Tarsophlebiidae

The family was established by Handlirsch (1906) on the basis of the type genus Tarsophlebia Hagen, 1866.[5]

Tarsophlebiopsis mayi was recognized as valid by most authors until the most recent revision of the family by Fleck et al. (2004),[6] who demonstrated that the holotype of this taxon most probably is only an aberrant specimen of Tarsophlebia eximia.[1][2]

The family contains 2 described genera with totally 9 valid species:[7]

• Genus †Tarsophlebia Hagen, 1866[8] (type genus; synonym: Tarsophlebiopsis Tillyard, 1923)[9]
  Diagnosis: cubito-anal areas of forewings and hindwings with four rows of cells or less; 16 or less postnodal crossveins; less than ten secondary antenodal crossveins in hindwing; and IR1 relatively short.
  • Tarsophlebia eximia (Hagen, 1862)[10] - (type species; type locality: Eichstätt, Bavaria, Germany; type horizon: Upper Jurassic, Malm zeta 2b, Lower Tithonian / 150 mya, Hybonotum Zone, Solnhofener Plattenkalk Formation; holotype: no. AS VI 44a,b at BSPG Munich Museum; size: forewing 37.1, hindwing 34.7 mm; synonyms: Tarsophlebia major Handlirsch, 1906, Tarsophlebia longissima Handlirsch, 1906, Tarsophlebiopsis mayi Tillyard, 1923[9])

Tarsophlebia minor, Upper Jurassic, Solnhofen Plattenkalk, female with ovipositor, specimen at BMMS

• • Tarsophlebia minor Fleck et al., 2004[1] - (type locality: Eichstätt, Bavaria, Germany; type horizon: Upper Jurassic, Malm zeta 2b, Lower Tithonian / 150 mya, Hybonotum Zone, Solnhofener Plattenkalk Formation; holotype: no. 55 at MCZ; size: hindwing 26 mm)
• Genus Turanophlebia Pritykina, 1968[11]
  Diagnosis: wings with more dense venation; presence of more than 25 postnodal crossveins (11-16 in Tarsophlebia); six or more rows of cells between CuA and posterior hindwing margin (less than five rows in Tarsophlebia); more than 10 secondary antenodal crossveins in hindwing (less than 10 in Tarsophlebia); IR1 longer than in Tarsophlebia; presence of long secondary longitudinal not zigzagged veins in area between IR2 and RP2.
  • Turanophlebia anglicana Fleck et al., 2004[1] - (type locality: Clockhouse (Butterley) Brickworks, England; type horizon: Lower Cretaceous, Upper Hauterivian / 130 mya, Lower Weald Clay; holotype: no. BMB 018531 at Booth Museum in Brighton / UK; size: hindwing 39.7 mm)
  • Turanophlebia liaoningensis Fang & Zheng, 2022[12] - (type locality: Huangbanjigou Village, western Liaoning, China; type horizon: Lower Cretaceous, Yixian Formation)
  • Turanophlebia martynovi Pritykina, 1968[11] - (type species; type locality: Karatau-Mikhailovka, Chimkent region, Southern Kazakhstan; type horizon: Upper Jurassic, Callovian- Kimmeridgian or Oxfordian-Kimmeridgian / 163 mya, Karabastau Formation; holotype: no. PIN 2554/210 at Palaeontological Institute Moscow; size: hindwing 41.0 mm)
  • Turanophlebia mongolica Fleck et al., 2004[1] - (type locality: Bon-Tsagaan, Bayanhongor Aimak, Central Mongolia; type horizon: Lower Cretaceous, Hauterivian-Barremian / 129 mya, Dzun-Bain Formation; holotype: no. PIN 3559/69 at Palaeontological Institute Moscow; size: hindwing 41.0 mm)
  • Turanophlebia neckini (Martynov, 1927)[13] - (type locality: Karatau-Karabastau, Chimkent region, Southern Kazakhstan; type horizon: Upper Jurassic, Callovian- Kimmeridgian or Oxfordian-Kimmeridgian / 163 mya, Karabastau Formation; holotype: no. PIN 2452/3 at Palaeontological Institute Moscow; size: hindwing 39.0 mm; originally described in the genus Tarsophlebia)
  • Turanophlebia? sibirica Pritykina, 1977[14] - (type locality: Baissa, Vitim River, Buryat Republik, Siberia / Transbaikalia, Russia; type horizon: Lower Cretaceous, Neocomian-Berriasian / 145-140 mya or Barremian-Aptian / 125 mya, Zaza Formation; holotype: no. PIN 1989/1258 at Palaeontological Institute Moscow; estimated size: hindwing 46.9, based on a comparison of the distance between wing base and end of MP, which is 24.7 mm in T. sibirica and 21.6 mm in T. martynovi that has a total hindwing length of 41.0 mm)
  • Turanophlebia sinica Huang & Nel, 2009[15] - (type locality: Western Liaoning, PR China; type horizon: Lower Cretaceous, Barremian-Aptian / 125-121 mya, Yixian Formation; holotype: no. NIGP 148201 at Nanjing Institute of Geology and Palaeontology; size: forewing 49.5 and hindwing 45.5 mm)
  • Turanophlebia vitimensis Fleck et al., 2004[1] - (type locality: right side of Vitim River downstream Romanovka village, Eravna district, Buryat Republic, Siberia / Transaikalia, Russia; type horizon: Lower Cretaceous, Neocoman / Berriasian / 145-120mya or Barremian-Aptian / 125 mya, Zaza Formation; holotype: no. PIN 2361/1 at Palaeontological Institute Moscow; estimated size: hindwing 47.0 mm)

