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.com09/27/2021 05:05:25AM via free access
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.com09/27/2021 05:05:25AM via free access
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.com09/27/2021 05:05:25AM via free access
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 AA1+2 and AA3+4 ending on posterior wing margin and ‘IN-’ by Kukalová-Peck (1985); AA1+2 with probably two distal branches; probably two short branches of AP and with intercalaries between each veinal neighbours. Measurements Wing length fragment 44 mm, estimated wing length about 50 mm, wing width in the widest part 19 mm.
Holotype Specimen No. 1/2014, in collection of J. Kukalová–Peck (Ottawa, ON, Canada) awaiting deposition at the Canadian Museum of Nature, Ottawa, ON, Canada. A nearly complete fore wing, well preserved except of the anal area. Antero-medial part of the wing is slightly distorted. Etymology The specific epithet honours Prof. Frank Morgan Carpenter, who worked particularly on Paleozoic insect fauna in the Museum of Comparative Zoology at Harvard Univer- sity, Cambridge, MA, USA.
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Figs 2–5. (2, 3) Carrizoneura carpenteri gen. nov. et sp. nov., holotype No. 1/2014, Kukalová–Peck coll., Canadian Museum of Nature, Ottawa, Ontario, Canada. (2) Photograph of hindwing venation; (3) detail of anal area. (4, 5) Carrizodiaphanoptera permiana gen. nov. et sp. nov., holotype No. 2/2014, Kukalová– Peck coll., Canadian Museum of Nature, Ottawa, Ontario, Canada. (4) Detail of basal veinal splits; (5) photograph of wing venation. Scale bars: 2, 5 = 5 mm, 3, 4 = 2 mm.
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Type locality Carrizo Arroyo, NM, USA.
Type strata Carboniferous/Permian transition, Red Tanks Member, Bursum Formation, insect ho- rizons belong to Early Permian (Asselian) (see Lucas and Krainer, 2004, Lucas et al. 2016).
Discussion Carrizoneura gen. nov. bearing the combination of main diagnostic characters of Syn- tonopteridae, i.e. venation with prominent corrugation of main veins, presence of in- tercalary veins with opposite polarity between the branches of main veins forming triads, MA with a strong anterior curve at its base, coalesced with RP for a distance; anal area with the most anterior branch of AA with a strong curve (‘zigzag’) and touch- ing CuP in one point (Kukalová-Peck 1997, Prokop et al. 2010). Family Syntonopteridae comprise four genera as Syntonoptera Handlirsch, 1911, Lithoneura Carpenter, 1938, Gallolithoneura Garrouste et al., 2009 and Anglolitho- neura Prokop et al. 2010. In addition other incomplete fossil from middle Permian (Gaudalupian) of Lodève Basin in France was described and placed in Syntonopteridae gen. et sp. indet. (Prokop and Nel 2011). First, Gallolithoneura known from the Permian (Gaudalupian) of Var (France) is a poorly known genus based on a mid-wing portion, which differs in CuA having fewer branches in comparison to Carrizoneura (Garrouste et al. 2009). Anglolithoneura known from Langsettian (Lancashire, UK) shares with Carrizoneura considerably broad pectinate CuA differing in separation of branches posteriorly, MA only touching with RP, broadly developed MP with both main branches secondarily forked and presence of M-CuA strut (Prokop et al. 2010). Lithoneura Carpenter, 1938 known by three species (L. lameerei Carpenter, 1938, L. mirifica Carpenter, 1944, L. carpenteri Richardson, 1956) from Moscovian of Mazon Creek (Illinois, USA) shares with Carrizoneura oval wing shape and pattern of veins ScP, RA, RP, MA and MP as present in, e.g., L. mirifica, but differs in less devel- oped CuA having three terminal branches only (Carpenter 1992, fig. 28). Syntonoptera Handlirsch, 1911 known from Moscovian of Mazon Creek, IL, USA is fragmentary fossil, which shares broad area of CuA, MP deeply branched on the level of first branching of CuA like inCarrizoneura , differing mainly in more developed MA with five terminal branches (Carpenter 1987). The venation of Carrizoneura gen. nov. is quite similar to that in the Moscovian genera Lithoneura and Syntonoptera (both older by about 9 million years) and thus represents the youngest unambiguous record of the family Syntonopteridae apart from fragmentary Gallolithoneura and poorly preserved syntonopterid from Lodève Basin in southern France (Garrouste et al. 2009, Prokop & Nel 2011).
