Phobaeticus 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 : 127-134

6.3 Genus Phobaeticus Brunner v. Wattenwyl, 1907

103. Phobaeticus foliatus (Bragg, 1995) , ♀ HT, Midden-East-Borneo ( RMNH) .

104. Phobaeticus foliatus (Bragg, 1995) , ♂ Midden-East-Borneo ( RMNH)

105. Phobaeticus mucrospinosus spec. nov., ♀ HT, W-Sumatra, Solok ( RMNH)

106. Phobaeticus sinetyi Brunner v. Wattenwyl, 1907, ♀ PLT, S-India, Madras Prov., Shembagonor (NHMW) 107. Phobaeticus sinetyi Brunner v. Wattenwyl, 1907, ♂ LT, S-India, Madras Prov., Shembagonor (NHMW)

108. Phobaeticus magnus nom. nov., ♀ N-Thailand, Lemgao Petchabun (coll. FH, No. 0130-1)

109. Phobaeticus magnus nom. nov., ♂ N-Myanmar, Highland of Tenasserim (coll. FH, No. 0130-2)

110. Phobaeticus redtenbacheri ( Dohrn, 1910) , ♀ Sabah, Mount Kinabalu 1500 m ( ZMUH)

Type species: Phobaeticus sobrinus Brunner v. Wattenwyl, 1907: 184, pl. 7: 1a & b, by subsequent designation of Brock, 1996: 28.

Phobaeticus Brunner v. Wattenwyl, 1907: 183, pl. 7: 1a–b (in part).

Brock, 1996: 28 (in part).

Seow-Choen, 1997: 120, figs. 102–103.

Seow-Choen, 1998a: 40.

Brock, 1999: 136 (in part).

Seow-Choen, 2000: 38.

Otte & Brock, 2005: 268 (in part).

Baculolonga Hennemann & Conle, 1997a: 347 . (Type species: Cladoxerus serratipes Gray, 1835 , by original designation). [Synonymised by Otte & Brock, 2005: 268]

Bragg, 2001: 377 (in part).

Bacteria, Burmeister, 1838: 565 (in part).

Bactridium, Rehn, 1920: 242 .

Cladoxerus Gray, 1835: 42 (in part).

De Haan, 1842: 131 (in part).

Eucarcharus, Carl, 1913: 12 .

Günther, 1928: 218.

Otte & Brock, 2005: 136 (in part).

Heteronemia, Otte & Brock, 2003: 290 (in part).

Lobophasma Günther, 1935b: 138 (in part). (Type species: Eucarcharus rex Günther, 1928 , by original designation of Günther, 1935b: 139) syn. nov.

Günther, 1953: 555.

Otte & Brock, 2005: 178.

Nearchus Redtenbacher, 1908: 448 . (Type species: Nearchus maximus Redtenbacher, 1908 , by subsequent designation of Bragg, 1995: 273) syn. nov.

Dohrn, 1910: 409.

Günther, 1935a: 10.

Bragg, 1995b: 273, 274 (in part).

Brock, 1999: 131, 181.

Bragg, 2001: 388.

Otte & Brock, 2005: 209.

Pharnacia, Kirby, 1904a: 359 (in part—only species No’s 1, 1a, 2).

Redtenbacher, 1908: 449 (in part).

Karny, 1923: 240 (in part).

Günther, 1932: 315.

Brock, 1995: 95 (in part).

Phibalosoma, Westwood, 1859: 77 (in part).

Kaup, 1871a: 19.

Kaup, 1871b: 37.

Wood-Mason, 1877a: 161 (in part).

Wood-Mason, 1877b: 346 (in part).

Phryganistria Stål, 1875: 6 , 63 (in part).

Tirachoidea Brunner v. Wattenwyl, 1893: 83 (in part).

Kirby, 1904a: 359 (in part—only species No. 3).

Karny, 1923: 240 (in part).

Hypermergethes Brunner v. Wattenwyl, in litt.

