratum.</b> Spring site, Middle Fork of the Flathead River, Pinnacle, Montana, USA. Coal Creek Member of the Kishenehn Formation, early Middle Eocene, 46.2±0.4 or 43.5±4.9 mya (Constenius, 1996).</p> <p> <b>Description.</b> Apical portion (≈ 70%) of a fossil damselfly wing, hyaline, 21.48 mm long and 5.13 mm and 6.35 mm wide at nodus and widest point respectively (Fig. 5 B); pterostigma four cells and 2.60 mm long, 0.54 mm wide; pterostigmal brace oblique; distance between wing apex and pterostigma and pterostigma and nodus 3.22 and 13.4 mm respectively; post-pterostigmal cell area consists of three single cells proximally, four smaller cells distally and five double cells in between; nodal crossvein and subnodal crossvein oblique; anterior nodal bracket apparently thickened; 12 postnodal crossveins aligned with postsubnodal crossveins except for two most distal; ‘‘lestine’’ oblique vein ‘‘O’’ present three cells distal of base of RP 2; base of IR1 four cells and 3.18 mm distal of RP 2 origin, IR1 underlies a single row of cells that, proximal to the pterostigma, are higher than wide and overlies a row of nine single cells that distally expand through three double cells and, more distally, three rows of cells; base of RP 2 three cells and 2.13 mm distal of subnodus; IR2 and PR3/4 appear to originate closer to arculus than nodus, although their origins are not preserved (IR2 extends basally to edge of fossil at which point it is slightly less than halfway (0.44) between RP 1 and RP 3/4); IR2 underlies a row of at least 18 single cells basally, followed by a gradual increase in number of secondary veins until there are eight small cells at wing margin; RP 3/4relatively straight and underlies a row of single cells, although a short two-cell-long supplementary longitudinal sector appears at level of pterostigma; MA relatively straight proximal of RP 2, only very slightly zigzagged thereafter, and underlies a row of cells that transitions to double cells just distal of IR1 origin and then gradually to 17 small cells at wing margin; MP relatively straight, underlies a single row of cells; CuA zigzagged throughout and terminates three cells short of end of MP. Broad cubital field 1.78 mm in height and contains two supplementary longitudinal sectors that transition to a single supplementary sector through six cells and then disappear ten cells prior to termination of MP (Fig. 6 B); cubito-anal field 2.10 mm in height and with a single row of cells.</p> <p>The same slab also contains a single fossil of the dipteran family Chironomidae.</p>Published as part of <i>Greenwalt, Dale E. & Bechly, Günter, 2014, A re-description of the fossil damselfly Eolestes syntheticus Cockerell, 1940 (Odonata: Zygoptera: Eolestidae n. fam.) with description of new taxa from the Eocene of North America, pp. 138-156 in Zootaxa 3887 (2)</i> on pages 140-143, DOI: 10.11646/zootaxa.3887.2.2, <a href="http://zenodo.org/record/251053">http://zenodo.org/record/251053</a&gt

A re-description of the fossil damselfly Eolestes syntheticus Cockerell, 1940 (Odonata: Zygoptera: Eolestidae n. fam.) with description of new taxa from the Eocene of North America

Greenwalt, Dale E., Bechly, Günter (2014): A re-description of the fossil damselfly Eolestes syntheticus Cockerell, 1940 (Odonata: Zygoptera: Eolestidae n. fam.) with description of new taxa from the Eocene of North America. Zootaxa 3887 (2): 138-156, DOI: http://dx.doi.org/10.11646/zootaxa.3887.2.

Eolestes ramosus Greenwalt & Bechly, 2014, n. sp.

