Home page >>> History of Insects SUPERORDER DICTYONEURIDEA Handlirsch, 1906

(= Palaeodictyopteroidea)

by N.D. Sinitshenkova

(a) Introductory remarks. Concept of the superorder is original differing much from previous accounts (cf. Rohdendorf 1969, Rohdendorf & Rasnitsyn 1980, Kukalová-Peck 1991). The reason is the accumulation of groups combining characters of different orders and even former superorders and thus obscuring their boundaries (those assigned here to Psychroptilidae and Frankenholziina in Dictyoneurida and to Eubleptina in Mischopterida). This made unescapable re-arranging the group as a whole.

(b) Definition. Body size large to very large, sometimes medium (20-400 mm wingspan). Head with sucking beak formed by mandibular, maxillary, and hypopharyngeal stylets, and sheathed by labrum (dorso-basally), maxillary palps, and sometimes by labium. Prothorax and abdominal segments usually with lateral paranota (pronotal paranota can be circular), sometimes bearing marginal spines, or lost. Wings costalised (with C, SC, and R thick and running relatively close each other), with single convex M vein (either MA or MA stock + MA1) forming fore margin of M system (unlike mayflies and dragonflies with entire MA convex with concave intercalaries). Costal margin straight to gently convex, costal space narrow, RS predominantly pectinate, MA poor in branching. Hind wing with anal area at most moderately enlarged. Flight variable: fast but not manoeuvrable, often of biplane type (with widely overlapping fore and hind wings) in the largest dictyoneuridans; more manoeuvrable and versatile in respect of flight speed in moderately large forms with wide wing bases; slow and highly manoeuvrable, approaching hovering ability in the case of those with a narrow wing base (Wootton & Kukalová-Peck 2000). Legs hardly much used in walking, often with clinging adaptations, particularly in fore pair (coxae shifted or inclined forward, tarsi sometimes 2-segmented with single thick claw, etc.). Ovipositor short to moderately long, cutting (probably using to place eggs into plant tissues), with sheath shorter than stylets, bearing long stylus. Cerci long, multisegmented. Development gradual, terrestrial (phytophilous), with imaginal moults. Adult and most probably immature feeding by sucking ovule contence of gymnosperm plants.

(c) Synapomorphies. Wings costalised, with fore M vein (either simple MA or MA stock + MA1) convex; possibly: prothoracic paranota winglet-like (unless inherited from ancestral Hypoperlida Strephocladina and supposing circular paranota in Homoioptera Brongniart a reversal).

(d) Range. Latest Early Carboniferous (early Namurium A) through the Late Permian; widespread but common only in Eurameria and, in Early Permian, in westernmost Angaraland (the Urals).

(e) System and phylogeny. The superorder is hypothesised (see Fig. 109) to descend from Heterologopsis-like forms (Fig. 134) which may deserve the state of the order per se, but because of insufficient knowledge are considered as unplaced Dictyoneuridea. Otherwise the superorder is taken composed of three orders, with Diaphanopterida and Mischopterida forming a sister group in respect to Dictyoneurida. Hence it is supposed that the permanently outstretched wing position has been acquired independently by Dictyoneurida and Mischopterida.

Homoeodiction Martynov (see Fig. 73) from the Late Permian of Kargala in the Urals and Klebsiella Meunier from the Late Carboniferous of Commentry in France are similar to dictyoneurideans in the convex MA but otherwise differ in their venation and so are considered Scarabaeones incertae sedis [2.2]. ORDER DICTYONEURIDA Handlirsch, 1906

(= Palaeodictyoptera Goldenberg, 1854)

(a) Introductory remarks. One of the leading and most popular groups of Palaeozoic insects, spectacular because of their large size, long and wide, often brightly patterned wings, and long cerci (Figs. 135, 136). Concept of the order is essentially modified comparing Rohdendorf & Rasnitsyn (1980), particularly concerning taxonomy. Primary data are reviewed and catalogued by Handlirsch 1906-1908, 1922), Rohdendorf (1962), Carpenter (1967a), Kukalová (1969a, b, 1970), Brauckmann (1991), and completed by Sharov & Sinitshenkova (1977), Sinitshenkova (1979, 1980a, 1980b, 1981, 1992a), Carpenter (1983, 1992a), Pinto (1994a), Brauckmann & Gröning (1998), Brauckmann et al. (1996)

