Pharnacia ponderosa Stål, 1877

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 : 112-117

Pharnacia ponderosa Stål, 1877

( Figs. 89–90, 145–146, 194–196, 269, 290–291, 318, 376, 427, 436–438)

Pharnacia ponderosa Stål, 1877a: 40 . LT, ♂: Ins. Phil., Semper; Pharnacia ponderosa Stål [darker brown specimen] (NHRS); PLT, ♂: Ins. Phil., Semper; Pharnacia ponderosa Stål (NHRS) . Kirby, 1904a: 359. Redtenbacher, 1908: 455. Sjöstedt, 1933: 6. Brock, 1996: 27. Brock, 1999: 181. [Designation of LT] Bresseel, 2004: 15, figs. 3–5. Otte & Brock, 2005: 265. Lit & Eusebio, 2008: 120, figs. 2a, d, 3a & table 1.

Pharnacia magdiwang Lit & Eusebio, 2008: 117 , figs, 2c, 2f, 3a, 4, 5. HT, ♀: ex Mangifera indica, Sibuyan Island, Brangay Ipil, Magdiwang, 31.X.2006, S.R. Rabo (UPPC, No. PHA-00130); AT, ♂: reared from egg of holotype, died 15.IV.2007, UPLB MNH Central Lab. (UPPC, No. PHA-00132); PT, ♂, ♀: same data as AT (UPPC, No’s PHA-00131 & 00133); PT, 10 eggs: laid by HT (UPPC). syn. nov.

[Not: Pharnacia ponderosa, Kirby, 1896: 451 ; Misidentification, = Phasmotaenia sanchezi ( Bolívar, 1897) ]

Further material: [45 ºº, 14 ♂♂, 2 nymphs, eggs]:

PHILIPPINES:

1 ♀, 3 ♂♂, 2 ♀♀ (nymphs), 4 eggs: Philippinen, Marinduque Id. , ex. Zucht F. Hennemann, 1992 (coll. FH, No’s 0192-1 to 6 & 0192-E1) ; 3 ♀♀, 1 egg: Philippinen, Mindoro Island, Mt. Halcon , leg. Mohagan 2.IV.–5.V.1996 (coll. FH, No’s 0192-27 to 29 & 0192-E3) ; 1 ♀: Philippinen, N-Luzon Id., Mountain Province, Nueva Viscaya , Balite , leg. I.O. Lumawig VII.1996 (coll. FH, No. 0192-30) ; 15 ♀♀, 4 ♂♂: Philippinen, Panay Island, Mt. Nangtud , 1500 m, leg. Mohagan VII. 1997 (coll. FH, No’s 0192-7 to 25 & 0192-E2) ; 1 ♀: without data (coll. FH, No. 0192-26); 1 ♂, 1 ♀: ex Zucht; R. Galunder ( Nümbrecht ) III.2008, Philippinen (coll. FH, No’s 0192-31 & 32) ; 1 ♂, 1 ♀: ex Zucht. F. Hennemann 2008, Herkunft: Philippinen (coll. FH, No’s 0192-33 & 34); 5 ♀♀: Philippinen, Panay Island, Mt. Nangtud , 1500 m, leg. Mohagan VII.1997 (coll. OC) ; 1 ♀: Philippinen, Mindoro, Mt. Halcon , leg. Mohagan 15.–26.V.1996 (coll. OC) ; 1 ♀: Philippines, Luzon , from UP Los Baños, without data, X. 1995 (coll. OZ, No. 0084-2) ; 1 ♀: Philippines, Sibuyan Id. , Romblon Prov., Magdiwang , Pawala River , Tampayan , Camp New St. Gallen area , 20.–30.VII.1986, 100– 400 m, Roland A. Müller leg. (coll. OZ, No. 0084-3) ; 1 ♀ (nymph): Philippines, Cebu Id., Minglanilla , Camp 7, 300 m, 21./ 22.V.1991, Roland A. Müller leg., Coll. R. A. Müller (coll. OZ, No. 0084-4) ; 1 ♂: Zucht O. Zompro, Philippines, Marinduque Id. (coll. OZ, No. 0084-1) ; 1 ♂: Philippinen, Aroroy ( MHNG) ; 1 ♀: Mt. Makiling , Luzon, Baker ( ANSP) ; 1 ♀: Mt. Isarog, Tiaong, camarines, Sur , Elev. 2000 feet, May 20, 1924, M.A. Mariano, P.C.A. ( ANSP) ; 1 ♀, 1 ♂: Island Sibuyan, Baker ( ANSP) ; 1 ♂: N.W. Panay, 20773 ( ANSP) ; 1 ♀: Los Baños, P.I., August 1928 J.R. Arena, P.C.A. ( ANSP) ; 2 ♀♀: Philippines: Sierra Madre, Quezon, E. Luzon , VI.1999, I.O. Lumawig; BMNH (E) 2005-98 ( BMNH) ; 1 ♀: Philippines: Panay Is. , 12.IV.1998; BMNH (E) 2005-98 ( BMNH) ; 1 ♀: Philippines: Panay Is. , IV.1998, BMNH (E) 2005-98 ( BMNH) ; 2 ♀♀: Philippines: Mountain Prov. , N. Luzon, VII.96, I. Lumawig, BMNH (E) 2005-98 ( BMNH) ; 1 ♂, 1 ♀: Philippines , Sorsogon, S. Luzon, VIII.1998, I. Lumawig; BMNH (E) 2005-98 ( BMNH) ; 1 ♀, 9 eggs: Philippines, Marinduque Island, Reared by H. Probst , March 1990; BMNH (E) 2005-98 ( BMNH) ; 1 ♂: Canarines S. Libuanan, Maranedo ( MNCN) .

