Venupherodes venustula (Audinet-Serville, 1838) Audinet-Serville, 1838

Frank H. Hennemann, Oskar V. Conle & Daniel E. Perez-Gelabert, 2016, Studies on Neotropical Phasmatodea XVI: Revision of Haplopodini Günther, 1953 (rev. stat.), with notes on the subfamily Cladomorphinae Bradley & Galil, 1977 and the descriptions of a new tribe, four new genera and nine new species (Phasmatodea: “ Anareolatae ”: Phasmatidae: Cladomorphinae), Zootaxa 4128 (1), pp. 1-211 : 167-172

Venupherodes venustula (Audinet-Serville, 1838) n. gen., n. comb.

( Figs. 322–333 , 345 , 364 , 374 , 384–387 )

Platycrana venustula Audinet-Serville, 1838: 242 . HT, ♀: Cuba [MNHN—lost]. NT (by present designation), ♀: 800; Havaniense (M. Leay) Westw., Cuba, Otto [MNHU].

Diapherodes venustula, Westwood, 1859: 84 .

Saussure, 1872: 185.

Bolivar, 1888: 140.

Redtenbacher, 1908: 435.

Rehn, 1909: 200.

Moxey, 1972: 117 (in litt.).

Otte & Brock, 2005: 121.

Rabaey & Simoens, 2007: 6, figs. [Culture report]

Diapherodes (?) venustula, Kirby, 1904a: 362 .

Phasma (Diapherodes) venustulum, de Haan, 1842: 109 .

Phasma havaniense Westwood, 1859: 34 , pl. 22: 7 (♀). HT, ♀: Ins. Havannah; Type—Westwood, Phasma havaniense, Cat. Phasm. 1859 , p. 24, pl. 22 f. 7; Pleseophyllum Havaniense ; Type Orth: 615, Phasma havaniense Westw., Hope Dept. Oxford [OXUM, No. 615]. [Synonymised by Saussure 1871–1872: 185] Moxey, 1972: 117 (in litt.). Otte & Brock, 2005: 121.

Haplopus juvenis Redtenbacher, 1908: 430 , pl. 20: 4 (♂). LT (by present designation), ♂: Coll. Br. v. W., ex. Coll. Sommer, Cuba; det. Br. v. W., Haplopus juvenis (specimen with open wings) [NHMW, No. 795]; PLT, ♂: Coll. Br. v. W., ex. Coll. Sommer, Cuba; det. Br. v. W., Haplopus juvenis [NHMW, No. 795]; PLT, ♂: Coll. Br. v. W., ex. Coll. Sommer, Cuba; det. Br. v. W., Haplopus juvenis , 77.; 7647, jamaicensis , Cuba [NHMW, No. 795]; PLT, ♂: Coll. Br. v. W., ex. Coll. Sommer, Cuba; det. Br. v. W., Haplopus juvenis ; 7647; 7647 Phasma jamaicensis Fabr. Drur. , Spectrum viride Stoll , capta T. Müller, Cuba, Phas. Inv. Nr. 795 [NHMW, No. 795]. n. syn.

Rehn, 1909: 200.

Moxey, 1972: 117 (in litt.). [Listed as a synonym of Diapherodes venustula (Audinet-Serville, 1838) —unpublished] Otte & Brock, 2005: 151.

Aplopus juvenis, Brock, 1998a: 57 .

Further material [78 ♂♂, 53 ♀♀, 3 nymphs, eggs]: CUBA:

