ISSN 00310301, Paleontological Journal, 2013, Vol. 47, No. 2, pp. 162–165. © Pleiades Publishing, Ltd., 2013. Original Russian Text © N.D. Sinitshenkova, 2013, published in Paleontologicheskii Zhurnal, 2013, No. 2, pp. 35–38.
New Mayflies (Insecta: Ephemerida = Ephemeroptera) from the Upper Permian Locality of Isady, Northern European Russia N. D. Sinitshenkova Borissiak Paleontological Institute, Russian Academy of Sciences, Profsoyuznaya ul. 123, Moscow, 117997 Russia email:
[email protected] Received April 12, 2012
Abstract—Two new species of the family Protereismatidae, Alexandrinia vitta sp. nov. and A. ipsa sp. nov., are described from the Upper Permian locality of Isady (Severodvinian Stage, Poldarsa Formation). The third new species Misthodotes dubius sp. nov., belongs to the family Misthodotidae. Keywords: Perlida, Protereismatidae, Alexandrinia, Misthodotidae, Misthodotes, Upper Permian, Isady, North of European Russia DOI: 10.1134/S0031030113020135
INTRODUCTION The mayflies described below were collected in the rich Upper Permian locality of Isady (= Mutovino), situated on the left bank of the Sukhona River, 1.8 km downstream of the Mutovino landscape unit, opposite the village of Isady (alternative English spelling: Issady), Velikoustyugskii District, Vologda Region. The insectbearing deposits belong to the Severodvin ian Stage, Kichuga Member, Pondarsa Formation, dated to the Tatarian Permian. Mayflies of only two families occur in this locality: Protereismatidae Sellards, 1907 and Misthodotidae Tillyard, 1932, characterized by almost homonomous wings. One species of the former family, Alexandrinia gigantea Sinitshenkova et Vassilenko, 2012, was described from Isady earlier, as well as one species of the latter family, Misthodotes tshernovae Sinitshenkova et Vassilenko, 2012 (Sinitshenkova and Vassilenko, 2012). Specimens of three new species have been found among the new materials: Alexandrinia vitta sp. nov. and A. ipsa sp. nov. (Protereismatidae), and Mis thodotes dubius sp. nov. (Misthodotidae). It is from Isady that the latest records of protereismatid mayflies are known; moreover, they include both the largest (A. gigantea) and the smallest known members of this family. Both new species of Alexandrinia Sinitshenk ova et Vassilenko, 2012 are considerably smaller than all hitherto known species of the genus; the wing vena tion of A. vitta is similar to that of A. gigantea, and the wing venation of A. ipsa is similar to that of the North American species A. directa (Carpenter, 1979), origi nally described in the genus Protereisma Sellards, 1907 as P. directum (Carpenter, 1979). In both new species of Alexandrinia, the closely set bases of the pedicels of
R, RS, and MA have been found, which is atypical of the hitherto known species. The possibility cannot be excluded that this trait is a result of the diminution of the wing: the wings of both species are 1.5–2 times shorter than those of North American members of the genus and 2–3 times shorter than in A. gigantea. The species diversity of mayflies in Isady is remark able: among the 12 known fossils, five species have been identified to date. This number of species is com parable to that found in the Early Permian deposits of North America, where the two families, Protereisma tidae and Misthodotidae, are also found together, and the number of known mayfly fossils is over one hun dred (Carpenter, 1933, 1979). Examination of the specimens under a TESCAN VEGA XMU scanning electron microscope revealed the presence of strong small spines on the anterior margins of the wings of A. vitta sp. nov. and A. ipsa sp. nov. Similar microsculpture along the wing margin was described in the wings of P. latum Sellards, 1907 and A. directa (Carpenter, 1979), on the veins of which one can also discern setae, which cannot be found on the wings of the mayflies from Isady, possibly because of their poor preservation quality. The family Misthodotidae is known from the Early Permian to Middle Triassic. The Permian species from North America (five species), Western Europe (one species), and Perm Region, Russia (two species) belong to the genus Misthodotes (Chernova, 1965; Kinzelbach and Lutz, 1984); from the Triassic of France, the genus Triassodotes Sinitshenkova, Mar chalPapier, GrauvogelStamm et Gall, 2005 and one species of this genus were described (Sinitshenkova et al., 2005). The new species M. dubius sp. nov., found