<i>Eolestes ramosus</i> n. sp. <p>(Figures 5 C, 6C, 7)</p> <p> <b>Holotype.</b> USNM 559047, National Museum of Natural History, Washington, D.C.</p> <p> <b>Type locality and stratum.</b> Disbrow Creek site, Middle Fork of the Flathead River, Pinnacle, Montana, USA. Coal Creek Member of the Kishenehn Formation, early Middle Eocene, 46.2±0.4 or 43.5±4.9 mya (Constenius, 1996).</p> <p> <b>Etymology.</b> Species name <i>ramosus</i> from the Latin word ramosus (branching), an indication of the branching of the supplementary longitudinal sectors in the MP-CuA field.</p> <p> <b>Diagnosis.</b> Differs from <i>E. syntheticus</i> by the following characters: 1) CuP origin at instead of distal of the separation of AA’ and AA’’; 2) three supplementary longitudinal sectors in the cubital field vs. two; 3) 2–4 rows of cells between distal parts of IR2 and RP 3/4 vs. 1–2.</p> <p> <b>Description.</b> An intact fossil damselfly wing, hyaline, 27.1 mm long and 6.06 mm and 7.13 mm wide at nodus and widest point respectively (Fig. 5 C); pterostigma four cells and 3.05 mm long, 0.61 mm wide; pterostigmal brace oblique; distance between wing apex and pterostigma, pterostigma and nodus, nodus and arculus and arculus and base 2.43, 12.13, 5.8 and 3.66 mm respectively; petiole length relative to distance from petiole to nodus = 0.44; post-pterostigmal cell area consists of two single cells proximally, three smaller cells distally and five double cells in between; two antenodal and two antesubnodal crossveins. Ax1 opposite separation of AA’ & AA’’ and separated from Ax2 by 1.30 mm; Ax2 opposite arcular crossvein just distal to RP origin and anterior arcular crossvein; RP separates from RP +MA just below RA+ RP so that RP +MA very short; supplementary antenodal crossveins absent; posterior arcular crossvein separating from discoidal cell 0.33 mm below RA; nodus and subnodal bracket not preserved, subnodal crossvein apparently oblique; 14 postnodal crossveins exactly aligned with postsubnodal crossveins with exception of last four; ‘‘lestine’’ oblique vein ‘‘O’’ present four cells distal of base of RP 2; base of IR1 four cells and 2.7 mm distal of RP 2 origin, IR1 zigzagged through first three cells, underlies a single row of cells that, proximal to pterostigma, are higher than they are wide and overlies a row of seven single cells that distally expand through six double cells and subsequently, an area of reticulated cells with two and three secondary longitudinal veins; base of RP 2 two cells and 1.46 mm distal of subnodus and six cells and 5.41 mm distal of origin of IR2; IR2 and PR3/4 originate much closer to arculus than nodus. IR2 arched abruptly toward RP at base with crossvein opposite, one cell and 0.86 mm from origin of RP 3/4; IR2 underlies a row of 18 single cells basally, followed by a gradual increase in number of secondary veins until there are 11 small cells at wing margin; RP 3/4 originates one cell and 1.30 mm from arculus and underlies a row of single cells that transitions from cells longer than high to higher than long before double cells appear just proximal to beginning of pterostigma; MA slightly zigzagged after a point about midway between arculus and nodus, and underlies a row of cells that transitions to double cells at level of IR1 origin and then gradually to 17 small cells at wing margin; MP underlies a single row of cells that gradually transitions from square-shaped to cells higher than wide; CuA a prominent vein that leaves subdiscoidal cell 0.29 mm below MP and transitions to a zigzagged pattern at about level of origin of RP 2; it underlies a broad cubital field 2.20 mm in height that starts with a single cell below base of MP and quickly transitions through one, two and then three secondary longitudinal veins and then back to two secondary veins at wing margin (Fig. 6 C); cubito-anal field consists of a single row of cells (12 distal of subnodus) that, except for the first, are higher than wide, and terminates at a point halfway between nodus and pterostigma; cubito-anal field 2.85 mm in height; CuP origin at separation of AA’ & AA’’ (Fig. 6), directly below Ax1; petiole well defined, 2.51 mm long (from base to separation of AA’ and AA’’) and 9.3% of wing length; discoidal cell closed basally, 1.50 mm long (end of posterior arcular vein to origin of MP) and 0.73 mm wide (origin of arcular vein to origin of MAb) with an acute posterior internal angle of 27 degrees (Note: given the curved nature of the posterior portions of MAb and MP & CuA, the angle is that between lines that align with the majority of these two veins.); distal side of discoidal cell (MAb) nearly perpendicular to RA (7 degrees from vertical relative to RA), and 1.11 mm in length; anterior, posterior and basal (posterior arculus) sides 0.52 mm, 1.45 mm and 0.39 mm long; Ratio of lengths of anterior and posterior sides of discoidal cell = 0.36; subdiscoidal cell elongate with no fusion of CuP & AA’ to posterior wing margin distal of CuP; MP only partially preserved but with apparently no or very little arch as it leaves discoidal cell at angles of 93 and 76 degrees from MAb and vertical respectively.</p>Published as part of <i>Greenwalt, Dale E. & Bechly, Günter, 2014, A re-description of the fossil damselfly Eolestes syntheticus Cockerell, 1940 (Odonata: Zygoptera: Eolestidae n. fam.) with description of new taxa from the Eocene of North America, pp. 138-156 in Zootaxa 3887 (2)</i> on page 145, DOI: 10.11646/zootaxa.3887.2.2, <a href="http://zenodo.org/record/251053">http://zenodo.org/record/251053</a&gt

