Tirachoidea cantori ( Westwood, 1859 ), 2008

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 : 226-232

Tirachoidea cantori ( Westwood, 1859) stat. rev.

( Figs. 133–134, 157–158, 212–214, 272, 401)

Phibalosoma cantori Westwood, 1859: 74 , pl. 37, fig. 1, 1a–b (♂), pl. 38, fig. 1, 1a–b (♀). LT, ♂: Cantor, Malacca; Type, Westwood. Phibalosoma cantor i, Type Orth. 621, Phibalosoma cantori Westw. Hope Dept. Entom. ; Tirachoidea cantori (Westwood) Lectotype ♂ det. P.D. Brock, 1995 (OXUM, Nr. 621); PLT, ♀, Cantor, Malacca; Type, Westwood. Phibalosoma cantor i, Type Orth. 621, Phibalosoma cantori Westw. Hope Dept. Entom. ; Tirachoidea cantori (Westwood) ♀ Paralectotype P. D. Brock, 1995 (OXUM, Nr. 621).

Wood-Mason, 1877a: 161

Wood-Mason, 1877b: 347.

Tirachoidea cantori, Brunner v. Wattenwyl, 1893: 83.

Kirby, 1904a: 359.

Karny, 1923: 240.

Rehn, 1904: 70.

Brock, 1995: 95. [Designation of LT]

Tay & Seow-Choen, 1996: 186.

Pharnacia cantori, Redtenbacher, 1908: 480 (in part—only ♀, → ♂ from Java = Tirachoidea biceps Redtenbacher ). Brock, 1996: 27.

Brock, 1999: 132 (in part—only description of ♂, Figs. 88a–d (♂ and ♀ )).

Seow-Choen, 2000: 35, pl. 94 (♂, ♀).

Otte & Brock, 2005: 264.

[Not: Pharnacia cantori?, Bragg, 2001: 398 . Bornean specimens are Tirachoidea herberti spec. nov., see below]

Further material: [12 ♀♀, 26 ♂♂, eggs]:

PENINSULAR MALAYSIA:

3 ♀♀: W-Malaysia, Perak, Tapah Hills , ca. 600 m, via M.K.P. Yeh VIII. 1994 (coll. FH 0038-1 bis 3) ; 22 eggs: ex Zucht F. Hennemann 1993, urspr.: W-Malaysia (coll. FH 0038-ED) ; 1 ♂: Cameron Highlands , 2000ft., 19.6.1972, C.C.Chua; BM 1975-604 ( BMNH) ; 2 ♀♀: Perak, Rothschild , Bequest, BM 1939-1 ( BMNH) ; 2 ♀♀: Malaisie, Perak, Cameron Highl. , 1200–1600 m, leg. Pfanner ( MHNG) ; 1 ♀: Zoo Frankfurt , Zucht 18.12.84 ( SMFM) ; 1 ♂: Malaysia, Cameron Highlands , leg. A. E. Selner, Augsburg 41990 ( ZSMC) ; 1 ♂, 2 ♀♀, eggs: Captive reared by M. Herbert, culture stock from West Malaysia (coll. MH) .

S-THAILAND:

1 ♂: Peninsular Siam, Nakon Sri Tamorat, Khao Ram 1200–3000ft., 3. 1922 H. M. Pendlebury; Nr. 717 ( RMNH) .

S-MYANMAR:

2 ♂♂: S-Myanmar, Mergui District , Tenasserim, 20.III.1994 , leg. Lehmann (coll. FH, 0038-4 & 5); 6 ♂♂: S-Myanmar, Mergui District , Tenasserim, VI.1996 , leg. Lehmann (coll. FH, 0038-6 to 11); 14 ♂♂: S-Myanmar, Tenasserim, leg. Lehmann VI.1996 (coll. OC); 1 ♀: Tenasserim, Mekane, 90 km, E. of Móulmein , 200m, 2.-8.XI.1934 ( RMNH) .

NO DATA:

1 ♀: without data ( BMNH) .

