Phobaeticus kirbyi 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 : 153-157

Phobaeticus kirbyi Brunner v. Wattenwyl, 1907

( Figs. 114–115, 171–172, 237–239, 345–346, 361, 382, 418)

Phobaeticus kirbyi Brunner v. Wattenwyl, 1907: 185. HT, ♀: Borneo (BMNH). Brock, 1996: 29. Seow-Choen, 1998a: 41. Seow-Choen, 1998b: 89. Otte & Brock, 2005: 269.

Pharnacia kirbyi, Bragg, 1995a: 27 , fig. 1. [Description and illustration of egg]

Baculolonga kirbyi, Hennemann & Conle, 1997: 348 . Bragg, 2001: 379, figs. 142 (♂, ♀) & 143 (egg).

Pharnacia pilicornis Redtenbacher, 1908: 455 . HT, ♂: Brunei, Borneo, Staudinger; Collectio Br. v. W.; det. Br. v. W. Pharnacia pilicornis ; 20.854 (NHMW, No. 865). [Synonymised with Pharnacia sagitta by Günther, 1932: 315]

Phobaeticus pilicornis, Brock, 1998a: 50 . [As synonym of Ph. kirbyi ] Otte & Brock, 2005: 269. [As synonym of Ph. kirbyi ]

Pharnacia sagitta Redtenbacher, 1908: 454 , pl. 22: 2. HT, ♀ (subadult nymph): Coll. Br. v. W., Borneo, Doben; det. Br. v. W. Pharnacia sagitta ; 18.444; 65 (NHMW, No. 866). [Synonymised by Bragg, 1995a: 27]

Günther, 1932a: 315.

Günther, 1943: 155.

Phobaeticus sagitta, Brock, 1998a: 55 . [As synonym of Ph. kirbyi ]

Otte & Brock, 2005: 269. [As synonym of Ph. kirbyi ]

Pharnacia maxima, Kirby, 1904a: 359 . [Misidentification relating to the HT of Ph. kirbyi Brunner von Wattenwyl ]]

Pharnacia serratipes, Kirby, 1896: 448 & 450. [Misidentification relating to the HT of Ph. kirbyi Brunner von Wattenwyl ]

Redtenbacher, 1908: 455 (in part - only the specimen of Kirby, 1896: 448, the HT of Ph. kirbyi Brunner v. Wattenwyl).

Hypermergethes herberti Brunner v. Wattenwyl, in litt. ♂: Borneo; Hypermergethes herberti Br. det. (ETHZ).

Further material: [22 ºº, 20 ♂♂, 8 nymphs, 3 eggs]:

BORNEO, BRUNEI:

1 ♂ (penultimate instar nymph), 1 ♀ (penultimate instar nymph): Brunei, Seria , Badas, leg. Hennemann & Herbert VIII. 1994 (coll. FH, No’s 0160-1 & 2) ; 1 ♂, 1 ♀: Brunei, Badas , leg. M. Herbert IV.1995 (coll. MH, No’s 10-4 & 10-5) ; 1 ♀, 1 ♂: N-Borneo, Brunei, Capt. Waterstraat, Van der Poll, Coll. Bought, Janson 1909-463 & 464 ( OXUM) .

BORNEO, SARAWAK:

4 ♂♂, 4 ♀♀, 3 eggs: NO-Sarawak, Niah National Park , leg. Hennemann & Conle 2.-3. VIII. 1996 (coll. FH, No’s 0160-4 to 11 & E) ; 1 ♀ (penultimate instar nymph): NW-Sarawak, Mt. Serapi , ca. 200 m, leg. Hennemann & Conle VII.1996 (coll. FH, No’s 0160-3) ; 1 ♀, 3 ♂♂: Sarawak, Batu Niah , XII.1980, A. Harman leg., BM 1982-387 ( BMNH) .

