Cryptophyllium icarus gen. et, 2021

Cumming, Royce T., Bank, Sarah, Bresseel, Joachim, Constant, Je ́ ro ̂ me, Tirant, Stephane Le, Dong, Zhiwei, Sonet, Gontran & Bradler, Sven, 2021, Cryptophyllium, the hidden leaf insects - descriptions of a new leaf insect genus and thirteen species from the former celebicum species group (Phasmatodea, Phylliidae), ZooKeys 1018, pp. 1-179 : 1

Cryptophyllium icarus gen. et sp. nov. Figures 5H , 8I , 8J , 9I , 32 , 33 , 34 , 35 , 36

Material examined.

Holotype ♂: "Coll. I.R.Sc.N.B., Ex breeding Tim Bollens, 2018, Coll. I.R.Sc.N.B., Vietnam, Lam Dong prov., Bidoup-Nui Ba N.P., 12°26'N, 108°30'E, 21-25.vii.2014, Leg. J. Constant and J. Bresseel, GTI Project, I.G.: 32.779". Deposited in the Royal Belgian Institute of Natural Sciences (RBINS).

Paratypes: (26 ♀♀, 49 ♂♂, 78 eggs) • 1 ♂: "Coll. I.R.Sc.N.B., Ex breeding Tim Bollens, 2018, Coll. I.R.Sc.N.B., Vietnam, Lam Dong prov., Bidoup-Nui Ba N.P., 12°26'N, 108°30'E, 21-25.vii.2014, Leg. J. Constant and J. Bresseel, GTI Project, I.G.: 32.779" [vomer dissected] (RBINS) • 5 ♀♀, 4 ♂♂: "Coll. I.R.Sc.N.B., Ex breeding Tim Bollens, 2018, Coll. I.R.Sc.N.B., Vietnam, Lam Dong prov., Bidoup-Nui Ba N.P., 12°26'N, 108°30'E, 21-25.vii.2014, Leg. J. Constant and J. Bresseel, GTI Project, I.G.: 32.779" (RBINS) • 1 ♀, 1 ♂: "Coll. I.R.Sc.N.B., Ex breeding Tim Bollens, 2018, Coll. I.R.Sc.N.B., Vietnam, Lam Dong prov., Bidoup-Nui Ba N.P., 12°26'N, 108°30'E, 21-25.vii.2014, Leg. J. Constant and J. Bresseel, GTI Project, I.G.: 32.779" (VNMN) • 1 ♀, 5 ♂♂: "Coll. I.R.Sc.N.B., Ex breeding Tim Bollens, 2019, Coll. I.R.Sc.N.B., Vietnam, Lam Dong prov., Bidoup-Nui Ba N.P., 12°26'N, 108°30'E, 21-25.vii.2014, Leg. J. Constant and J. Bresseel, GTI Project, I.G.: 32.779" (RBINS) • 1 ♀, 1 ♂: "Coll. I.R.Sc.N.B., Ex breeding Bruno Kneubühler, 2017, Coll. I.R.Sc.N.B., Vietnam, Lam Dong prov., Bidoup-Nui Ba N.P., 12°26'N, 108°30'E, 21-25.vii.2014, Leg. J. Constant and J. Bresseel, GTI Project, I.G.: 32.779" (RBINS) • 6 ♀♀, subadult ♀, ♂: "Coll. I.R.Sc.N.B., Vietnam, Lam Dong prov., Bidoup-Nui Ba N.P., 12°26'N, 108°30'E, 21-25.vii.2014, Leg. J. Constant and J. Bresseel, GTI Project, I.G.: 32.779" (RBINS) • 1 ♂; "Vietnam, Lam Dong, Bao Loc" (molecular sample SLT005 within our analysis) (Coll SLT) • 3 ♂♂; "ex Zucht