NO DATA: 1 ♀: 94.138 ( BMNH) .

Diagnosis: Closely related to Ph. palawanica spec. nov. from Palawan and Ph. borneensis spec. nov. from Borneo. From the first ♀♀ differ by: the considerably broader and more robust body and legs; relatively shorter body segments; much broader lateral lobes of abdominal tergite VII; different shape of the anal segment and longer cerci. The eggs differ by the larger dimensions and different shape of the micropylar plate, which has the three extensions relatively longer and more slender.

From the Bornean Ph. borneensis spec. nov. it differs by: the larger size; more robust body and broader lateral lobes of abdominal tergite VII of both sexes; deeper posteromedian excavation of the anal segment and more slender cerci of ♀♀; considerably shorter and smaller cerci and much broader, posteriorly broadly rounded semi-tergites of the anal segment of ♂♂. The eggs differ by the larger dimensions, more decidedly indented polar-area and relatively larger micropylar plate, which has the three extension longer and the anterior one slightly elevated and truncate at the apex.

Etymology: From the Latin “ponderosus” (= weighty) and referring to the robust body of this species.

Description: The colouration is described from preserved wild specimens and live captive reared specimens.

♀♀ ( Fig. 89): Large (body length 170.0–205.0 mm), very robust and massive species of cigar-like appearance (maximum body width 14.0–18.0 mm). General colour of body and legs variable, ranging from straw over pale to dark brown, more rarely mid to dull green. Colouration either plain, or body more or less decidedly mottled with pale greyish, yellowish or whitish speckles and legs to a variable degree furnished with annulations and markings of the same colour. Head usually with six dark, ± decided, sub-parallel longitudinal lines. Abdominal tergite III sometimes paler than remaining tergites and with distinct pale greenish or whitish mottling. Spination of the legs dark greyish green, the spines of the medioventral carina of the meso- and metafemora ochre. Antennae mid to dark brown with ventral surface very dark brown to black. Eyes dark greyish or reddish brown.

Head (Fig. 290): Prominent, oval, 1.6x longer than wide with vertex slightly convex. Vertex with a ± distinctly impressed coronal-line. Between the bases of antennae with a small but distinct curved impression. Eyes of moderate size, higher than long, gently convex with anterior margin slightly angled; length contained almost 5x on that of cheek. Antennae reaching almost 2/3 the way along the mesonotum, with 30 segments. Antennomeres slightly increasing in length with the apical 5–7 segments conspicuously shortened. Scapus dorsoventrally flattened, slightly oval in dorsal aspect and about 2.3x longer than wide. Pedicellus less than half the length of scapus, slightly longer than wide, cylindrical. Third antennomere slightly longer than pedicellus.

Thorax: Pronotum 2/3 the length and distinctly narrower than head, posterior margin broader than anterior margin, about 1.2x longer than wide. Anterior margin raised and anterior half of segment with a prominently impressed median line. Median transverse depression distinct but short and at best covering half the width of segment. Mesothorax less than 1.8x longer than head and pronotum combined, decidedly constricted anteriorly, very gently swollen pre-medially and slightly widened at posterior margin. Mesonotum with a very faint longitudinal median line. Metanotum 2/5 the length of mesonotum, almost 2x longer than wide and parallelsided. Meso- and metasternum smooth.