(b) Definition. Body stout, large to very large (up to more than 50 cm wingspread). Integument rather thick, finely granulated. Head with sucking beak directed obliquely forward and formed by mandibular, maxillary, and hypopharyngeal stylets, and sheathed by labrum (dorso-basally), and maxillary palps. Beak moderately (Spilapteridae) to very long (Eugereonidae). Prothoracic paranota either circular (Fig. 137) or, usually, forming winglets. Wings spread permanently (not folding over abdomen at rest). Both wing pairs similar except minor venational details and wider base in hind wing, with either rich archedycteon or numerous cross-veins (exceptionally cross-veins less common, forming transverse rows), sometimes with evident nygmata (in both adult and nymphal wings of Homoiopteridae, Lithomanteidae, Breyeriidae), often brightly patterned (with dark stripes in Spilapteridae, Lamproptilidae, Fouqueidae). Longitudinal veins straight or gently curved, rarely with CuP and A more strongly curved (Calvertiellidae, Breyeriidae, Eugereonidae); occasionally merging for a distance and then more sharply bent. C often running submarginally, leaving narrow precostal space. Because of permanently spread wings, adult able only to fly and to cling to plant for feeding, oviposition and rest. Otherwise as in superorder [].

(c) Synapomorphies. Head prognathous; wings permanently spread; M5 lost.

(d) Range. The Early Carboniferous (Namurium A) through the Late Permian (Kazanian), with highest abundance during Middle and especially Late Carboniferous; widespread in the north hemisphere but common only in Eurameria, solitary records also in Argentina, Tasmania, East Siberia and Mongolia. Most probably the distribution results from the dictyoneurid thermophily.

(e) System and phylogeny of the order (Fig. 138) are based in part on Riek (1976c), otherwise are original. Unfortunately the group is possible to be classified relying almost entirely on the venational characters.

Two suborder and 8 superfamilies are proposed here. Dictyoneurina Handlirsch, 1906, stat. nov. includes 6 superfamilies of the typical palaeodictyoneuridans with wings rich in venation, wide including basally so as fore wing broadly overlapping hind one, and best fitted for soaring flight. Eugereonoidea Handlirsch, 1906, stat. nov. (Dictyoptiloidea sensu Riek 1976c, but Dictyoptilidae have been synonymised with Eugereonidae by Carpenter 1964, and the synonymisation has not been challenged yet) is the largest one with its 11 constituent families (Eugereonidae, Fig. 139, Archaemegaptilidae, Graphiptilidae, Jongmansiidae, Lithomanteidae, Lycocercidae, Megaptilidae, Polycreagridae, Protagriidae, Synarmogidae, Tchirkovaeidae). Characteristic for the group are wing length 2.5 times width, SC reaching C near the wing apex, CuA and MA with at most short apical fork, branching MP and CuP, and the archedictyon replaced by numerous cross-veins or sparse between more spaced veins.

Dictyoneuroidea Handlirsch, 1906, stat. nov. with 3 families [Dictyoneuridae (including Palaeoneura giligonensis Hong, described in Neuburgiidae, a synonym of Spilapteridae: Hong 1985b), Peromapteridae, Saarlandiidae) similar to Eugereonoidea but differing in wing length 3.5 times width, dense archedictyon and often narrow wing apex.

Homoiopteroidea Handlirsch, 1906, stat. nov. (consisting of Homoiopteridae Figs. 137, 140, and Heolidae) and the closely related superfamily Spilapteroidea Brongniart, 1893, stat. nov. with its 6 families (Spilapteridae, Figs. 135, 136, 141, Aenigmatidiidae, Fouqueidae, Homothetidae, Lamproptilidae, Mecynostomatidae) differ from Eugereonoidea in multibranched MA. They differ each other in the veins arching gently in the very wing base of Spilapteroidea vs. not bent in Homoiopteroidea.

Unlike the four above groups, Breyerioidea Handlirsch, 1904, stat. nov. and Calvertielloidea Martynov, 1932, stat. nov., have SC short (scarsely longer than half wing length) and meeting R (except Cryptoveniidae). The former superfamily includes 3 families (Breyeriidae, Stobsiidae, Cryptoveniidae) and the latter only two (Calvertiellidae and Mongolodictyidae which is unusual in having R seemingly falling into SC). Calvertielloidea differ in the strikingly arching longitudinal veins, in simple CuP, and in CuA lost its base and thus seemingly originating from M.

