NOMINA CIRCUMSCRIBENTIA INSECTORUM
Holopandictyoptera Kluge 2010
NOMEN: Holopandictyoptera Kluge 2010 [N.J. Kluge. Circumscriptional names of higher taxa in Hexapoda. – Bionomina, 2010, No.1: 15–55]. PDF
ORIGINAL LISTED MEMBERSHIP (Kluge 2010): Cryptovipositoria + Palaeoblattariae
NON-MONOSEMANTIC CIRCUMSCRIPTIONAL SYNONYMS (with uncertain position of †Palaeoblattariae):
= Panisoptera Crampton 1919 s.l.
= Blattopteroidea Martynov 1924 s.l.
= Isopteria Crampton 1938 s.l.
= Dictuopteroida Kevan 1976 s.l.
TYPIFIED NAMES IN USE:
Classification of Holopandictyoptera:
The taxon consisting of praying mantids, cockroaches, and termites, is often called “Dictyoptera” or “Dictuoptera”, with “Leach 1815” indicated as the author of both. In fact Leach (1815) used the name Dictuoptera for an order consisting of a single genus, Blatta (i.e. what the current classification has as Neoblattariae), placing the praying mantids with the Orthoptera, and termites with the Neuroptera.
The name Dictyoptera (verbatim, ‘lacewings’) was independently given to various groups of insects: Dictyoptera Clairville 1798 is a junior circumscriptional synonym of Neuroptera Linnaeus 1758. Brullé (1832) established the new order Dictyoptera to accommodate amphibiotic insects with incomplete metamorphosis (Odonata, Ephemeroptera, and Plecoptera), formerly placed in the order Neuroptera; junior circumscriptional synonyms of Dictyoptera Brullé 1832 are Arkiptera Laporte 1834 and Amphibiotica Gerstaecker 1863. The preoccupied name Dictyoptera Dohrn 1866, suggested for a peculiar group of Paleozoic insects, was replaced with its objective synonym, Palaeodictyoptera Goldenberg 1877. Besides, Enderlein (1904) applied the name “Dictyoptera” to the taxon consisting of cockroaches and praying mantids, and Crampton (1915) — to the taxon containing praying mantids only.
The name Pandictyoptera Crampton 1917 is the oldest name whose original circumscription fits the taxon consisting of praying mantids, cockroaches, and termites, and should be used as its valid name. Names Panisoptera Crampton 1919, Blattopteroidea Martynov 1924, and Isopteria Crampton 1938 are its junior circumscriptional synonyms, each can be also used where priority is not an issue.
Based on extant taxa alone, the name Pandictyoptera and its circumscriptional synonyms Panisoptera, Blattopteroidea and Isopteria in their original circumscriptions fit the taxon consisting of praying mantids, cockroaches and termites. However, the Paleozoic and Mesozoic Palaeoblattariae Scudder 1879, undoubtedly related to the true cockroaches, do not fit the taxon Pandictyoptera as defined based solely on the extant fauna. Palaeoblattarians, together with true cockroaches, termites, and praying mantids, form a holophyletic taxon, Pandictyoptera s.1., with Pandictyoptera s.str. as a subordinated taxon. As long as the extant fauna is concerned, or the context is other than taxonomic or morphological, the difference between Pandictyoptera s.l. and Pandictyoptera s.str. can be ignored. In some situations, though, this difference should be stressed: for example, the statement “all the females have an outer genital chamber” would be true for Pandictyoptera s.str., but not for Pandictyoptera s.l. To avoid confusion, I propose two new names: Holopandictyoptera for Pandictyoptera s.l. (including Palaeoblattariae with their long ovipositor and no outer genital chamber) and Cryptovipositoria for Pandictyoptera s.str., with ovipositor reduced and hidden (completely or partly) in the outer genital chamber formed by the enlarged abdominal sternum 7. Thus, the name Dictyoptera and its monosemantic circumscriptional synonyms Panisoptera, Blattopteroidea, and Isopteria are non-monosemantic circumscriptional synonyms of Holopandictyoptera and Cryptovipositoria. Holopandictyoptera and Cryptovipositoria, on the other hand, are not circumscriptional synonyms and belong to different taxa subordinated one to another within the following classification of Pandictyoptera:
1. Holopandictyoptera = Pandictyoptera s.l. (typified name: Blatta/fg)
1.1. Plesiomorphon Palaeoblattariae (typified name: Mylacris/fg)
1.2.1. Raptoriae (typified name: Mantis/fg)
1.2.2. Oothecophora (typified name: Blatta/fg)
18.104.22.168. Plesiomorphon Neoblattariae (typified name: Blatta/fg)
22.214.171.124. Isoptera (typified name: Termes/fg)
The new name Oothecophora Kluge 2010 is proposed for the taxon Neoblattariae + Isoptera, separated from Raptoriae based on the following autapomorphic characters: (1) Median ocellus absent. (2) Ovipositor strongly reduced and completely hidden in the outer genital chamber, where the ootheca is formed (Isoptera and some Neoblattariae do not form oothecae); eggs are packed in two rows (this is true for Neoblattariae and Isoptera-Hemiclidoptera, while in Isoptera-Crypticlidoptera eggs are not packed). (3) Six abdominal ganglia (vs. seven in Raptoriae).