Abdomen: Median segment slightly shorter than metanotum, 1.5x longer than wide and roughly rectangular with the lateral margins gently concave. Segments II–VI indistinctly increasing in length, II about 1.3x, VI 1.5x longer than wide; III and IV broadest segments. Tergite VII slightly shorter than VI, the lateral margins gradually widened to form a ± distinct, rounded or roundly triangular lobe in posterior half, which may project by as much as half of the body width. VIII narrower than VII, 2x longer than wide, constricted medially and strongly convex; anterior margin slightly broader than posterior margin. IX a little more than half the length of VIII, roughly quadrate, strongly convex. Anal segment slightly longer than IX and with a fine longitudinal median carina. Posterior margin with a deep, almost semi-circular median excavation, the outer angles each forming a rounded lobe (Fig. 195). Supraanal plate rounded, keeled and not extending beyond posterior margin of anal segment. Cerci obtuse, moderately elongate, oval in cross-section, slightly tapered towards apex and reaching about to apex of anal segment. Gonapophyses elongate, slender, up-curving and variable in length; either staying slightly beyond or considerably projecting over anal segment. Subgenital plate strongly keeled, boat-shaped and slightly projecting over apex of anal segment (Fig. 194).

Legs: All rather short and robust, the meso- and metafemora slightly swollen sub-basally. Profemora slightly longer than pro-, and mesothorax combined, mesofemora very slightly shorter than mesothorax, metafemora projecting over posterior margin of abdominal segment IV and hind tarsi at best slightly exceeding apex of abdomen. Anterodorsal carina of profemora with 17–23 distinct, pointed serrations; posteroventral carina with a similar number of considerably smaller teeth. All carinae of protibiae very indistinctly dentate. Meso- and metafemora with all carinae rather minutely dentate; teeth more numerous on ventral carinae (Fig. 318). Dorsal carinae of mesofemora sometimes with a slightly enlarged, triangular tooth about 1/3 off the base; occasionally there is a further much smaller, triangular tooth some 2/3 the way along posterodorsal carina and a small tooth 1/3 off the base anterodorsal carina. Metafemora usually without conspicuously enlarged teeth, but rarely with a slightly enlarged, triangular tooth 1/3 off the base on posterodorsal carina. Ventral surfaces of meso- and metafemora slightly convex and with a fine medioventral carina, set with 6–8 rather distinct and pointed spines; 2–4 largest. Ventral carinae of meso- and metatibia very densely but minutely serrate, more sparsely and minutely on dorsal carinae. Occasionally there is a slightly enlarged or even lobe-like, triangular tooth about 1/3 off the base on posterodorsal carina. Probasitarsus with a uniformly raised but dorsal carina, all carinae smooth and about as long as remaining tarsomeres combined. Meso- and metabasitarsi with all carinae minutely dentate, dorsal carina very slightly raised and as long as following three tarsomeres combined.

♂♂ ( Fig. 90): Medium-sized to large (body length 111.0– 134.5 mm) and robust species (maximum body width 4.5–5.5 mm) with long alae (56.0– 67.5 mm). Colouration variable. Body, legs, tegmina and costal region of alae uniformly yellowish, greenish or greyish pale to dark brown; more rarely dull green. Head occasionally with a broad dark brown marking or postocular band. Meso- and metasternum often dull green with several white, elongate spots. Anterior margin of tegmina and alae with a bold white exterior and a more indistinct green interior longitudinal line. Tegmina occasionally with a short, white diagonal band, which begins at the posterior margin and terminates in the centre of tegmen. Anal region of alae transparent greyish brown with darker veins. Abdominal tergite IX with a ± decided longitudinal white patch close to lateral margins. Antennae dark brown with ventral surfaces darker to black. Eyes greyish or reddish mid to dark brown.

Head (Fig. 291): Generally as in ♀♀, but eyes more prominent, circular and projecting hemispherically; their length contained out 2.5x in that of cheek. Antennae with 30 segments and slightly projecting over posterior margin of abdominal tergite II. Antennomeres distinctly increasing in length with the apical 7–8 antennomeres conspicuously shortened. Scapus und pedicellus as in ♀♀.

Thorax: Pronotum as in ♀♀, 1.3x longer than wide. Mesonotum 2.1x longer than head and pronotum combined, parallel-sided and very slightly broadened at posterior margin. Meso- and metasternum smooth. Teg- mina oval, almost 2x longer than wide, constricted basally and with a distinct , roundly conical central hump; projecting over posterior margin of metanotum. Alae reaching about half way along tergite VI.