Eolestes Cockerell 1940

Genus <i>Eolestes</i> Cockerell, 1940 <p>(Figures 1, 2, 3, 4, 5, 6, 7)</p> <p> <b>Type species.</b> <i>Eolestes syntheticus</i> Cockerell, 1940.</p> <p> <b>Diagnosis.</b> Same as for monotypic family.</p> <p> <b>Comment.</b> This genus should not be confused with the invalid junior homonym <i>Eolestes</i> Bown & Schankler, 1982 for an Eocene insectivorous mammal, which was replaced by the valid name <i>Auroralestes</i> Holroyd, Bown & Schankler, 2004.</p>Published as part of <i>Greenwalt, Dale E. & Bechly, Günter, 2014, A re-description of the fossil damselfly Eolestes syntheticus Cockerell, 1940 (Odonata: Zygoptera: Eolestidae n. fam.) with description of new taxa from the Eocene of North America, pp. 138-156 in Zootaxa 3887 (2)</i> on page 140, DOI: 10.11646/zootaxa.3887.2.2, <a href="http://zenodo.org/record/251053">http://zenodo.org/record/251053</a&gt

Bolitophila rohdendorfi Greenwalt & Blagoderov 2019, sp. nov.

<i>Bolitophila rohdendorfi</i> sp. nov. <p>(Figures 7, 12.2)</p> <p>urn:lsid:zoobank.org:act: AF0CFA8D-7F46-4A39-8CAC- 3FEB29 BE9DAD</p> <p> <b>Etymology</b>. The specific epithet is a patronym for Prof. Boris Rohdendorf (1904–1977), Russian entomologist and palaeontologist.</p> <p> <b>Holotype</b>. PIN 964 /1318; deposited in the Palaeontological Institute of Russian academy of Sciences (PIN), Moscow, Russia.</p> <p> <b>Type Horizon and Locality.</b> Baltic amber of unknown provenance (Late Eocene). The collection 964 was purchased by PIN in 1948.</p> <p> <b>Diagnosis.</b> R 2+3 straight, ending in R 1; C ends far beyond tip of R 5; <i>tb</i> present, but very short. Stigma weak. Proctiger longer than gonocoxites; gonostyli tapering towards apex, with two sclerotized teeth.</p> <p> <b>Description.</b> Measurements: body: 5.7 mm, wing: 5.1 mm (Figure 7.1). Head with brown face, wider than long, with triangular ventral membranous area above clypeus. Clypeus small, dark brown, triangular. Scape cylindrical as long as wide, pedicel spherical, slightly narrower than scape. Antennal pubescence long, hairs approximately 1.5x the width of flagellomeres; 6 th flagellomere about 10x longer than wide (Figure 7.2). Thorax uniformly brown; mesonotum with a narrow black longitudinal stripe in anterior part. Halteres slightly longer than thorax. Legs: t1/bt1 = 1:1.12, t2/bt2 = 1:0.78, t3/bt3 = 1:0.8. Tibial spurs about as long as tibia width at apex. Wings (Figure 7.4) hyaline, with weakly defined stigma at apex of R 1. Sc ending just beyond the base of Rs1, costa extending beyond R 5 halfway distance to M 1. R 2+3 almost straight, ending on C just distad of tip of R 1. Crossvein <i>rm</i> slightly shorter than M3 section. Crossvein <i>tb</i> (basal bart of M 3+4) present, short, transverse. Crossvein <i>sc-r</i> at the level of <i>tb</i>. R 5, M 1, M 2 and M 3+4 almost parallel. A 1 weakened at apex. Abdomen with tergite 9 very short, trapezoid, as wide as tergite 8; cerci large, round; proctiger longer than gonocoxites; gonostyli tapering, with two black distal teeth (Figure 7.3, 12.2).</p> <p> <b>Remarks.</b> <i>Bolitophila rohdendorfi</i> <b>sp. nov.</b> differs from other species of the genus in the structure of genitalia: the combination of long proctiger and tapering gonostyli bearing two sclerotised teeth.</p>Published as part of <i>Greenwalt, Dale E. & Blagoderov, Vladimir A., 2019, Review of the fossil record of Bolitophilidae, with description of new taxa and discussion of position of Mangas Kovalev (Diptera: Sciaroidea), pp. 546-560 in Zootaxa 4567 (3)</i> on pages 548-552, DOI: 10.11646/zootaxa.4567.3.6, <a href="http://zenodo.org/record/2599061">http://zenodo.org/record/2599061</a&gt

