Phryganistria grandis Rehn, 1906

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 : 204-209

Phryganistria grandis Rehn, 1906

( Figs. 126, 186–187, 280–281, 289, 397)

Phryganistria grandis Rehn, 1906: 279 , figs. 1 & 2 (♂). HT, ♂: Tonkin, Indo-China ; Phryganistria grandis Rehn , Type (ANSP).

Otte, 1978: 78.

Hennemann et al., 2008: 24.

Phobaeticus grandis, Günther, 1935b: 124 .

Brock, 1996: 28.

Otte & Brock, 2005: 268.

Phibalosoma acanthopus, Wood-Mason, 1877a: 161 .

Wood-Mason, 1877b: 346.

Phobaeticus fruhstorferi Brunner v. Wattenwyl, 1907: 184 (in part – only PLT’s in NHMW). PLT, ♂: Tonkin, Montes Mahokon, April. Mai, 2–3000m, H. Fruhstorfer; Collectio Br. v. W.; det. Br. v. W. Phobaeticus fruhstorferi ; 24.418 (NHMW, No. 307); PLT, 2 ♂♂: Central-Tonkin, Chiem-Hoa, Aug. Sept., H. Fruhstorfer; Collectio Br. v. W.; det. Br. v. W. Phobaeticus fruhstorferi (NHMW, No. 307); PLT, ♀ (penultimate instar nymph): Tonkin, Than-Moi, H. Fruhstorfer; Collectio Br. v. W.; 24.419, det. Br. v. W. Phobaeticus fruhstorferi (NHMW, No. 307). [Not: LT, ♀: Mt. Cariani, D. Tornatore 1898, Phobaeticus fruhstorferi Br. , Type, Birma (MCSN)— erroneous synonym of Günther, 1935b: 124, see comments below]

Brock, 1996: 28.

Phobaeticus frühstorferi, Vanschuytbroeck & Cools, 1981: 16 .

? Phobaeticus fruhstorferi, Chen, 1993: 87 . [Record from South China (Guangxi), see comments below]

Further material: [22 ºº, 21 ♂♂, 1 egg]:

VIETNAM:

