Lonchodinae Brunner

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 : 73-75

5.4 Subfamily Lonchodinae Brunner v. Wattenwyl, 1893

Type genus: Lonchodes Gray, 1835: 19 .

Lonchodidae Brunner v. Wattenwyl, 1893: 80.

Lonchodinae Rehn, 1904: 38 .

Kirby, 1904a: 317.

Günther, 1953: 548.

Bradley & Galil, 1977: 180.

Bragg, 2001: 431.

Otte & Brock, 2005: 30.

Prisomerinae Karny, 1923: 236 .

Lonchodinae Brunner v. Wattenwyl, 1893 is a very large subgroup of the anareolate Phasmatodea which presently contains 37 genera (→ see list below) and more than 300 described species. It shows a wide geographical range and extends from the Seychelles and India as far east as to New Guinea, the Solomon Islands and northern Australia. The most northern disperseals are Japan and central China.

Günther (1953: 560) characterized Lonchodinae by the following characters: anal segment of ♂♂ split and bilobed, or at least with two finger-shaped posterior processes; median segment considerably shorter than the metanotum; antennae long and filiform, usually as long as, or longer than body; ventral carinae of meso- and metafemora not distinctly serrate; both sexes apterous. Bradley & Galil (1977) placed Lonchodinae in Heteronemiidae and currently, Lonchodinae is treated as a subfamily of the Diapheromeridae Kirby, 1904 ( Zompro, 2001a: 192) . Several features of the genital exosceleton of Lonchodinae however clearly prove this subfamily is not a member of Diapheromeridae , but is closely related to the Old World subfamily Clitumninae of ( Phasmatidae s. l.). These are in particular the longitudinally split anal segment of ♂♂ which consists of two interiorly dentate (= “Dornenfeld”) and reduced or lacking external vomer, both of which correspond to Clitumninae (→ 5.). The profemora of Diapheromeridae are triangular in cross-section with the medioventral carina distinct, ledge or lamella-like and ± conspicuously displaced towards the anteroventral carina, ♀♀ have a praeopercular organ on sternum VII and the eggs have an open internal micropylar plate and conspicuous opercular structures, but the genital structures are strikingly distinct. ♂♂ of Diapheromeridae have the anal segment consisting of a single plate and never tectiform or split, with the posterior margin at best notched medially. The external vomer is either well developed and seen to be a sclerotised, roughly triangular or hooklike organ, or it is functionally replaced by strongly elongated often highly specialized cerci which enable ♂♂ to grasp the ♀ abdomen during copulation. Furthermore, as at least Diapheromerinae are restricted to the New World close relationship to Lonchodinae is rather unlikely. As a result, the striking genital exosceleton of the ♂♂ obviously places Lonchodinae near Clitumninae , hence it must be regarded a sub-ordinate taxon of Phasmatidae sensu lato rather than Diapheromeridae .

Although several features confirm a close relationship between Lonchodinae and Clitumninae (→ see below), Lonchodinae is well distinguished by the longer, filiform antennae which consist of more than 50 segments (exception Pericentrus Redtenbacher, 1908 ) and exceed half of the body length, as well as the lack of dentations or serrations of the legs.

Günther (1953: 560) divided Lonchodinae into two tribes, and in discussing the tribe Lonchodini , noted “Es liegt für diese Tribus nahe, an eine engere Verwandtschaft zwischen ihr und den Necrosciinae zu denken, zu denen sie in ähnlicher Beziehung stehen könnten, wie die Baculini [= Clitumnini ] zu den Pharnaciini in der subfam. Phasminae [= Phasmatinae ]” [For this tribe it seems appropriate to consider a closer relationship to Necrosciinae , to which they might stand in a similar relation as do Baculini to Pharnaciini in the subfamily Phasminae ]. For the second tribe of Lonchodinae , Neopromachini, Günther (1953: 561) supposed a possible relationship to the subfamily Eurycanthinae (family Phasmatidae ). Bradley & Galil (1977: 182) misinterpreted Günther’s Neopromachini and erroneously changed the name to Menexenini based on Menexi Brunner v. Wattenwyl, 1893. However, Günther had definitely placed the Menexenus Stål, 1875 in the tribe Lonchodini . Zompro (2001b) transferred three of the five genera which Günther placed in Neopromachini to Eurycanthinae (these are: Neopromachus Giglio-Tos, 1912 , Brachyrtacus Sharp, 1898 and Eupromachus Brunner v. Wattenwyl, 1907). As these include the type genus of Neopromachini, Neopromachus, Zompro transferred Neopromachini to Eurycanthinae but did not accept it as a separate tribe. Up to date Lonchodinae was not subdivided into tribes, but based on differences of genital structures Hennemann (2007: 24) united several genera in a generic group provisionally termed the “ Neohirasea -complex”.

Lonchodinae is quite heterogeneous both in egg morphology and several features of the insects, and so far no author has attempted to subdivide it into tribes, an exercise long overdue. The morphology of the eggs and genitalia of the insects indicate it will become necessary to subdivide Lonchodinae into at least three distinct tribes but this should be subject to a complete revision of Lonchodinae on the generic level. However, as the “ Neohirasea -complex” differs from all remaining genera of Lonchodinae , this group certainly represents a distinct tribal unit within the subfamily. Due to some genera (e.g. Mortites Günther, 1935 ) are probably misplaced in Lonchodinae and belong to other subfamilies, and the status of the “ Neohirasea -complex” is questionable (→ Neohiraseini trib. nov., see below), Lonchodinae as presently treated appears to be nonmonophyletic. Moreover, two predominantly New Guinean genera currently contained in the subfamily Necrosciinae ( Leprocaulinus and Phenacocephalus ) obviously belong in Lonchodinae : Lonchodini and are here transferred. This is emphasized by numerous features like the longitudinally split anal segment of ♂♂ which consists of two movable semi-tergites, lack of an external vomer, laterally compressed meso- and metafemora, dorsally lobed two basal tarsomeres, lamellate dorsal carina of the protibiae, and ovoid eggs with a stalked capitulum and an open micropylar plate. All of these features indicate relation to e.g. Hermagoras , Austrocarausius or sections of Carausius . Affinity of Leprocaulinus to the subfamily Lonchodinae was already suggested by Clark-Sellick (1997a: 106 & 1998: 216) who included Leprocaulinus in a “ Carausius - complex” of Lonchdinae, but did not formally transfer the genus. Examination of Cladomimus from the Loyalty Islands has shown this genus to be misplaced in Pharnaciini and to belong in Lonchodinae : Lonchodini . The elongated and apically pointed anal segment suggests it is close to the Australian Hyrtacus . The laterally compressed meso- and metafemora, dorsally lobed two basal tarsomeres and lamellate dorsal carina of the protibiae show affinity to e.g. Leprocaulinus , Phenacocephalus or Hermagoras and clearly place Cladomimus in the Lonchodinae . Carl (1915: 192) stated the antennae of the holotype of Cladomimus griseus Carl, 1915 (the only specimen known of the entire genus) to be shorter than the fore legs and to consist of only 25 segments, but examination of the specimen has proven them to be broken. They must therefore be assumed to have been considerably longer when complete.