Mangas brevisubcosta Greenwalt & Blagoderov 2019, sp. nov.

<i>Mangas brevisubcosta</i>, sp. nov. <p>(Figure 10)</p> <p>urn:lsid:zoobank.org:act: 113812FA-36E3-4F79-827F-603E42D0DFBB</p> <p> <b>Etymology</b>. The species epithet refers to relatively short subcostal vein of the species and is to be treated as a noun in apposition.</p> <p> <b>Holotype.</b> PIN 5026 /265, part and counterpart (counterpart incorrectly marked 5026/286, another piece is marked with the same number), partial impression of a male; in the collection of Palaeontological Institute of the Russian academy of Sciences, Moscow.</p> <p> <b>Type locality and horizon.</b> Khasurty; Western Transbaikalia, Buryatia, Zakamensk District, in the middle reaches of the River Khasurty 10 km south of the village of Tsakir (50°21’N 103°37’E); Gusinoe Ozero Group, Lower Cretaceous..</p> <p> <b>Description.</b> Wing length 4.5 mm. Antennae slightly shorter than body, flagellomeres with length 6x the width. Median and cubital veins bare. Three palpomeres visible, antepenultimate ovate, length 4x the width, apical two bacilliform, combined lengths equals that of the antepenultimate. Costa extends beyond R 5 for at least 1/3 distance to M 1. Sc short, ending on C before the level of base of Rs1; <i>sc-r</i> at the level of <i>m-cu</i>. Veins R 5, M 1, and M 2 parallel. Section M3 relatively long: section M3/M forks ratio, 1:3.5; M3/M2 sections ration, 1:1.9.</p> <p> <b>Remarks.</b> Similar to <i>M. kovalevi</i> but differs in the shorter Sc ending in the C before the level of base of Rs1, the more distal position of <i>sc-r</i>, the relative shape of veins R 5, M 1, and M 2 and the relatively longer section M3.</p>Published as part of <i>Greenwalt, Dale E. & Blagoderov, Vladimir A., 2019, Review of the fossil record of Bolitophilidae, with description of new taxa and discussion of position of Mangas Kovalev (Diptera: Sciaroidea), pp. 546-560 in Zootaxa 4567 (3)</i> on pages 556-558, DOI: 10.11646/zootaxa.4567.3.6, <a href="http://zenodo.org/record/2599061">http://zenodo.org/record/2599061</a&gt

Bolitophila warreni Greenwalt & Blagoderov 2019, sp. nov.