1 ♂, 1 ♀: Tonkin, Chiem-Hoa , leg. Fruhstorfer VIII–IX. ( SMNS) ; 1 ♂: Tonkin, Than Moi , leg. Fruhstorfer ( ZMUH) ; 1 ♀: Tonkin, Chiem Hoa , leg. Fruhstorfer 15.IV. 1902 ( ZMUH) ; 2 ♂♂, 2 ♀♀: Tonkin, Chiem-Hoa , Aug. Sept., leg. H. Fruhstorfer ( MHNG) ; 1 ♂: Tonkin, Than-Moi , Juni-Juli, H. Fruhstorfer ; 1 ♂: Tonkin , coll. Le Moult ; 1 ♀: Tonkin, Chiem Hoa , Aug. – Sept., H. Fruhstorfer ( ISNB) ; 3 ♂♂, 2 ♀♀: Tonkin , Juni – Juli, H. Fruhstorfer ; 1 ♀: Tonkin, Chiem Hoa , Aug. – Sept., H. Fruhstorfer ( RMNH) ; 1 ♂: Central-Tonkin, Chiem-Hoa, Aug. Spet. , leg. H. Fruhstorfer, L. 845 ( SMTD) ; 1 ♀: Tonkin , Fruhstorfer ( SMTD) ; 1 ♂: Tonkin , Than Moi, Fruhstorfer ( MNHU) ; 1 ♂: Tonkin , det. Willemse ( MNHU) ; 1 ♂, 1 ♀: Central-Tonkin, Chiem-Hoa , Aug. / Sept., H. Fruhstorfer ( ETHZ) ; 1 ♂: 27, Central-Tonkin, Chiem-Hoa , Aug. Sept., H. Fruhstorfer ( SMFM) ; 1 ♀ + 1 egg (ex ovipositor), 1 ♂: Cochinchina, Mt. Bavi , 1935, S. Massey eff., BM 1937-658 ( BMNH) ; 1 ♀: Cochinchina , Tuyen Quang, 1936, S. Massey eff., BM 1937-658 ( BMNH) ; 1 ♂: Tonkin , 1935-448, Phryganistria sarmentosa BMNH (BMNH) ; 1 ♀: Tonkin , 1935-448, Phryganistria fruhstorferi (BMNH) ; 1 ♀: Vietnam : 55 km NNW Tam Dao, 26.VII.–10.VIII.1998, A. Napolov, BMNH (E) 2005-98, Phobaeticus grandis det. P. Brock 2005 ( BMNH) ; 1 ♂: Museum Paris, Haut-Tonkin et Bas-Yunnan , Entre Mau-Hoa , Huong-Num (Prés Lao-Kay) et Bau-Hau- Khun Lieut, Lesourt 1905 ( MNHN) ; 1 ♂: Museum Paris , Tonkin Hajiank, H. Donchier 1924 ( MNHN) ; 1 ♀: Museum Paris , Tonkin N., Reg. Ma-Giang, Siebens Olivier 1916 ( MNHN) ; 1 ♀: Museum Paris, Haut-Tonkin, Reg. De Bac Ken , Ma-Giang Guam-Ba et Yen Minh, F. de Broissia 1918 ( MNHN) ; 1 ♂, 1 ♀: Tonkin ( ZSMC) ; 1 ♂: Tonkin, Than Moi , Juni – Juli, H. Fruhstorfer ( ZSMC) ; 1 ♀: Tonkin, Chiem Hoa , Aug. – Sept., H. Fruhstorfer ( ZSMC) ; 1 ♂, 1 ♀: Central Tonkin , Chiem Hoa, Aug. Sept. H. Fruhstorfer ( MNCN) .

MYANMAR:

1 ♀: Upper Burma , Nam Tamai Valley, 22.9.1938, R. Kaulbach leg., BM 1938-741, att. 3000ft., Lat. N 27°42`, Long 97°54`( BMNH) .

NO DATA:

1 ♂: no data ( MNHU) ; 1 ♂, 1 ♀: no data ( ZSMC) ; 1 ♀: no data ( SMFM) ; 1 ♀: no data ( BMNH) .

Diagnosis: Very closely related and certainly the adelphotaxon of Ph. heusii (Hennemann & Conle, 1997) , with which it is sympatric in Northern Vietnam. It however frequently differs by: the more slender body; relatively longer and more slender legs and tarsi as well as the slightly more globose head of both sexes. ♂♂ furthermore differ by: the plain brown profemora which have the posteroventral carina unarmed; relatively smaller eyes; more narrow apices of the semi-tergites of the anal segment and longer, more slender cerci which almost reach to the apex of the anal segment (Fig. 281). ♀♀ may be distinguished by: the lack of a posteromedian tubercle on abdominal tergites II–VI; longer and more slender lobe-like spines of the praeopercular organ (Fig. 280) and much deeper posteromedian excavation of the anal segment (Fig. 289). Eggs differ from those of Ph. heusii by the smaller dimensions, slightly more ovoid capsule and relatively smaller micropylar plate (Figs. 186–187).

Etymology: “ Grandis ” (lat. = large) refers to the large size of ♂♂.

Description: ♀♀: Long to very long (body length 234.5–253.0 mm) and slender species (maximum body width 8.0–9.0 mm) with the body surface slightly glossy. General colouration of body and legs plain mid to dark brown, ochracheous, brownish green or dull green, the abdomen in particular furnished with several small blackish speckles. Armature of legs dark greyish green with black points. Antennae dark brown.

