Phyllium (Phyllium) celebicum de Haan, 1842

Hennemann, Frank H., Conle, Oskar V., Gottardo, Marco & Bresseel, Joachim, 2009, On certain species of the genus Phyllium Illiger, 1798, with proposals for an intra-generic systematization and the descriptions of five new species from the Philippines and Palawan (Phasmatodea: Phylliidae: Phylliinae: Phylliini) 2322, Zootaxa 2322 (1), pp. 1-83 : 11-16

Phyllium (Phyllium) celebicum de Haan, 1842

( Figs. 1–5 , 13–17, 24–25 , 85 , 95–96, 113, 139)

Phasma (Phyllium) celebicum de Haan, 1842: 111 . HT, ♀: Celebes, Tondano (RMNH – not traced).

Phyllium celebicum, Kirby, 1904: 423 (in part).

Redtenbacher, 1906: 176 (in part).

Willemse, 1945: 319, fig. 3 (illustration of HT ♀).

Phyllium (Phyllium) celebicum, Klante, 1976: 63 (in part).

Otte & Brock, 2005: 273 (in part).

Hennemann, Conle & Zhang, 2008: 42. [Notes on identity of Chinese records]

[Not: Phyllium celebicum, Gray, 1843: 121 , fig. e (♀). Philippines, = Ph. (Ph.) ericoriai n. sp.]

[Not: Phyllium celebicum, Westwood, 1859: 173 , pl. 40: 6 (♂). Philippines, = Ph. (Ph.) ericoriai n. sp.]

[Not: Phyllium celebicum, Wood-Mason 1875: 218 , pl. 16 (♀ & nymph). Myanmar, = Ph. (Ph.) westwoodii Wood- Mason , 1875]

[Not: Phyllium celebicum, Rehn & Rehn, 1933: 413 , pl. 17: 5 (♀). Philippines, = Ph. (Ph.) ericoriai n. sp.]

[Not: Phyllium celebicum, Grösser, 1990: 199 & 200, = Ph. (Ph.) westwoodii Wood-Mason, 1875 ]

[Not: Phyllium celebicum, Hennemann, 1992: 31 . Thailand, = Ph. (Ph.) westwoodii Wood-Mason, 1875 ]

[Not: Phyllium celebicum, Liu, 1993: 207 , figs. 10 & 11 (♀). Specimens from China (Guangxi, Yunnan, Guizhou and Hainan) are an as yet unnamed species close to Ph. (Ph.) westwoodii Wood-Mason, 1875 ]

[Not: Phyllium celebicum, Bragg, 1996: 110 , the specimen stated to be the HT is clearly a distinct species]

[Not: Phyllium (Phyllium) celebicum, Grösser, 2001: 64 , figs. 16, 41–44 & 82–87. Thailand, = Ph. (Ph.) westwoodii Wood-Mason, 1875 ]

[Not: Phyllium (Phyllium) celebicum, Zompro & Grösser, 2003: 136 & 138, fig. 12: 3 (egg). Thailand, = Ph. (Ph.) westwoodi Wood-Mason, 1875 ]

[Not: Phyllium (Phyllium) celebicum, Sorpongpaisal & Thanasinchayakul, 2006: 22 . Northern Thailand = Ph. (Ph.) westwoodii Wood-Mason, 1875 ]

[Not: Phyllium (Phyllium) celebicum, Chen & He, 2008: 368 . S-China, = Ph. (Ph.) westwoodii Wood-Mason, 1875 ]

[Not: Phyllium (Phyllium) celebicum, Grösser, 2008: 103 , figs. 24, 34, 52–55, 122–127, 187 & 195. Thailand, = Ph. (Ph.) westwoodii Wood-Mason, 1875 ]