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in Isady, is similar to another species described from the same locality, M. tshernovae. SYSTEMATIC PALEONTOLOGY Order Ephemerida (=Ephemeroptera) Suborder Protereismatina Family Protereismatidae Sellards, 1907 Genus Alexandrinia Sinitshenkova et Vassilenko, 2012 Alexandrinia vitta Sinitshenkova, sp. nov. Plate 7, figs. 1–4
(a)
E t y m o l o g y. From the Latin vitta (ribbon). H o l o t y p e. PIN, no. 3840/1707, positive and negative impressions of basal half of forewing, well preserved; Isady locality; Upper Permian, Severodvin ian Stage, Poldarsa Formation. D e s c r i p t i o n (Fig. 1a). In the forewing, strong small spines are visible along the anterior margin; the costal and subcostal areas are almost equal in width. The precostal vein is clearly pronounced, almost straight, runs along the middle of the costal area, and joins C somewhat proximal to its fusion with R. The bases of the stems R, RS, and MA are set closely at the base of the wing, so that they seem fused over a consid erable distance; MA diverges somewhat basal to the bifurcation of MP; RS runs on very closely to R. MP divides considerably basal to the short fork of CuA. CuP and A1 are smoothly Sshaped; the gemination of these veins is visible near the wing margin. At least 13 simple straight parallel crossveins diverge from A1 towards the wing margin; these crossveins are twice as long as the distances between them. M e a s u r e m e n t s, mm. Length of forewing frag ment, 9.7; total forewing length, about 17. C o m p a r i s o n. The structure and number of the crossveins diverging from A1 towards the wing margin make the new species similar to A. gigantea, from which it is clearly distinguished by the less than half as large size and the fusion of the bases of the stems R, RS, and M in the basal part of the wing. M a t e r i a l. Holotype. Alexandrinia ipsa Sinitshenkova, sp. nov. Plate 7, figs. 5–6
E t y m o l o g y. From the Latin ipsa (herself). H o l o t y p e. PIN, no. 3840/1708, positive and negative impressions of basal half of forewing, well preserved; Isady locality; Upper Permian, Severodvin ian Stage, Poldarsa Formation. D e s c r i p t i o n (Fig. 1b). In the forewing, strong small spines are visible along the anterior margin; the costal area basally is wider than the subcostal area more distally they are almost equal in width. The pre costal vein is clearly pronounced, slightly curved, located considerably closer to the costal one than to the subcostal one, and joins C somewhat proximal of its fusion with SC. The bases of the stems R, RS, and MA are set closely at the base of the wing; the bases of PALEONTOLOGICAL JOURNAL
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Fig. 1. Mayflies of the genus Alexandrinia: (a) A. vitta sp. nov., holotype PIN, no. 3840/1707, basal portion of forewing; (b) A. ipsa sp. nov., holotype PIN, no. 3840/1708, basal portion of forewing; Isady locality; Upper Permian. Scale bar, 2 mm.
RS and MA are fused and visibly diverge only in the distal quarter of the wing. CuP and A1 are smoothly curved. At least 10 simple straight parallel crossveins diverge from A1 towards the wing margin; these cross veins are 5–6 times as long as the distances between them. The base of A1 is connected with A2 by cross veins, the distances between which are almost as long as these crossveins. At least five almost straight, non branching crossveins diverge from A2 towards the wing margin; these crossveins are more than three times as long as the distances between them. M e a s u r e m e n t s, mm. Length of wing frag ment, 9.0; total wing length, about 12. C o m p a r i s o n. The structure of the anal area, with crossveins diverging from A1 and A2 towards the wing margin, makes the new species similar to A. directa, from which it is clearly distinguished by the considerably smaller size (the wing length of A. directa is 26 mm) and simple, nonbranching crossveins diverging from the anal veins (in A. directa, some of
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SINITSHENKOVA Plate 7
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Explanation of Plate 7 Figs. 1–4. Alexandrinia vitta sp. nov., holotype PIN, no. 3840/1707: (1) basal portion of forewing; (2) base of forewing; (3) frag ment of venation showing closely set stems of R, RS, and MA; (4) strong small spines along anterior wing margin. Figs. 5–6. Alexandrinia ipsa sp. nov., holotype PIN, no. 3840/1708: (5) basal portion of forewing; (6) base of forewing. Fig. 7. Misthodotes dubius sp. nov., holotype PIN, no. 3840/1706, forewing; Isady locality; Upper Permian.