Head: Globose, oval in dorsal aspect, roughly 1.4x longer than wide and vertex gently convex. Coronal line distinct, impressed and on each side accompanied by a pair of faint dark brown, longitudinal lateral lines. Posterior margin of head capsule with three short, longitudinal impressions. Between the bases of antennae with an oval, slightly transverse depression followd by two slightly raised, mid brown areas and a pair of very faint impressions. Eyes rather small, circular and moderately convex; their length contained about 3.3x in that of cheek. Antennae reaching to posterior margin of mesonotum, with the antennomeres increasing in length towards apex of antennae. Scapus dorsoventrally compressed, rectangular, almost 3x longer than wide. Pedicellus slightly spherical and about 1/3 the length of scapus.

Thorax: Pronotum shorter and narrower than head, 1.6x longer than wide, posterior margin slightly broader than anterior margin. Anterior margin arcuate and followd by a transverse furrow. Anterior half with a faint trace of an impressed longitudinal median line. Median transverse depression distinct, slightly curved but not reaching lateral margins of segment. Mesothorax almost 3x longer than head and pronotum combined. Mesonotum parallel-sided except for being slightly widened posteriorly. Metanotum parallel-sided, about 4x longer than wide and 3/5 the length of mesonotum. Pleurae and sterna unarmed.

Abdomen: Median segment indistinctly longer than wide, roughly ¼ the length of metanotum and slightly trapezoidal, with posterior margin broader than anterior margin. Segments II more than 2x longer than median segment, II–VI increasing in length, II 2x, VI more than 3x longer than wide, all parallel-sided and of uniform width. Tergite VII about ¾ the length of VI with lateral margins elevated to form a slight, rounded lobe posteriorly. Praeopercular organ formed by a pair of very prominent, long, carinate, back-curving, lobe-like spines at posterior margin of sternum VII. Tergite VIII half the length of VII, constricted anteriorly and 2x longer than wide. IX indistinctly longer than wide, slightly more than half the length of VIII, parallel-sided. Anal segment slightly longer than IX, with a very faint, longitudinal median carina and a deep, narrow triangular excavation posteromedially; posterolateral angles triangular, slightly in-curving and swollen (Fig. 289). Supraanal plate very small, rounded and strongly keeled dorsally. Cerci small, sub-oval in cross-section and tapered towards a narrow apex; finely bristled. Subgenital plate moderately convex, strongly keeled, boatshaped and projecting over apex of abdomen by at least 1/3 the length of anal segment (Fig. 280).

Legs: All long and moderately slender. Profemora almost as long as head, pro- and mesonotum combined, mesofemora slightly shorter than mesothorax, metafemora reaching about half way along abdominal segment V and metatibiae slightly projecting over apex of abdomen. Anterodorsal carina of profemora armed with 11– 17 prominent triangular serrations which decrease in size towards apex of femur, posterodorsal carina set with 5–9 minute teeth. Posteroventral carina with 13–20 small to medium-sized but acute, triangular teeth. Medioventral carina of protibiae sparingly and rather minutely dentate, remaining carinae unarmed. Two outer ventral carinae meso- and metafemora armed with 12–20 triangular teeth of average size. Dorsal carinae each with only 2–6 teeth, the basal one of the posterior carina sometimes conspicuously enlarged and triangular. Medioventral carina armed with a longitudinal row of 6–12 minute spines. All carinae of meso- and metatibiae serrate, although considerably more sparsely on dorsal carinae. Posterodorsal carina often with a ± prominently enlarged, triangular tooth pre-medially and gently rounded apically. Probasitarsus a little longer than remaining tarsomeres combined, dorsal carina very Anacardiaceae raised, all carinae unarmed. Meso- and metabasitarsi as long as following three tarsomeres combined, dorsal carina gradually raised towards the apex, ventral carinae minutely dentate.

♂♂ ( Fig. 126): Long to very long (body length 171.5–232.0 mm) and moderately slender species (maximum body width 7.0– 8.5 mm), with body surface smooth and glabrous. General colouration of body and legs, ferruginous, reddish or yellowish pale to mid brown. Greater portion of the mesonotum except anterior and posterior margin as well as the anterior ¾ of the metanotum covered by a washed malachite green, bluish green or greyish marking; may be very faint in some specimens. Two outer ventral carinae of meso- and metafemora dull green. Complete armature of legs tipped with black. Eyes reddish brown. Antennae dark reddish brown to dark brown.