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Superorder Palaeodictyopterida Bechly, 1996 Order Diaphanopterodea Handlirsch, 1919 Family Diaphanopteridae Handlirsch, 1906
Type genus Diaphanoptera Brongniart, 1893
New emended diagnosis Wings elongated broadest in distal third with pointed apex, sparse simple straight or slightly oblique crossveins; ScP slightly bowed ending in RA beyond mid-wing; RP diverging from RA before mid-wing sending off 6–8 pectinate branches, the stem of M and the basal portion of MA coherent with radius and RP from the base till after the RP diverges from RA, then MA fuses shortly to RP and reaches posterior margin as a simple vein or with terminal twig (see Figs 7–8); MP fuses basally temporarily to CuA, diverges from it by forming a broad conspicuous arch and then forks dichotomously about 4-5×, the stem of Cu is short and adhering to the stem of M, CuA immediately temporarily connected to the stem of M and to the basal portion of MP and then separates as a simple vein, CuA runs close and in parallel to CuP, cubital field narrow, short basal oblique cross-vein opposite to a narrowing of area between CuA and CuP.
Remarks The re-examination of holotypeD. munieri Brongniart, 1893 revealed different basal separation of the vein MA in contrast to the previous assumption of Carpenter (1992), Kukalová-Peck & Brauckmann (1990), and Béthoux and Nel (2003). The vein MA is behind the divergence from MP markedly connected to the stem of R and RP separat- ing well behind division of RA and RP.
Genus Carrizodiaphanoptera gen. nov.
Type species Carrizodiaphanoptera permiana sp. nov. by present description
Diagnosis Diaphanopterodean genus based on venation pattern; anterior wing margin serrate, ScP straight ending on RA about midwing; veins RA + RP together with stems M and Cu basally slightly curved; RA simple straight, RP pectinate ending with seven branches; MA partly connected to stem of R and RP diverging well behind division of RA and RP; MA ending with two terminal branches; MP basally coalesced with CuA, pectinate end- ing with four main branches, first distally bifurcated; CuA simple; CuP simple diverges from CuA very close to wing base running parallel to CuA, strong slightly oblique cross- vein present between CuA and CuP behind division of MP and CuA.
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Fig. 6. Carrizodiaphanoptera permiana gen. nov. et sp. nov., holotype No. 2/2014, Kukalová–Peck coll., Canadian Museum of Nature, Ottawa, Ontario, Canada. Line drawing of wing venation. Scale bar rep- resents 5 mm.
Etymology Composite name after Carrizo (name of locality) and Diaphanoptera (type genus of family Diaphanopteridae); feminine in gender.
Carrizodiaphanoptera permiana sp. nov. (Figs 4–6)
Description Wing slender broadest in about midwing, membrane originally hyaline with several pigmented yellow areas or spots close to wing base, costal margin serrate, rather strong, nearly straight; ScP running straight parallel to the wing margin ending on RA about midwing 13.5 mm from wing base; several transversal simple cross-veins irregularly spaced in area between ScP and costal wing margin, one cross-vein just opposite end of ScP; RA + RP and stems M + Cu basally very close, connected and slightly curved; division of RA and RP 10.5 mm from the wing base before ending of ScP; one simple cross-vein in area between RA and RP; convex RA simple and straight running towards wing apex; RP pectinate with seven branches covering whole wing apex, first diverges 16.3 mm from wing base and terminally twigged; division RP + MA 7.5 mm from wing base; convex MA running coherent with RA + RP and further with RP, diverging from RP 1.3 mm behind division of RA and RP + MA, MA ending on posterior wing margin with two short branches; concave MP diverges from CuA 1.7 mm behind divi- sion of RA + RP + MA and MP + CuA, MP ending with three or four main branches; long bowed cross-vein present between MA and MP below connection of RP and MA; convex CuA simple; simple CuP diverges from CuA close to wing base; strong slightly oblique crossvein present between CuA and CuP behind division of MP and CuA; anal veins basally well separated from stem of Cu, convex AA diverged 6.1 mm from wing base into AA1+2 and AA3+4, both secondary branched ending with four main branches on posterior wing margin, concave AP pectinate ending with six main offshots on posterior wing margin.
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Measurements Wing length fragment 24.2 mm, estimated wing length about 27 mm, wing width in the widest part 7.6 mm.