Abdomen: Median segment 2x longer than metanotum, 3x longer than wide and slightly narrowing towards the posterior. Segments II–IV slightly increasing, V–VII decreasing in length, IV longest segment and about 3.5x longer than wide. Tergite VII ¾ the length of VI and posterolaterally dilated into a distinct , rounded lobe, which laterally extends by as much as half the width of segment. Tergites VIII and IX prominently swollen. VIII broadest segment and conspicuously widening towards the posterior, posterior margin almost 2x wider than VI. IX about as long as VIII, slightly narrowing towards the posterior, gently constricted medially and strongly convex. Anal segment slightly shorter than IX. Semi-tergites very broad, 1.3x longer than high and rounded apically (Fig. 196). Interior surfaces densely covered with minute hooked spines. Cerci small, moderately elongate, cylindrical, gently in-curving and reaching to apex of anal segment. Poculum strongly convex, cup-like and with a blunt, conical hump at the angle; reaching about ¾ the way along tergite IX.

Legs: Generally as in ♀♀ but relatively longer, much more slender and with spination slightly more decided. Profemora longer and mesofemora as long as than combined length of head, pro-, and mesonotum, metafemora almost reaching posterior margin of abdominal tergite V and metatibiae reaching about to apex of abdomen. Ventral surfaces of meso- and metafemora not conspicuously convex, dorsal carinae without enlarged teeth. Meso- and metatibiae occasionally with an enlarged triangular tooth some 1/3 off the base on a posterodorsal carina. Tarsi generally as in ♀♀ but more slender and segments relatively longer.

Eggs (Figs. 145–146): Large, capsule slightly laterally compressed, oval in cross-section, longer than high. Seen laterally the polar-area has a ± distinct impression (depends on locality). Capsule surrounded by a blunt dorsoventral keel, which begins and ends at the operculum. General colouration of capsule dull blackish brown or grey, micropylar plate greyish brown, operculum black and capitulum dark reddish brown. Capsule surface minutely but densely granulated. Two broad oval impression near operculum anterodorsally- and ventrally and two anteroventral depressions near polar end of capsule. Micropylar plate gently raised from capsule and with a slightly darker brown outer margin. Generally shaped like a bold inverted “Y” with all three extensions rather long. Anterior extension slightly broadened and truncate at the apex, the two posterior extensions with apices rounded. Median line short, not extending over micropylar plate. Micropylar cup small, placed in centre of posteromedian gap of the plate. Operculum flat, oval and with a knob-like capitulum on a short stalk in its centre.

Measurements [mm]: length including capitulum 5.6–5.9, length 4.9–5.1, width 3.1–3.3, height 4.0–4.1, leng of micropylar plate 3.3.

Variation: In accordance to its wide distributional range in the Philippines Ph. ponderosa shows quite some deal of intraspecific variation. Features that underlie considerable variation are the size, width and shape of the lateral lobes of abdominal tergite VII, armature of the dorsal carinae of the mid and hind legs and colouration. Most of these features appear to vary from locality to locality. A 200.0 mm ♀ from N-Luzon (Mountain Province) in coll. FH (No. 0192-30) has the lateral lobes of tergite VII very broad, truncate and elevated towards the posterior (Fig. 269) and exhibits two conspicuously enlarged teeth about one third the way along the antero- and posterodorsal carina of the mesofemora and a distinct triangular pre-medial dorsal lobe on the meso- and metatibiae. In contrast, enlarged teeth or lobes on the legs are completely lacking or just very poorly developed and the lateral lobes of abdominal tergite VII considerably smaller and rounded in specimens from Panay, Mindoro or Marinduque. These two features appear to be typical for specimens from Luzon only. Slight variation is also seen in the shape of the semi-tergites of the anal segment in ♂♂, which have the apex very broad in specimens from Panay or Luzon (Fig. 196) but comparatively more narrow in specimens from Marinduque. ♀♀ from Mindoro Island are comparatively smaller (173.0–182.0 mm, coll. FH) than ♀♀ from Luzon, Marinduque, Panay or Sorsogon (184.0–204.0 mm, coll. FH). Captive reared ♂♂ from Marinduque are typical for having a distinct blackish postocular marking on the cheeks and a whitish, diagonal line along the posterior margin of the tegmina, both features lacking in ♂♂ from other localities. Wild ♀♀ are mostly various shades of brown, more rarely green, and often prettily mottled with white or pale brown. Captive reared ♀♀ that were raised on alternative foodplants in Europe, are exceptionally mid to dull green .

Not only the insects but also the eggs show slight variation, those from the islands of Panay and Mindoro being somewhat smaller with the polar-area less distinctly impressed than eggs from Luzon or Marinduque. Furthermore, slight variation is observed in the shape of the micropylar plate.