Material examined [11 ♀, 1 ♂, eggs]: SULAWESI: 1 ♀, eggs: Indonesien, S-Sulawesi, Tiulapolu, leg. Jasmin III.2008 (coll. FH, No. 0634-3 & E) ; 2 ♀: Indonesien, S-Sulawesi, Bugadidi 2008, via Sigetake Suzuki (coll. FH, No’s 0634-1 & 2) ; 1 ♂, 2 ♀: ex Zucht F. Hennemann 2009, F1-Generation, Herkunft: S- Sulawesi, Bugadidi, leg. Jasmin 2008 (coll. FH, No’s 0634-4 to 6); 1 ♀: Indonesien, Süd-Sulawesi , Bugadidi, X.2002 (coll. OC) ; 1 ♀: Sulawesi (coll. OC) ; 2 ♀: Sulawesi , 10.2002 (coll. OC) ; 1 ♀ + 1 egg (ex ovipositor): Südost-Sulawesi , Buton, 6.2001 (coll. OC, No. 01826) ; 1 ♀: Süd-Sulawesi , Bugadidi, 10.2002 (coll. OC) .

AMBON: 1 ♀: Amboina, Felder; Phyllium celebicum ♀ deHaan Brunner det. ( MNHU) .

NO DATA: 1 ♀: no data ( NHMW, No. 293) .

Comparison: Very close to Ph. (Ph.) westwoodii Wood-Mason, 1875 from continental SE-Asia but distinguished by a number of characters of the insect and egg-morphology. ♀ differ by: the more elongate and slender meso-praescutum which is about 1.4x longer than wide (only 1.2x in westwoodii ); larger more acutely angular exterior lobe (angle <90°) and more numerous and prominent teeth of the interior lobe of the profemora ( Figs. 13–16 ); longer, more slender and differently structured antennae ( Fig. 85 ), which have the apical antennomere more elongate and tapered and III with only 38–44 teeth in the pars stridens (50–56 in westwoodii ); shorter alae which are just a little longer than half the length of tegmina (about ¾ the length in westwoodii ), and shorter less acutely pointed subgenital plate ( Fig. 24 ). ♂♂ also differ by: the longer and more slender mesonotum; larger and more angulate exterior lobe and more distinct armature of the interior lobe of the profemora ( Fig. 17 ); single terminal hook of the vomer (two hooks in westwoodii ) as well as the different shape of the abdomen, which has segments V–VI roughly parallel-sided and IV and VII angulate (oval in westwoodii ), and presence of two additional eye-like spots in abdominal segment VI. In addition to features mentioned for adults, nymphs of both sexes can frequently be distinguished from those of westwoodii by having four conspicuous eye-like spots on the abdomen. Eggs are also similar to those of westwoodii but differ by: the larger dimensions; more convex lateral surfaces of the capsule, which have the circular pits deeper and more conspicuous; longer micropylar plate and shorter hairy structures of the capsule and operculum (Figs. 95–96).

11. ♀ PT: Philippines (Marinduque Id.) [coll. FH, No. 0672-1]

12. ♂ PT: Philippines (Marinduque Id.) [coll. FH, No. 0672-3]

Ph. (Ph.) celebicum was frequently confused with Philippine specimens here described as Ph. (Ph.) ericoriai n. sp. but both sexes differ from this species by the more slender and elongate meso-praescutum; less distinct armature of the interior lobe of the profemora and more slender just gently rounded interior lobe of the mesofemora, which is at best equal in width to the exterior lobe. ♀ furthermore differ by: the more acutely angular exterior lobe (angle <90°) of the profemora ( Figs. 13–16 ); having 38–44 teeth on pars stridens of antennomere III (27–28 in ericoriai , Fig. 85 ); sub-parallel abdominal segments V and VI and gradually tapered to gently concave segment VIII. ♂♂ are well distinguished by the more slender abdomen, which does not have segment VI considerably expanded, two additional eye-like spots in abdominal segment VI and larger more angular exterior lobe of the profemora. The eggs differ by lacking the spectacular high longitudinal lamellae of the capsule and having a conical operculum (Figs. 95–96)