Holotype No. 2/2014, in collection of J. Kukalová–Peck, to be placed in the Canadian Museum of Nature, Ottawa, ON, Canada; imprint and counter-imprint of a nearly complete wing, venation well preserved, apex missing; membrane originally hyaline with sec- ondary pigmented yellow areas or spots caused possibly due to taphonomy reasons.
Etymology Named after the Permian period.
Type locality Carrizo Arroyo, NM, USA.
Type strata Carboniferous/Permian transition, Red Tanks Member, Bursum Formation, insect ho- rizons belong to Early Permian (Asselian) (see Lucas and Krainer, 2004, Lucas et al. 2016).
Discussion The pattern of wing venation with basally close RA + RP and stem of M distinctly curved, separation of veins MA and MP beyond this curvature and also alternation of convex and concave branches, presence of prominent horizontal cross-vein between CuA and CuP well correspond to order Diaphanopterodea (Carpenter 1992). This group is unique within Palaeoptera because their members were able to fold wings backward along the abdomen in the rest due to specific arrangement of articular scle- rites (autapomorphy) different from flexing mechanism in Neoptera as described by Kukalová (1974). Béthoux and Nel (2003) revised Diaphanoptera species and proposed new diagnosis for the family Diaphanopteridae. They considered as doubtful assignment of genus Philiasptilon Zalessky, 1932 comprising two species P. maculosum Zalessky, 1932 and P. huenickeni Pinto and Pinto de Ornellas, 1978 mainly due to absence of a short ba- sal cross-vein between CuA and CuP as shared by other diaphanopterid families like Elmoidae Tillyard, 1937 and Parelmoidae Rohdendorf, 1962 (Pinto and Pinto de Or- nellas, 1978, Zalessky, 1932). Béthoux and Nel (2003) propose the exclusion of genus Diaphterum Pinto and Adami-Rodrigues, 1999 (monotypic) from Diaphanopteridae due to absence of narrow area between CuA and CuP (Pinto and Adami-Rodrigues, 1999). We support their conclusions resulting in attribution of sole genus Diaphanop- tera within Diaphanopteridae (Béthoux and Nel, 2003).
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Carrizodiaphanoptera gen. nov. exhibits rather similar pattern of venation to Diaphanoptera Brongniart, 1893 known by two species as D. munieri Brongniart, 1893. and D. vetusta Brongniart, 1893, both from Pennsylvanian (Gzhelian, Stephanian B/C in European subdivision) of Commentry in France. First, Carrizodiaphanoptera and Diaphanoptera share considerably large size of wings with a short ScP ending to RA in about midwing, pectinate RP with numerous branches, MP with 4 main branches, and well differentiated anal area.Carrizodiaphanoptera differs mainly by more slender wing shape with different coloration, pattern of numerous crossveins forming rectan- gular or pentagonal cells along posterior wing margin, very basal separation of veins CuA and CuP, numerous terminal branches of MA and MP, and MA basally diverging from MP connected to the RA + RP and RP instead of rather short connection with RP as considered in Diaphanoptera by Carpenter (1963) followed by Béthoux and Nel (2003). However, the latter character is erroneously interpreted by the previous authors due to different polarity of supposed vein MA behind the division with MP and contact with RP (see Fig. 8 close up photograph to compare with Béthoux and Nel, 2003: fig. 1.2). According to the polarity of veins, we assume that the vein MA first diverges from MP and secondary diverges from RP running to the posterior wing margin exactly as present in Carrizodiaphanoptera. The prominent crossvein between MA and MP (supposed emergence of MA by previous authors) is clearly concave or neutral polarity. Therefore, it seems that there is no reason to consider as MA, but we should admit that the above mentioned part of MA is rather weakly preserved and easily overlooked. Moreover, such pattern of MA is also present in other diaphanop- terodean family Asthenohymenidae (see Carpenter 1992).