BORNEO, SABAH:

4 ♀♀, 3 ♀♀ (nymphs): Brit. Nord-Borneo, Kinabalu-Gebirge, 16. XII. 1912 ( SMNS) ; 1 ♂, 1 ♀: Sandakan, C.V.Creagh, 96-197; 1 ♀ (nymph): B.N. Borneo, nr. Kinabalu, Kabayau, 600m, 9.V.1929 from H.M. Pendlebury collection ( RMNH) ; 1 ♀ (nymph): Murmet-Territory, N-Borneo, Higgins leg. 1871 ( RMNH) ; 1 ♀: Borneo Exped. Mr. Morvet, Boeleit, Dadop. ( RMNH) , 1 ♂: No. 193 (k / 52), XL.VIII. ( RMNH) ; 1 ♀: Flalek Papan, Kamp., Juli 1912 ( RMNH) , 1 ♂: B.N. Borneo, Mt. Kinabalu , Kian, 3000 ft., April 1929 ( RMNH) ; 1 ♀: NO-Borneo, S.- Peleben, VII.-VIII.1935, Baron Plessen leg. ( RMNH) ; 3 ♂♂: Brit. N-Borneo, Darvel-Bay , Pagel S.V. leg., det. K. Günther „ Phasgania pilicornis “ (MNHU) ; 1 ♂: N. Borneo, Kina Balu 1500 m, leg. Rolle 25. VI.1904 ( ZMUH) ; 1 ♀, 1 ♂: Nord-Borneo, Waterstraat ( ETHZ) ; 2 ♂♂: Labuan, Brit. N. Borneo, Pharnacia pilicornis Redt . ♂, Det. Hebard 1917, Hebard Cln. ( ANSP) ; 1 ♀: Labuan, Brit. N. Borneo; Phobaeticus n. sp. I, Det. Hebard 19; Hebard Cln. ( ANSP) ; 1 ♀: Labuan, Brit. N. Borneo; Pharnacia pilicornis Redt . ♀, Det. Hebard 1917; Hebard Cln. ( ANSP) .

BORNEO, KALIMANTAN:

1 ♀: Borneo, Exped. Dr. Nieuwenhuis, Long-Bloe, XII.1898 ( RMNH) ; 1 ♀: Borneo Exped. Mt. Morvet, Bukit Dadap [data difficult to read] ( RMNH) ; 1 ♀: E-Borneo, 125m, Tabang, 21.VIII.1956; det. P.E.Bragg ( RMNH) .

BORNEO:

1 ♂: Borneo , Hypermergethes herberti Br. det. ( ETHZ) ; 1 ♂, 1 ♀: Borneo , Phryganistria serratipes ( MNHN, coll. Finot, Box-No. 280) ; 1 ♂: Wattan-Borneo , Purchased and pres.´73 by Mrs. F.W. Hope ( OXUM) ; 1 ♀: Borneo ( ZSMC) .

NO DATA:

2 ♀♀: ( MNCN) .

Diagnosis: Well characterised and distinguished from other members of the genus by the strongly globose head of both sexes (Fig. 307–308), very short subgenital plate of ♀♀ which stays clearly beyond the apex of the abdomen (Fig. 237) and characteristic colouration especially of the head of ♂♂ (Fig. 308) which have the posteroventral carina of the profemora entirely unarmed (Fig. 346). The eggs clearly distinguish Ph. kirbyi from all remaining members of the genus by the distinct radial ridges of the dorsoventral keel of the capsule (Figs. 171–172).

Closely related to Ph. serratipes ( Gray, 1835) from Peninsular Malaysia, Singapore and Sumatra, and the three other Bornean species Ph. redtenbacheri ( Dohrn, 1910) , Ph. mjoebergi (Günther, 1935) and Ph. chani Bragg spec. nov.. From Ph. chani Bragg spec. nov. it furthermore differs by: the smaller size; relatively more robust body; less elongate body segments and lack of any enlarged teeth on the mesofemora of ♀♀, as well as the considerably shorter alae and slender, finger-like semi-tergites of the ♂♂ (Fig. 239). ♀♀ of Ph. redten- bacheri (Dohrn) and Ph. mjoebergi (Günther) at once differ from Ph. kirbyi by the strongly elongated, lanceolate subgenital plate, which considerably extends over the apex of the abdomen. ♂♂ of Ph. kirbyi are similar to those of Ph. redtenbacheri (Dohrn) but can in addition to the characteristic features mentioned above be distinguished by: the longer alae and apically tapered and acute semi-tergites of the anal segment.

