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New Insects from the Earliest Permian of Carrizo Arroyo (New Mexico, USA) Bridging the Gap Between the Carboniferous and Permian Entomofaunas

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New Insects from the Earliest Permian of Carrizo Arroyo (New Mexico, USA) Bridging the Gap Between the Carboniferous and Permian Entomofaunas Insect Systematics & Evolution 48 (2017) 493–511 brill.com/ise New insects from the earliest Permian of Carrizo Arroyo (New Mexico, USA) bridging the gap between the Carboniferous and Permian entomofaunas Jakub Prokopa,* and Jarmila Kukalová-Peckb aDepartment of Zoology, Faculty of Science, Charles University, Viničná 7, CZ-128 43 Praha 2, Czech Republic bEntomology, Canadian Museum of Nature, Ottawa, ON, Canada K1P 6P4 *Corresponding author, e-mail: [email protected] Version of Record, published online 7 April 2017; published in print 1 November 2017 Abstract New insects are described from the early Asselian of the Bursum Formation in Carrizo Arroyo, NM, USA. Carrizoneura carpenteri gen. et sp. nov. (Syntonopteridae) demonstrates traits in hindwing venation to Lithoneura and Syntonoptera, both known from the Moscovian of Illinois. Carrizoneura represents the latest unambiguous record of Syntonopteridae. Martynovia insignis represents the earliest evidence of Mar- tynoviidae. Carrizodiaphanoptera permiana gen. et sp. nov. extends range of Diaphanopteridae previously restricted to Gzhelian. The re-examination of the type speciesDiaphanoptera munieri reveals basally coa- lesced vein MA with stem of R and RP resulting in family diagnosis emendation. Arroyohymen splendens gen. et sp. nov. (Protohymenidae) displays features in venation similar to taxa known from early and late Permian from the USA and Russia. A new palaeodictyopteran wing attributable to Carrizopteryx cf. arroyo (Calvertiellidae) provides data on fore wing venation previously unknown. Thus, all these new discoveries show close relationship between late Pennsylvanian and early Permian entomofaunas. Keywords Ephemeropterida; Diaphanopterodea; Megasecoptera; Palaeodictyoptera; gen. et sp. nov; early Asselian; wing venation Introduction The fossil record of insects from continental deposits near the Carboniferous-Permian boundary is important for correlating insect evolution with changes in climate and in plant ecosystems. One of the most prolific localities is Carrizo Arroyo situated in central New Mexico, USA, with an approximately 100-m-thick section of the latest Pennsylvanian (late Gzelian) to Early Permian (early Asselian or equivalent Wolfcam- pian) fauna and flora (Lucas et al. 2016, Schneider et al. 2016). Other well-known © Koninklijke Brill NV, Leiden, 2017 DOI 10.1163/1876312X-48022160 Downloaded from Brill.com09/25/2021 01:28:06PM via free access <UN> <UN> 494 J. Prokop and J. Kukalová-Peck / Insect Systematics & Evolution 48 (2017) 493–511 insect localities slightly older than Carrizo Arroyo come from Europe as Wettin in Germany and Commentry in France (both Gzhelian) or somewhat younger as Obora in the Czech Republic (Sakmarian–Artinskian). Insects occur in two freshwater lay- ers inter-bedded by marine deposits of the Bursum Formation (Red Tanks Member) of early Asselian age. The entomofauna was first collected almost forty years ago by Kukalová-Peck & Peck (1976) and later extended by Lucas and Krainer (2004), Ras- nitsyn et al. (2004) and by Schneider et al. (2004). However, a number of specimens housed in private and institutional collections are still awaiting description. The fol- lowing insect groups, almost entirely represented by wings, were described in pre- ceding studies: Archaeognatha, Odonatoptera, Diaphanopterodea, Palaeodictyoptera, Megasecoptera, Orthoptera, Caloneurodea, “Grylloblattida”, Blattodea, “Hypoper- lida”, Psocodea, Hemiptera, “Miomoptera”, Mecopterida (Kukalová-Peck and Peck 1976; Shear et al. 1992; Rowland 1997; Rasnitsyn et al. 2004; Schneider et al. 2004; Nel et al. 2012). The material described here was collected by the second author (JKP) in 1976 with the assistance of legendary Czech collectors Mr. Karel Havlata and Mrs. Anna Havlatová (both sampled for decades together with JKP famous Early Permian Moravian localities in the Boskovice furrow). In this contribution, we describe new taxa of Hydropalaeoptera (Ephemeropterida: Syntonopteroidea) and of †Rostropalae- optera (Diaphanopterodea, Megasecoptera, Palaeodictyoptera). Material and methods The fossil specimens preserved in fine grained greenish grey sediment were observed under a stereomicroscope WILD M5A in a dry state. The venation pattern line draw- ings were made directly through a stereomicroscope with a coupled camera lucida. Photographs were taken by digital camera Canon D550 with lenses MP-E 65 mm and EF 50 mm. Original photographs were processed using the image-editing software Adobe Photoshop CS and for some images were used stacking software Helicon Focus Pro 6.2.2. Type