Notice Rohdendorf (1962: 77) included the genus Tchirkovaea Zalessky, 1931 in Diapha- nopteridae on the basis of the wing venation. Sinitshenkova (1979) established the family Tchirkovaeidae and attributed it to Palaeodictyoptera comprising two genera as Tchirkovaea Zalessky, 1931 and Paimbia Sinitshenkova, 1979 demonstrating shared characters in wing venation pattern among members of Megasecoptera, Palaeodicty- optera and Diaphanopterodea. This attribution was also followed by Carpenter (1992) and Béthoux and Nel (2003). Our direct examination of venation in Paimbia ultima Sinitshenkova, 1981 exhibits some characterics shared by members of Diaphanop- teridae, such as the presence of bowed veins of ScP, RA + RP and M connected close to the wing base, but stem Cu is well separated (Fig. 9). Veins CuA and CuP are simple with oblique crossveins close to the wing base as present in some members of Diaphanopterodea. Finally, Paimbia share with Diaphanoptera the common presence of nygmata as circular spots or punctures on wing membrane (Figs 9, 10). However, these structures are well known in some members of Holometabola like Hymenoptera, Neuroptera, Mecoptera, and Trichoptera, but scarcely also reported in some members of Palaeodictyoptera (Forbes 1924, 1943; Engel et al. 2013). Family Martynoviidae Tillyard, 1932 Genus Martynovia Tillyard, 1932
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Figs 7–10. (7, 8) Diaphanoptera munieri Brongniart, 1893, holotype No. R51214, Department of Pal- aeontology coll., Museum national d’Histoire naturelle, Paris, France. (7) Photograph of wing venation pattern (immersed under film layer of ethylalcohol); (8) detail of basal part (immersed under film layer of ethylalcohol). (9, 10) Paimbia ultima Sinitshenkova, 1981, holotype No. 3115–192, Palaeontological Institute of RAS coll., Moscow, Russia; (9) Wing venation pattern with position of nygmata; (10) detail of nygma. Scale bars: 7, 9 = 5 mm, 8 = 2 mm.
Martynovia insignis Tillyard, 1932 (Fig. 11)
Material Specimen No. 3/2014, in collection of J. Kukalová–Peck, to be deposited in the Cana- dian Museum of Nature, Ottawa, ON, Canada, imprint and counter-imprint of fore wing, venation well preserved, apex missing.
Discussion The fore wing bears hyaline membrane with pigmentation in apical part of area be- tween RA and costal wing margin. It shares with other Martynoviidae the following characters: broad costal area in basal part, ScP terminating on RA about midwing, RA + RP and veinal stems of M and Cu basally coalesced, RP with four terminal branches forming large part of wing apex, and MA, MP, CuA and CuP represented by simple veins. Veinal pattern suggests attribution to the genus Martynovia Tillyard, 1932 having the vein MA diverges from MP close to origin of RA and RP and shortly connected to RP. The genus comprises four speciesM. insignis Tillyard, 1932, M. halli
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Beckemeyer 2004, M. longipennis Carpenter, 1947, M. protohymenoides Tillyard, 1932 known from early to middle Permian of the Wellington Formation, Kansas and Okla- homa (USA), Obora in Boskovice furrow, Czech Republic and Salagou Formation in Lodève Basin, France (Tillyard 1932, Carpenter, 1947, Beckemeyer 2004, Dostál and Prokop 2009, Prokop et al. 2015). Fore wing specimen No. 3/2014 is closest to M. insignis (Tillyard, 1932) from Kansas and Oklahoma (USA), which now also in- cludes M. aff. insignis from the Guadalupian of Lodève Basin, France (Béthoux et al. 2003). Furthermore, it represents the oldest record of genus Martynoviia from the low- ermost Permian supplementing the previous discovery of Phaneronera Carpenter, 1947 (Phaneroneura rineharti Sinitshenkova, 2004) known from the same locality (Rasnit- syn et al. 2004). Order Megasecoptera Brongniart, 1885 Family Protohymenidae Tillyard, 1924
Arroyohymen gen. nov.
Type species Arroyohymen splendens sp. nov. by present description.
Diagnosis Prominent pterostigmal area darkly pigmented, lens-like narrow field enclosing ante- riorly by ScP and posteriorly by RA; simple MA with long connection to RP adhered shortly behind division of RA and RP.
Etymology Composite name after Arroyo (a watercourse or dry creek from Spanish) and hymen (membrane from Greek); masculine in gender. Arroyohymen splendens sp. nov. (Figs 12, 13, 15)
Description Fore wing and hindwing nearly homonomous, slender, petiolate, broadest in distal third, membrane originally hyaline; hindwing slightly shorter and broader than fore wing; anterior wing margin basally bulging forming enlarged costal area, costal area become very narrow at divergence of veins RA and RP; ScP probably running close RA ending in wing apex; RA running closely parallel to anterior wing margin diverging 2.2 mm from apex, ScP delimit anteriorly darkly sclerotized area (pterostigma) posteriorly enclosed by RA; concave RP diverges from RA 3.5 mm close to wing base ending with two terminal branches; M + CuA basally connected, simple MA with long connection to RP adhered shortly behind division of RA and RP; concave MP and CuP simple; convex CuA simple; convex anal vein AA ending with two posterior branches running closely parallel to posterior wing margin.