Diagnosis: ♀ ( Figs. 1–2, 4–5 , 139 ). Medium-sized to large for the subgenus (body length 75.5–92.0 mm) with a broad and angulate abdomen (maximum width 30.0–39.0 mm), large and acutely angular exterior lobes of the profemora, and developed alae which are roughly half as long as the tegmina (26.0–32.0 mm). Colouration variable, ranging from green over yellow and orange to pale brown, body and legs to variable degree furnished with brown speckles or mottling. Meso-praescutum and dorsal portion of mesopleurae mid to dark brown. Antennae and tarsi reddish brown, the latter with the base of each segment pale cream. Head broad with the cheeks convex and distinctly widened towards posterior; vertex smooth. Antennae rather slender and elongate (3.8–4.0 mm), shorter than postocular section of head capsule. Apical antennomere (IX) about 1.5x longer than wide, distinctly longer than VIII and gradually tapered towards tip. Pars stridens on antennomere III with 38–44 teeth ( Fig. 85 ). Pronotum roundly trapezoidal and narrowed towards the posterior with anterior margin about 2.3x broader than posterior margin; the latter rounded. Centrally with a small but conspicuous T-shaped impression. Meso-praescutum very gently converging towards the posterior, elongate and about 1.4x longer than wide. Lateral margins irregularly armed with low tubercles of variable sizes. Disc very slightly tectiform longitudinally and evenly but irregularly granulose and rugose. Anterior margin with a short and rather low, C-shaped transverse ridge. Mesopleurae narrow and set with a few spiniform tubercles in anterior half, the posterior half prominently diverging with the lateral margin just minutely tuberculose. Mesosternum slightly tectiform longitudinally and set with a variable number of granules and small tubercles in anterior portion; posterior half smooth. Tegmina at least reaching half way along abdominal segment VII. Alae developed, slightly more than half the length of tegmina and slightly projecting over posterior margin of abdominal segment IV (26.0–31.0 mm). Abdominal segments II–III gradually widened, IV acutely angulate and V–VI very slightly narrowing. VII roundly angulate (roughly 90°) and occasionally forming a small, rounded lobe ( Figs. 1–2, 4–5 ). V widest segment. Subgenital plate rather short, acutely triangular and very slightly projecting over posterior margin of tergite IX ( Fig. 24 ). Profemora with a large and acutely angular exterior lobe (angle <90°), complete outer margin minutely dentate. Armature of interior lobe prominent but variable ( Figs. 13–16 ); width about 2/3 that of exterior lobe. Interior lobe of protibiae distinct and roundly triangular (angle roughly 90°). Interior lobe of mesofemora very gently rounded and narrower or at best equal in width to exterior lobe; the latter angular medially. Protarsus about 2/3 the length of corresponding tibia, probasitarsus roughly 4.5x longer than wide.

♁♁ ( Fig. 3 ). Medium-sized to large for the subgenus (body length 62.0 mm) with a moderately broad and medially parallel-sided abdomen (maximum width 19.8 mm), four conspicuous eye-like spots on the abdomen, and distinctly angulate exterior lobes of the profemora. Colouration green and to a variable degree furnished with reddish brown speckles and markings, large parts of mesopleurae brown. Antennae pale straw, the two apical segments reddish brown. Vertex smooth. Antennae consisting of 24 segments and ± reaching to posterior margin of abdominal segment V. Pronotum and mesothorax structured like in ♀. Mesopleurae narrow in anterior half but distinctly diverging in posterior half, lateral margins minutely tuberculose. Mesosternum minutely granulose anteriorly and smooth posteriorly; gently tectiform. Tegmina reaching about half way along abdominal segment III, alae reaching roughly half way along segment IX. Abdominal segment II very slightly narrowing, III widening, IV roundly angulate (angle about 110°), V and VI roughly parallelsided, VII widely rounded and VIII–X gradually narrowing. VI and VII widest segments. Segments V and VI each with two conspicuous eye-like spots, those on V considerably larger and with a broad brown outer margin. Vomer broadly triangular, slightly transverse and with a singly, short and rather blunt terminal hook; slightly curved to the left. Poculum moderately convex, with the posterior margin gently angulate and slightly projecting over posterior margin of tergite IX; over complete length with a fine longitudinal median carina ( Fig. 25 ). Exterior lobe of profemora large and angulate (angle about 100°) with the outer margin minutely dentate. Interior lobe with 6–7 teeth of variable sizes, IV most prominent ( Fig. 17 ). Protibiae with a widely triangular interior lobe.