Etymology: Dedicated to William Forsell Kirby (1844–1912), a well known English entomologist and folklorist, who first recorded this species as “ Pharnacia serratipes ”( Kirby, 1896: 448) .

Description: The colouration is described from colour photos of live adult insects taken at Batu Niah (NE-Sarawak) in August 1996.

♀♀ ( Fig. 114, 418): Long to very long (body length 187.0–317.0 mm) and slender species (average body width 4.0–5.0 mm) with a very short subgenital plate. Variable in size, colouration and leg armature. General colouration of body and legs mid to dull green or mid to dark brown, either uniformly coloured or with darker and paler speckles. Occasionally brown specimens occur which have a prominent white marking on the vertex, a bold white longitudinal median stripe on the pronotum, bold white patches and markings on the entire body, or a white longitudinal median line running along the full length of the dorsal body surface; in such specimens the meso- and metafemora with a broad whitish transverse sub-basal band. Armature of the legs dull red with black points. Eyes reddish brown. Antennae dark reddish brown and black ventrally, two basal segments of paler colour.

Head (Fig. 307): Strongly globose, indistinctly longer than wide and conspicuously broader than thorax; vertex convex. A slightly depressed area between the bases of the antennae. Eyes rather small, cylindrical and strongly convex; their length contained about 2.3x in that of cheeks. Antennae filiform and reaching to posterior margin of mesonotum, consisting of about 35 antennomeres. Scapus slender, 3x longer than wide, dorsoventrally flattened and narrowed towards the base. Pedicellus 1/3 the length of scapus, cylindrical. Following antennomeres first increasing then decreasing in length towards apex of antennae.

Thorax: Pronotum 1.7x longer than wide, shorter and distinctly narrower than head, the posterior portion somewhat broader than the anterior. Median transverse depression distinct very gently curved, but not reaching lateral margins of segment. Mesothorax about 3.6x longer than head and pronotum combined, mesonotum parallel-sided. Metanotum 4.5x longer than wide, rectangular and about 1/3 the length of mesonotum. Meso and metasternum simple.

Abdomen: Median segment slightly shorter than metanotum and slightly widened in posterior portion. Segments II–VI slightly increasing in length, II less than 4x, VI almost 5x longer than wide. VII slightly shorter than VI and very gently widened posteriorly. Tergite V with a scale-like posteromedian tubercle. Praeopercular organ formed by two prominent, sickle-shaped lobes near posterior margin of sternum VII (Fig. 361). Tergites VIII–X narrower than previous, VIII about half as long as VII and slightly narrowed in anterior half, IX half as long as VIII and indistinctly longer than wide. Anal segment conspicuously longer than IX, with a fine longitudinal median carina and a deep triangular posteromedian excavation; outer angles considerably deflexed and roughly triangular (Fig. 238). Supraanal plate very short, rounded. Subgenital plate longitudinally keeled, boat-shaped but short and only reaching half way along anal segment (Fig. 237).

Legs: All very long and slender but very variable in length. Profemora about as long as head, pro- and mesothorax combined, mesofemora slightly shorter than mesonotum, metafemora reaching about 1/3 the way along abdominal segment V and metatibiae very slightly projecting over apex of abdomen. Anterodorsal carina of profemora with 16–28 prominent and acutely triangular serrations (Fig. 345), posterodorsal carina very indistinct, unarmed. Armature of the posteroventral carina variable, either completely smooth, set with 3.4 minute teeth, or sub-basally with a ± prominent triangular lobe-like tooth and 3–5 smaller teeth along the length of femur. All carinae of mid and hind legs quite regularly set with small, pointed teeth (smaller on dorsal carinae). Medioventral carina very indistinct and set with a few minute spines in the median section of femur. Posterodorsal carina of meso- and metatibiae forming a slight rounded lobe at the apex; occasionally with a slightly enlarged tooth some 2/5 off the base, and rarely with another enlarged tooth 1/5 off the apex. Probasitarsus very elongate and longer than remaining tarsomeres combined, dorsal carina slightly elevated, all carinae smooth. Meso- and metabasitarsi a little longer than following three tarsomeres combined, the dorsal carina slightly but gradually raised towards the apex, all carinae minutely dentate.