Head: Generally as in ♀♀ but slightly more decidedly narrowed posteriorly. Posterior margin with four faint impressions, which anteriorly continue as four sub-parallel, slightly impressed longitudinal lines. Eyes rather small, circular and projecting hemispherically; their length contained about 3x in that of cheek. Antennae projecting over posterior margin of median segment, otherwise as in ♀♀.

Thorax: Pronotum slightly shorter but distinctly narrower than head, 1.5x longer than wide, anterior portion narrowed, otherwise as in ♀♀. Mesothorax 3.6x longer than head and pronotum combined and slightly swollen posteriorly, mesonotum parallel-sided except for being slightly broadened at posterior margin. Metanotum less than 2/3 the length of mesonotum and slightly constricted medially, more than 7x longer than wide. Pleurae and sterna smooth.

Abdomen: Median segment about 1/4 the length of metanotum, 1.8x longer than wide and slightly trapezoidal being gently widened towards the posterior. Segment II about 1.7x longer than median segment, II–VII slightly constricted medially. III–IV subequal in length and slightly longer than II, almost 6x longer than wide. V–VI subequal and slightly longer than III–IV, almost 6.5x longer than wide. VII about 2/3 the length of VI and broadened posteriorly. Tergite VIII half the length of VII, strongly convex and gradually widening towards posterior margin. IX strongly convex and slightly shorter than VIII, anterior margin distinctly broader than posterior margin. Anal segment laterally compressed, strongly tectiform and carinate dorsally. Semi-tergites roughly triangular, very broad anteriorly and strongly tapered in apical portion, the apices narrow and rounded (Fig. 281). Interior surfaces apically swollen and covered with several minute, black and back-curving teeth. Cerci elongate, sub-oval in cross-section, gradually tapered towards apex and roughly reaching apex of anal segment. Poculum strongly convex with a distinct transverse carina and a blunt rectangulate extension anteriorly; reaching about 2/3 the way along tergite IX.

Legs: All very long and slender, meso- and metafemora robust, conspicuously swollen and considerably broader than corresponding tibiae. Profemora longer than combined length of head, pro- and mesonotum, mesofemora a little longer than mesothorax, metafemora almost reaching posterior margin of abdominal segment V and metatibiae projecting considerably over apex of abdomen. Anterodorsal carina of profemora with 14–20 small triangular serrations, posterodorsal and posteroventral carinae unarmed. All carinae of protibiae unarmed. Two outer ventral carinae of meso- and metafemora armed with 12–20 moderately sized, triangular teeth, each sub-apically with a prominent, acutely dagger-like spine. Medioventral carina with a longitudinal row of 5–10 very minute spines. Dorsal carinae smooth except for 1–3 minute teeth. Ventral carinae of meso- and metatibiae densely, dorsal carinae rather more sparsely dentate. Probasitarsus very elongate and slender, about 1.5x longer than remaining tarsomeres combined, all carinae smooth. Mesobasitarsi about as long as, metabasitarsi a little longer than remaining tarsomeres combined; ventral carinae with a few minute teeth in basal half.

Eggs (Fig. 188–189): One fully developed egg was removed from the ovipositor of the ♀ from Cochinchina ( Mount Bavi ) in BMNH (accession No. BM 1937-658) and served for the following description .

Medium-sized, capsule oval in lateral aspect and slightly laterally compressed, oval in cross-section. Capsule with a very faint keel on dorsal and ventral surfaces and polar-area, beginning and ending at operculum. Capsule surface slightly glossy and minutely granulated (70x). General colouration of capsule and micropylar plate creamish dark brown, operculum black. Micropylar plate elongate and covering about 2/3 the length of capsule, tapered towards anterior end and with a wide gap posteromedially. Outer margin of plate black. Micropylar cup placed in the posterior gap of plate, small and dark reddish brown. Median line on a distinctly raised hump, almost reaching polar-area. Operculum oval and slightly convex. Capitulum large, dark brown, irregularly conical and with several large lateral impressions. Stalk obvious, whitish.