F. Hennemann 2019, Herkunft: Z-Vietnam, Lâm Dông Prov., Dam Rông, Bidoup Nui Ba NP, leg. Bresseel and Constant VII.2014 [ex coll. FH]" (Coll SLT) • 1 ♀; "VIETNAM, Lam Dong Prov., Dam Rong, Bidoup-Nui Ba NP, Leg. Bresseel and Constant VII.2014. Ex breeding F. Hennemann 2015-2017. Ex coll. F. H. Hennemann (Germany); Coll RC 18-226" (Coll RC) • 4 ♂♂; "ex Zucht F. Hennemann 2019, Herkunft: Z-Vietnam, Lâm Dông Prov., Dam Rông, Bidoup Nui Ba NP, leg. Bresseel and Constant VII.2014 [ex coll. FH]", Coll RC 20-099-20-102 (Coll RC) • 1 ♂; "VIETNAM, Dak Lak Prov., May 2018. Coll RC 18-408" (Coll RC) • 1 ♂; "VIETNAM, Lam Dong Prov., Dam Rong, Bidoup-Nui Ba NP, Leg. Bresseel and Constant VII.2014. Ex breeding F. Hennemann 2015-2017. Ex coll. F. H. Hennemann, Germany; Coll RC 18-225" (Coll RC) • 1 ♂; "VIETNAM, Lam Dong Prov., Bao Loc, October 2016, Coll RC 16-248" (Coll RC) • 1 ♂; "VIETNAM, Lam Dong Prov., Bao Loc, May 2016, Coll RC 17-264" (Coll RC) • 3 ♂♂; "VIETNAM, Lam Dong Prov., Bao Loc, July 2017", Coll RC 17-266, 17-267, 17-268 (Coll RC) • 1 ♂; "VIETNAM, Lam Dong Prov., Bao Loc, June 2017. Coll RC 17-265" (Coll RC) • 2 eggs: "VIETNAM, Lam Dong Prov., Dam Rong, Bidoup-Nui Ba NP, Leg. Bresseel and Constant VII.2014. Ex breeding F. Hennemann 2015-2017. Ex coll. F. H. Hennemann, Germany"; Coll RC 18-238 and 18-239 (Coll RC) • 20 eggs: "VIETNAM: Lam Dong Prov., Bidoup Nui Ba N.P., culture from Maxime Ortiz (France), 2018". Coll RC 18-343-18-362 (Coll RC) • 7 ♀♀, 1 ♂; "VIETNAM: Lam Dong Prov., Bidoup Nui Ba N.P., bred by Bruno Kneubühler (Switzerland), circa 2016" (Coll OC) • 3 ♀♀, 7 ♂♂, 28 eggs: "ex Zucht F. Hennemann 2018, Herkunft: Z-Vietnam, Lâm Dông Prov., Dam Rông, Bidoup Nui Ba NP, leg. Bresseel and Constant VII.2014" [coll. FH, No’s 0896-1 to 10, E1] (Coll FH) • 1 ♀, 7 ♂♂, 1 ♂ nymph n5, 28 eggs; "ex Zucht F. Hennemann 2019, Herkunft: Z-Vietnam, Lâm Dông Prov., Dam Rông, Bidoup Nui Ba NP, leg. Bresseel and Constant VII.2014" [coll. FH, No’s 0896-11 to 19, E2], (Coll FH) • 1 ♀: "ex Zucht F. Hennemann 2020, Herkunft: Z-Vietnam, Lâm Dông Prov., Dam Rông, Bidoup Nui Ba NP, leg. Bresseel and Constant VII.2014" [coll. FH, No’s 0896-20], (Coll FH) • 3 ♀♀, 4 ♂♂; "Vietnam, Lâm Dông Prov., Dam Rông, Bidoup Nui Ba, bred by Maxime Ortiz, France, circa 2020" (Coll MO).

Remarks.

