Abstract
The family Phasmatidae Gray, 1835 is reviewed and the subfamily Phasmatinae shown to be polyphyletic. Based on features of the exosceleton of the insects, egg-morphology and copulation habits a new arrangement of Phasmatidae is proposed. The monophyly of Lanceocercata Bradler, 2001 is confirmed but this name shown to be a synonym of Phasmatidae, hence Lanceocercata is here referred to as Phasmatidae sensu stricto. Six subfamilies belong in Phasmatidae sensu stricto all of which share several common and supposedly apomorphic characters: Phasmatinae, Tropidoderinae, Extatosomatinae (stat. nov.), Xeroderinae, Pachymorphinae and “Platycraninae”. The other two subfamilies contained in Phasmatidae sensu Bradley & Galil, 1977 (Eurycanthinae and Cladomorphinae) are not cosely related and here regarded as subfamilies of Phasmatidae sensu lato. The subfamily Phasmatinae sensu Bradley & Galil, 1977 is shown to be polyphyletic. The two tribes Pharnaciini and Clitumnini (= Baculini Günther, 1953) are removed from Phasmatinae and shown to be closely related to each other. They are transferred to the here established subfamily Clitumninae, a subordinate clade of Phasmatidae sensu lato. The subfamily Lonchodinae is closely related to Clitumninae, hence removed from Diapheromeridae and transferred to Phasmatidae sensu lato. The tribes Achriopterini and Stephanacridini (formerly in Phasmatinae) are shown to be not closely related to either Phasmatinae sensu stricto, Clitumninae or Lonchodinae, and provisionally must be treated as tribes of Phasmatidae sensu lato (incerte sedis). A re-arrangement of Phasmatidae sensu stricto is proposed along with determinating keys to all subfamilies and their tribes. The subfamilies Phasmatinae, Tropidoderinae and Extatosomatinae stat. nov. are re-described and discussed in detail. Full lists of genera are provided for each tribe. Only three of seven tribes formerly in Phasmatinae remain in the subfamily, this is Phasmatini, Acanthomimini and Acanthoxylini. The subfamily Tropidoderinae contains three tribes: Tropidoderini, Monandropterini and Gigantophasmatini trib. nov. The tribe Extatosomatini Clark-Sellick, 1997 is removed from Tropidoderinae and raised to subfamily level (Extatosomatinae stat. nov.). Several genera are transferred to other tribes or subfamilies. Didymuria Kirby, 1904 is removed from Tropidoderini, since it differs by having a closed internal micropylar plate in the eggs (open in all Tropidoderini). It here remains as a genus incerte sedis of Tropidoderinae and its systematic position clearly deserves further clarification. Gigantophasma Sharp, 1898 from the Loyalty Islands is removed from Pharnaciini, and becomes the type genus of the tribe Gigantophasmatini trib. nov.. Anophelepis Westwood, 1859 is removed from “Platycraninae” and shown to belong in Phasmatinae: Acanthomimini. The two Australian genera Arphax Stål, 1875, and Vasilissa Kirby, 1896 are removed from Acanthoxylini and provisionally transferred to Acanthomimini, but their position remains as yet debatable. Echetlus Stål, 1875 is misplaced in “Platycraninae” and shown to be a likely member of Phasmatinae. The two Brazilian species Echetlus evoneobertii Zompro & Adis, 2001 and Echetlus fulgens Zompro, 2004b are obviously misplaced and belong in the New World Diapheromeridae: Diapheromerinae: Diapheromerini. The subfamily Pachymorphinae is briefly discussed and considered polyphyletic. Two genera of Pachymorphinae: Gratidiini Bragg, 1995 (Parapachymorpha Brunner v. Wattenwyl, 1893 and Cnipsomorpha Hennemann et al., 2008) are transferred to Clitumninae: Medaurini trib. nov. The genus Gongylopus Brunner v. Wattenwyl, 1907 is transferred from Pachymorphinae: Gratidiini to Clitumninae: Clitumnini. The subfamily Xeroderinae is briefly discussed and shown likely to be polyphyletic, due to it contains two fundamentally different types of genitalia in the males. Only the genera Xeroderus Gray, 1835 and perhaps Epicharmus Stål, 1875 clearly belong in Phasmatidae sensu stricto. Both, the Pachymorphinae and Xeroderinae certainly deserve more detailed investigation to clarify their systematic positions with confirmation. Two generic groups are recognized within Clitumnini (subfamily Clitumninae). Due to differing by genital features and egg-morphology Medaura Stål, 1875 and Medauroidea Zompro, 2000 are removed from Clitumnini and transferred to the newly described Medaurini trib. nov.. The new tribe furthermore contains two genera formerly included in Pachymorphinae: Gratidiini