The Isoptera are generally accepted to have originated from within the Neoblattariae. However, no circumscriptional name has been proposed yet for the holophyletic taxon Neoblattariae + Isoptera. Hennig (1969, 1981) applied to it the typified name Blattodea, traditionally used for cockroaches alone.
The plesiomorphon Palaeoblattariae Scudder 1879 includes extinct taxa only whose long, functional ovipositor separate them from the Oothecophora (and Cryptovipositoria in general). Originally, Scudder (1879) included in the Palaeoblattariae the extinct forms placed in genera Mylacris Scudder 1868, Archimylacris Scudder 1868, and the newly described genera Lithomylacris, Necymylacris, Etoblattina, Anthracoblattina, Gerablattina, Hermatoblattina, Progonoblattina, Oryctoblattina, and Petrablattina. The Neoblattariae Scudder 1895 originally encompassed all Recent cockroaches plus the extinct Mesozoic genera Neorthroblattina Scudder 1885 and Scutinoblattina Scudder 1885, and originally excluded, besides the palaeoblattarian taxa listed above, the extinct forms arranged in genera Paromylacris Scudder 1885, Microblattina Scudder 1895, Spiloblattina Scudder 1884, Poroblattina Scudder 1885, and Leptoblattina Woodward 1887, all placed by Scudder (1895) with the Palaeoblattariae. All these extinct forms were originally described based on wing venation only; palaeoblattarian specimens with ovipositors were discovered later, but for most species Scudder originally included in the Palaeoblattariae there is still no evidence of ovipositor or lack thereof.
The name Palaeoblattida Brongniart 1893 was introduced as emendation of Palaeoblattariae Scudder 1879 and originally applied to a family consisting of the same set of taxa as Scudder’s Palaeoblattariae, to which Brongniart added a number of new species. He also provided drawings of the Anthracoblattina ensifer and Etoblattina sp. females with long ovipositors (Brongniart 1893: Pl.XLIII (32): Fig. 2–4).
To avoid creating new names, I suggest to accept the original circumscriptions of both Palaeoblattariae Scudder 1897 and Palaeoblattida Brongniart 1893 as fitting the plesiomorphon excluding Cryptovipositoria, and the original circumscription of Neoblattariae Scudder 1895 as fitting a taxon within Cryptovipositoria-Oothecophora, until proven otherwise.
Brongniart C. 1893. Recherches pour servir à l’histoire des insectes fossiles des temps primaires précédées d’une étude sur la nervation des ailes des insectes. Saint-Etienne: 1–493.
Brullé M. 1832. Des animaux articulés. In: Expédition scientifique de Morée, Section des Sciences physiques, 3 (1), Zoologie, Section 2. Paris: 1–400.
Clairville J. de 1798. Entomologie helvétique ou catalogue des insectes de la Suisse rangés d’après une nouvelle méthode. Helvetische Entomologie oder Verzeichniss der Schweizerischen insecten nach einer neuen Methode Geordnet. Tome 1. Zurich (Orell, Fussli & Co.): 1–149.
Crampton G. C. 1915. The thoracic sclerites and the systematic position of Grylloblatta campodieformis Walker, a remarkable annectant, “Orthopteroid” insect. // Entomological News, 26 (10): 337–350.
Crampton G. C. 1917. A phylogenetic study of lateral head, neck and prothoracic regions in some Apterygota and lower Pterygota. // Entomological News, 28 (9): 398–412.
Crampton G.C. 1919. Notes on phylogeny of the Orthoptera. // Entomological News, 30 (2): 42–48; (3): 64–72.
Crampton G. C. 1938. The interrelationships and lines of descent of living insects. // Psyche, 45 (4): 165–181.
Dohrn A. 1866. Eugereon Boeckingi, eine neue Insectenform aus dem Todtliegenden. // Palaeontographica, 13: 333–340, pl. 41.
Enderlein G. 1904. Läusse-Studien. Über die Morphologie, Klassification und systematische Stellung der Anopluren nebst Bemerkungen zur Systematik der Insectenordungen. // Zoologischer Anzeiger, 27 (4): 121–147.
Hennig W. 1969. Die Stammesgeschichte der Insecten. Frankfurt am Main (Krammer): 1–436.
Hennig W. 1981. Insect phylogeny. Translated and edited by A. C. Pont, revision notes by D. Sclee. Chichester, New York, Brisbane & Toronto (John Wiley & Sons): 1–514.
Leach W.E. 1815. Classification. // In: D. Brewster, Edinburgh Encyclopedia, vol. 9, Entomology: 76–162.
Martynov A.V. 1924. [On the two kinds of insect wings and their evolution]. [Russian zoological Journal], 4 (1–2):155–185. [In Russian].
Scudder S. H. 1879. Palaeozoic cockroaches: a complete revision of the species of both worlds, with an essay toward their classification. // Memoirs of the Boston Society of natural History, 3 (1), No.3: 23–133, 6 pl.
Scudder S.H. 1895. The fossil cockroaches of North America. // Proceedings and Transactions of the royal Society of Canada, 12 (4): 147–153.