Description (ď, º): Long to very long, mostly very elongate and extremely slender Pharnaciini with distinct sexual dimorphism. Head longer than wide and to almost spherical; vertex flat or rounded, smooth. Antennae of ♀♀ longer than mesonotum, of ♂♂ as long as or longer than head and complete thorax combined. Mesothorax very elongate and slender, 2.3–3.5x longer than head and pronotum combined, parallel-sided. Mesonotum usually smooth but occasionally covered with some minute granules or blunt spines in one species. Mesosternum with a distinct longitudinal median carina; occasionally median carina also present on metasternum. Median segment at least ½ the length to slightly longer than the metanotum. ♂♂ either completely apterous, with rudimentary or fully developed alae. Alae at beast reaching half way along abdominal tergite V. Tegmina (if present) elongate, spatulate and strongly constricted towards the base, central hump very indistinct; at least 2.5x longer than wide. Tergite VII of ♀♀ either parallel-sided or with a ± prominent posterolateral lobe; in one case tergites II–IX each bear a dentate lateral lobe. Sternum VII of ♀♀ with a ± distinct praeopercular organ, formed by a pair of more or less prominent tubercles, spines or triangular lobes. Cerci cylindrical or oval in cross-section, short in ♀♀, but elongate, constricted and incurving apically in ♂♂. Semi-tergites of ♂♂ anal segment strongly elongated, slender and conspicuously finger-like; occasionally down-curving. Subgenital plate of ♀♀ strongly keeled, either not reaching or only slightly projecting over the apex of anal segment, or very elongate, lanceolate and projecting over anal segment by as much as the combined length of abdominal tergites VII–X. Poculum of ♂♂ strongly convex and with a ± conical, backward pointing central spine at the angle. All legs very long and slender, carinae more or less distinctly serrate and / or occasionally armed with single enlarged teeth or triangular lobes; in one case with numerous prominent foliaceous appendages. Anterodorsal carina of profemora distinctly raised with prominent saw-like serrations (♀♀ in particular); serrations usually black in ♂♂. Meso- and metafemora with a fine and ± spinose median carina. Mesofemora distinctly longer than metanotum and median segment combined. Probasitarsus of ♀♀ at least as long as remaining tarsomeres combined usually distinctly longer; much longer than remaining tarsomeres in ♂♂. Meso- and metabasitarsus longer than following three tarsomeres combined, in ♀♀ dorsal carina simple or ± distinctly raised and rounded; never considerably elevated in ♂♂.

Eggs (Figs. 169–185): Medium-sized to large. Capsule strongly laterally compressed and elliptical in cross-section, basically lens-shaped and with a prominent keel on dorsal and ventral surfaces and polar-area. Keel with a depression at micropylar plate (sometimes faint) and at polar-area. Occasionally the keel may be considerably elevated, forming flap-like extensions. Capsule surface either almost smooth and shiny, minutely punctured or distinctly sculptured. External micropylar plate short, at best covering 2/3 of capsule length, usually transverse, bilobed and generally heart or kidney-shaped. Operculum elliptical and with a distinct conical, hat or knob-like capitulum on a short stalk.

Variation: In Phobaeticus Brunner v. Wattenwyl, 1907 several morphological characters of the insects and eggs are subject to considerable intrageneric variability. This e.g. concerns to: the shape and length of the subgenital plate of ♀♀; shape of the semi-tergites of the anal segment of ♂♂; shape of the head; relative length of the median segment; armature of the legs and length of the tegmina and alae in ♂♂. Even the eggs show considerable variability e.g. concerning to the sculpturing of the capsule and shape of the micropylar plate. All these features serve well for the distinction of species within the genus.