and transferred here, Cnipsomorpha Hennemann et al., 2008 and Parapachymorpha Brunner v. Wattenwyl, 1893. Phryganistria Stål, 1875 is removed from Clitumnini and transferred to Pharnaciini. Nesiophasma Günther, 1934 is shown to belong in the tribe Stephanacridini. The Australasian subfamily Lonchodinae Brunner v. Wattenwyl, 1893 has formerly been included in Diapheromeridae Zompro, 2001 (= Heteronemiidae by Bradley & Galil, 1977). However, numerous features of the genitalia and egg morphology show close relation to the Oriental subfamily Clitumninae instead. Thus, Lonchodinae is here transferred to the family Phasmatidae (sensu lato). Within Lonchodinae the new tribe Neohiraseini trib. nov. is recognized and contains the five genera formerly placed in the “Neohirasea-complex” of that subfamily, namely Andropromachus Carl, 1913, Neohirasea Rehn, 1904, Pseudocentema Chen, He & Li, 2002, Qiongphasma Chen, He & Li, 2002 and Spinohirasea Zompro, 2001. It differs from all other Lonchodinae (= tribe Lonchodini) by the well developed vomer of males and the lack of a capitulum in the eggs. The genus Cladomimus Carl, 1915 was previously misplaced in Clitumninae: Pharnaciini and is here transferred to Lonchodinae: Lonchodini. It appears to be close to the Australian Hyrtacus Stål, 1875. Leprocaulinus Uvarov, 1940 and Phenacocephalus Werner, 1930 are removed from the subfamily Necrosciinae and transferred to Lonchodinae: Lonchodini. Extensive research on the genera which belong to the tribe Pharnaciini Günther, 1953 and taking features of the genital exosceleton and egg-morphology into account, has shown this tribe to be polyphyletic. Based on such features two generic groups are easily recognized within Pharnaciini sensu Günther, 1953. Males of the first group have a longitudinally split anal segment, which consists of two separate, more or less elongate semi-tergites and forms a clasping apparatus, the vomer is strongly reduced or lacking, the profemora have a prominent, lamellate medioventral carina which is strongly displaced towards the anteroventral carina and the eggs have an open internal micropylar plate with a clear median line. Only the genera falling into this group remain in Pharnaciini. Males of the second group in contrast have an anal segment which is not split, but possess a clearly visible, well sclerotised, triangular or hook-like external vomer, an indistinct medioventral carina on the profemora and eggs with a closed internal micropylar plate. Most of the genera which fall into the second group are here transferred to the tribe Stephanacridini Günther, 1953, this is Diagoras Stål, 1877b, Eucarcharus Brunner v. Wattenwyl, 1907, Phasmotaenia Návas, 1907 and Sadyattes Stål, 1875. A detailed discussion of the differences between Pharnaciini and Stephanacridini is provided along with distinguishing keys, illustrations and maps showing the distinct geographic distributions. The five genera that belong in Pharnaciini are: Baculonistria gen. nov., Pharnacia Stål, 1877a, Phobaeticus Brunner v. Wattenwyl, 1907 (= Baculolonga Hennemann & Conle, 1997a, = Lobophasma Günther, 1934b syn. nov. , = Nearchus Redtenbacher, 1908 syn. nov. ), Tirachoidea Brunner v. Wattenwyl, 1893 stat. rev. and Phryganistria Stål, 1875. Pharnacia annulata Redtenbacher, 1908 and Pharnacia enganensis Redtenbacher, 1908 were misplaced in Pharnacia Stål, 1877 (tribe Pharnaciini) and are transferred to the genus Sadyattes Stål, 1875 (tribe Stephanacridini, comb. nov.). Phobaeticus kuehni Brunner v. Wattenwyl, 1907 is removed from Phobaeticus Brunner v. Wattenwyl, 1907 (Phasmatinae: Pharnaciini) and shown to belong in Nesiophasma Günther, 1934c (tribe Stephanacridini, comb. nov.). Phobaeticus incertus Brunner v. Wattenwyl, 1907 (= Nearchus grubaueri Redtenbacher, 1908 syn. nov.) is unlikely to belong in Pharnaciini and here only retained in the original genus Phobaeticus Brunner v. Wattenwyl, 1907 with doubt, it may belong in Nesiophasma Günther, 1934c (tribe Stephanacridini). Based on a total of almost 700 examined specimens, the Oriental tribe Pharnaciini Günther, 1953 is revised at the species level. The new genus Baculonistria gen. nov. (Type species Baculonistria alba (Chen & He, 1990) comb. nov.), is described to contain three species from Central and Eastern China. Tirachoidea Brunner v. Wattenwyl, 1893 was erroneously synonymised with Pharnacia Stål, 1877 and is here re-established as a valid genus (stat. rev.). All five genera are re-diagnosed and differentiated, their systematic position within