1 ♀: Coll. Br. v. W., Kuba, Saussure; det. Br. v. W. Diapherodes venustula ; 11.165; 11.165 Diapherodes venustula Serv. (Havaniensis) ♀ [ NHMW, No. 840]; 1 ♀: Diapherodes venustula, Serv. ; Cuba (Lembeye) [ MNCN]; 1 ♀: Diapherodes venustula , Cuba, Gundlach [ MNCN]; 1 ♂ (penultimate instar): Cuba, Cabrara [ MNCN]; 1 ♀, 2 ♂♂: Cuba, achété par M. Poey; Diapherodes venustula Serv. ( MHNG); 2 ♀♀, 2 ♂♂: Cuba, Mr. H. de Saussure; Diapherodes venustula Serv. [ MHNG]; 1 ♂: Cuba, coll. Guerin; Diapherodes venustula Serv. [ MHNG]; 1 ♂, 1 ♀ (nymph): Cuba; Diapherodes venustula Serv. [ MHNG]; 2 ♂♂: 836; venustula (Westw.) , Platycr. venustula Serv. , Cuba, Otto [ MNHU]; 1 ♀, 1 ♂: ex Zucht: K. Rabaey ( Belgien), Herkunft: W-Cuba, leg. I. Fritzsche, VII.2006 [coll. FH, No’s 0599- 1 & 3]; 1 ♂: ex Zucht: F. Hennemann, urspr.: CUBA, leg. I. Fritzsche, VII.2006 [coll. FH, No. 0599-2]; 13 ♀♀, 18 ♂♂, 1 ♂ (penultimate instar), eggs: ex Zucht: F. Hennemann 2007/2008, Herkunft: W-Cuba, Pinar del Rio Provinz, leg. I. Fritzsche 2005 [coll. FH, No’s 0599-4 to 35 & E]; 1 ♀, 2 ♂♂: ex Zucht F. Hennemann 2008/09, Herkunft: W-Cuba, Pinar del Rio Provinz, leg. I. Fritzsche 2005 [coll. FH, No’s 0599-36 to 38]; 1 ♂: Sierra Rangel, de Rio—1934; Roberts Coll.; wO172 [ ANSP]; 1 ♂, 1 ♀: Somorrostro, Havana, Aug. 20, 35, Schramuzza, y. Brunner [ ANSP]; 1 ♂, 1 ♀: Marianao, Cuba, Aug. (From. T. Barbouy) [ ANSP]; 30 ♀♀, 46 ♂♂, eggs: ex Zucht O. Conle, Herkunft: W-Cuba, Pinar del Rio Provinz [coll. OC].

Description: ♀ ( Figs. 322–324 , 384 , 386–387 ). Small (body length including subgenital plate 65.0–80.0 mm) and robust Haplopodini (maximum body width at abdominal segment IV 8.0–10.0 mm) with a strongly broadened mesothorax and medially swollen abdomen. Colour of body and head bright apple green with the ventral body surface dull green. Sometimes specimens occur which have certain parts of the dorsal body surface furnished with irregular washed brownish or blackish markings ( Figs. 324 , 387 ); more rarely blackish markings are seen on the ventral surface of the abdomen. Rather frequently specimens have the head dull brownish green or almost entirely brown. Anterior margin of pronotum yellow. Anterior portion of the lateral margins of mesonotum ochre, meso- and metapleurae ochre with a broad, white longitudinal stripe along ventral margin. Coxae reddish brown. Cerci pale to dark brown. Front legs greenish brown with the basal portion of the femora pale green. Mid and hind legs pale to mid green with the base of the femora slightly brownish. Basitarsi green, remaining tarsomeres brown. Scapus and pedicellus mid to dark brown. Remainder of antennae pale brown and each antennomere with a fine transverse dark band at the apex; this turning to black towards apex of antennae. Eyes reddish brown with pale cream markings.

Head: Globose, indistinctly longer than wide, and considerably broader than prothorax. Vertex convex, with a few small, scattered granules and in centre mostly armed with a blunt, conical dextral tubercle; the sinistral one faint or obsolete ( Fig. 326 ). Eyes convex, almost circular and of moderate size; their length contained about 1.8x in that of cheeks. Antennae slender and hardly reaching posterior margin of median segment; with about 45 antennomeres. Scapus dorsoventrally compressed, about 2x longer than wide and constricted at the base. Pedicellus cylindrical and about half as long as scapus.