The ♀♀ subgenital plate varies considerably in length and shape. It is either short and does not reach the posterior margin of the anal segment (e.g. Ph. kirbyi Brunner v. Wattenwyl, 1907), roughly reaches to (e.g. Ph. serratipes ( Gray, 1835) or Ph. sinetyi Brunner v. Wattenwyl, 1907) or slightly projects over the same (e.g. Ph. lumawigi Brock, 1997 ), or is conspicuously lanceolate and projects over the apex of the abdomen by as much as the combined length of tergites VII–X (e.g. Ph. magnus nom. nov. or Ph. redtenbacheri ( Dohrn, 1910)) . The semi-tergites of the anal segment of ♂♂ vary in shape from being slender and straight with the apex strongly elongated and finger-like (e.g. Ph. serratipes ( Gray, 1835) and Ph. redtenbacheri ( Dohrn, 1910)) to rather short, broad and roundly rectangular in lateral aspect (e.g. Ph. chani Bragg spec. nov.). The head ranges in shape from elongate-oval, being distinctly longer than wide with the vertex flat (e.g. Ph. serratipes ( Gray, 1835) and Ph. chani Bragg spec. nov.), to globose and almost spherical (e.g. Ph. kirbyi Brunner v. Wattenwyl, 1907 or Ph. palawanensis spec. nov.). The mesonotum is usually destitute of spines or granules as typical for Pharnaciini but one species, Phobaeticus foliatus (Bragg, 1995) , has the mesonotum armed with several prominent spines. The median segment varies in length from being only half as long as the metanotum ( Ph. sinetyi Brunner v. Wattenwyl, 1907) to being considerably longer than the metanotum, especially in ♂♂ (e.g. Ph. palawanensis spec. nov.). ♂♂ are either apterous (e.g. Ph. sobrinus Brunner v. Wattenwyl, 1907 or Ph. sinetyi Brunner v. Wattenwyl, 1907), have vestigial tegmina and alae ( Ph. palawanensis spec. nov.), brac- hypterous (e.g. Ph. serratipes ( Gray, 1835) and Ph. kirbyi Brunner v. Wattenwyl, 1907) or are enabled of active flight with the alae well developed and covering most of the abdomen (e.g. Ph. hypharpax ( Westwood, 1859) or Ph. magnus nom. nov). The legs shows a very wide range of armature. They are only very sparingly dentate or densely serrate, may bear single enlarged teeth or lobes or may even be covered with numerous large, foliaceous appendages ( Ph. foliatus ( Bragg, 1995b)) . The medioventral carina of the meso- and metafemora is either smooth, minutely spinulose or armed with several prominent spines. Abdominal tergite VII of ♀♀ is either almost parallel-sided, posterolaterally elevated to form a more or less distinct, rounded or roundly triangular lobe, or in one case ( Ph. rex ( Günther, 1928)) is protruded into a large, irregularly dentate appendix which laterally extends by as much as the length of the complete segment. As an extreme, Ph. foliatus ( Bragg, 1995b) exhibits a more or less prominent, dentate lateral lobe on tergites II–X in ♀♀ and II–VI in ♂♂.

The egg capsule is generally restricted to the typical lens-like shape but shows variation concerning to the shape and structure of the dorsoventral keel which surrounds the capsule, shape of the micropylar plate, operculum and capitulum as well as the sculpturing of the capsule surface. Unfortunately, up to now fully developed eggs of only nine species are known, which does obviously not allow to describe the complete existing variability that occurs in eggs of Phobaeticus .

Diagnosis ( Table 2): Phobaeticus Brunner v. Wattenwyl is easily distinguished from all other genera of Pharnaciini by: the strongly laterally compressed, keeled and lens-shaped egg capsule; short, mostly bilobed, heart or kidney-shaped external micropylar plate which usually covers less than 2/3 of the capsule length as well as the elongate, slender and finger-like semi-tergites of the anal segment and elongate, apically constricted and in-curving cerci of ♂♂. It is very closely related to Phryganistria Stål, 1875 , with which it shares the distinct praeopercular organ of ♀♀. Apart from the typical features mentioned above, Phobaeticus differs from Phryganistria by: the longer median segment of both sexes which is longer or at least ½ the length of the metanotum, the keeled mesosternum of ♂♂ and relatively shorter antennomeres.

Certain species of Phobaeticus may at first glance show striking similarity to the more slender taxa of Pharnacia Stål, 1877 but apart from the distinguishing features mentioned above, Phobaeticus differs from that genus by: the presence of a prominent praeopercular organ and antennae of ♀♀, being longer than the mesonotum; not conspicuously elevated carinae of the basitarsi of ♂♂ as well as the shorter and differently shaped micropylar plate of the eggs (shaped like an inverted “Y” in Pharnacia ). Furthermore, the subgenital plate of ♀♀ is never strikingly elongated and lanceolate in Pharnacia Stål and the median segment of ♀♀ is at least 2/3 the length of the metanotum.