13–16. Ph. (Ph.) celebicum de Haan, 1842 , left profemur of ♀

17. Ph. (Ph.) celebicum de Haan, 1842 , left profemur of ♂

18. Ph. (Ph.) ericoriai n. sp., left profemur of ♀ PT [coll. FH, No. 0672-1]

19. Ph. (Ph.) ericoriai n. sp., left profemur of ♂ PT [coll. FH, No. 0672-3]

20–21. Ph. (Ph.) westwoodii Wood-Mason, 1875 , left profemur of ♀ [coll. FH]

22–23. Ph. (Ph.) westwoodii Wood-Mason, 1875 , left profemur of ♂♂ [coll. FH]

24. Ph. (Ph.) celebicum de Haan, 1842 , apex of abdomen (ventral view) ♀ [coll. FH, No. 0634-2]

25. Ph. (Ph.) celebicum de Haan, 1842 , apex of abdomen (ventral view) ♂ [coll. FH, No. 0634-4]

26. Ph. (Ph.) westwoodii Wood-Mason, 1875 , apex of abdomen (ventral view) ♀ [coll. FH, No. 0542- 17]

27. Ph. (Ph.) westwoodii Wood-Mason, 1875 , apex of abdomen (ventral view) ♂ [coll. FH, No. 0542- 14].

Nymphs: Newly hatched nymphs are very similar to those of Ph. westwoodii but slightly larger (body length 13–14 mm) with a proportionally broader abdomen (width 2.0 mm). General colour mahogany with white spots on the legs. The meso- and metafemora each have two conspicuous white spots sub-basally. Abdominal segments II–VII each with a bold pale green marking.

Eggs (Figs. 95–96). Of moderate size, strongly angular and with distinct parallel longitudinal keels, two dorso-laterally and three ventrally. Capsule about 1.6x longer than wide, the dorsal surface flattened, the lateral surfaces gently convex and the dorsal surface tectiform; polar-area slightly concave. Almost entire capsule surface irregularly covered with low moss-like extensions resembling small palm-trees; these particularly numerous and most prominent along the longitudinal keels of the capsule and operculum. Capsule otherwise set with several deep circular pits, roughly arranged in longitudinal rows. Micropylar plate elongate, about ¾ the length of capsule and tapered towards both ends, posterior portion very narrow; outer margin set with hairy structures. Micropylar cup placed roughly in centre of plate and furnished by moss-like structures. Operculum roundly conical, round in cross-section and slightly displaced towards ventral egg surface; opercular angle about 25°. Lateral surfaces with several deep impressions. Colouration mid to dark brown, the moss-like structures of a slightly paler colour.

Measurements [mm]: length (including operculum) 5.0–5.1, length 4.5–4.6 mm, width 2.8–3.0 mm, height 2.9–3.1 mm, length of micropylar plate 3.6–3.7 mm.

Variation: This species shows slight variation in the size, armature of the interior lobe of the profemora and shape of abdominal segment VII of ♀, which is either angulate or protruded into a slightly rounded posteriorly extending lobe. The colouration of ♀ is strongly variable, specimens being either green and to a variable degree furnished with brown speckles and mottling, or yellow to pale brown with darker brown mottling ( Figs. 1–2 ). More rarely orange specimens occur. Due to the lack of specimens nothing is known about the variation of ♂♂.