All other described taxa are synonyms of the type species Tarsophlebia eximia.[1][2]

The monophyly of Tarsophlebiidae is strongly supported by the following set of derived characters (autapomorphies):
hindwings with hypertrophied subdiscoidal cell that is developed as "pseudo discoidal cell"; fusion of veins MAb+MP+CuA for a considerable distance before separation of MP and CuA in hindwing; vein AA strongly bent at insertion of CuP-crossing; extremely acute distal angles of forewing discoidal and subdiscoidal cell. The body characters "distinctly prolonged legs, with very long tarsi" and "male cerci with paddle-like distal expansions" are known from one species of the genus Tarsophlebia (T. eximia) and Turanophlebia (T. vitimensis) respectively, and thus belonged to the common ground plan of all Tarsophlebiidae. The extremely prolonged female ovipositor could be a further synapomorphy for the family, but it is only known from T. eximia and from a single specimen of T. minor at the Solnhofen museum.[1]

Bechly (1996) and Nel et al. (2004: Appendix 2) proposed a long list of putative synapomorphies that demonstrate a closer relationship of Tarsophlebiidae with crown group Odonata:
presence of a costal triangle as broad and strong sclerotisation of the basal costal margin; the distal discoidal vein MAb (= distal side of discoidal cell) and the subdiscoidal vein (origin of the CuA on MP) are aligned and dorsally enforced by a strong sclerotisation, so that this structure appears to cross the vein MP and the concave fold along this vein (formation of a “discal brace” sensu Carle 1982); this discal brace is aligned with the arculus in the ground plan (only retained in some taxa with an open discoidal cell and in the forewings of Epiophlebia); the midfork (first fork of RP and base of IR2) is shifted basally, with the RP3/4 generally arising basal of the subnodus (reversed in some Coenagrionoidea) and RP2 arising close to the subnodus (in the ground plan); more derived type of nodus, with a kink in ScP; the oblique basal brace (still present in Protanisoptera and Protozygoptera) is transformed into a transverse “basal bracket” Ax0 which looks like a primary antenodal crossvein; presence of two strong primary antenodal crossveins Ax1 and Ax2 (also present in some Protozygoptera, e.g. specimen PIN 1/276 from the Upper Permian of Russia); pterostigma distinctly braced by an oblique postsubnodal crossvein beneath the basal margin of the pterostigma; presence of a tracheated lestine oblique vein 'O' between RP2 and IR2 (secondarily absent in some taxa); in the median space (= basal space) the convex vestige of the Media-stem (“vestigial CuA” sensu Fraser 1957) is suppressed since it is fused with the cubital stem to a common medio-cubital-stem (the alleged presence of this vestige in the holotype of Tarsophlebiopsis is a teratology or an artefact of preservation), convergent to some Protanisoptera, Triadophlebiomorpha, and Protozygoptera.