Comments: Stål (1877a: 40) originally described Pharnacia ponderosa from two ♂♂ with the unprecise location “ Philippinen ” in NHRS of which Brock (1999: 181) designated the darker brown specimen as the LT. The fact that both specimens have a triangular pre-medial dorsal lobe on the meso- and metafemora suggests both specimens are from Luzon (see comments on variation above). Already Redtenbacher (1908: 455) doubted that the ♀ which Kirby (1896: 451) described to have 7.0 mm long tegmina was the corresponding ♀ of Stål’s species. Günther (1933: 159) correctly stated this specimen to be a ♀ of Phasmotaenia sanchezi ( Bolívar, 1897) , which was confirmed by examination.

Although Ph. ponderosa has a wide distributional range in the Philippines and is quite frequently encountered on most of the larger islands of the archipelago (see distribution below), being rather abundant in some localities, the ♀ and egg have so far remained formally undescribed .

Pharnacia magdiwang Lit & Eusebio, 2008 was described upon both sexes and the eggs from Magdiwang in the Romblon Province, Sibuyan Island. Lit & Eusebio (2008: 117) distinguished this species from Ph. ponderosa Stål, 1877 mainly on the basis that both sexes lack an obvious pre-medial dorsal lobe on the meso- and metatibiae and slight differences of the egg-morphology. Based on examination of specimens from various Philippine islands the pre-medial dorsal lobe of the meso- and metatibiae is here shown to underlie considerable intraspecific variability (see comments on variation above) and to be frequently well developed only in specimens from Luzon, the only island from which Lit & Eusebio (2008) had specimens of Ph. ponderosa at hand for comparison with their Ph. magdiwang . As stated above, even the eggs of Ph. ponderosa show slight variation depending on the locality, thus the differences summarized by Lit & Eusebio (2008: 117) for the eggs of Ph. magdiwang lie well within the range of variation of Ph. ponderosa . The “distinct sculpturing” and “fontanelle-like depression” of the head in Ph. magdiwang mentioned as distinctive features upon Ph. ponderosa by Lit & Eusebio (2008: 117) are questionable. Measurements and length relations of the insects generally correspond to other specimens of Ph. ponderosa examined for the present study (→ Table 10). The fact that Sibuyan Island shows a very high degree of endemicity and is hence often recognized as a separate biogeographical subregion of the Philippines (see Heaney & Regalado, 1998), can also not support to the validity of Ph. magdiwang , since Ph. ponderosa is known from most of the larger islands that surround Sibuyan (e.g. Luzon, Marinduque, Mindoro, Panay and Cebu) and the genus Pharnacia Stål, 1877 itself is apart from the Philippines distributed throughout entire Sundaland. Consequently, as none of the distinguishing features mentioned by Lit & Eusebio, 2008: 117) for Ph. magdiwang Lit & Eusebio, 2008 holds true this species is obviously a junior synonym of Ph. ponderosa Stål, 1875 (syn. nov.).

In the Philippines Lit & Eusebio, 2008: 121) stated Ph. ponderosa to feed on Canarium ovatum (Burseraceae) , Mangifera indica (Anacardiaceae) , Terminalia microcarpa (Combretaceae) an guava ( Psidium guajava , Myrtaceae ). In captivity in Europe it accepts oaks ( Quercus robur , Q. petraea , Q. rubra & Q. ilex , Fagaceae ), bramble ( Rubus spp. , Rosaceae ), eucalyptus ( Eucalyptus spp. , Myrtaceae ), Salal ( Gaultheria shallon , Ericaceae ) and rose ( Rosa spp. , Rosaceae ) as alternative foodplants. Culturing in Europe has been attempted several times already from stock originating from various localities but none was maintained for more than a few generations. A current culture-stock from an unknown locality however is rather easy to rear in large and well ventilated cages. Eggs have an incubation time of 4–5 months and nymphs reach maturity in about 6–8 months. A ♀ produces an average of 8– 10 eggs per day and a total of 400–700 during her lifetime.

Distribution (Fig. 376): Philippines (endemic): Aroroy Id., Cebu Id. (Minglanilla 300 m), Luzon Id. (Mount Makiling; Sorsogon; Sierra Madre, Quezon; Los Baños; Camarines Sur Province, Mount Isarog: Tia-

ong 2000 ft. & Mountain Province: Nueva Viscaya, Balite), Marinduque Id., Mindoro Id. (Mount Halcon), Panay Id. (Mount Nangtud), Sibuyan Id. (Romblon Province: Magdiwang, Pawala River, Tampayan 100–400 m & Brangay Ipil) & Libuanan Id.

Number of specimens examined: 61

* according to Lit & Eusebio (2008: 119, 120)