Phantasca Redtenbacher, 1906 : 111 Phasma phantasma Westwood, 1859 : 126 Pterolibethra Günther, 1940 : 498 Pterolibethra heteronemia Günther, 1940 : 499 Phantasca Pterolibethra Günther, 1940 Heteronemiinae Diapheromerinae Diapheromerini Clonistria Pterolibethra Heteronemiinae Diapheromerinae Diapheromerini Clonistria Bacteria Phasma Studies on neotropical Phasmatodea XVII: Revision of Phantasca Redtenbacher, 1906, with the descriptions of six new species (Phasmatodea: Diapheromeridae: Diapheromerinae) Hennemann, Frank H. Conle, Oskar V. Bellanger, Yannick Lelong, Philippe Jourdan, Toni European Journal of Taxonomy 2018 2018-05-15 435 1 62 8NXFX Redtenbacher, 1906 Redtenbacher 1906 [653,1017,706,732] Insecta Diapheromeridae Phantasca GBIF Animalia Phasmida 2 3 Arthropoda genus    Phantasca Redtenbacher, 1906: 111(→ Phasmini) ( typespecies: Phasma phantasma Westwood, 1859: 126, pl. 12, fig. 5a–b ( ♂), by subsequent designation of Zompro 2001: 223).   Pterolibethra Günther, 1940: 498( typespecies:  Pterolibethra heteronemia Günther, 1940: 499by original designation) syn. nov.   Phantasca– Günther 1940: 500 (  Pterolibethra Günther, 1940listed as a synonym). — Bradley & Galil 1977: 180 (→ Heteronemiinae: Libethrini). — Zompro 2001: 223, fig. 32 (designation of typespecies, → Diapheromerinae: Diapheromerini: „  Clonistria-group“). — Bragg 2001: 641. — Zompro 2004a: 318. — Zompro 2004b: 142. — Otte & Brock 2005: 264. — Jourdan, Lelong & Bellanger 2014: 490.   Pterolibethra– Bradley & Galil 1977: 180 (→ Heteronemiinae: Libethrini). — Zompro 2001: 225 (reestablished as a valid genus, → Diapheromerinae: Diapheromerini: „  Clonistria-group“). — Bragg 2001: 643. — Zompro 2004a: 320. — Otte & Brock 2005: 294. — Conle, Hennemann & Gutiérrez 2011: 56. non  Bacteria– Chopard 1911: 343 (in part). non  Phasma– Westwood 1859: 117 (in part).    Description Small to medium sized and mostly very slender Diapheromerinae(body length ♂♂<75.0 mm, females <130.0 mm), with strong sexual dimorphism; females apterous, males much more slender and very delicate winged insects. Body of uniform width and smooth or, with mesothorax at best sparsely granulose. Colour of females mostly green, grey or various shades of brown. Males usually ranging from green to brown but may be fairly colorful; often, with conspicuous dark markings or stripes on head, pronotum, mesonotum and abdominal tergites VII–X or, with annulated legs or colorful apices and/or bases of femora and tibiae. Anal region of alae of males transparent but dark brown, with the basal portion orange in a single species. Head longer than wide,, with vertex flattened or at best gently convex; unarmed. No ocelli. Gula covering more than ¼ of cervical membrane. Eyes large and projecting hemispherically in males. Antennae long and filiform, at least ¾ the length of body in males and more than ⅔ the length of body in females. Scapus compressed dorsoventrally and longer than wide; rectangular. Pedicellus cylindrical and antennomere III considerably longer than preceding or following three antennomeres. Pronotum longer than wide and narrower than head. Mesothorax very elongate,> 2× as long as than head and pronotum in females and up to 3.5× as long as head and pronotum combined in males. Mesosternum simple. Tegmina of males slender, squamiform,, with apical portion angular and basal portion strongly narrowed. Alae of males of moderate length and reaching to abdominal segment IV or V; anal region considerably longer than wide. Median segment> ⅔ the length of metanotum in females and several times longer than metanotum in males. Abdominal segment II of males shorter than median segment. II–VII longer than wide and roughly of uniform width in both sexes. VI in females occasionally, with a pair of swellings. Sterna smooth but VII of females, with an indistinct praeopercular organ formed by a node or wart-like posteromedian swelling. Stigmae of tergum VIII close to anterior margin of segment. Epiproct of females small, scale-like and at best slightly projecting from apex of anal segment. Cerci of females straight, cylindrical and variable in length. Gonapophyses VIII slender and longer than IX and at best reaching to apex