Cryptophyllium icarus sp. nov. was first collected in 2014 by Joachim Bresseel (RBINS) and Jérome Constant (RBINS) from Bidoup Nui Ba N.P., Vietnam. Several adult females and a subadult male nymph were found on small trees between the side of the road and next to a high cliff with primary forest on top (Fig. 32C, D ). This species was successfully brought into the phasmid breeding community which allowed observation of morphological variation in the females and description of the adult male, egg, and freshly hatched nymph morphology (Fig. 33 ). Interestingly, a significant change in abdominal shape occurred when this species was reared in captivity vs. the wild collected females. The wild collected paratype females (Fig. 32A ) has a slenderer abdomen, with segments VI and VII notably converging towards the posterior with only a slight bend near the posterior of segment VII, vs. individuals reared in captivity (Fig. 33A ) which instead develop a rather boxy abdominal shape, with segments VI and the majority of VII parallel-sided and ending in a distinct lobe on the posterior of VII. Interestingly all wild caught females were slender and all bred females have been boxy; at this time we are unsure what mechanism is promoting this polymorphism within this species. Hopefully additional field work in the region will reveal additional observations to review.

An additional unique feature for this species is the coloration of the freshly hatched nymphs, which do not match well with the coloration of other known congenerics. This could be due to our lack of knowledge as freshly hatched nymph coloration is known only for nine species (Fig. 9 ). Molecularly, we found Cryptophyllium icarus sp. nov. to be sister species to Cryptophyllium limogesi sp. nov. whose freshly hatched nymph morphology we do not know. It is possible that these may share similar coloration, but at the present Cryptophyllium icarus sp. nov. with their solid red abdomen and dark extremities (Fig. 9I ) appear quite unique within the Cryptophyllium gen. nov. as presently known.

Differentiation.

Female morphology is superficially similar to Cryptophyllium daparo sp. nov. and Cryptophyllium chrisangi comb. nov. due to the tapered abdominal shape, obtusely rounded profemoral exterior lobes, and similarly shaped and textured thorax. Cryptophyllium icarus sp. nov. can be differentiated from both species by the highly reduced alae not reaching abdominal segment II (Fig. 33C ), vs. the other species which both have well-developed alae.

Males with their thorax shape and spination, the spade-shaped abdomen, shorter tegmina, and exterior profemoral lobe which is thinner than the interior, are morphologically similar to Cryptophyllium bankoi sp. nov. and Cryptophyllium rarum comb. nov. males. From both species, Cryptophyllium icarus sp. nov. can be differentiated by the width of the profemoral lobe which in Cryptophyllium icarus sp. nov. is only ca. 2 × as wide as the profemoral shaft, vs. the other species which are at least 2½ or 3 × as wide as the profemoral shaft. Additionally, males of Cryptophyllium rarum comb. nov. are notably larger than the largest Cryptophyllium icarus sp. nov. males.

Distribution.

Only presently known from southern Vietnam, from the provinces of Lam Dong and Dak Lak.

Description.

Female. Coloration. Coloration is variable, as when it was bred in captivity females were almost uniformly pale green, with only slightly orange/tan areas along the profemoral lobes, antennae, eyes, thorax, and terminal abdominal margins (Fig. 33A ). In contrast, the paratype female when she was found in the wild was quite colorful with a vibrant green body with highlights of red throughout the antennae, legs, thorax, tegmina, and abdomen (Fig. 32A ), much darker and more plentiful than on captive bred specimens.