♂♂ ( Fig. 115, 418): Moderately sized to large (body length 132.0–166.0 mm) and very slender for the genus (2.0–3.0 mm) with shortened alae (43.0– 49.5 mm) and a very characteristic colouration. Body and legs mid green (often greenish brown or pale brown when preserved). Head bright yellow with a distinct , broad black lateral marking on the cheeks. Tegmina and costal region of alae pale greenish brown, anterior margin broadly white. Anal region of alae transparent grey with brown veins. Dorsal serrations of the profemora orange or red with the posterior edges black; other leg armature black. Eyes yellow to ochre. Antennae black, except for a green to greenish brown scapus and pedicellus.

Head (Fig. 308): Strongly globose and generally as in ♀♀, but eyes much more prominent and projecting hemispherically; their length contained only about 1.5x in that of cheeks. Antennae projecting over posterior margin of median segment. Scapus dorsoventrally flattened, gently narrowed towards the base and almost 4x longer than wide. Scapus cylindrical and less than 1/3 the length of scapus.

Thorax: Pronotum 1.8x longer than wide, with a distinct median transverse depression and the anterior 1/ 3 of segment conspicuously constricted. Mesothorax about 3.5x longer than head and pronotum combined. Mesosternum with a distinct longitudinal median keel (very faint on metasternum). Metanotum short, only half the length of median segment. Tegmina slender, strongly narrowed in basal half and with a faint central hump. Alae slightly projecting over posterior margin of abdominal tergite III

Abdomen: Median segment 6x longer than wide. Abdominal segments II–VI 8x and VII 6x longer than wide. Posterior margins of tergites VI–V slightly raised, on V forming a transverse, scale-like posteromedian hump. VIII about 2x longer than wide, about half the length of VII and slightly widened towards the posterior. IX slightly shorter than VIII and constricted medially. Anal segment almost as long as VII, strongly tectiform and laterally compressed. Semi-tergites elongate, tapered towards a slender apex and slightly down-curving (Fig. 239). Interior surfaces apically with a blunt longitudinal keel set with several dark brown teeth. Cerci long, slender and in-curving, the apex distinctly tapered. Poculum strongly convex, cup-like and with a blunt, conical central hump.

Legs: All very long and slender, profemora longer than head, pro- and mesothorax combined, mesofemora a little longer than pro- and mesonotum combined, metafemora reaching to posterior margin of abdominal segment V and metatibiae projecting considerably over apex of abdomen. Anterodorsal carina of profemora armed with ± 20 prominent, acutely triangular teeth (Fig. 346), posteroventral carina unarmed (rarely with 2– 3 very minute teeth). All carinae of mid and hind legs densely armed with minute pointed teeth (more sparsely on dorsal carinae). Ventral surfaces of meso- and metafemora smooth. Probasitarsus very elongate, unarmed and conspicuously longer than remaining tarsomeres combined. Meso- and metabasitarsus a little longer than remaining tarsomeres and with all carinae minutely serrate.

Eggs (Figs. 171–172): The following description is based on three eggs laid by wild specimens from Batu Niah, NE-Sarawak. Due to Bragg (1995a & 2001) had a sample of 28 eggs at hand for examination, his measurements are given in addition to the ones taken from the three examples in the first author’s collection (coll. FH, No. 0160-E).

Capsule basically lens-shaped, laterally compressed, slightly longer than high. General colouration of capsule and operculum creamish pale to mid brown. Whole capsule surrounded by a prominent dorsoventral keel, beginning and ending at the operculum and only interrupted at polar-area and the micropylar plate. Polar impression distinct. Keel with an irregular outer margin and laterally furnished with distinct, impressed radial ridges, each basing in a deep circular impression where they join with the main capsule. Micropylar plate bilobed, strongly constricted medially and forming an oval lobe on each side of the dorsal median line. Outer margin with a fine reddish brown line. Micropylar cup small and placed at polar end of plate. Operculum slightly convex, oval and with an irregularly knob-shaped, black capitulum in the centre.

Measurements [mm]: Length (including capitulum) 6.0–6.4, length 5.3–5.6, height 4.4–4.8, width 3.1– 3.2, length of micropylar plate 1.9–2.1.

