Eur. J. Entomol. 106 (1): 145-152, 2009 | DOI: 10.14411/eje.2009.017

Tackling the "Protoblattoidea problem": Revision of Protoblattinopsis stubblefieldi (Dictyoptera; Late Carboniferous)

Olivier BÉTHOUX1,2, Klaus-Dieter KLASS2, Jörg W. SCHNEIDER1
1 Freiberg University of Mining and Technology, Institute of Geology, Department of Palaeontology, Bernhard-von-Cotta Str. 2, D-09596 Freiberg, Germany; e-mail: obethoux@yahoo.fr; joerg.schneider@geo.tu-freiberg.de
2 Senckenberg Natural History Collections of Dresden, Museum of Zoology, KGnigsbrücker Landstraße 159, D-01109 Dresden, Germany; e-mail: klaus.klass@senckenberg.de

The forewing morphology of the fossil species †Protoblattinopsis stubblefieldi Laurentiaux, 1953 (Dictyoptera; Late Carboniferous) is re-investigated. Unlike previously stated, in this species, the stems of R and M are distinct and CuA has branches. The occurrence of the putative plesiomorphies "long ScP" and "differentiated R sectors" (i.e., RA and RP) in †P. stubblefieldi indicates a position basal to all extant Dictyoptera and many members of its paraphyletic stem. Unlike representatives of the families †Archimylacridae and †Blattinopsidae, but similar to Blattodea s. str., †P. stubblefieldi lacks a M-CuA arculus. The comparison of the branching pattern of the median system in †P. stubblefieldi and †Archimylacridae suggests that the arculus of the latter is composed of posterior branches of M. It is noted that the distal location of the point of origin of CuA and CuP, the sigmoid course of the anterior stem of CuA, the strong curvature of CuP, and the occurrence of a short fusion between R/RP and the anterior branch of M are similarities with the representatives of the genus Polyphaga Brullé, 1835, belonging to Dictyoptera s. str. However, these similar character states are unlikely to be homologous in these taxa, mainly because the hypothesis that Polyphaga is the sister group of the remaining Dictyoptera s. str. (or even Blattodea s. str.) is not strongly supported. Owing to its character state combination, the species †P. stubblefieldi is unique within the "Protoblattoidea waste-basket". Comprehensive studies of wing morphology in stem-Dictyoptera are needed to clarify whether it might be closely related to the lineage from which Dictyoptera s. str. evolved.

Keywords: Dictyoptera, fossil, forewing venation, Late Carboniferous, Protoblattinopsis, Polyphaga, arculus, phylogeny

Received: August 14, 2008; Revised: January 12, 2009; Accepted: January 12, 2009; Published: March 10, 2009  Show citation

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BÉTHOUX, O., KLASS, K., & SCHNEIDER, J.W. (2009). Tackling the "Protoblattoidea problem": Revision of Protoblattinopsis stubblefieldi (Dictyoptera; Late Carboniferous). EJE106(1), 145-152. doi: 10.14411/eje.2009.017
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References