Morphology. Head. Head capsule longer than wide, vertex relatively smooth, with the only notable feature the posteromedial tubercle which is not notably broad but is significantly raised above the head capsule (Fig. 34E ). Frontal convexity broad and stout, with a lumpy surface, and with several setae throughout (Fig. 34E ). Compound eyes slightly protruding from the head capsule, not notably large, only taking up slightly <1/4 of the length of the lateral head capsule margins (Fig. 34E ). Ocelli absent. Antennal fields slightly wider than the first antennomere (Fig. 34C ). Antennae. Antennae consisting of nine segments, with the terminal segment about as long as 2½× the preceding segments lengths (Fig. 34C ). Antennomeres I-VIII sparsely marked with small transparent setae, the terminal antennomere and the distal margin of antennomere VIII has darker, shorter, and denser setae than the other segments (Fig. 34C ). Thorax. Pronotum with a slightly concave anterior margin and slightly convex lateral margins, which converge to a straight posterior margin that is half the width of the anterior margin (Fig. 34E ). The pronotum surface and moderately formed pronotum rims are only slightly lumpy, lacking significant granulation, with only a prominent pit in the center, and slight furrows anterior, posterior, and lateral to the pit (Fig. 34E ). Prosternum with moderate granulation, mesosternum anterior half and lateral margins with moderate granulation (Fig. 34B ). Metasternum relatively smooth, lacking notable nodes. Prescutum slightly longer than wide, lateral rims with four larger nodes and four or five nodes interspersed throughout (Fig. 34E ). Prescutum anterior rim prominent but not strongly protruding, surface marked throughout with irregular granulation, no prominent singular sagittal spine present (Fig. 34F ). Prescutum surface covered densely by small tubercles and numerous nodes, with those along the sagittal plane the largest (Fig. 34E ). Mesopleura beginning slightly posterior to the anterior margin of the prescutum and evenly diverging; lateral margin with five larger tubercles, and five or six smaller tubercles interspersed unevenly throughout (Fig. 34E ). Face of the mesopleura slightly wrinkled, with a distinct pit near the anterior margin and one near the center (Fig. 34E ). Wings. Tegmina long, reaching ½ way through abdominal segment VII. The subcosta (Sc) is the first vein in the forewing and runs parallel with the tegmina lateral margin for the first half of the vein, then bends gently and runs to the to the lateral margin of the wing where it terminates ca. ⅓ through the length. The radius (R) spans the central portion of the forewing with two subparallel branched veins; radius 1 (R1) terminates slightly proximal to the midline, and the radial sector (Rs) terminates ca. ⅔ of the way through the wing length. There is a weak continuation of the radius following the prominent Rs branching which continues on as a short and thinner R-M crossvein that does not solidly connect the two veins as it reaches the media. The media (M) is simply bifurcate with both the media anterior (MA) and media posterior (MP) terminating on the posterior ¼ of the wing. The cubitus (Cu) is also bifurcate, branching near the posterior ⅕ of the wing into the cubitus anterior (CuA) and cubitus posterior (CuP) which both terminate at or very near the wing posterior apex. The first anal vein (1A) is simple and fuses with the cubitus early on, halfway between the branching of the R1 and the Rs. Alae are highly reduced, not reaching the anterior margin of abdominal segment II. Abdomen. Abdominal segments II through the anterior half of IV diverge uniformly towards the posterior. The posterior half of IV and segment V are parallel-sided and are the widest portion of the abdomen. Segment VI and VII are variable depending on the environmental conditions, with wild collected females (such as in the paratype; Fig. 32A ) having segments VI and VII strongly converging and VII ending with a small bulge on the posterior margin with a final width about the same as the anterior margin of segment VIII, vs. captive bred individuals generally having VI and VII subparallel and ending with VII having a distinct lobe which is notably wider than the anterior margin of segment VIII (Fig. 33A ). Segments VIII-X converge uniformly to the rounded apex. Genitalia. Subgenital plate starts at the anterior margin of segment VIII, is long and narrow reaching ca. ½ onto segment X (Fig. 34H ). Gonapophyses VIII are long and significantly broad, each about as wide as the subgenital plate projection, with their tips reaching the apex of segment X, gonapophyses IX are smaller and slender, hidden below the gonapophyses VIII (Fig. 34H ). Cerci flat, not strongly cupped, with a heavily granular surface and a setae throughout (Fig. 34H ). Legs. Profemoral exterior lobes notably wider than the interior lobe with a rounded obtuse angle (Fig. 34D ). Exterior lobe margin is not marked by teeth and is instead rather smooth or at most slightly granular (Fig. 34D ). Profemoral interior lobe ca. 2 × as wide as the greatest width of the profemoral shaft, with an obtuse angle, and marked with five prominent teeth arranged in a two-one-two pattern with large looping gaps between the teeth (Fig. 34D ). Mesofemoral exterior lobe arcs from end to end in a slightly bent lobe slightly weighted to the distal ½ and marked with two or three small serrate teeth on the distal ½ only. Interior and exterior lobes are of a similar width, or the exterior is slightly wider. Mesofemoral interior lobe arcs smoothly end to end, is marked with six or seven serrate teeth only on the distal half of the arc, and is about as wide as the mesofemoral shaft. Metafemoral interior lobe arcs end to end, but is slightly wider on the distal half, and has six or seven serrate teeth on the distal half of the lobe only. Metafemoral exterior lobe is thin and smooth, hugging the metafemoral shaft and lacks dentation. Protibiae interior lobe spans the entire length of the protibiae and is at ca. 2 × as wide as the protibial shaft. The lobe is distinctly triangular with the broadest point distal to the midline (Fig. 34D ). Pro-, meso-, and meta- tibiae lacking exterior lobes; meso-, and meta- tibiae lack interior lobes as well.