Diagnosis: Closely related to T. siamensis spec. nov. but distinguished by: the relatively shorter mesothorax; less globose vertex and black ventral surface of the antennae of both sexes; broader body (maximum width 15.0–18.0 mm); deeper and more triangular posteromedian excavation of the anal segment (Fig. 213); smaller

133. Tirachoidea cantori ( Westwood, 1859) , ♀ West Malaysia, Perak, Tapah Hills (coll. FH, No. 0038-2) 134. Tirachoidea cantori ( Westwood, 1859) , ♂ S-Myanmar, Mergui District , Tenasserim (coll. FH, No. 0038-6) 135. Tirachoidea herberti spec. nov., ♀ HT, Brunei, Badas ( BMNH)

136. Tirachoidea herberti spec. nov., ♂ PT, Brunei, Badas (coll. MH)

137. Tirachoidea biceps ( Redtenbacher, 1908) , ♀ Java, captive reared (coll. FH, No. 0193-6)

138. Tirachoidea biceps ( Redtenbacher, 1908) , ♂ Java, captive reared (coll. FH, No. 0193-8)

139. Tirachoidea siamensis spec. nov., ♂ PT, N-Thailand, Dansai Loei (coll. FH, No. 0142-2)

140. Tirachoidea westwoodii ( Wood-Mason, 1875) , ♂ N-Myanmar, Highland of Tenasserim (coll. FH, No. 0211-1) 141. Tirachoidea inversa (Brunner v. Wattenwyl, 1907), ♀ Java (HT of Ph. semilunaris Redtenbacher, NHMW )

and roundly triangular posterolateral lobes of abdominal tergite VII (Fig. 272); less distinct praeopercular organ; smooth abdominal tergites V–VII and dull green armature of the legs of ♀♀; as well as the ventro-apically elevated or slightly hook-like semi-tergites of the anal segment (Fig. 214); shorter meso- and metabasitarsi and more regular and dense serrations of the legs of ♂♂. The eggs are easily distinguished from those of T. siamensis spec. nov. by the creamish brown, almost spherical egg capsule (Figs. 157–158). In addition to these morphological features, T. cantori is distributed more southward than siamensis spec. nov. which is restricted to the northern and central portions of Thailand.

Also related to T. biceps ( Redtenbacher, 1908) but at once distinguished by lacking the prominent conical elevation of the vertex, relatively shorter mesothorax and more robust body and legs both sexes. ♀♀ furthermore differ by the relatively shorter mesothorax, broader and rounded posterolateral angles of the anal segment and more prominent ventral armature of the meso- and metafemora, ♂♂ by the broader semi-tergites of the anal segment, less acute central spine of the poculum and white or pale yellow instead of apple green anterior margin of the alae. The eggs are considerably larger than those of T. biceps and have the micropylar plate relatively longer and more slender.

Etymology: Dedicated to Dr. Theodore Cantor, member of the East India Company, who first collected this species in 1840.

Description: ♀♀ ( Fig. 133): Large to very large (body length 190.0– 246.5 mm), very robust species (maximum body width 15.0–18.0 mm). All legs of moderate length, robust and heavily spinose. General colouration of body either dull green with numerous darker speckles or greyish, greenish or reddish brown with irregular darker and paler speckles, often with washed whitish areas on abdomen. Back of head with four ± decided dark brown longitudinal lines and a shallow dark postocular line. Abdominal tergites with 1–3 irregular pairs of slightly darker patches in posterior portion. Ventral surface of abdomen with ± distinct white mottling. Mesonotum with a longitudinal row or small dark brown speckles on a whitish background along lateral margins. Meso- and metasternum dark green or brown and set with a few small, oval yellow spots. Legs dull green, brown or slightly aubergine. All carinae of mid and hind legs and entire spination green with black points (may be yellowish green on tibiae). Antennae brown with ventral surfaces of all antennomeres except scapus and pedicellus black. Eyes reddish brown.

Head: About 1.8x longer than wide, oval, vertex roundedly convex and with a pair of ± distinct tubercles or small, blunt spines (the sinistral one smaller). Between the bases of antennae with a deep, transverse impression and between the eyes a larger, slightly raised and smooth region. Eyes rather large, higher than long with the anterior margin triangularly elevated; their length contained length contained about 3.5x in that of cheek. Antennae reaching to posterior margin of mesonotum, segments first distinctly increasing then decreasing in length towards apices of antennae; consisting of about 37 antennomeres. Scapus dorsoventrally flattened, 2.7x longer than wide and parallel-sided, ventrally with a longitudinal median furrow. Pedicellus about 1/3 the length of scapus, cylindrical and indistinctly longer than wide. III longer than pedicellus, IV strongly shortened.