  1. BETHOUX O. 2005: Wing venation pattern of Plecoptera (Neoptera). Illiesia 1: 52-81
  2. BETHOUX O. 2007: Emptying the Paleozoic wastebasket for insects: members of a Carboniferous "protorthopterous family" reassigned to natural groups. Alavesia 1: 41-48
  3. BETHOUX O. 2008: Groundplan, nomenclature, homology, phylogeny, and the question of the insect wing venation pattern. Alavesia 2: 219-232
  4. BETHOUX O. & BRIGGS D.E.G. 2008: How Gerarus lost its head: stem-group Orthoptera and Paraneoptera revisited. Syst. Entomol. 33: 529-547 Go to original source...
  5. BETHOUX O. & NEL A. 2001: Venation pattern of Orthoptera. J. Orthopt. Res. 10: 195-198 Go to original source...
  6. BETHOUX O. & NEL A. 2002: Venation pattern and revision of Orthoptera sensu nov. and sister groups. Phylogeny of Palaeozoic and Mesozoic Orthoptera sensu nov. Zootaxa 96: 1-88 Go to original source...
  7. BETHOUX O. & NEL A. 2004: Some Palaeozoic "Protorthoptera" are "ancestral" orthopteroids: major wing braces as clues to a new split among the "Protorthoptera". J. Syst. Palaeontol. 2: 285-309 Go to original source...
  8. BETHOUX O. & WIELAND F. in press: Evidence for Carboniferous origin of the order Mantodea (Insecta: Dictyoptera) gained from forewing morphology. Zool. J. Linn. Soc
  9. BOLTON H. 1922: A monograph of the fossil insects of the British Coal Measures. Part II. Palaeontol. Soc. 74: 81-156 Go to original source...
  10. BOLTON H. 1925: Insects from the Coal Measures of Commentry. British Museum (Natural History), London, 56 pp
  11. BRULLE G.A. 1835: Orthopteres et Hemipteres. In Audouin J.V. & Brulle G.A. (eds): Histoire Naturelle des Insectes Traitant de leur Organisation et de leurs Moeurs en General. Vol. 9. F.D. Pillot, Paris, pp. 1-230
  12. GRIMALDI D. 2003: A revision of Cretaceous mantises and their relationships, including new taxa (Insecta: Dictyoptera: Mantodea). Am. Mus. Nov. 3412: 1-47 Go to original source...
  13. GRIMALDI D. & ENGEL M.S. 2005: Evolution of the Insects. Cambridge University Press, New York, 755 pp
  14. HANDLIRSCH A. 1906: Die fossilen Insekten und die Phylogenie der rezenten Formen. Ein Handbuch fuer Palaeontologen und Zoologen. Wilhelm Engelmann, Berlin, 1430 pp
  15. HANDLIRSCH A. 1919: Revision der palaeozoischen Insekten. Denkschr. Kais. Akad. Wiss., Math.-Naturwiss. Kl. 96: 511-592
  16. HANDLIRSCH A. 1937: Neue Untersuchungen ueber die fossilen Insekten mit Ergaenzungen und Nachtraegen sowie Ausblicken auf phylogenetische, palaeogeographische und allgemein biologische Probleme. I Teil. Ann. Naturhist. Mus. Wien 48: 1-140
  17. HENNIG W. 1969: Die Stammesgeschichte der Insekten. Kramer, Frankfurt am Main, 436 pp
  18. HENNIG W. 1981: Insect Phylogeny. Wiley & Sons, New York, xxii + 514 pp
  19. HORNSCHEMEYER T. & STAPF H. 2001: Review of Blattinopsidae (Protorthoptera) with description of new species from the Lower Permien of Niedermoschel (Germany). N. Jb. Geol. Palaeontol. Abh. 221: 81-109 Go to original source...
  20. INWARD D., BECCALONI G. & EGGLETON P. 2007: Death of an order: a comprehensive phylogenetic study confirms that termites are eusocial cockroaches. Biol. Lett. 3: 331-335 Go to original source...
  21. KLASS K.-D. & MEIER R. 2006: A phylogenetic analysis of Dictyoptera (Insecta) based on morphological characters. Entomol. Abh. 63: 3-50.
  22. KUKALOVA J. 1959: Remarks to the systematic position of the family Blattinopsidae Bolton, 1925. Rozpr. Csl. Akad. Ved, Rada MPV 69: 1-27
  23. KUKALOVA-PECK J. 1991: Fossil history and the evolution of hexapod structures. In Naumann I.D., Crane P.B., Lawrence J.F., Nielsen E.S., Spradbery J.P., Taylor R.W., Whitten M.J. & Littlejohn M.J. (eds): The Insects of Australia. A Textbook for Students and Researchers. Melbourne University Press, Melbourne, pp. 141-179