<i>Bolitophila warreni</i> sp. nov. <p>(Figures 1–6, 12.1)</p> <p>urn:lsid:zoobank.org:act: D7D7EBC6-FA7D-4C0D-8999-210F4C71D276</p> <p> <b>Etymology</b>. The specific epithet is derived from the paternal name of Jill Warren, a middle school science teacher from Corvallis, Montana who assisted in the collection of specimens and whose Science Olympiad team won state championships in 2010, 2012 and 2013.</p> <p> <b>Holotype</b>. USNM 595138; deposited in the Department of Paleobiology, National Museum of Natural History (NMNH), Smithsonian Institution, Washington, District of Columbia.</p> <p> <b>Type Horizon.</b> Middle Eocene Coal Creek member, Kishenehn Formation.</p> <p> <b>Type Locality</b>. Disbrow Creek site, Middle Fork of the Flathead River (Pinnacle, Montana).</p> <p> <b>Diagnosis.</b> R 4 sigmoid, ending in R 1; C ends beyond tip of R 5; <i>tb</i> present, but very short. Stigma very weak. Parameres longer than aedeagus; gonostyli tapering towards apex, simple.</p> <p> <b>Description.</b> Adult male (Figure 1), body length 4.66 mm. Head black, 0.43 mm long, 0.62 mm high. Eyes not visible. Maxillary palp 0.3 mm long. Antenna brown, at least 3.2 mm long (estimated, middle flagellomeres not preserved), 58 mm and 35 mm wide (first and terminal flagellomeres respectively); flagellomeres ~ 8x longer than wide (Figures 1 and 2). Thorax 1.04 mm long. Mesonotum black/dark brown dorsally, lateroteregite, anepisternum and katepisternum light brown, anepimeron yellow; coxae yellow (Figure 2). Haltere very lightly pigmented, spatulate, approximately 0.9 mm long (visible portions 0.73 and 0.39 mm for right and left haltere respectively), 35 mm and 183 mm wide at midpoints of stem and knob respectively. Wing length 4.6 mm, width 1.33 mm (Figure 3). L/W ratio = 3.46. Wing membrane with microtrichia only. Stigma faint, at the apex of R 1. Vein R 2+3 slightly sigmoid, terminating in costa (Figure 4.1). Sc ending at the level of base of Rs1, <i>sc-r</i> at the level of middle of <i>m-cu</i>. Cell r 1 parallel-sided proximad of R 2+3. C extending only slightly beyond R 5. R 5 and M 1 slightly converging at the apex, M 1 and M 2 rather parallel along most of their extension, except at the very tip, M2 strongly diverging from M1; M 4 and CuA curved basally at apex. Cross-veins r-m and m-cu, and the very base of M3+4 (<i>tb</i>) present, oblique. Length of <i>r-m</i> 0.5x the section M3 and equal to the length of <i>m-cu</i>. A 1 present but distal portion not visible due to folding of the wing. Wing surface covered with microtrichia, anterior margin of wing with spiniform setae approximately 50 mm long. All six legs present, setose (Fig. 1). Length of tibial spurs on fore, mid and hind legs 35, 95 and 155 mm, respectively (Fig. 5). Abdomen brown, setose, length 3.51 mm (including terminalia), 0.45 mm wide at distal tip of gonocoxites. Length of segments 7 and 8 respectively 0.8x and 0.6x of that of segment 6. Male genitalia with tergite 9 and cerci short; gonocoxites 0.45 mm wide, 0.15 mm long; gonostyli with long setae on lateral surface, evenly tapering towards apex and very slightly curved inwards, without visible teeth or branches. Parameres longer than aedeagal complex (Figure 6, 12.1).</p> <p> <b>Remarks.</b> The species seems to be close to <i>B. hybrida</i> (Meigen) group but differs from all known species in combination of shape of gonostyli, length of parameres, presence of the distinct base of M 3+4 (<i>tb</i>) and C running beyond the tip of R 5.</p>Published as part of <i>Greenwalt, Dale E. & Blagoderov, Vladimir A., 2019, Review of the fossil record of Bolitophilidae, with description of new taxa and discussion of position of Mangas Kovalev (Diptera: Sciaroidea), pp. 546-560 in Zootaxa 4567 (3)</i> on pages 547-548, DOI: 10.11646/zootaxa.4567.3.6, <a href="http://zenodo.org/record/2599061">http://zenodo.org/record/2599061</a&gt