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Figs 11–14. (11) Martynovia insignis Tillyard, 1932, specimen No. 3/2014, Kukalová–Peck coll., Canadian Museum of Nature, Ottawa, ON, Canada, photograph of wing venation. (12, 13) Arroyohymen splendens gen. nov. et sp. nov., holotype No. 2927/20639, San Diego Natural History Museum coll., San Diego, CA, USA. (12) Photograph of wing venation; (13) detail photograph of wing apex with pterostig- ma. (14) Carrizopteryx cf. arroyo Kukalová-Peck & Peck, 1976, specimen No. 4/2014, Kukalová–Peck coll., Canadian Museum of Nature, Ottawa, Ontario, Canada. Photograph of fore wing venation. Scale bars: 11, 15 = 3 mm, 12= 2 mm, 13 = 1 mm. For abbreviations, see Material and methods.
Measurements Fore wing length fragment 9.8 mm, estimated wing length about 10.5 mm, wing width in the widest part 2.0 mm; hindwing length fragment 9.4 mm, estimated wing length about 10 mm, wing width in the widest part 2.2 mm.
Holotype No. 2927/20639, San Diego Natural History Museum collection (San Diego, CA, USA), imprint and counter-imprint of fore- and hindwing with venation well discern- able, the forewing posterior part is missing.
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Fig. 15. Arroyohymen splendens gen. nov. et sp. nov., holotype No. 2927/20639, San Diego Natural His- tory Museum coll., San Diego, CA, USA. Line drawing of wing venation. Scale bar = 2 mm.
Type locality Carrizo Arroyo, NM, USA.
Type strata Carboniferous/Permian transition, Red Tanks Member, Bursum Formation, insect hori- zons belong to Early Permian (Asselian) (see Lucas and Krainer, 2004, Lucas et al. 2016).
Discussion Arroyohymen gen. nov. can be attributed to family Protohymenidae due to the presence of ScP and RA running parallel to the anterior wing margin in distal wing half, with long anastomosed veins MA with RP and CuA with MP as occurred in Protohymen Tillyard, 1924 and Permohymen Tillyard, 1924 (see Carpenter 1947, fig. 12). Another significant character is the presence of unique darkly colored pterostigmal area un- known in latter two genera. Protohymenidae are known by three genera (Protohymen Tillyard, 1924, Ivahymen Martynov, 1932, Permohymen Tillyard, 1924) from Early Permian of Kansas and Oklahoma (USA) and Late Permian of Arkhangelsk Region (Russia). Arroyohymen differs mainly by the presence of long connection between MA and RP whereas it is shorter in all other genera and a specific narrow lens-like field of darkly colored pterostigmal area enclosed between ScP and RA.
Order Palaeodictyoptera Goldenberg, 1877 Family Calvertiellidae Martynov, 1931
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Genus Carrizopteryx Kukalová-Peck & Peck, 1976 Carrizopteryx cf. arroyo Kukalová-Peck & Peck, 1976 (Figs 14, 16)
Description Fore wing broadest shortly before mid-wing, with a slightly bowed anterior margin, undulated posterior margin and posteriorly pointing apex; membrane hyaline, cross veins dense and almost regular in the cubital and anal area and along the anterior margin, short irregular intercalar sectors bordering the posterior margin, the rest of wing filled with network; ScP ending on RA at mid-wing; RA + RP and veinal stems M, Cu basally closely adjacent and diverged as trifurcation just behind the separation of RA, RP sends off five main pectinate branches, first branch distally forked emerging about midwing; MA simple separates from MP 0.5 mm behind the trifurcation, MP forked three times, CuA simple and S-shaped, CuP simple, strongly curved, adjacent at length to AA1 and separated from it only close to the posterior margin, cubital field very broad with a weak irregular intercalar sector following the curvature of CuP (an autapomorphy), anal area unusually broad with eight convex anal veins connected by numerous simple straight crossveins.
Measurements Fore wing length fragment 37.8 mm, estimated fore wing length about 40 mm, wing width in the widest part 14.7 mm.