Thorax: Pronotum distinctly narrower and shorter than head, trapezoidal, and widening towards posterior margin. Anterior margin slightly raised and with a distinct pit at each outer lateral angle. Median transverse depression gently curved and just not reaching lateral margins of segment. Mesothorax about 1.8x longer than head and pronotum combined, slightly constricted anteriorly and conspicuously broadened at posterior margin. Mesosternum smooth except for a very indistinct longitudinal carina along lateral margins and two parallel longitudinal median ridges which terminate shortly before the apex. Metanotum distinctly shorter than mesonotum, about 1.7x longer than wide and slightly constricted medially. Metasternum smooth. Tegmina and alae present as very small (ca. 1.0 mm), sclerotised scale-like structures.

Abdomen: Median segment a little shorter than metanotum, slightly trapezoidal with anterior margin rounded and about 1.4x longer than wide. Tergite II about as long as median segment, as long as wide and slightly broadening towards the posterior. III–VI slightly increasing in length, III indistinctly longer than wide, VI about 2.2x longer than wide. VII slightly shorter than VI, lateral margins gradually dilated and forming a roundly triangular lobe posteriorly, which projects by about 1/3 the width of the segment (Fig. 272). Sternites II–VI smooth. Praeopercular organ formed by a pair of very shallow and low, converging ridges close to posterior margin of sternum VII. Tergite VIII about as long as VI and longer than XI and X combined , distinctly narrower than previous, gently constricted medially, strongly convex and about 3x longer than wide. IX strongly convex, indistinctly longer than wide and about 1/3 the length of VIII. Anal segment longer than IX, with a distinct median carina and a broad and deep, more or less triangular median excavation, outer angles forming a narrow, rounded lobe (Fig. 213). Supraanal plate short, rounded and with a distinct longitudinal keel. Cerci short, oval in cross-section, gently up-curving and tapered at the apex. Gonapophyses elongate, up-curving and slightly projecting over apex of anal segment. Subgenital plate elongate, strongly keeled, boat-shaped and slightly projecting over anal segment (Fig. 212).

Legs: All of moderate length, very robust and heavily spinose, basal half of meso- and metafemora distinctly swollen with the two outer ventral carinae gently elevated. Profemora about as long as pro- and mesonotum combined, mesofemora about as long as mesothorax, metafemora reaching about one third the way along abdominal segment V and hind tarsi just not reaching apex of abdomen. Anterodorsal carina of profemora armed with 15–18 prominent and pointed serrations. Posterodorsal carina with 6–7 acute triangular teeth. Posteroventral carina armed with 9–12 pointed spines. Anterodorsal carinae of protibiae with low but regular, wide-spaced serrations, posteroventral carina with ± 20 rather small and low serrations. The posterodorsal carina occasionally with an triangular, tooth-like lobe just before the middle. Dorsal carinae of meso- and metafemora sparingly dentate, each usually with a ± enlarged triangular tooth or lobe 1/3 off the base. Two outer ventral carinae with 10–15 very prominent, long and pointed spines which decrease in size towards the apex. Medioventral carina becoming increasingly acute but rounded towards the base and armed with 6–8 very prominent, slightly hooked spines; the three basal ones largest. Dorsal carinae of meso- and metatibiae sparingly but acutely dentate. Posterodorsal carina occasionally with a ± prominent triangular lobe just before the middle and forming a rounded, acutely dentate lobe at the apex. Ventral carinae densely armed with prominent, slender and pointed spines, those of the median carina largest. Probasitarsus with all carinae slightly elevated and about as long as remaining segments combined. Meso- and metabasitarsi with all carinae bearing 2–4 small teeth and the dorsal carina gently rounded; about as long as following three tarsomeres combined.