Measurements according to Bragg (1995b: 29 & 2001: 383) [mm]: Length 5.26–5.84, height 4.68–5.06, width 3.12–3.31, length of capitulum 0.71–0.84 (dehydrated).

Comments: Kirby (1896: 450) described the presumed ♀ of Ph. serratipes ( Gray, 1835) based on a tentatively identified specimen from Borneo in BMNH. For this specimen Kirby quoted a body length of “ 1 foot 10 lines “ which converts to 328 mm. This was given as 330 mm by Brunner v. Wattenwyl (1907: 185) and subsequent authors ( Bragg, 1995a: 28, 2001: 383). Kirby (1896: 448) referred to a second specimens and stated “Another specimen, from a doubtful locality, measures only 10 ½ inches in total length”. This latter specimen is not traced in BMNH. Kirby (1896: 448) already commented that he was not certain whether his Bornean specimens really represented Gray’s species and subsequently ( Kirby, 1904a: 359) decided them to be Pharnacia maxima ( Bates, 1865) which however is a junior synonym of Ph. serratipes (Gray) (syn. nov.). Brunner v. Wattenwyl (1907: 185) recognised that Kirby’s Bornean specimens were a distinct species and based on these described Phobaeticus kirbyi . Consequently, and as the second specimen is not traced the unique ♀ in BMNH must be considered the HT of Brunner v. Wattenwyl’s species ( Bragg 1995a: 28).

Redtenbacher (1908) described two species in the genus Pharnacia Stål, 1877 which are junior synonyms of Ph. kirbyi . Pharnacia pilicornis Redtenbacher, 1908 was described from a unique ♂ from Brunei in NHMW and synonymised with Pharnacia sagitta Redtenbacher, 1908 by Günther (1932: 315). Pharnacia sagitta Redtenbacher, 1908 was described from a single subadult ♀ nymph and synonymised with Ph. kirbyi by Bragg (1995a: 27).

Bragg (1995 a, 2001: 379) provided detailed descriptions of both sexes and the eggs, information on the synonymy and distribution of Ph. kirbyi as well as illustrations of the ♂, genitalia and egg. Bragg stated the length of the body and extremities to be considerably variable in specimens from different localities in Borneo. The ♀♀ from Mt. Kinabalu in SMNS and specimens in ANSP are remarkably shorter than all other recorded material.

For 95 years Ph. kirbyi was regarded “the longest insect in the world”, based on Kirby’s specimen in BMNH which had erroneously been identified as Pharnacia serratipes ( Gray, 1835) with a quoted body length of 330 mm. Re-measuring of the specimen has shown the actual body length to be only 317.0 mm. The overall length is 499 mm if the legs are included. Bragg (1995a: 28, 2001: 383) listed an even longer specimen with an overall length of 546 mm (body length 283 mm) from Kuala Belalong ( Brunei: Temburong District).

As can be seen from the numerous records, Ph. kirbyi is widely distributed in Borneo and quite abundant in certain localities. Bragg (1995a: 27 & 2001: 384) reported the species to be exceptionally common along the path leading from the Park Head Quarters to the Great Cave in Niah National Park (NE-Sarawak), with more than 40 adult specimens and numerous mating pairs encountered in only one night during August 1992. The authors encountered similar numbers at the same locality in August 1996.

Distribution (Fig. 382): Borneo, endemic. Sabah (Sandakan [ Bragg, 2001: 379]; Sepilok [ Bragg, 2001: 379]; Ulu Dusun [ Bragg, 2001: 379]; Labuan; Sepitang; Kabayau 600 m [ Bragg, 2001: 379]; Boeleit; Murmet Territory; Flalek Papan & Mount Kinabalu up to 1500 m), Brunei (Badas, swamp forest & Temburong District: Kuala Belalong 60 m), Sarawak (Niah National Park (= Batu Niah); Kuching [ Bragg, 2001: 379]; 3km NE of Sungai Arip, 20 km SW of Tatau [ Bragg, 2001: 379]; Baram River [ Bragg, 2001: 379] & Mount Serapi 200 m), Kalimantan (Balekpapan; Mount Muret: Bukit Dadap; Long Bloe Oe; Bengen River: Tabang 125 m & Wattan).

Number of specimens examined: 53