Measurements of paratype females [mm]. Length of body (including cerci and head, excluding antennae) 81.8-94.1, length/width of head 8.0-9.7/6.4-7.4, antennae 3.3-4.2, pronotum 4.6-5.7, mesonotum 6.7-9.1, length of tegmina 45.4-57.5, length of alae 6.0-7.2, greatest width of abdomen 30.7-34.4, profemora 17.9-21.6, mesofemora 13.8-15.9, metafemora 16.2-18.6, protibiae 11.1-11.8, mesotibiae 10.1-10.7, metatibiae 13.6-13.7.

Male. Coloration. Coloration description based on live captive reared males (Fig. 33B ) and the male nymphs found in the wild (Fig. 32B ). Captive males are generally mostly pale green throughout with the margins of the legs, thorax, and abdomen a reddish brown, and slightly transparent eye spots on abdominal segment V. The wild caught nymphs had these areas typically darker, more dark red than reddish brown and dark eyespots on abdominal segment V.

Morphology. Head. Head capsule about as long as wide, with a vertex that is weakly granular; posteromedial tubercle small but notable and slightly raised above the head capsule (Fig. 35D ). Frontal convexity stout with a few short setae near the apex. Compound eyes large and bulbous, occupying ca. ⅖ of the head capsule lateral margins and significantly protruding from the head capsule (Fig. 35D ). There are three well-developed ocelli between and slightly posterior to the compound eyes. Antennae (including the scapus and pedicellus) consists of 23 segments. The scapus and pedicellus are bare, all other segments are covered in dense, thin, pale setae that are as long as or longer than the antennae segment is wide. The terminal three segments have shorter darker setae. Thorax. Pronotum anterior margin is distinctly concave and lateral margins are slightly convex and converge to a straight posterior margin that is slightly> ½ width of the anterior rim (Fig. 35D ). Anterior and lateral margins of the pronotum have moderate rims and the posterior margin lacks a rim (Fig. 35D ). Face of the pronotum is slightly lumpy, has a distinct sagittal furrow, a pit just posterior to the center, a moderate perpendicular furrow just anterior to the central pit, and has a distinct pit on each side near the anterior margin (Fig. 35D ). The prosternum surface is slightly granular. The mesosternum surface is marked densely with prominent nodes, with the largest along the sagittal plane and more prominent on the anterior margin, posterior margin with less prominent and slightly smaller nodes. Prescutum slightly longer than wide, with lateral margins slightly converging to the posterior which is ca. ¾ the width of the anterior rim (Fig. 35D ). Lateral rims with nine or ten nodes of slightly varying size, none very large or prominent, but each marked with a single stiff seta protruding from the tip. The surface of the prescutum is notably granulose along the sagittal plane with lateral surfaces rather smooth (Fig. 35D ). Prescutum anterior rim slightly granulose with no distinct central tubercle (Fig. 35E ). Mesopleura narrow, only gradually diverging from the anterior to the posterior (Fig. 35D ). Lateral margin granulose throughout, with only four or five slightly larger than the rest, but not significantly larger. The largest nodes along the mesopleura have a singular seta protruding from them like those on the prescutum margins. Face of the mesopleura smooth but slightly wrinkled and with two faint pits, one on the anterior margin and one near the middle of the mesopleura. Wings. Tegmina short, only reaching the anterior margin of abdominal segment III. Tegmina wing venation: the subcosta (Sc) is the first vein and terminates the earliest, ca. ⅖ of the way