Thorax: Pronotum about as long but distinctly narrower than head and strongly tapered towards the posterior; anterior margin about 1.3x wider than posterior margin and median portion very gently constricted. Transverse median sulcus very distinct, slightly curved but just not reaching lateral margins of segment. Anterior half with a pair of prominent, blunt spines and close to posterior margin with two small tubercles; otherwise smooth ( Fig. 326 ). Mesothorax about 1.5x longer than head and pronotum combined and strongly constricted anteriorly; posterior margin about 3x wider than anterior margin. Mesonotum roughly 2x longer than wide and distinctly lyriform, being strongly broadened towards about 2/3 of its length and gently constricted in the posterior portion. Lateral margins strongly rounded in medial portion and overlapping the mesopleurae. Entire dorsal surface of mesonotum with a distinct longitudinal median keel, close to the anterior margin with a small pair of ± distinct tubercles ( Fig. 326 ) and with a few rounded granules along the lateral margins; otherwise smooth. Lateral margins very minutely denticulate. Mesopleurae with a fine longitudinal keel close to anterior margin and with a median row of granules. Metapleurae slightly tectiform with an acute longitudinal median keel, unarmed. Mesosternum smooth or with a very few scattered granules, metasternum smooth. Metanotum less than 1/3 the length of mesonotum and roughly quadrate; dorsal surface with a faint longitudinal median keel.

Abdomen: Median segment slightly longer than metanotum, a little longer than wide and very gently widening towards the posterior. Dorsal surface with a faint longitudinal median keel, otherwise smooth. Abdomen swollen medially with II–III widening, V–X narrowing and IV broadest segment ( Fig. 322 ). Tergites II–IV transverse with IV almost 2x wider than long, VII slightly longer than wide. Tergites II–IX each with four distinct, longitudinal and roughly parallel carinae; two close to the median line and one on each side about half way towards the lateral margin. The two median carinae are melted together to form a single blunt carina on VIII–X. Sternites II–VII smooth. Praeopercular organ formed by three small, wart-like granules close to posterior margin of sternum VII; these roughly arranged as a triangle ( Fig. 345 ). Tergum VIII about ¾ the length of VII and strongly convex. IX slightly wider than long and about 2/3 the length of VIII. Anal segment about as long as IX, flattened and gradually tapered towards the posterior; posterior margin rounded and with a small median notch and lateral margins somewhat emarginated medially ( Figs. 328–329 ). Epiproct very small, slightly projecting over anal segment, semicircular and with a keel dorsally. Cerci mostly hidden underneath anal segment; finely bristled. Subgenital plate longitudinally keeled and tapered towards an acute apex; projecting over apex of abdomen by about 1.5x the length of the anal segment ( Figs. 328–329 ).

Legs: All fairly short, profemora and mesofemora shorter than mesothorax and metafemora slightly projecting over posterior margin of abdominal segment IV. Two outer ventral carinae of meso- and metafemora each with 1–2 small but pointed sub-apical spines. Medioventral carina of profemora with 1–2 small sub-apical spines, of meso- and metafemora with five moderately strong spines, which increase in size towards the apex of femur. Dorsal carina of the probasitarsus very gently rounded, all basitarsi roughly as long as following two tarsomeres combined.

♂ ( Figs. 325 , 385–386 ). Fairly small (body length 54.5–68.0 mm) and moderately slender insects with well developed alae (29.5–35.0 mm) and a very characteristic colouration. General colour of head and dorsal body surface mid to dull green. Ventral body surface creamish mid brown, sometimes with a slight greenish wash. Head with a bold white longitudinal stripe along the cheeks ( Fig. 327 ). Pronotum ochre with the central portion green. Mesonotum mid to dull green with the anterior and lateral margins broadly ochre to mid brown. Greatest parts of metapleurae bright apple green. Tegmina and costal region of alae mid to dull green with the anterior margin brown and towards the interior bordered by a longitudinal whitish stripe. Central hump of tegmina sometimes dull yellow. Anal region translucent orange. Eyes reddish brown with pale cream markings. Antennae generally as in ♀♀. Fore legs brownish green, mid and hind legs mid to dull green with the bases of the femora pale green. Tarsi greenish brown with the basitarsus green. In addition to the typical colour-form here described, considerably darker specimens frequently occur amongst captive reared specimens. These have the mid and hind legs, tegmina and costal region of the alae very dark green and the mesothorax almost entirely brown, even the metapleurae are dull instead of apple green. It is not known whether such varieties also occur in the wild.