of subgenital plate. Gonoplacs distinct and equal in length to slightly longer than gonapophyses IX. Subgenital plate small, short and carinate longitudinally,, with apex ± acutely pointed; variable in length but at best reaching to apex of abdomen. Terminalia of males, with conspicuous specialisations. Tergum and sternum VIII partly to almost fully fused, with lateral margins of tergum VIII, often no more than indicated by a fine fissure. Sternum VIII ± enlarged and fused, with sternum IX, which is reduced and has its borders at best indicated by an obsolete fissure. Poculum small, scaphiform and variable in shape,, with apex specialised. Tergum IX, with lateral margins approaching each other ventrally and partly or completely fused, with each other; in the latter case forming a closed tube. Anal segment much shorter than tergum IX, variable in shape and with posterior margin ± distinctly notched medially; ventral surface of posterior margin, with a pair of small, ± developed thorn pads. Vomer increasingly reduced, variable in size, ranging from fairly distinct to very small and triangular in shape,, with a single terminal hook. Cerci very variable in size and shape and of specific structure; ranging in length from being no longer than anal segment to more than twice the length of that segment, and in shape from straight, over gently upcurving or arcuate to strongly incurving and hook-like. Legs of both sexes elongate and ± slender (males in particular) and entirely unarmed. Profemora compressed and curved basally, trapezoidal in cross-section,, with dorsal carinae strongly approaching and anterodorsal carina distinctly raised; medioventral carina moderately distinct and somewhat displaced towards anteroventral carina. Meso- and metafemora and all tibiae trapezoidal in cross-section,, with a fine longitudinal median carina. In females the two outer ventral carinae of meso- and metafemora occasionally, with a sub-apical rounded lobe or medioventral carina of meso- and metatibiae protuded into a rounded lobe sub-basally. Cerci slender and without specializations. Basitarsus strongly elongated (males in particular) and considerably longer than following three tarsomeres combined; trapezoidal in cross-section and sulcate dorsally. Eggs: Capsule ovoid, longer than wide and ± compressed laterally. Surface ranging from almost smooth and glossy over minutely punctured to slightly rugulose; ± glossy. Micropylar plate elongate, slender and roughly parallel-sided,, with only posterior portion very gently widened;> ¾ the length of capsule. Micropylar cupe near posterior end of plate. Median line distinct and variable in length. Internal plate open,, with a short median line that is distinctly separated from the plate itself. Operculum oval and inserted into capsule at a right angle or at best, with an angle of 10°. In centre, with a low, open and net- or sponge-like capitulum. Colour mid to very dark brown.    Diagnosis The relationships of  Phantascaare not yet fully clarified, but in several aspects a relationship to sections of  LibethraStål, 1875( strigiventrisspecies group in particular) and certain Jamaican species of  ClonistriaStål, 1875(e.g.,  C. annulipes Rehn & Hebard, 1938,  C. bicoloripes Rehn & Hebard, 1938,  C. latebricola Rehn & Hebard, 1938,  C. monticola Rehn & Hebard, 1938) appear likely (see remarks below). From representatives of the strigiventrisspecies group of  Libethra, both sexes of  Phantascamay be distinguished by the much longer median segment, which is several times longer than the metanotum in males and at least ⅔ the length of the metanotum in females. Males also differ by the presence of wings, smaller poculum, partially to almost fully fused abdominal tergum and sternum VIII and the not conspicuously node-like lateral swellings of the anal segment, females can readily be distinguished by the unspecialised apex of the subgenital plate (deeply median notched and bifid in  Libethra). The eggs are averagely less elongate than in  Libethraand have the operculum inserted into the capsule roughly at a right angle (inserted at an angle of about 70° in  Libethra) From the above mentioned Jamaican species of  ClonistriaStål, 1875representatives of  Phantascaare readily distinguished by the considerably longer median segment and presence of wings in males. While the genital morphology is very similar in most aspects, males of  Phantascahave the stigmae of abdominal tergum VIII very close to the anterior margin (somewhat displaced towards the posterior in the mentioned species of  Clonistria), the poculum averaging smaller, and tergum and sternum VIII partly to almost fully fused. Differences between the females are as yet difficult to define, since only the female of one of the Jamaican species of  Clonistriais known.    