Fig. 2. Misthodotes dubius sp. nov., holotype PIN, no. 3840/1706, forewing; Isady locality; Upper Permian. Scale bar, 2 mm.
such crossveins are branching; Carpenter, 1979, p. 263, textfig. 1). M a t e r i a l. Holotype. Family Misthodotidae Tillyard, 1932 Genus Misthodotes Tillyard, 1932 Misthodotes dubius Sinitshenkova, sp. nov. Plate 7, fig. 7
E t y m o l o g y. From the Latin dubius (doubtful). H o l o t y p e. PIN, no. 3840/1706, positive and negative impressions of almost complete forewing, well preserved; Isady locality; Upper Permian, Severodvinian Stage, Poldarsa Formation. D e s c r i p t i o n (Fig. 2). The forewing is more than three times as long as wide. The costal and sub costal areas in the basal half of the wing are almost equal in width; RS divides for the first time somewhat basal to the middle of the wing and reaches the wing margin in five branches. The forks of MA and MP are wide; MA divides visibly distal to the fork of MP, which, in its turn, divides approximately in the middle of its length. An oblique crossvein diverges from the anterior branch of MA and from the intercalary MP and reaches the wing margin. CuA, CuP, and A1 are simple; crossveins are rather scant. M e a s u r e m e n t s, mm. Forewing length, 6.0. C o m p a r i s o n. The structure of the media makes the new species similar to M. tshernovae, from which it is clearly distinguished by the narrower wing; the smaller number of veins in the RS system distin guish it from all known species of the genus, in which RS reaches the wing margin in seven branches. M a t e r i a l. Holotype. PALEONTOLOGICAL JOURNAL
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ACKNOWLEDGMENTS The author is grateful to R. Rakitov (PIN), who studied the specimens under a TESCAN VEGA XMU scanning electron microscope and found microsculp ture on the wings of the mayflies. This study was supported by the Russian Founda tion for Basic Research, project no. 100401713, and by the Program of the Presidium of the Russian Acad emy of Sciences “Origin of the Biosphere and Evolu tion of Geobiological Systems.” REFERENCES Carpenter, F.M., The Lower Permian Insects of Kansas: Part 6. Delopteridae, Protelytroptera, Plectoptera and a New Collection of Protodonata, Odonata, Megasecoptera, Homoptera, and Psocoptera, Proc. Am. Acad. Arts and Sci., 1933, vol. 68, no. 11, pp. 411–503. Carpenter, F.M., Lower Permian Insects from Oklahoma: Part 2. Orders Ephemeroptera and Palaeodictyoptera, Psyche, 1979, vol. 86, nos. 2–3, pp. 261–280. Chernova, O.A., On Some Fossil Mayflies (Ephemeroptera, Misthodotidae) from the Permian Sedi ments of the Urals, Entomol. Obozr., 1965, vol. 44, no. 2, pp. 253–361. Kinzelbach, R. and Lutz, H., Eine neue Eintagsfliege Mis thodotes stapfi n. sp. aus dem Rotliegenden des NaheGebietes, Paläontol. Z., 1984, vol. 58, nos. 3/4, pp. 247–253. Sinitshenkova, N.D., MarchalPapier, F., GrauvogelStamm, L., and Gall, J.C., The Ephemeroptera (Insecta) from the Grès à Voltzia (Early Middle Triassic) of the Vosges (NE France), Paläontol. Z., 2005, vol. 79, no. 3, pp. 377–397. Sinitshenkova, N.D. and Vassilenko, D.V., The Latest Record of Mayflies of the Family Protereismatidae Sellards (Ephemerida = Ephemeroptera) and a New Species of the Family Misthodotidae in the Upper Permian of Europe, Paleontol. J., 2012, vol. 46, no. 1, pp. 61–65.
