Paoliidae Handlirsch, 1906

Prokop, Jakub, Krzemiński, Wiesław, Krzemińska, Ewa & Wojciechowski, Dariusz, 2012, Paoliida, a Putative Stem-Group of Winged Insects: Morphology of New Taxa from the Upper Carboniferous of Poland, Acta Palaeontologica Polonica 57 (1), pp. 161-173 : 163-164

Family Paoliidae Handlirsch, 1906

Type genus: Paolia Smith, 1871 .

Composition.—Genera included by Carpenter (1992) supplemented by Brauckmann (1984) and reviewed by Rasnitsyn (2002b) and Prokop and Nel (2007): Holasicia Kukalová, 1958 ; Kemperala Brauckmann, 1984 ; Mertovia Prokop and Nel, 2007 , Olinka Kukalová, 1958 ; Paolia Smith, 1871 ; Paoliola Handlirsch, 1919 ; Pseudofouquea Handlirsch, 1906 ; Sustaia Kukalová, 1958 ; Zdenekia Kukalová, 1958 (a list of species is provided in the Appendix 1).

Comments.—The order Paoliida Handlirsch, 1906 (= Protoptera Sharov, 1966) is a small group of pterygote insects comprising ten genera and twelve species. Paoliids are known only from the continents belonging to former Laurussia (North America, East USA; Europe, Belgium, Czech Republic, England, Germany, the Netherlands, Wales), and from a relatively short period in the early Upper Carboniferous (Namurian B to Westphalian A [= Langsettian]).

The possible affinities of paoliid insects have been widely discussed. The family Paoliidae was created by Handlirsch (1906), who attributed this taxon to Palaedictyoptera (an extinct palaeopteran order) and proposed close relationships with spilapterids. Originally two species, Paolia vetusta Smith, 1871 and Paoliola gurleyi ( Scudder, 1885) were included on the basis of rich branching of cubital and anal veins forming a network along the posterior wing margin (Handlirsch 1906). Before that the first mentioned species was placed amongst protolocustids (i.e., stem−Caelifera) by Brongniart (1883), only to be somewhat later transferred by Scudder (1885) to protophasmids (i.e., stem−Phasmatodea) together with Paolia gurleyi . The latter species was illustrated for the first time by Melander (1903).

Carpenter (1954) re−assigned paoliids together with the oldest pterygotes ( Ampeliptera Pruvost, 1927 and Stygne Handlirsch, 1906 ) to the Palaeodictyoptera . At a later stage Kukalová (1958a), in her extensive work on fossil insects from the Upper Silesian Coal Basin in Czech Republic included this family in the order “ Protorthoptera ” (Cacurgoidea), currently considered as paraphyletic group (e.g., Béthoux and Nel 2005). Sharov (1962) first considered Paoliidae to fall into “ Paraplecoptera ” (currently treated as stem group of Grylloblattodea ) and later transferred them in the infraclass “Archaeoptera”, acknowledged by him as a basal stem group of Pterygota ( Sharov 1966: 115). He noticed the position of wing in imago of Paoliidae directed backwards at certain angle to the body at the rest, a position similar to many Palaeozoic larvae and, according to him, distinguishing this group from the Neoptera. Sharov (1966) also reviewed other characters in comparison to Neoptera, such as absence of any folds along cubital and anal veins. A new or− der Protoptera was created by him on the basis of fore and hind wings being homonomous, the absence of an anal fan and a specific position of the wings at the rest. Furthermore, Sharov (1966) noticed in the Paoliidae also an archedictyon (= dense pattern of reticulated crossveins), widely considered as a plesiomorphic character. He believed that the group might have given rise to the whole clade of winged insects (Pterygota), including Palaeoptera and Neoptera. Carpenter (1992: 100) considered the Paoliidae , together with Homoeodictyidae and Thoronysididae , as the most basal Protorthoptera , all having reticulate venation as well as concave MP in forewings. Further rearrangement into “hemipteroid lineage” was done by Kukalová−Peck and Brauckmann (1992) on the basis of the presence of an “arculus” between veins MP and CuA and accompanied by a simple CuP (or only terminal twigging), a deeply incised claval fold with tendency

http://dx.doi.org/10.4202/app.2010.0064

to form anal loops, and a pointed anal lobe as occurs in modern hemipteroids. Kukalová−Peck and Brauckmann (1992) proposed to include into paoliid line the following families: Paoliidae , Eucaenidae , Strephocladidae , Blattinopsidae , Synomaloptilidae , Cymbopsidae , and unassigned taxa: Limburgina antigua Laurentiaux, 1950 (assigned to Protorthoptera incertae sedis by Béthoux and Nel, 2002) and Heterologopsis ruhrensis Brauckmann and Koch, 1982 (later transferred in Archaeorthoptera sensu Béthoux and Nel, 2002). Haas and Kukalová (2001) considered hindwings of Paoliidae to represent the ancestral hemipteroids. Finally Rasnitsyn (1976, 2002a: fig. 1) postulated that paoliids are a basal stem group of the Pterygote lineage as was previously proposed by Sharov (1966), but without denoting any distinct synapomorphies.