Remarks There has been uncertainty about the systematic position of  Phantasca Redtenbacher, 1906, mainly because the genus was precedingly only known from the males. Redtenbacher (1906: 111)originally established  Phantascafor three small and very slender, winged species described by Westwood (1859), namely  Phasma phantasma( Westwood, 1859),  Ph. puppeius( Westwood, 1859)and  Ph. valgius( Westwood, 1859), and placed the genus in his areolate tribe Phasmini. Günther (1940: 500)recognized that none of these species exhibits an area apicalis and suggested a close relationship to  LibethraStål, 1875and the strigiventrisspecies-group of that genus in particular (see Günther 1932: 258; Conle, Hennemann & Gutiérrez 2011: 205). Zompro (2001: 220)placed  Phantascain his “  Clonistria-group” of the subfamily Diapheromerinae: Diapheromeriniand stated the genus to “show similarity” to  ClonistriaStål, 1875( typespecies:  Clonistria bartholomaeaStål, 1875) and  LibethroideaHebard, 1919( typespecies:  Libethroidea inusitataHebard, 1919) in being slender and lacking conspicous metamorphosis of the body. While this assumption is fairly meaningless, the generic diagnosis of  Phantascapresented by Zompro (2001: 223)was only based on the males and is inaccurate in some aspects. The mentioned characters “Metathorax half as long as metathorax [should be: mesothorax] and 3 times as long as median segment” and “[abdominal segment] II less than two times as long as median segment” are not true, since the median segment is several times longer than the metanotum.  Günther (1940: 498)described  Pterolibethra( typespecies:  Pterolibethra heteronemia Günther, 1940) for the two newly described species  P. heteronemiaand  P. poecilopteraand stated that it was closely related to  LibethraStål, 1875, but in the appendix of the same publication ( Günther 1940: 500) he synonymised  Pterolibethra, with  Phantasca.This synonymy was revised by Zompro (2001: 225), who re-establised  Pterolibethraas a valid genus on the basis that the genitalia of  Phantascaresembled those of  Clonistriaand those of  Pterolibethrawere similar to those of  Libethra. However, the “similarities” postulated were not defined in any more detail by Zompro and examination of all concerned species does in fact not show any differences in the genital morphology that would distinguish  Pterolibethrafrom  Phantasca. Hence, the synonymy established by Günther (1940: 500)is correct and  Pterolibethrais consequently re-synonymised, with  Phantascaherein (syn. nov.). Zompro (2001: 226)furthermore stated that Günther was in error about the length of the median segment of  Pterolibethra heteronemiaand that it was obvious that the median segment measured only two-thirds of the metanotum. In fact, however, the median segment of the concerned species is several times longer than the metanotum, these two segments showing a similar length relationship as in all other species of  Phantasca. The true systematic position and relationships of  Phantascaare still not fully clarified, but the conspicuous genital morphology of the males suggests close relationships to sections of the South American  LibethraStål, 1875(the strigiventrisspecies group in particular), certain species of the exceptional Caribbean  ClonistriaStål, 1875from the Greater Antilles as well as some species of as yet unknown generic position that are distributed throughout southern Braziland  Paraguay. Furthermore, the often partly to almost fully fused abdominal tergum und sternum VIII, increasingly approaching and sometimes partly ventrally fused