Comments: Examination has shown the ♀ in RMNH, stated to be the HT of Ph. celebicum de Haan by Bragg (1996: 110), cannot be the type of de Haan’s celebicum and clearly is a distinct species. Although this specimen is from Sulawesi, it obviously differs from the original description by the gradually tapered abdominal segments IV–VII, much more slender exterior lobe and differently armed interior lobe of the profemora and lack of alae. De Haan (1842: 111) stated the abdomen to be distinctly angulate and the alae to be rather well developed and about half as long as the tegmina, as shown in the illustration of the HT provided by Willemse (1945: 319, fig. 3). Extensive search for de Haan’s HT in the RMNH collection in April 2006 could not trace a specimen that matches with the original description and illustration by Willemse (1945) and hence, the specimen must be presumed lost. The possibly necessary designation of a neotype should however await confirmation concerning the depository and existence of the HT of Ph. (Ph.) celebicum de Haan.

♀ from South Sulawesi in the two first author’s collections (coll. FH & coll. OC) match perfectly in every aspect with the original description of Ph. celebicum and illustration of the HT provided by Willemse (1945), and all possess the typically half-sized alae. Hence, these specimens do without any doubt represent the same species. The previously unknown ♂ and egg are here described and illustrated for the first time.

There was much confusion around Ph. (Ph.) celebicum since its description in 1842 with the species frequently misinterpreted and confused with Ph. (Ph.) westwoodii Wood-Mason, 1875 by almost all subsequent authors except Willemse (1945) and Klante (1976). Localities in fact relating to Ph. (Ph.) westwoodii were first confused by Redtenbacher (1906: 176), who considered the distributional range of celebicum to include Sulawesi, Ambon, the Philippines, Myanmar, Laos, Vietnam and the Seychelles. However, most of these records are obviously based on misidentified material, all records from the Philippines representing a distinct species described as new below ( Ph. (Ph.) ericoriai n. sp.). Records from the Asian mainland ( Thailand, Myanmar, Laos, Vietnam and China) and erroneously affiliated with Ph. (Ph.) celebicum by Redtenbacher (1906: 176), Liu (1993: 207) and Grösser (2001: 64) almost exceptionally appear to relate to Ph. (Ph.) westwoodii , which has already been pointed out for the Chinese records by Hennemann, Conle & Zhang (2008: 42).

Culture-stock of Ph. (Ph.) celebicum was collected by a native insect-supplier near Bugadidi, S-Sulawesi in 2008 and eggs were sent to Sigetake Suzuki (Hokkaido, Japan) who passed on some of them to the first author. The species has proven fairly easy to rear in captivity with both nymphs and adults readily accepting bramble ( Rubus fruticosus , Rosaceae ) and oak ( Quercus robur , Fagaceae ) as alternative foodplants. At temperatures of approximately 25°C nymphs reached maturity in about 6–7 months with the mortality rate during the first instar rather low. ♀ produce an average of 3– 4 eggs per day and a total of about 300 eggs in an entire life-time. Eggs of the captive reared F1-generation were distributed to various breeders throughout Europe in 2009.

Distribution ( Fig. 113 ): N-Sulawesi (Tondano [type-locality]); S-Sulawesi (Bugadidi [coll. FH, coll. OC], Buton [coll. OC] and Tiulapolu [coll. FH]) and Ambon [MNHU].

* according to Klante (1976: 63)

** the ♀ from Ambon in MNHU has the following remarkable dimensions: body length 92.0 mm, length of alae 49.0 mm. The alae are considerably longer than in all other examined specimens .

Figures

FIGURES 1–5. Phyllium (Ph.) celebicum de Haan, 1842 [scale = 10 mm]

FIGURES 13–27. Left profemur and genitalia of members of the “celebicum-group” [scale = 2 mm].

FIGURES 85–94. Right antenna of ♀, interior view [scale = 1 mm]

FIGURES 137–140. Live Insects

FIGURES 113–118. Maps showing the distribution of Phyllium spp.