There are also several insufficiently known groups possible to be classified only as Dictyoneurina incertae sedis (Archaeoptilidae, Lithoptilidae, "Mecynoptera" tuberculata Bolton).

Suborder Frankenholziina Guthörl, 1962, stat. nov. is less diversified covering only 10 families in 2 superfamilies. Unlike Dictyoneurina and similarly to Mischopterida its wings are narrow, especially basally, indicating more active and manoeuvreable flight. Frankenholzioidea Guthörl, 1962, stat. nov. (including Frankenholziidae, Dictyoneurellidae, Caulopteridae, Ancopteridae, Fig. 142, and Psychroptilidae) is less advanced and shows only incipient modifications in the above direction. These changes are better manifested in Arcioneuroidea Kukalová, 1975, stat. nov. (Arcioneuridae, Elmoboriidae, Hanidae, Fig. 143, Eubrodiidae fam. nov. [Eubrodia Carpenter, 1967, type genus; the family differs from all other Frankenholziina in having the unique combination of simple MA and CuA, and regular archediction covering the hole wing], Eukulojidae).

Excluded from the order are Eohymen Martynov (Caloneurida [], Boltonocosta Carpenter, and Hypermegethes Handlirsch (Hypoperlida []), Merlebachia Waterlot (Scarabaeones incertae sedis probably related to Paoliidae [2.2]), and Bardapteron G. Zalessky (incertae sedis).

(f) History. The oldest (Namurian A, Arnsbergian) Delitzshala bitterfeldensis (Spilapteridae) from Germany is the only Early Carboniferous winged insect (Fig. 136). The Namurian (the latest Early and earliest Middle Carboniferous) dictyoneurids are found in West Europe and represented by 11 species belonging to Breyeriidae, Dictyoneuridae, Graphiptilidae, Homoiopteridae (Fig. 137), Lithomanteidae, and Spilapteridae. The post-Namurian (Westphalian) Middle Carboniferous representatives were much more numerous and diverse, being represented by ca.140 species of 21 families in fossil sites of West Europe and North America, with leading families being Dictyoneuridae and Spilapteridae. Still more diverse and abundant they were in the Late Carboniferous (Stephanian), this time predominantly in West Europe (20 families, dominated by Dictyoneuridae and Spilapteridae, but also in North America (12 families, dominated by Dictyoneuridae), Siberia (3 families, with most abundant Tchirkovaeidae, see Fig. 71), China, Argentina and Tasmania, each with a single family. Most of the Namurian families have persisted in the Westphalian, and all the Westphalian ones have survived till Stephanian. This distribution pattern, and particularly the low diversity of Siberian assemblage of Chunya strongly dominated by the sole species Tchirkovaea guttata M. Zalessky indicates that the group was thermophilous and rather uniform and stable over the equatorial belt in the Carboniferous.

Much rarer were the Early Permian dictyoneurids. 18 species are found there, mostly belonging to the families inherited from the Late Carboniferous. These are Dictyoneuridae, Spilapteridae, Calvertiellidae and Homoiopteridae represented in Europe (Obora in Czechia and Tshekarda at the Urals, Fig. 141), in North America (Elmo in Kansas, Fig. 135) and Siberia (Fatyanikha at Tunguska River). Several more families have appeared there (all in Europe) for the first time. These families (Ancopteridae, Arcioneuridae, Caulopteridae, and Hanidae, all from Obora in Czechia, Figs. 142, 143) represented not only taxonomic, but also morphological and bionomic innovation. They had narrow wings, often with festooned hind margin and surely practised different type of flight comparing normal Dictyoneurida.

Particularly poor were the Late Permian dictyoneurid fauna. Ufimian beds of Inta Formation in Pechora Basin (North Russia) have yielded 2 species of Spilapteridae (Fig. 144). Another North Russian locality Soyana (Arkhangelsk Region) of the Kazanian age has given 1 species of Calvertiellidae (Fig. 145) and 4 more species of the highly unusual family Eukulojidae: dipterous, with wide, poorly veined wings (Fig. 146). One species of Mongolodictyidae is known from Bor-Tologoy (possibly the Tatarian of Central Mongolia).

The Permian decline of the order seems possible to attribute in part to the decreasing abundance of the food plants (Sharov 1973), in part to increasing pressing from the flying predators dragonflies.

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