Head: Generally as in ♀♀ but vertex slightly less convex and the two cephalad tubercles more distinct ( Fig. 327 ). A small C-shaped impression between the bases of the antennae. Eyes of moderate size, circular and strongly convex; their length contained less than 1.5x in that of cheeks. Antennae moderately thickened and ± reaching to posterior margin of abdominal segment III. Otherwise as in ♀♀.

Thorax: Pronotum slightly shorter and narrower than the head, slightly trapezoidal and gradually narrowed towards the posterior; posterior margin rounded. Median transverse depression very distinct, gently curved and almost reaching lateral margin of segment; armature as in ♀♀ but posterior pair of tubercles indistinct to obsolete ( Fig. 327 ). Mesothorax elongate, roughly 2x longer than head and pronotum combined, cylindrical and slightly widened at the posterior. Anterior margin of mesonotum usually with one or two pairs of blunt spine-like tubercles or spines ( Fig. 327 ); sometimes lacking. Surface otherwise with a few irregular, minute granules. Mesopleurae with a few small granules arranged in a longitudinal row, metapleurae as well as meso- and metasternum smooth. Tegmina oval, reaching about 1/3 the way along median segment and with a shallow central hump. Alae reaching to abdominal tergum VI.

Abdomen: Median segment considerably longer than metanotum, gently narrowed towards the posterior; smooth. Segments II–VII, elongate, cylindrical, parallel-sided and gradually decreasing in length; II almost 4x and VII just 2.6x longer than wide. Tergites II–VI and sternites II–VII smooth; tergites VI–X with a fine longitudinal median carina. VIII–X indistinctly broader than previous. VIII ¾ the length of VII and very slightly widened towards the posterior. IX a little shorter than VIII, slightly longer than wide and very gently constricted medially. Anal segment 3/5 the length of IX slightly widened towards the posterior and the posterior margin rounded with a broad triangular median indentation ( Fig. 331 ). Epiproct very small and hidden underneath anal segment. Vo m e r large, flattened and broadly triangular with a very short terminal hook ( Fig. 364 ). Cerci about 2/3 the length of anal segment, slender, strongly laterally compressed and very gently in-curving. Poculum moderately convex, cup-like, acutely carinate longitudinally and with the posterior margin notched medially; reaching about ¾ along tergum IX ( Figs. 330 , 364 ).

Legs: Generally as in ♀♀ but relatively longer and more slender. Profemora about equal in length to mesothorax, mesofemora shorter than mesothorax, metafemora reaching about 2/3 the way along abdominal segment IV.

Nymphs: Newly hatched nymphs are slender and have a body length of ± 16.0 mm. The body, fore legs and antennae are greenish brown, the head, mid and hind legs bright green and the tarsi brown. Later instars vary from dull green over various shades of brown, often irregularly furnished with pale or dark mottling. The legs are pale green to straw. It is interesting to note that nymphs generally resemble those of various Diapherodes and Haplopus - species, exhibiting rounded lateral lobes on abdominal tergum VII which disappear in the adult stage. Furthermore, the typical body shape of ♀♀ does not occur before the final ecdysis, even penultimate instar nymphs having a rather slender body.

Variability: In adults insects considerable variability is limited to the size and colouration. Slight variability is also seen in the length/width-relation of the mesothorax of ♀♀ as well as the size of the spines on the vertex, pronotum and anterior portion of the mesonotum (the latter ones may be missing). In addition to the usual plain green specimens ones may occur that are washed with dark brown, or have certain body-parts brownish ( Fig. 316 ).