lateral margins of tergum IX, as well as the small poculum in males of  Phantascaindicate a relationship to  Caribbiopheromera Zompro, 2001( typespecies:  Caribbiopheromera jamaicana Zompro, 2001), which does obviously not belong in a close relationship to the “  Diapheromeragroup” of Diapheromerinae: Diapheromeriniwhere it was originally placed by Zompro ( Bradler 2009: 102). In addition to the genital morphology of males, a relationship between  Phantascaand the aforementioned genera is also supported by the morphology of the genitalia of females and egg morphology. However, any broader discussion on the placement and relationships of  Phantascaappears premature at this point and apart from a better knowledge of the as yet generically unassigned but possibly closely related species that occur throughout southern Braziland  Paraguay(see above) warrant a comprehensive phylogenetic study that incorporates all of the mentioned genera and support by molecular analyses. These are the subject of forthcoming studies conducted by the authors and will hopefully provide clarification of the true systematic position of  Phantasca. Examination of the genital morphology of the male (the only sex known) of  Pterolibethra laeta Conle, Hennemann & Gutiérrez, 2011from Colombiahas shown it to be misplaced. Although the poculum is comparatively small, the short abdominal tergum VIII, which is strictly separated from sternum VIII, the large vomer and broad, dorsoventrally flattened and hook-like cerci place this species in the genus  Jeremiodes Hennemann & Conle, 2007, hence now  Jeremiodes laetus(comb. nov.).    Distribution North Brazil,  French Guiana, east Peru, east Ecuadorand southeast Colombia. The genus is very likely to also be represented in Suriname, but there are no records so far.    Species included 1.  Phantasca adiposa sp. nov.(  French Guiana) 2.  Phantasca amabile sp. nov.( Ecuador: NapoProv.) 3.  Phantasca femorata sp. nov.(  French Guiana) 4.  Phantasca guianensis sp. nov.(  French Guiana) 5.  Phantasca montana( Redtenbacher, 1908) comb. nov.(east Peru: Dept. Loreto) 6.  Phantasca heteronemia( Günther, 1940)comb. rev. (east Peru: Dept. Loreto) 7.  Phantasca nigrolineata sp. nov.(  French Guiana) 8.  Phantasca phantasma( Westwood, 1859)(northeast Brazil&  French Guiana) 9.  Phantasca poeciloptera( Günther, 1940)comb. rev. (southeast Colombia& East Peru) 10.  Phantasca puppeia( Westwood, 1859)(northeast Brazil(Tapajós)) 11.  Phantasca quadrilobata( Chopard, 1911) comb. nov.(  French Guiana) 12.  Phantasca ruboligata sp. nov.(  French Guiana) 13.  Phantasca valgius( Westwood, 1859)(not known)    Key to the species of  Phantasca  Males    1. Cerci short, arcuately upcurving and not projecting over posterior margin of anal segment …………2  – Cerci long and projecting distinctly over posterior margin of anal segment …………………………3   2. Apex of cerci blunt; poculum large,, with posterior margin labiate and angled downward ( Fig. 68); posteriormarginofanalsegment,withawidetriangularmedianexcavation;head,withoutadistinctpale marking on frons ( Fig. 67) …………………………………………  P. phantasma( Westwood, 1859)  – Apex of cerci acutely pointed; poculum small, cup shaped ( Fig. 96),, with the posterior margin obtusely angulate and dorsally directed; posterior margin of anal segment, with a small median indentation ( Fig. 95); head, with a large, distinctly pale cream to whitish marking on frons ( Fig. 92) ………… ……………………………………………………………………  P. quadrilobata( Chopard, 1911)   3. Cerci angled inward and ± hook-shaped ……………………………………………………………4  – Cerci straight or at best gently incurving ……………………………………………………………6   4. More or less plain green or brown insects; anal fan of alae transparent ………………………………5  – Very colourful insects, with head, most of legs and cerci red ( Fig. 71); anal fan of alae dark brown, with the basal half bright orange ( Fig. 74); SE Colombia& E Peru…  P. poeciloptera( Günther, 1940)   5. Very slender, bright green insects; scapus, coxae and bases of all tibiae bright red; base of alae, with a black marking interiorly; posterior margin of poculum slightly narrowed and gently bilobate ( Fig. 114);  French Guiana………………………………………………………  P. ruboligata sp. nov.  – Stocky, dull greenish brown; abominal sternites II–V black; base of alae, without a dark marking; posterior margin of poculum widened, obtusely angular and preceded by a conspicuous narrowing ( Fig. 44); E Peru…………………………………………………  P. heteronemia( Günther, 1940)   6. Head ± unicoloured,, without conspicuous dark markings …………………………………………7  – Head, with distinct and well defined dark markings or lines ………………………………………9   7. Very delicate insects; head indistinctly wider than prothorax ………………………………………8  – Comparatively stocky; head globose and almost 2× as wide as prothorax ( Fig. 8) …  P. adiposa sp. nov.   8. All longitudinal veins of tegmina and alae marked, with brown ( Fig. 120) ……………………… …………………………………………………………………………  P. valgius( Westwood, 1859)  – Tegmina and costal region of alae plain green ( Fig. 78) …………  P. puppeia( Westwood, 1859)   9. Head, with a single bold central black marking or longitudinal stripe ………………………………10  – Head, with various dark markings …………………………………………………………………11   10. Colourful insects, green, with bases of tibiae red and apices of femora and tibiae black; costal region of alae, with a dark brown longitudinal stripe interiorly ( Fig. 15); posterior margin of anal segment bilobate, with a deep and narrow median incision; E Ecuador………  P. amabile sp. nov.  – Greenish brown insects; no dark longitudinal stripe on costal region of alae; posterior margin of anal segment only, with a very slight median indentation ( Fig. 59);  French Guiana…  P. nigrolineata sp. nov.   11. Anal segment ± cucullate; cerci short and projecting beyond apex of abdomen by no more than half the length of the anal segment ……………………………………………………………………12  – Anal segment flattened and almost parallel-sided, the posterior margin, with a wide triangular excavation ( Fig. 30); cerci very long and projecting beyond apex of abdomen by almost the length of the anal segment;  French Guiana…………………………………………  P. guianensis sp. nov.   12. Head elongate, cheeks parallel-sided and with two fine dark longitudinal postocular stripes ( Fig. 21); pronotum, with a fine longitudinal black line along lateral margins; poculum, with lateral margins roundly emarginated and apical half strongly narrowed ( Fig. 24);  French Guiana……… ………………….........................………………………………………………  P. femorata sp. nov.  – Head ovoid, vertex, with two bold diverging dorsal stripes ( Fig. 47); pronotum, without a dark lateral line; poculum cup-shaped, with lateral margins straight and posterior portion not conspicuously narrowed; East Peru……………………………………………  P. montana( Redtenbacher, 1908)   Females*     1. Cerci at best equal in length to anal segment ………………………………………………………2  – Cerci lanceolate and distinctly longer than the anal segment ( Figs 106–108) …  P. ruboligata sp. nov.   2. Cerci elongate and> ¾ the length of the anal segment; femora and tibiae, without lobes ……………3  – Cerci <⅔ the length of the anal segment; femora or tibiae, with lobes ………………………………4   3. Very slender insects ( Figs 48–49), abdominal segments II–VII> 2× as long as wide; femora slender; subgenital plate flattened ( Fig. 54) ………………………………………  P. nigrolineata sp. nov.  – Stockyinsects ( Figs1–2), abdominalsegmentsII–VIIindistinctlylongerthanwide; femorabroadened; subgenital plate strongly convex medially ( Fig. 4) ……………………………  P. adiposa sp. nov.   4. Abdominal tergum VI smooth dorsally ………………………………………………………………5  – AbdominaltergumVI, withapairoftransverselycarinateswellings…  P.quadrilobata( Chopard,1911)   5. Large insects (body length> 12 cm); abdominal tergum VI slender; meso- and metatibiae, with a rounded sub-basal lobe on medioventral carina ……………………………  P. guianensis sp. nov.  – Smaller insects (body length < 12 cm); lateral margins of abdominal tergum VI roundly deflexed; meso- and metafemora, with a rounded sub-apical lobe on both outer ventral carinae and these carinae gently deflexed sub-basally<( Fig. 20) …………………………………  P. femorata sp. nov.  * The females of  P. amabile sp. nov.,  P. heteronemia( Günther, 1940),  P. montana( Redtenbacher, 1906),  P. phantasma( Westwood, 1859),  P. poeciloptera( Günther, 1940),  P. puppeia( Westwood, 1859)and  P. valgius( Westwood, 1859)are not known.