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Raphidioptera: Raphidiidae) TranSacTionS of The KanSaS Vol. 114, no. 1-2 acadeMy of Science p. 77-87 (2011) A new snakefly from the Eocene Green River Formation (Raphidioptera: Raphidiidae) Michael S. engel Division of Entomology (Paleoentomology), Natural History Museum, and Department of Ecology and Evolutionary Biology, 1501 Crestline Drive – Suite 140, University of Kansas, Lawrence, Kansas 66049-2811 [email protected] Agulla protomaculata, new species (Raphidiidae: Raphidiinae), is described and figured from a series of males and females preserved as fine compressions in middle Eocene shale from the Green River Formation of Colorado. The specimens are exquisitely preserved, complete with integumental color patterns. The species is compared with other Tertiary snakeflies. Keywords: Neuropterida, Raphidioptera, Raphidiidae, taxonomy, paleontology, Tertiary. inTroducTion be undertaken on the North American species of Raphidioptera and it is possible some new The snakeflies (Raphidioptera) of North species will yet be discovered in the mountains America are one of the lesser encountered of Mexico. orders and, along with the equally infrequent Grylloblattodea (Notoptera) and Timematodea Most North American fossil snakeflies (Phasmatodea), are restricted to the western documented to date have come from the part of the continent, albeit more ‘widespread’ prolific Eocene-Oligocene deposits around than the latter two groups and typically more Florissant in central Colorado, with single southerly than the ice crawlers. Presently species recorded from all other localities (Table there are 31 recognized species of North 2). Interestingly, the most abundant material of American Raphidioptera, mostly of the Tertiary snakeflies occurs in the middle Eocene family Raphidiidae but with some interesting deposits of the Green River Formation in Utah, Inocelliidae (Carpenter, 1936, 1958, 1960; Wyoming, and Colorado and yet this species Woglum and McGregor, 1964; Aspöck and has not been characterized to date. In fact, Aspöck, 1970, 1976, 1978, 1996; Aspöck, unlike most deposits, snakeflies are moderately 1973, 1975, 1982, 1987, 1988; Aspöck abundant relative to most Neuropterida (pers. et al., 1991, 1992, 1994a, 1994b; Aspöck obs.). Herein I provide a formal description for and Contreras-Ramos, 2004) (Table 1). the species found in the Green River deposits Phylogenetic studies have suggested that while (Figs. 1–8). The Green River Formation is the New World genera are monophyletic, they exposed in various sedimentary basins in do not necessarily together comprise a clade northeastern Utah, southern Wyoming, and and may be a grade basal to the Palearctic northwestern Colorado. Insects are abundant in lineages (e.g., Haring et al., 2011) suggesting the Parachute Member within the Piceance and that the former fauna comprises relatively Uinta Basins of these deposits (Wilson, 1978; plesiomorphic elements. Accordingly, the fossil Grande, 1984; Dayvault et al., 1995). The record of Raphidioptera in North America stratigraphy and depositional environment of is of considerable interest for phylogenetic the Green River Formation is summarized by and biogeographic studies on the order. Franczyk et al. (1992), Young (1995), and Hail Considerable phylogenetic work remains to and Smith (1997). 78 Engel Table 1. Checklist of extant North American Raphidioptera. Transactions of the Kansas Academy of Science 114(1-2), 2011 79 Table 2. North American fossil snakeflies (Raphidioptera), updated from Engel (2009). Those placed in Raphidia should be considered of dubious generic assignment and likely belong to Agulla or Alena; similarly, the single species of Fibla could belong to Negha. Format for the description and general SySTeMaTic PaleonTology terminology follows that of Engel (2003, 2009), Aspöck et al. (1991), Jepson and Order Raphidioptera Navás Jarzembowski (2008), and Jepson et al. (2009), Suborder Raphidiomorpha Willmann with emendations to the application of some Infraorder Neoraphidioptera Engel veins and cell names. Herein I have considered Family Raphidiidae Latreille those cells bounded between MP and CuA and Subfamily Raphidiinae Latreille branches of MP to be “discoidal” (rather than Genus Agulla Navás “medial”), reserving the term “medial” for Agulla protomaculata, new species those cells between what is interpreted herein (Figs. 1–9) as MA and MP, and reserving “radial” for cells bounded between R and Rs. I’ve followed Diagnosis — The new species can be Jepson and Jarzembowski (2008) and Jepson distinguished by the following combination et al. (2009) in using the term “discal” for the of features: head with extensive yellow cell between Rs and MA. The system adopted, maculations (Figs. 1–3, 5, 7), legs typically for better or for worse, is outlined in figure 9. with some yellow to light brown coloration Photomicrographs were prepared by the author (e.g., Figs. 1, 5, 6); proximal section of using a Nikon D1x digital camera attached to forewing costal space broader than in other an Infinity K-2 long-distance microscopic lens. North American Paleogene species except Metrics for the holotype are provided in the Raphidia exhumata Cockerell; forewing description, with the range of values from the Sc terminating on forewing margin more paratype series provided within parenthetical proximad, separated from pterostigmal base marks. by distance more than pterostigmal length (distance as long as or typically shorter than 80 Engel Figs. 1–2. Holotype female (part) of Agulla protomaculata Engel, new species (USNM 31487). 