material is housed in the Kukalová-Peck collection, awaiting deposition in the Canadian Museum of Nature, Ottawa, ON, Canada. Specimen No. 2927/20639 is deposited in the collection of The Natural History Museum in San Diego, CA, USA. We follow the wing venation nomenclature in general of Kukalová-Peck (1991). The venational symbols used here specified as follows: symbols in capitals denote the longi- tudinal veins (ScA/ScP, subcostal anterior/posterior; RA/RP, radial anterior/posterior; MA/MP, medial anterior/posterior; CuA/CuP, cubital anterior/posterior; AA/AP, anal anterior/posterior); ‘IN-’, concave vein between AA1+2 and AA3+4. Other symbols used for morphological structures: ng, nygma. Systematics and higher divisions follow the concept of Ephemeropterida: Syntonopteroidea sensu Prokop et al. (2010), Pal- aeodictyopteroida sensu Bechly (1996, 2014), Riek (1976), and Sinitshenkova (2002). Systematic Palaeontology Order Syntopterida Kukalová-Peck, Peters & Soldán, 2009 Downloaded from Brill.com09/25/2021 01:28:06PM via free access <UN> <UN> J. Prokop and J. Kukalová-Peck / Insect Systematics & Evolution 48 (2017) 493–511 495 Superfamily Syntonopteroidea Handlirsch, 1911 Family Syntonopteridae Handlirsch, 1911 Included genera after Prokop et al. (2010): Anglolithoneura Prokop et al., 2010, Carri- zoneura gen. nov. (present study), Gallolithoneura Garrouste et al., 2009, Syntonoptera Handlirsch, 1911 (type genus); Lithoneura Carpenter, 1938. Carrizoneura gen. nov. Diagnosis Based on wing venation, costal area relatively narrow, venation with numerous triads along the posterior wing margin; RA and RP are basally separate and placed at the same level as the stem of M (MA and MP); RP pectinate having three main branches, first two secondary bifurcated, all separated by long intercalary veins; MA strongly bowed partially adjacent at length to RP, bearing a long broad fork and intercalary vein IMA; MP forks at the same level as divergence of MA and RP bearing one intercalary vein IMP; CuA basally close to stem of M, CuA anteriorly pectinate ending with five main branches and intercalary veins, first branch on the level of division of MP; CuP simple; groove IN- appears to start at the wing base, forming a constriction of the anal area between AA1+2 and convex AA3+4, AA1+2 touching CuP in one point near wing base; a constriction of the area between AA3+4 and first branch of concave AP at the same point. Etymology Composite name after Carrizo (name of locality) and suffixneura (meaning veins); feminine in gender. Carrizoneura carpenteri gen. nov. et sp. nov. (Figs 1–3) Description Wing venation (?hindwing) originally with dark smoky coloration and rather thick membrane; pattern of sparse transversal cross-veins with partial dense network of small cells present; venation having prominent corrugation of main longitudinal veins to- gether with number of intercalary veins emerging between the terminal branches; main longitudinal veins basally covered by prominent dense setae (Fig. 2); concave ScP ba- sally adjacent to RA nearly parallel to anterior wing margin ending 3.9 mm from wing apex; area between ScP and costal margin with several simple cross-veins; simple RA di- verges from ScP in about 1/3 of the wing length ending close to wing apex; RP diverges from RA about 10 mm from wing base running towards wing apex, distally pectinate with three main branches, first and second secondarily terminally bifurcated; branches of RP interlay with intercalary veins forming prominent corrugation along posterior wing margin; field between RA and RP bears numerous simple straight cross-veins; con- vex MA emerging from M-stem nearly at the same level as that of RA and RP, directed towards RP and touching it for a short distance of 4.1 mm; MA deeply branched ending Downloaded from Brill.com09/25/2021 01:28:06PM via free access <UN> <UN> 496 J. Prokop and J. Kukalová-Peck / Insect Systematics & Evolution 48 (2017) 493–511 Fig. 1. Carrizoneura carpenteri gen. nov. et sp. nov., holotype No. 1/2014, Kukalová–Peck coll., Canadian Museum of Nature, Ottawa, Ontario, Canada. Line drawing of hindwing venation. Scale bar represents 5 mm. with two curved branches and concave intercalary vein IMA between them; concave MP deeply bifurcated nearly at same level as division that of MA and RP, ending with two main branches and intercalary convex vein IMP between them; convex CuA run- ning basally very close to stem of M, anteriorly pectinate with five branches and four intercalary veins ICuA between the them, first branch emerges on the level of division of MP; CuP simple, strongly concave and weakly distally curved; anal area with convex AA1+2 touching CuP in one point near wing base (see detail photograph Fig. 3); a con- striction of the area between AA1+2 and convex AA3+4, and a constriction of the area between AA3+4 and first branch of concave AP at the same point; a strongly concave longitudinal vein between
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