Pharnaciini discussed, and complete synonymic and species-listings as well as distribution maps and determination keys to the insects and eggs are provided. Detailed descriptions, diagnoses, synonymic listings, illustrations, material listings, distribution maps and measurements are provided for all 42 valid species. The type material of a further two species appears to be lost. Seven new species are described: Pharnacia borneensis spec. nov. from Borneo; Pharnacia palawanica spec. nov. from Palawan, Phobaeticus mucrospinosus spec. nov. from Sumatra, Phobaeticus palawanensis spec. nov. from Palawan, Tirachoidea herberti spec. nov. from Borneo, Tirachoidea siamensis spec. nov. from Thailand and S-Vietnam and Phobaeticus chani Bragg spec. nov. from Borneo. Phobaeticus chani Bragg spec. nov. is the world’s longest known insect with a maximum body length of 357 mm and an overall length of 567 mm in the female. Twelve new synonymies were discovered: Bactridium grande Rehn, 1920 = Phobaeticus serratipes (Gray, 1835) syn. nov.; Pharnacia rigida Redtenbacher, 1908 = Phobaeticus sumatranus Brunner v. Wattenwyl, 1907, syn. nov.; Clitumnus irregularis Brunner v. Wattenwyl, 1907 = Phibalosoma tirachus Westwood, 1859, syn. nov.; Pharnacia magdiwang Lit & Eusebio, 2008 = Pharnacia ponderosa Stål, 1877 syn. nov.; Pharnacia spectabilis Redtenbacher, 1908 = Phibalosoma hypharpax Westwood, 1859, syn. nov.; Pharnacia semilunaris Redtenbacher, 1908 = Eucarcharus inversus Brunner v. Wattenwyl, 1907, syn. nov.; Pharnacia chiniensis Seow-Choen, 1998c = Pharnacia biceps Redtenbacher, 1908, syn. nov.; Nearchus grubaueri Redtenbacher, 1908 = Phobaeticus incertus Brunner v. Wattenwyl, 1907, syn. nov.; Phibalosoma maximum Bates, 1865 = Cladoxerus serratipes Gray, 1835, syn. nov.; Phobaeticus lambirica Seow-Choen, 1998a = Eucarcharus rex Günther, 1928, syn. nov.; Phobaeticus sichuanensis Cai & Liu, 1993 = Baculum album Chen & He, 1990, syn. nov. and Phobaeticus beccarianus Brunner v. Wattenwyl, 1907 is shown to represent the previously unknown female of Phobaeticus sobrinus Brunner v. Wattenwyl, 1907 (syn. nov.) Lectotypes are designated for: Nearchus redtenbacheri Dohrn, 1910, Pharnacia biceps Redtenbacher, 1908, Pharnacia ingens Redtenbacher, 1908, Pharnacia heros Redtenbacher, 1908, Phibalosoma westwoodi Wood-Mason, 1875, Phobaeticus sinetyi Brunner v. Wattenwyl, 1907, and Phobaeticus sumatranus Brunner v. Wattenwyl, 1907. A neotype is designated for Nearchus maximus Redtenbacher, 1908 and Phobaeticus magnus nom. nov. introduced as a replacement name for Nearchus maximus Redtenbacher, which is a junior homonym of Phibalosoma maximum Bates, 1865.The previously unknown males of Pharnacia heros Redtenbacher, 1908, Phobaeticus ingens (Redtenbacher, 1908), Tirachoidea jianfenglingensis (Bi, 1994), Pharnacia sumatrana (Brunner v. Wattenwyl, 1907), Phryganistria fruhstorferi (Brunner v. Wattenwyl, 1907) and Tirachoidea westwoodii (Wood-Mason, 1875) as well as the females of Pharnacia ponderosa Stål, 1877a and Pharnacia tirachus (Westwood, 1859) are described and illustrated for the first time. A brief description on the basis of colour photos of the so far unknown male of Pharnacia kalag Zompro, 2005 are presented. Detailed descriptions and illustrations are provided for the eggs of 24 species. The eggs of the following 18 species are described and illustrated for the first time: Phobaeticus magnus nom. nov., Pharnacia borneensis spec. nov., Pharnacia palawanica spec. nov., Pharnacia ponderosa Stål, 1877a, Pharnacia sumatrana (Brunner v. Wattenwyl, 1907), Pharnacia tirachus (Westwood, 1859), Phobaeticus hypharpax (Westwood, 1859), Phobaeticus chani Bragg spec. nov., Phobaeticus incertus Brunner v. Wattenwyl, 1907, Phobaeticus magnus nom. nov., Phobaeticus philippinicus (Hennemann & Conle, 1997a), Phobaeticus sinetyi Brunner v. Wattenwyl, 1907, Phryganistria grandis Rehn, 1906, Phryganistria virgea (Westwood, 1848), Tirachoidea biceps (Redtenbacher, 1908), Tirachoidea herberti spec. nov., Tirachoidea jianfenglingensis (Bi, 1994) and Tirachoidea siamensis spec. nov.. Several species were originally placed in or subsequently transferred into wrong genera by various authors. Consequently, numerous taxa are here transferred or re-transferred to other genera, which results in 22 new or revised combinations or status of genera and species (comb. nov. / stat. rev. / stat. nov.). A list of the taxonomic changes made in this revision is provided in the summary ( 9.2), which in all lists 70 nomenclatural changes.