Bechly (1996) proposed that several unique symplesiomorphic features of all Tarsophlebiidae indicate that this family represents the sister group of all Recent Odonata.[4] These features are the basally open discoidal cell in the hindwing (instead of closed) which implies an incomplete arculus, the (meanwhile disputed) presence of four tarsomeres of equal length (instead of only three), and the very primitive condition of the male secondary genital apparatus (viz ligula orimentary; vesicula spermalis still very short and flat with a very wide porus) without any intromittent organ. Bechly therefore considered the similarities of Tarsophlebiidae and Epiprocta mentioned by Nel et al. (1993), viz the less separated and relatively large eyes, the presence of two cephalic sutures, and the small leg spines (also present in Meganisoptera[3]), as symplesiomorphies.

However, based on a cladistic study of 14 characters Fleck et al. (2004) again suggested that Tarsophlebiidae might rather be the sister group of the clade Epiprocta that includes Epiophlebiidae and Anisoptera. Nevertheless, this result has a very low statistical support and might as well be an artefact of the parsimony computer algorithm, because none of the 14 characters represents an unambiguous synapomorphy for Tarsophlebiidae and Epiprocta.[1]

Huang & Nel (2009) presented convincing evidence from a new fossil Tarsophlebiidae from China that the number of tarsomeres is only three as in modern odonates, but that the first tarsomere is about twice as long as the others.[15] Bechly (1996) already discussed this possibility and suggested that such a long basal tarsomere, compared to the short basal tarsomere in protodonates and all Recent odonates, could still be a plesiomorphic state that could have resulted by the fusion of two basal tarsomeres.[4] Huang & Nel (2004) state that there is no evidence for such a fusion in the fossils and that the elongation of the first segment could as well be a derived feature (autapomorphy) of Tarsophlebiidae.

The two alternative phylogenetic positions of Tarsophlebiidae

Hypothesis of Fleck et al. (2004):

Hypothesis of Bechly (1996, 2007):

Next to nothing is known about the ecology and behavior of Tarsophlebiidae, but it probably was quite similar to Recent odonates. With their very long ovipositor the females probably inserted their eggs into mud in or close to water, similar to modern Cordulegastridae. It is remarkable that two other groups of Mesozoic odonates (viz Steleopteridae and Aeschnidiidae) had a similarly elongated ovipositor.

The fossil record of this family ranges from Upper Jurassic of Kazakhstan and Germany to the Lower Cretaceous of England, Transbaikalia, Mongolia and China. There are no records of Tarsophlebiidae from any fossil locality outside the Eurasiatic region.[7]

The first tarsophlebiid fossils were specimens of Tarsophlebia eximia from the Upper Jurassic Solnhofen Plattenkalk of Germany described as "Heterophlebia eximia" and Euphaea longiventris by Hagen (1862).[1] Larger revisions of this family have been provided by Nel et al. (1993) and Fleck et al. (2004).[1][2]

• Carpenter FM (1992). Superclass Hexapoda. Volume 3 of Part R, Arthropoda 4; Treatise on Invertebrate Paleontology. Geological Society of America & The University of Kansas. ISBN 978-0-8137-3019-6.
• Fraser FC (1957). "A reclassification of the order Odonata, based on some new interpretations of the venation of the dragonfly wing". Handbook of the Royal Zoological Society of New South Wales. 12: 1–133.
• Grimaldi D, Engel MS (2005). Evolution of the Insects. Cambridge University Press. ISBN 978-0-521-82149-0.
• Rasnitsyn, AP, Quicke, DLJ (2002). History of Insects. Kluwer Academic Publishers. ISBN 978-1-4020-0026-3.{{cite book}}: CS1 maint: multiple names: authors list (link)

Wikimedia Commons has media related to Tarsophlebiidae.

• Die Fossilien von Solnhofen (photos of Solnhofen fossils of Tarsophlebia)
• Tree of Life Project (Odonata page)
• Hyperbolic phylogenetic tree of Odonata (by G. Bechly)
• Phylogenetic Systematics of Odonata - Tarsophlebiidae (by G. Bechly, 2007)