♂♂ ( Fig. 134): Of moderate size (body length 119.0– 152.5 mm) and rather robust for the genus with very long alae (77.5–83.0 mm). General colouration of body and legs yellowish, greenish or greyish dark brown, often with a few small white spots on meso- and metanotum. Mesonotum usually with a slight greyish wash. Mesopleurae with a fine longitudinal orange line along upper margin. Meso- and metasternum dark brown, abdominal sternites I–VIII with ± distinct white mottling and a sepia brown transverse streak posteriorly. Abdominal tergite usually with an irregular white lateral marking. Tegmina and costal region of alae either plain yellowish, greenish or greyish brown, anterior margin of tegmina and basal quarter of anterior margin of alae with a bold, longitudinal whitish line. Anal region of alae transparent greyish brown with sepia brown veins. Outer ventral carinae of meso- and metafemora dull green, spination of legs dark orange with black points. Cheeks with a shallow dark brown postocular streak. Eyes reddish brown. Antennae mid to dark brown dorsally and except for scapus and pedicellus black ventrally.

Head: Generally as in ♀♀, but eyes much more prominent and projecting hemispherically; their length contained no more than 2.5x in that of cheek. Antennae projecting over posterior margin of tergite II, otherwise as in ♀♀.

Thorax: Pronotum distinctly narrower and shorter than head, gently widened towards the posterior, otherwise as in ♀♀ but relatively longer; 1.6x longer than wide. Mesothorax about 2x longer than head and pronotum combined and slightly broadened at posterior margin. Mesonotum parallel-sided and with a very faint longitudinal median line. Mesosternum with a distinct longitudinal median carina which terminates slightly before posterior margin. Metasternum just with a very faint longitudinal median line. Metanotum slightly longer than wide, considerably broader than mesonotum. Tegmina oval, narrowed towards the base and with a prominent and rather acute, conical central hump; reaching to posterior margin of metanotum. Alae varying in length, reaching at least half way along abdominal tergite VI and at best half way along tergite VII.

Abdomen: Median segment almost 3x longer than wide and 2x longer than metanotum, slightly gradually narrowing towards the posterior. Segments II–IV increasing in length,V–VII decreasing, IV longest and VII shortest segment. II 3.5x, IV more than 5x and VII only about 2.5x longer than wide, parallel-sided. Tergite VII slightly constricted medially and with a ± prominent, rounded posterolateral lobe. VIII slightly longer than VII, strongly swollen and broadening towards the posterior; almost 2x the width of II–VII. IX a little longer than VIII, almost 3x longer than wide and decidedly constricted medially. Anal segment shorter than IX, laterally compressed and strongly tectiform. Semi-tergites rather broad, slightly tapered with the apex rounded and ± decidedly truncate or hook-like ventrally (Fig. 214). Interior surfaces densely covered with minute black teeth in the apical portion. Cerci small but obtuse, sub-cylindrical and slightly in-curving. Poculum strongly convex, cup-like, posteriorly marginated and with a small, ± acute backward pointing central spine (Fig. 214).

Legs: All of moderate length and rather robust. Profemora slightly longer than head, pro- and mesonotum combined, mesofemora about as long as pro- and mesonotum combined, metafemora slightly projecting over posterior margin of abdominal tergite IV and hind legs slightly projecting over tip of abdomen. Armature similar as in ♀♀, but considerably less prominent but with the number of spines larger. Spines rather uniform in size on each carina. The meso- and metafemora in particular are much less heavily spinose and not considerably broadened basally. Posterodorsal carina of profemora only with 4–6 very minute teeth. The posterodorsal and posteroventral carina of the mesofemora may bear a few slightly enlarged teeth in the basal portion. Occasionally there may be abroad triangular pre-median lobe on the posterodorsal carina of the mesotibiae. Probasitarsus as long as remaining tarsomeres combined except claw, all carinae slightly elevated. Meso- and metabasitarsi with all carinae minutely dentate and the dorsal carina decidedly rounded; slightly longer than following three tarsomeres combined.

Eggs (Figs. 157–158): Large, capsule almost spherical. General colouration of capsule creamish pale brown, the lateral and ventral surfaces slightly darker. Micropylar plate pale greyish brown, operculum black, capitulum dull orange. Lateroventrally with a faint, slightly darker marking and slightly darker areas around micropylar plate. Capsule surface strongly shiny and very unevenly granulose (70x). Micropylar plate shaped like an inverted “Y” with outer margin dark brown and covering more than ¾ of capsule length. Apices of all extensions rounded. Micropylar cup placed in posteromedial gap of plate; small, cup-like and with a median carina. Median line slightly raised and reaching to apices of posterior extensions of plate. Polar-area with a circular dark brown spot and a very minute, blunt granule in the centre. Operculum slightly oval, convex and impressed in centre. Capitulum distinct, conical and on an obvious black stalk.