  24. LAMEERE A. 1922: Sur la nervation alaire des insectes. Bull. Cl. Sci. Acad. R. Belg. 8: 138-149
  25. LAMEERE A. 1923: On the wing-venation of insects. Psyche 30: 123-132 Go to original source...
  26. LAURENTIAUX D. 1953: Classe des Insectes (Insecta Linne, 1758). In Piveteau J. (ed.): Traite de Paleontologie. Masson et Cie, Paris, pp. 397-527
  27. LAURENTIAUX D. 1966: Sur un insecte Westphalien precurseur de la serie Blattinopsoide (Protorthopteres). Bull. Soc. Geol. Fr. 8: 827-833 Go to original source...
  28. LAURENTIAUX-VIEIRA F. & LAURENTIAUX D. 1980: Nouvelle contribution L la connaissance du genre Westphalien Manoblatta Pruvost (Blattaires, Archymylacridiens). Ann. Soc. Geol. Nord 99: 415-423
  29. LO N., BENINATI T., STONE F., WALKER J. & SACCHI L. 2007: Cockroaches that lack Blattabacterium endosymbionts: the phylogenetically divergent genus Nocticola. Biol. Lett. 3: 327-330 Go to original source...
  30. PELLENS R., D'HAESE C.A., BELLES X., PIULACHS M.D., LEGENDRE F., WHEELER W.C. & GRANDCOLAS P. 2007: The evolutionary transition from subsocial to eusocial behaviour in Dictyoptera: Phylogenetic evidence for modification of the "shift-in-dependent-care" hypothesis with a new subsocial cockroach. Mol. Phylogenet. Evol. 43: 616-626 Go to original source...
  31. PROKOP J. & NEL A. 2007: An enigmatic Palaeozoic stem-group: Paoliida, designation of new taxa from the Upper Carboniferous of the Czech Republic (Insecta: Paoliidae, Katerinkidae fam. n.). Afr. Invertebr. 48: 77-86
  32. RASNITSYN A.P. 2002: 2.2.2. Infraclass Gryllones Laicharting, 1781. The grylloneans (= Polyneoptera Martynov, 1938). In Rasnitsyn A.P. & Quicke D.L.J. (eds): History of Insects. Kluwer Academic Publishers, Dordrecht, pp. 254-262
  33. RASNITSYN A.P. 2007: On the discussion of the wing venation of (Archae) Orthoptera (Insecta). Palaeontol. J. 41: 341-344 Go to original source...
  34. REHN J.W.H. 1951: Classification of the Blattaria as indicated by their wings (Orthoptera). Mem. Am. Entomol. Soc. 14: 1-134
  35. SCHNEIDER J.W. 1984: Die Blattodea (Insecta) des Palaeozoikums. Teil II: Morphogenese der Fluegelstrukturen und Phylogenie. Freib. Forsch. (C) 391: 5-34
  36. SMART J. 1956: On the wing venation of Chaeteessa and other mantids (Insecta: Mantodea). Proc. Zool. Soc. Lond. 127: 545-553 Go to original source...
  37. STOROZHENKO S.Y. 1998: Systematics, Phylogeny and Evolution of Grylloblattid Insects (Insecta: Grylloblattida). Dal'nauka, Vladivostok, 207 pp. (in Russian)
  38. STOROZHENKO S.Y. 2002: 2.2.2.2.1. Order Grylloblattida Walker, 1914 (= Notoptera, Crampton, 1915, = Grylloblattodea Brues et Melander, 1932, + Protorthoptera Handlirsch, 1906, = Paraplecoptera Martynov, 1925, + Protoperlaria Tillyard, 1928). In Rasnitsyn A.P. & Quicke D.L.J. (eds): History of Insects. Kluwer Academic Publishers, Dordrecht, pp. 278-281
  39. VRSANSKY P., VISHNIAKOVA V.N. & RASNITSYN A.P. 2002: 2.2.2.1.1. Order Blattida Latreille, 1810. The cockroaches (= Blattodea Brunner von Wattenvill, 1882). In Rasnitsyn A.P. & Quicke D.L.J. (eds): History of Insects. Kluwer Academic Publishers, Dordrecht, pp. 263-270
  40. WALKER F. 1868: Catalogue of the Specimens of Blattariae in the Collection of the British Museum. Trustees of the British Museum, London, 239 pp Go to original source...
  41. WARE J.L., LITMAN J., KLASS K.-D. & SPEARMAN L.A. 2008: Relationships among the major lineages of Dictyoptera: the effect of outgroup selection on dictyopteran tree topology. Syst. Entomol. 33: 429-450 Go to original source...

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