Phasmatinae sensu Bradley & Galil, 1977

Hennemann, Frank H. & Conle, Oskar V., 2008, Revision of Oriental Phasmatodea: The tribe Pharnaciini Günther, 1953, including the description of the world's longest insect, and a survey of the family Phasmatidae Gray, 1835 with keys to the subfamilies and tribes (Phasmatodea: " Anareolatae ": Phasmatidae), Zootaxa 1906, pp. 1-316 : 22-23

Phasmatinae sensu Bradley & Galil, 1977

Various authors have already stated the Phasmatinae sensu Bradley & Galil, 1977 as well as the complete suborder “Anareolatae” are polyphyletic ( Bradler, 2000; Tilgner, 2002; Whiting et al., 2003 & Zompro, 2004a). From the seven tribes that Bradley & Galil (1977: 192) contained in Phasmatinae , namely Phasmatini , Acanthomimini , Acanthoxylini , Pharnaciini , Baculini (= Clitumnini ), Stephanacridini and Achriopterini , the latter four are obviously not closely related nor members of this subfamily. This is confirmed by striking differences concerning to the morphology of the genital exosceleton amongst the tribes included and was already pointed out by Bradler (2001), who placed Phasmatini , Acanthomimini and Acanthoxylini in his supposedly monophyletic group Lanceocercata (= Phasmatidae s. str.). Indeed, only these three tribes share several common characters which readily distinguish them from the other four tribes contained. These are the laterally compressed, foliaceous cerci, strongly tectiform anal segment of ♂♂ which forms a conspicuous clasping apparatus, lack of a praeopercular organ in ♀♀ and the derived copulation position in which ♂♂ grasp the base of the ♀♀ abdominal sternum VIII (= subgenital plate) instead of sternum VII (→ 4.).

A specialized anal segment which consists of two movable semi-tergites is also found in ♂♂ of Pharnaciini and Clitumnini , which just like the mentioned three tribes have the external vomer strongly reduced. However, these two tribes lack the foliaceous cerci, have the semi-tergites of the anal segment structured rather differently from those of Phasmatini , Acanthomimini and Acanthoxylini , and ♀♀ possess a more or less distinct often specialized praeopercular organ on sternum VII, used as an anchorage during the copulation by ♂♂ (→ 5.). Moreover, the profemora of Pharnaciini and Clitumnini differ considerably from those of the mentioned three tribes by having the medioventral carina very prominent, lamellate or ledge-like and strongly displaced towards the anteroventral carina (→ Figs. 46–47). Undoubtedly, Pharnaciini and Clitumnini are very closely related and together belong in a separate subfamily, the Clitumninae (→ 5.). The mentioned features indicate relationship between Clitumninae and Lonchodinae (→ 5.).

The tribes Stephanacridini and Achriopterini were shown to be closely related to each other ( Hennemann & Conle, 2004) and differ fundamentally from all other tribes contained in Phasmatinae by Bradley & Galil (1977: 192) in the morphology of the genital exosceleton (→ 4.7). ♂♂ of both tribes have a simple anal segment which consists of a single dorsal plate and have a well developed and sclerotised, hook-like external vomer. ♀♀ of Stephanacridini furthermore have long, filiform gonapophyses which extend well beyond the apex of the abdomen. Furthermore, both sexes lack conspicuously foliacous cerci, have them small and cylindrical to oval in cross-section, and can hence not belong in Phasmatinae or Phasmatidae s. str. (= Lanceocercata).

As discussed above only four of the six subfamilies that were included in Phasmatidae by Bradley & Galil (1977) are closely related and form the monophyletic taxon Phasmatidae s. str. (= Lanceocercata), this is Phasmatinae , Tropidoderinae , Xeroderinae and “ Platycraninae ”. The latter two subfamilies need further clarification since they appear to be polyphyletic (→ 4.6.2), as is Phasmatinae sensu Bradley & Galil (1977) . Only the three tribes Phasmatini , Acanthomimini and Acanthoxylini form a natural group and belong in Phasmatinae . This subfamily is obviously closely related to Tropidoderinae , and based on morphological features of the genital exosceleton, together with sections of Xeroderinae and “ Platycraninae ” forms a monophyletic group within the family Phasmatidae sensu lato (→ 4.).

The basalmost taxa of Phasmatidae s. str. (e.g. Extatosomatinae and certain Tropidoderini or Phasmatini ) presumably represent some of the evolutionary oldest extant Phasmatodea , exhibiting several features plesiomorphic for the Phasmatodea , e.g. a just slightly prognathous head, ocelli, a short mesothorax, long median segment and abdomen, and well developed wings. In several aspects genera like Podacanthus Gray, 1833 , Micropodacanthus Brock & Hasenpusch, 2007 ( Tropidoderinae : Tropidoderini ) or Paracyphocrania Redtenbacher, 1908 ( Phasmatinae : Phasmatini ) remarkably resemble fossil records of Lower Neoptera from the Tertiary supposedly belonging to Phasmatodea (e.g. Aeroplanoidea Tillyard, 1918 and Hagiphasmatidae Ren, 1997 , → 3.2). However, as there appear to be differences concerning the wing venation and the organization of the cubito-median veins in particular ( Nel et al., 2004) between these and Recent Phasmatodea , the phylogenetic relationship between these Tertiary orthopteroid phasmid-like insects and extant Phasmatodea still remains uncertain ( Tilgner, 2001).

In contrast to the hypothesis of Whiting et al. (2003) all subgroups of Phasmatidae s. str. show a clear evolution from winged to wingless forms and appear to have lost their wings during the corse of their evolution, rather than having them recovered secondarily. This is supported by various still questionable fossil records of Phasmatodea (→ 3.2), all of which are insects with well developed alae ( Zompro, 2004a: 323). Evolutionary trends observed in all subgroups of Phasmatidae s. str. are a shortening of the median segment, reduction and final loss of tegmina, alae and ocelli, increasing elongation of the mesothorax, as well as a decreasing crypsis and sexual dimorphism.