Comments: Brunner v. Wattenwyl (1907: 183) originally placed Phobaeticus in Clitumnini and characterized it as being apterous in both sexes. Consequently, he included species most of which were only known from the ♀♀. Only the apterous ♂♂ of Ph. sinetyi Brunner v. Wattenwyl, 1907 and Ph. fruhstorferi Brunner v. Wattenwyl, 1907 were known to Brunner v. Wattenwyl, but the latter is obviously not a member of the genus and here transferred to Phryganistria Stål, 1875 (→ 6.4). Ph. kuehni Brunner v. Wattenwyl, 1907 from Roma Island ( Indonesia) is also not a member of Phobaeticus and here transferred to the genus Nesiophasma Günther, 1934c ( Phasmatidae : Stephanacridini , → 5.6). Ph. incertus Brunner v. Wattenwyl, 1907 is of uncertain systematic position and here removed from Phobaeticus ; it may belong in Nesiophasma Günther, 1934c (→ see below). Brunner v. Wattenwyl was not aware that some of the corresponding and winged ♂♂ of the species he described in Phobaeticus were included in Pharnacia Stål, 1877 by Redtenbacher (1908: 449) in the third part of their well-known monography “Die Insektenfamilie der Phasmiden”. Brock (1996: 26) recognize the erroneous position of Phobaeticus and transferred it from Baculini Bradley & Galil, 1977 (= Clitumnini Brunner v. Wattenwyl, 1893) to Pharnaciini . From the species originally placed in Phobaeticus by Brunner v. Wattenwyl, Brock (1996: 28) designated Ph. sobrinus Brunner v. Wattenwyl, 1907 as the type species, since it is the only species that Brunner v. Wattenwyl illustrated (1907, pl. 7: 1a–b, ♂).

Brock (1996) proposed a re-arrangement of the genera Pharnacia Stål, 1877 and Phobaeticus Brunner v. Wattenwyl, 1907 and synonymised Tirachoidea Brunner v. Wattenwyl, 1893 with Pharnacia Stål. Brock (1996: 28) re-defined Phobaeticus as containing apterous ♂♂ as well as ones with shortened or fully developed alae. Consequently, and on the basis of lacking a conically swollen or bi-tuberculate vertex, Brock (1996) transferred several of the more elongate species which were included in Pharnacia Stål, 1877 by Redtenbacher (1908) to Phobaeticus . The convex, swollen and/or bi-tuberculate vertex is however not a distinctive feature between Phobaeticus Brunner v. Wattenwyl and Pharnacia Stål but a key feature that distinguishes Pharnacia Stål from Tirachoidea Brunner v. Wattenwyl, 1893 (→ 6.2 and 6.5). The latter genus is here removed from synonymy with Pharnacia Stål and re-established as a valid genus (→ 6.5., stat. rev.).

As genital features and egg-morphology were generally not taken into account Phobaeticus remained as yet polyphyletic. Examination of the genitalia has shown three of the species which Redtenbacher (1908) originally described in Pharnacia Stål are not members of the tribe Pharnaciini , this is Phobaeticus annulatus Redtenbacher, 1908 , Ph. enganensis Redtenbacher, 1908 and Ph. nigricornis Redtenbacher, 1908 . These three species belong to the genus Sadyattes Stål, 1875 now in Stephanacridini and are here transferred (comb. nov.). Based on egg-morphology like the typically Y-shaped micropylar plate and genital features such as lacking a conspicuous praeopercular organ in ♀♀ two species need to be re-transferred to Pharnacia Stål , this is Pharnacia heros Redtenbacher, 1908 and Phibalosoma tirachus Westwood, 1859 (stat. rev.). Another four species do not belong to either genus but are members of Phryganistria Stål, 1875 or Tirachoidea Brunner v. Wattenwyl (→ 6.4, 6.5).

There are obvious differences which clearly distinguish Phobaeticus from Pharnacia , such as the presence of a distinct praeopercular organ in ♀♀, the comparatively much more elongate and finger-like semitergites of the anal segment and elongate and apically constricted, in-curving cerci of ♂♂ (→ Table 1 and diagnosis above). Furthermore, eggs of Phobaeticus differ significantly from those of Pharnacia , being laterally compressed, ± lens-shaped, surrounded by a distinct dorsoventral keel and having the micropylar plate considerably shorter, not Y-shaped but bilobed, kidney or heart-shaped and usually covering less than half of the capsule length.