Material Specimen No. 4/2014, in collection of J. Kukalová–Peck, to be deposited in the Cana- dian Museum of Nature, Ottawa, ON, Canada; imprint and counter-imprint of the fore wing with well-preserved venation, wing base missing; membrane hyaline.
Fig. 16. Carrizopteryx cf. arroyo Kukalová-Peck & Peck, 1976, specimen No. 4/2014, Kukalová–Peck coll., Canadian Museum of Nature, Ottawa, Ontario, Canada. Line drawing of fore wing venation. Scale bar = 3 mm.
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Locality Carrizo Arroyo, NM, USA.
Age and layer Carboniferous/Permian transition, Red Tanks Member, Bursum Formation, insect ho- rizons belong to Early Permian (Asselian) (see Lucas and Krainer, 2004, Lucas et al. 2016).
Discussion The family Calvertiellidae is known from the Carboniferous and Permian deposits in the Czech Republic and France (Obora, Lodève), United States (Carrizo Arroyo, Elmo, Midco), the Russian Federation (Soyana, Tschekarda) and China (Xiaheyan) (see Fu et al. 2015). The present fore wing belongs to genus Carrizopteryx Kukalová- Peck & Peck, 1976 due to unique split of radiating veins RP, M, and CuA; RP with five main branches, MP with four branches, CuP temporary fused with AA.Carriz - opteryx is established on hindwing venation of Carrizopteryx arroyo Kukalová-Peck & Peck, 1976 (type species) known from the same locality (Kukalová-Peck & Peck 1976). Carrizopteryx comprises also C. lucasi Sinitshenkova, 2004 based on sole wing apex (possibly fore wing). However, the species is based on very fragmentary fossil lack- ing most of the diagnostic characters of the genus or even family. Sinitshenkova in Rasnitsyn et al. (2004) separated C. lucasi from C. arroyo on the basis of presence archedictyon with small cells in costal and radial areas. This character is very variable in palaeodictyopterans and corresponds to the intraspecific variability in our opinion. Moreover, the pattern in apex perfectly corresponding with our fossil (see Fig. 16). Therefore, we consider C. lucasi Sinitshenkova, 2004 as a junior synonym of Carriz- opteryx arroyo. The main difference between fore- and hindwing venation in calvertiellids occurs in form of anal area (see, e.g., Carpenter 1992, fig. 21-7ab). Therefore, we consider the present fossil clearly attributable with Carrizopteryx cf. arroyo providing additional information about fore wing venation of this taxon.
Conclusion Herein above described palaeopteran insects from Carizzo Arroyo display close relation to early Permian (Cisuralian) taxa known from Euramerica (Kansas and Oklahoma in the USA, Boskovice furrow in the Czech Republic) sharing the occurrence of the some species (e.g., Martynovia insignis) as well as close relation to taxa of Pennsylva- nian age such as Carrizoneura (Syntonopteridae) or Carrizodiaphanoptera (Diaphano- pteridae). This wide range distribution of some taxa across the transition is supported by the occurrence of neopteran Anthracoptilidae like Carrizocladus Rasnitsyn, 2004 with relatives known from Gzhelian and Cisuralian localities. Dimichelle et al. (2004)
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summarized the knowledge on flora showing similar picture by dominance of early Permian (Asselian) elements characterized by seasonal moisture limitation and rare occurrence of Gzhelian wetland elements. The suggested depositional environment can be characterized as stream banks microhabitats with limited moisture or persis- tently wet substrates on the adjacent floodplain.
Acknowledgements We are grateful to T. Demere (Natural History Museum, San Diego, CA, USA) for photographs of the specimen No. 2927/20639. Our express thanks go to A. Nel (Mu- séum national d’Histoire Naturelle, Paris) for access to the Commentry collections (Diaphanoptera munieri) and to A. Rasnitsyn (Palaeontological Institute of RAS, Mos- cow, Russia) for enabling study of the holotype Paimbia ultima. J. G. Peters (Florida A&M University, Tallahassee, FL, USA) graciously opened her unique collection of modern mayflies to JKP for five years of study. Prof. S.B. Peck (Carleton University, Ottawa, ON, Canada) reviewed the manuscript and offered many improvements. The authors are grateful to two anonymous reviewers for their comments and suggestions. JP gratefully acknowledges research support from the Grant Agency of the Czech Re- public No. 14-03847J.
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