Egg ( Figs. 332–333 ): Description as for the genus (see above). Measurements [mm]: Length 3.1–3.5, length (including operculum) 3.7–3.9, width 1.9–2.2, height 2.2–2.6, length of micropylar plate 1.1–1.2.

Comments: Audinet-Serville (1838: 242) described Platycrana venustula based on a single ♀ which he stated to be deposited in the “Collection de la Société entomologique de France ”. This is now held in MNHN but extensive search has not revealed a single specimen of this species in the entire collection. Therefore, the holotype must be regarded lost (Otte & Brock, 2005: 121) and a ♀ in MNHU is here selected as the neotype to guarantee stability of Audinet-Serville's taxon. Phasma havaniense Westwood, 1859 was described based on a ♀ from Havana in OXUM and was correctly synonymised with P. venustula by Saussure (1872: 185). The corresponding ♂♂ were described by Redtenbacher (1908: 429) as Haplopus juvenis which consequently becomes a synonym of V. venustula (n. syn.).

Culture-stock of V. venustula originating in the Pinar del Rio Province was imported to Germany by I. Fritzsche (Wernigerode, Germany) in 2005. This has proven fairly easy to rear in captivity and was included on the Phasmid Study Group culture-list as culture No. 283. A warm, rather dry and well ventilated climate is preferred in captivity. Eucalyptus ( Eucalyptus spp., Myrtaceae ), oaks ( Quercus robur , Q. rubra and Q. ilex , Fagaceae ), salal ( Gaultheria shallon , Ericaceae ) and St. John's Wort ( Hypericum patulum , Hypericaceae ) are accepted as alternative food plants. Eggs have a rather short incubation period of only 10–12 weeks at an average temperature of 25°C. About 3– 4 eggs are laid per day and ♀, each of which produces a total of 150– 200 eggs. Mating takes place exclusively at night and lasts for one to three hours.

Distribution ( Fig. 374 ): NW-Cuba (Havanna [OXUM]; Mariano [ANSP]; Somorrostro [ANSP]; Cabrera [MNCN];Sierra Rangel, Pinar del Rio [ANSP]; Pinar del Rio [coll. FH, coll. OC]).

Number of specimens examined: 140

Figures

FIGURES 322 – 325. Venupherodes venustula (Audinet-Serville, 1839) n. gen., n. comb .. 322. ♀: captive reared from NW- Cuba, Pinar del Rio [coll. FH, No. 0599 - 13]; 323. ♀: captive reared from NW-Cuba, Pinar del Rio [coll. FH, No. 0599 - 3]; 324. ♀: captive reared from NW-Cuba, Pinar del Rio [coll. FH, No. 0599 - 20]; 325. ♂: captive reared from NW-Cuba, Pinar del Rio [coll. FH, No. 0599 - 1].

FIGURES 326 – 333. Venupherodes venustula (Audinet-Serville, 1839) n. gen., n. comb .. 326. Head and pronotum of ♀: captive reared from NW-Cuba, Pinar del Rio [coll. FH, No. 0599 - 6]; 327. Head and pronotum of ♂: captive reared from NW- Cuba, Pinar del Rio [coll. FH, No. 0599 - 2]; 328. Apex of ♀ abdomen: captive reared from NW-Cuba, Pinar del Rio (lateral view); 329. Apex of ♀ abdomen: captive reared from NW-Cuba, Pinar del Rio (dorsal view); 330. Apex of ♂ abdomen: captive reared from NW-Cuba, Pinar del Rio (lateral view); 331. Apex of ♂ abdomen: captive reared from NW-Cuba, Pinar del Rio (dorsal view); 332. Egg: captive reared from NW-Cuba, Pinar del Rio (dorsal view) [coll. FH, No. 0599 - E]; 333. Egg: captive reared from NW-Cuba, Pinar del Rio (lateral view) [coll. FH, No. 0599 - E].