Redtenbacher (1908: 448) established Nearchus for two newly described species, N. maximus Redtenbacher, 1908 (= Phobaeticus magnus nom. nov.) from Laos and Thailand and N. grubaueri Redtenbacher, 1908 (= Ph. (?) incertus Brunner v. Wattenwyl, 1907 syn. nov.) from Peninsular Malaysia and Banka Island in the Straight of Malacca. Redtenbacher (1908: 448) stated the new genus to be identical to Pharnacia Stål except for having a conspicuously elongate and lancet-like subgenital plate in ♀♀ (apparently referring to the more elongate species which actually belong in Phobaeticus ). But as discussed above the length of the subgenital plate is an extremely variable feature within Phobaeticus and the closely related Phryganistria Stål, 1875 , and hence only worth for distinction at the species level. Detailed examination of N. maximus Redtenbacher , the type species of Nearchus , and comparison with species of Phobaeticus have not revealed any differences that can be regarded as being of generic value. Consequently, Nearchus Redtenbacher is here synonymised with Phobaeticus Brunner v. Wattenwyl (syn. nov.). A synonymy of Phobaeticus and Nearchus was already suggested by Seow-Choen (1998a: 40). Bragg (1995b: 237) designated N. maximus Redtenbacher, and Brock (1995: 94) subsequently selected N. grubaueri Redtenbacher (= Ph. (?) incertus Brunner v. Wattenwyl, 1907 syn. nov.) as the type species of Nearchus . Due to an earlier publication date the type selection of Bragg (1995b) has priority ( Bragg, 2001: 388).

Lobophasma was established by Günther (1935b: 139) to include Eucarcharus rex Günther, 1928 , Eucarcharus lobulatus Carl, 1913 and Eucarcharus inversus Brunner v. Wattenwyl, 1907 of which he designated E. rex Günther as the type species. Careful investigation of E. rex and comparison with species of Phobaeticus have shown this to belong in Phobaeticus , which consequently places Lobophasma as a junior synonym (syn. nov.). This is e.g. seen in the presence of a distinct praeopercular organ, slender body and shape of the eggs

111. Phobaeticus redtenbacheri ( Dohrn, 1910) , ♂ Sabah, Mount Kinabalu 1500 m (coll. FH, No. 0264-1) 112. Phobaeticus serratipes ( Gray, 1835) , ♀ West Malaysia, captive reared (coll. FH, No. 0030-10)

113. Phobaeticus serratipes ( Gray, 1835) , ♂ West Malaysia, captive reared (coll. FH, No. 0030-6)

114. Phobaeticus kirbyi Brunner v. Wattenwyl, 1907, ♀ NE-Sarawak, Batu Niah (coll. FH, No. 0160-6)

and micropylar plate. The large, foliaceous lateral appendages of abdominal tergite VII of Lobophasma rex , which Günther (1935b: 139) stated to be the main key feature of Lobophasma , are unique in the entire Pharnaciini , but are a considerably variable feature within Phobaeticus and therefore not of generic value. Furthermore, Lobophasma sensu Günther is seen to be polyphyletic since E. inversus Brunner v. Wattenwyl belongs in Tirachoidea Brunner v. Wattenwyl, 1893 (→ 6.5).

Baculolonga was established by Hennemann & Conle (1997a: 347) to contain Ph. serratipes ( Gray, 1835) , Ph. kirbyi Brunner v. Wattenwyl, 1907, Nearchus redtenbacheri Dohrn, 1910 and the newly described Baculolonga philippinica . The authors designated C. serratipes Gray as the type species and distinguished Baculolonga from Phobaeticus on the basis of egg-morphology and the longer median segment of ♂♂. However, in Brunner v. Wattenwyl’s illustration of the ♂ HT of the type species Phobaeticus sobrinus Brunner v. Wattenwyl (1907: pl. 7: 1a) the median segment appears shorter than it actually is, which was proven by subsequent examination of the specimen. As further investigation and comparison have shown the length of the median segment to be a very variable feature in Phobaeticus , Baculolonga was correctly synonymised with Phobaeticus by Otte & Brock (2005: 268).

Although the genus mostly contains very long and often extremely slender forms, the ♀♀ of certain more robust species (e.g. Ph. sinetyi Brunner v. Wattenwyl, 1907) may at first glance strongly resemble some of the more slender members of Pharnacia Stål, 1877 . The presence of a distinct praeopercular organ however readily distinguishes Phobaeticus -species from Pharnacia . Due to numerous single sexes and eggs are still unknown, the definition of species-groups or an intrageneric systematization remain difficult.

Distribution ( Fig. 84): So far recorded from India, Sri Lanka, Bangladesh, Myanmar, Laos, Thailand, Peninsular Malaysia, Singapore, Sumatra, Banka Id. prope Sumatra, Borneo, Palawan & the Philippines.