Measurements [mm]: length (incl. capitulum) 5.4, length 4.4, width 2.9, height 3.6, length of micropylar plate 2.7.

Comments: Brunner v. Wattenwyl (1907: 184) appears to have not been aware of Rehn’s 1906 publication, described Phobaeticus fruhstorferi from a series of specimens from N-Vietnam and Myanmar in NHMW and MCSN and did furthermore not recognize that the type-series consisted of two distinct species (see comments on Phryganistria fruhstorferi ). The ♀ from Birma (= Myanmar) in MCSN is clearly distinct from the remaining types from N-Vietnam in NHMW and here selected as the lectotype of Ph. fruhstorferi Brunner v. Wattenwyl, 1907. All paralectotypes (3 ♂♂, 1 ♀ penultimate instar nymph) in NHMW clearly represent Phryganistria grandis Rehn, 1906 . Günther (1935b: 124) had obviously not seen the specimen in MCSN and synonymised both species. However, as Günther (1935b) did not designate a lectotype his synonymization is invalid, and Ph. fruhstorferi Brunner v. Wattenwyl here treated as a valid species with the MCSN specimen selected as the LT.

Wood-Mason (1877a: 161) recorded a ♀ of what he believed to be Phibalosoma acanthopus Burmeister, 1838 (= Phobaeticus serratipes ( Gray, 1835)) from Moolai to Moolat in Upper Tenasserim, Northern Myanmar. Already Redtenbacher (1908: 453) doubted the identity of Wood-Mason’s specimen and designated it as a syntype of his newly described Pharnacia ingens . The brief diagnosis provided by Wood-Mason however clearly shows this ♀ to represent a distinct species and to be even generically different from the LT of Ph. ingens from the Malabar Coast of SW-India. The prominent, bifid praeopercular organ, short median segment, locality and body length which Wood-Mason stated to be 260.0 mm obviously place it in Phryganistria Stål, 1875 .

* including median segment

** according to Chen (1993: 87)

Chen (1993: 87) recorded specimens, which he believed to be Ph. fruhstorferi (Brunner v. Wattenwyl, 1907), from the Guangxi autonomous region in South China close to the border to Northern Vietnam and provided a brief description of the ♀♀ along with a set of measurements. Due to having a relatively longer median segment compared to both, Ph. grandis Rehn and Ph. fruhstorferi Brunner v. Wattenwyl ( Tables 31– 32) the exact identity of these Chinese specimens deserves confirmation. Although they are shorter than specimens of Ph. grandis from N-Vietnam (body length 207.0–214.0 mm) the distribution rather associates them with Ph. grandis or perhaps Ph. guangxiensis Chen & He, 2007 rather than Ph. fruhstorferi . Unfortunately, the specimens could not be examined for this study.

Almost every European museum contains specimens of Ph. grandis Rehn , most of which were collected by H. Fruhstorfer in Vietnam during the late 19 th century. According to the large number of specimens collected in only a short period of time this species appears to be quite abundant in certain areas of Northern and Central Vietnam.

Distribution (Fig. 397): Northern Vietnam (Mount Mahokon 2000–3000 m; Chiem-Hoa; Than Moi; Ma- Giang Region & Mau-Hoa: Huong-Num), Southern Vietnam ( Cochinchina: Mount Bavi & Tuyen Quang) and Northern Myanmar (Upper Tenasserim: Moolai to Moolat 4000–6000 ft. [ Wood-Mason, 1877a: 161] & Nam Tamai Valley 3000 ft.). South China (Guangxi)—with doubt [Chen, 1993: 87].

Number of specimens examined: 48