FIGURES 342 – 348. Ventral view of abdominal sternum VII of ♀♀ showing the praeopercular organ. 342. Haplopus sobrinus n. sp. ♀ HT: Cuba, Camagüey [ANSP]; 343. Parhaplopus navarroi n. gen., n. sp. ♀ PT: Dominican Republic, Santo Domingo Province, Santo Domingo [USNM]; 344. Parhaplopus cubensis (Saussure, 1868) n. gen., n. comb. ♀: Cuba, Havana [MNHN]; 345. Diapherodes venustula (Audinet-Serville, 1838) n. gen., n. comb. ♀: captive reared from NW-Cuba, Pinar del Rio [coll. FH, No. 0599 - 3]; 346. Diapherodes angulata (Fabricius, 1793) ♀ NT: Guadeloupe [MNHN]; 347. Diapherodes dominicae Rehn & Hebard, 1938 ♀ HT: Dominica [ANSP]; 348. Diapherodes gigantea gigantea (Gmélin, 1789) ♀: captive reared from Grenada [coll. FH, No. 0359 - 1].

FIGURES 357 – 364. Ventral view of anal segment of ♂♂ showing the vomer. 357. Haplopus bicuspidatus deHaan, 1842 ♂: captive reared from Dominican Republic [coll. FH, No. 0342 - 8]; 358. Haplopus brachypterus n. sp. ♂ HT: Dominican Republic, Santiago Province [USNM]; 359. Haplopus intermedius n. sp., ♂ HT: Dominican Republic, Valverde Province, nr. Maizal [USNM]; 360. Haplopus micropterus (St. Fargeau & Audinet-Serville, 1828) ♂: captive reared from Dominican Republic [coll. FH, No. 0243 - 90]; 361. Haplopus scabricollis Gray, 1835 ♂: Loggerhead Key [ANSP]; 362. Haplopus scabricollis Gray, 1835 ♂: Vanassa Island [USNM]; 363. Haplopus woodruffi n. sp. ♂ HT: Cayman Islands, Cayman Brac [FSCA]; 364. Venupherodes venustula (Audinet-Serville, 1838) n. gen., n. comb. ♂: captive reared from NW-Cuba, Pinar del Rio [coll. FH, No. 0599 - 1].

FIGURES 373 – 374. Maps. 373. Map of the West Indies showing the principle islands; 374. Known distribution of Aploploides Rehn & Rehn, 1938 [green] and Venupherodes n. gen. [red].

FIGURES 382 – 384. Live insects. 382. Diapherodes achalus (Rehn, 1904) ♀: Puerto Rico, Maricao [Ó Mel J. Rívera]; 383. Diapherodes achalus (Rehn, 1904) ♂: Puerto Rico, Maricao [Ó Mel J. Rivera]; 384. Venupherodes venustula (Audinet-Serville, 1838) n. gen., n. comb. ♀: captive reared from NW-Cuba, Pinar del Rio.

FIGURES 385 – 387. Live insects. 385. Venupherodes venustula (Audinet-Serville, 1838) n. gen., n. comb. ♂: captive reared from NW-Cuba, Pinar del Rio; 386. Venupherodes venustula (Audinet-Serville, 1838) n. gen., n. comb. ♀ and ♂: captive reared from NW-Cuba, Pinar del Rio; 387. Venupherodes venustula (Audinet-Serville, 1838) n. gen., n. comb. ♀♀ usual green colour-form and specimen with brown mottling: captive reared from NW-Cuba, Pinar del Rio.

FIGURES 315 – 321. Parhaplopus navarroi n. gen., n. sp .. 315. Apex of abdomen of ♀ PT (lateral view) [USNM]; 316. Apex of abdomen of ♀ PT (dorsal view) [USNM]; 317. Apex of abdomen of ♂ HT (lateral view) [USNM]; 318. Apex of abdomen of ♂ HT (dorsal view) [USNM]; 319. Right tegmen and ala of ♂ HT [USNM]; 320. Egg PT (dorsal view) [USNM]; 321. Egg PT (lateral view) [USNM].