Measurements [mm]: Length including capitulum 5.0–5.2, length 4.6–4.8, width 4.1–4.2, height 4.0–4.1, length of micropylar plate 3.8–3.9.

Comments: Westwood (1859: 74) originally described Phibalosoma cantori from a ♂ and ♀ in OXUM collected in Peninsular Malaysia and provided nice figures of both sexes ( Westwood, 1859: plate 37 (♂), 38 (♀ )). Brock (1995: 95) designated the ♂ in OXUM as the lectotype. Brunner v. Wattenwyl (1893: 83) included Westwood´s taxon and three further species in his new genus Tirachoidea for which subsequently Kirby (1904: 359) designated Phibalosoma cantori Westwood as the type species. Redtenbacher (1908) re-transferred these three species to Pharnacia Stål, 1877 and erroneously synonymised the two genera (→ see: comments on Pharnacia Stål and Tirachoidea Brunner v. Wattenwyl).

Examination of Westwood’s type specimens in OXUM clearly showed the ♀ PLT to exhibit a pair of granules on the vertex, which are not indicated in the illustration neither mentioned in the original description by Westwood, a fact which has caused confusion about the status of T. cantori (Westwood) and the genus Tirachoidea Brunner v. Wattenwyl. This error of Westwood has however already been pointed out by Wood- Mason (1877a: 347) who stated “I have examined with Professor Westwood the typical specimens of P. Cantori preserved in the Hopeian collection at Oxford and find that the insects have been legitimately united by their describer; the female having small equal blunt representatives of the conspicous cephalic tubercles seen in the male; these tubercles not having been represented by Professor Westwood in his figure of the former.”.

In his book on Malaysian Phasmatodea Brock (1999: 132 , fig. 88) reproduced the illustrations of both sexes of Tirachoidea cantori (Westwood) of Westwood (1859, pl. 37: 1, 38: 1). The photographs of the ♀ ( Brock, 1999, pl. 26) and egg (pl. 24) stated to be this species however show Pharnacia sumatranus (Brunner v. Wattenwyl). Brock (1999: 133) stated there to be a “second type of egg” which he briefly characterized as “[…] glossy whitish brown with a black operculum,…”. These are in fact the egg of Tirachoidea cantori (Westwood) . Seow-Choen (2000: 35) provided illustrations and characterizations for both species as well as a list of features which distinguish the insects and eggs. Seow-Choen (2000: 36) cited the following bodylengths for T. cantori : ♂♂ 130.0–145.0 mm, ♀♀ 190.0–230.0 mm.

Examination of the ♂ from Java which Redtenbacher (1908: 480) recorded from SMNS has shown this to be a typical specimen of Tirachoidea biceps ( Redtenbacher, 1908) . Therefore, “ Java ” can be regarded an erroneous record of T. cantori . The specimens which Bragg (2001: 398) listed from Borneo are not conspecific with T. cantori (Westwood) but all represent Tirachoidea herberti spec. nov., which is the only known representative of Tirachoidea Brunner v. Wattenwyl in Borneo (see comments on this species).

* according to Seow-Choen (2000: 36)

T. cantori used to be successfully cultured in Europe during the 1980’s from stock originating in the Tapah Hills (Peninsular Malaysia). It was included on the Phasmid Study Group species list as culture No. 58 “ Pharnacia sp. ”. In captivity in Europe the insects readily accepted bramble ( Rubus spp. , Rosaceae ) and oak ( Quercus spp. , Fagaceae ) as alternative foodplants. Seow-Choen (2000: 36) listed Mangifera indica (Anacardiaceaee) , Psidium guajava (Myrtaceae) , Rubus fruticosus and Rubus moluccanus (Rosaceae) to be natural foodplants of T. cantori in Peninsular Malaysia.

Distribution (Fig. 401): Peninsular Malaysia (Perak: Tapah Hills & Cameron Highlands), South Thailand Peninsular (Nakon Sri Tamorat, Khao Ram 1200–3000ft.) and South Myanmar (Mergui District: Tenasserim & Mekane, 90 km east of Móulmein, 200 m).

Number of specimens examined: 40