1. Photomicrograph of entire specimen (preserved wing length 7.42 mm). 2. Detail of head and pronotum (head length 1.53 mm). pterostigmal length in other Tertiary North cell not reaching apex of pterostigma (reaching American species, e.g., R. creedei Carpenter apex of pterostigma in R. creedei); discal cell and R. exhumata); pterostigma length 4–4.5x open (i.e., rs-ma absent; closed in R. exhumata width [similar to R. creedei and R. exhumata, and R. creedei) (Figs. 4, 9); rs-rs crossveins but distinctly longer than in R. tumulata absent (present in R. creedei); two medial cells (Scudder) and shorter than those of R. formed by presence of rs+ma-mp crossvein funerata Engel, R. mortua Rohwer, and Agulla (single medial cell, i.e., rs+ma-mp absent, mineralensis Engel]; R beyond pterostigma in R. creedei); and terminal branches of MP simple (branching extensively in R. exhumata relatively simple (longer and branched in R. and A. mineralensis) (Figs. 4, 9); second radial exhumata) (Figs. 4, 9). Transactions of the Kansas Academy of Science 114(1-2), 2011 81 Figs. 3–4. Photomicrographs of paratype male of Agulla protomaculata Engel, new species (USNM 58453). 3. Entire specimen (preserved wing length 7.25 mm). 4. Detail of forewing venation. Description — Total length (excluding brown, with tarsi, tibiae, and apical portions ovipositor in females) 8.33 mm (7.86–9.64 of femora light brown to yellow; head with mm); forewing length 7.42 mm (6.42–8.33 extensive maculations on otherwise dark mm), maximal width 1.92 mm (1.75–2.33 brown integument, particular with large yellow mm). Head length 1.53 mm (1.33–1.67 patches laterally and posterior to compound mm), width 1.0 mm (0.7–1.1 mm), slightly eyes (e.g., Figs. 2, 3), with medial stripe of longer to about as long as pronotum, gentling yellow (and of varying widths) running from tapering in width posteriorly, with weak tangent just posterior of compound eyes to neck at junction with pronotum. Antenna posterior of head (Figs. 1–3, 5) (sometimes incomplete in all specimens, at least 36 stripe has a thin line of dark brown dividing flagellar articles. Pronotal length 1.4 mm it; e.g., Fig. 7); small medial patch of yellow (1.36–1.53 mm), depth 0.53 mm (0.47–0.53 typically present on frons, and with basal and mm), width not determinable in holotype lateral border of clypeus yellow, medially on (0.53–0.67 mm), pronotum about as long as clypeus brown to dark brown; labrum brown to thorax. Integument generally brown to dark light brown; antennae brown; female ovipositor 82 Engel Figs. 5–6. Photomicrographs of males of Agulla protomaculata Engel, new species. 5. Entire specimen (part) of paratype USNM 58416 (preserved wing length 7.4 mm). 6. Entire specimen of non-type USNM 31775 (preserved wing length 7.5 mm). light brown to brown, weakly curved, length in most specimens; proximal section of costal 3.8 mm (3.47–3.8 mm). Forewing apparently space relatively broad (about twice width of hyaline, with pterostigma slightly infumate subcostal space at its broadest); Sc terminating Transactions of the Kansas Academy of Science 114(1-2), 2011 83 Figs. 7–8. Photomicrographs of Agulla protomaculata Engel, new species. 7. Entire specimen of paratype USNM 31752 (preserved wing length 7.0 mm). 8. Entire specimen of paratype USNM 52500 (forewing length 7.1 mm), ovipositor incomplete in this specimen and so appearing relatively short. on forewing margin near wing midlength near proximal third of wing length, coalesced (Figs. 4, 9), separated from pterostigmal base with MA for some distance before diverging; by distance more than pterostigmal length; rs-rs crossveins absent (Figs. 4, 9); Rs reaching pterostigma length 4–4.5x width; R beyond wing apex with four terminal branches; two pterostigma simple; second radial cell not medial cells formed by presence of rs+ma- reaching apex of pterostigma; discal cell open mp crossvein; ma-mp present, positioned (i.e., rs-ma absent) (Figs. 4, 9); stem of Rs before apical branching of MA; MA reaching shorter than first abscissa MA, Rs originating posterior border of wing apex with three 84 Engel Fig. 9. Terms for major forewing veins, crossveins, and cells (cell names are in italics) as interpret- ed and applied herein. Forewing of paratype specimen USNM 58453. terminal branches; MP with six terminal Holotype — Female; USNM 31487 (part branches along posterior wing border, terminal and counterpart); Green River Formation, branches of MP relatively simple; single Parachute Member, middle Eocene (Lutetian), mp-mp crossvein present; two discoidal cells northwestern Colorado; deposited in the present of approximately subequal size (Figs.
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