through the overall tegmina length. The radius (R) spans the entire length of the tegmina with the first radius (R1) branching ca. ⅓ of the way through the wing length and terminating just posterior to the middle of the wing, the second radius (R2) branches near the distal ⅓ of the wing, and then the radial sector (Rs) runs straight to the tegmina apex and terminates. The media (M) spans the entire length of the tegmina, terminating at the wing apex as the media anterior (MA) with the first media posterior (MP1) beginning and terminating near the tegmina mid length followed by the second media posterior (MP2) which begins ca. ⅔ of the way through the tegmina length and terminates near the posterior quarter of the wing. The cubitus (Cu) runs through the wing surface angled until it meets the margin ca. ⅓ of the way through the tegmina length and then runs along the margin as the two media posterior veins then meet it and fuse and the cubitus continues to run nearly to the wing apex. The first anal (1A) vein runs subparallel to the cubitus until it meets it slightly> ⅓ of the way through the tegmina length and fuses with it. Alae well-developed in an oval fan configuration, reaching to the anterior margin of abdominal segment IX or halfway through it. Alae wing venation: the costa (C) is present along the entire foremargin giving stability to the wing. The subcosta (Sc) spans ca. ⅔ of the wing length and is mostly fused with the radius in the beginning but terminates when it meets the costa. The radius (R) spans the entire wing and branches ca. ⅖ of the way through into the radius 1 (R1) and radial sector (Rs) which run slightly diverging for the first ⅓ of their length, parallel for the central portion until the terminal quarter where they converge and terminate on the wing margin near each other but not touching. The media (M) branches early, ca. ⅙ of the way through the wing into the media anterior (MA) and the media posterior (MP) which run parallel with each other until the distal ⅙ of the wing where the media posterior fuses with the media anterior which then run fused together to the wing margin. The cubitus (Cu) runs unbranched and terminates at the wing apex. Of the anterior anal veins, the first anterior anal (1AA) fuses with the cubitus near the point where the media branches into the media anterior and media posterior and then the first anterior anal branches from the cubitus ⅔ of the way through the wing length where it uniformly diverges from the cubitus until it terminates at the wing margin. The anterior anal veins two-seven (2AA-7AA) have a common origin and run unbranched in a folding fan pattern of relatively uniform spacing to the wing margin. The posterior anal veins (1PA-7PA) share a common origin separate from the anterior anal veins and run unbranched to the wing margin with slightly thinner spacing than the anterior anal veins. Abdomen. Segment II, parallel-sided, segment III and the anterior half of IV diverging, the posterior ½ of IV through segment V either parallel-sided or slightly diverging, VI through the apex converging to a blunt rounded apex. The margins of segments VIII-X have a line of setae along them (Fig. 35F ). Genitalia. Poculum broad and rounded, ending in a rounded apex that passes beyond the anterior margin of segment X (Fig. 35G ). Cerci long and slender, with ca. ½ of their length extending out from under the anal abdominal segment. The cerci are slightly cupped, with a granulose surface and numerous short setae throughout (Fig. 35F ). Vomer broad and stout with rounded sides converging to the apex which is armed with two upwards turning hooks, one at the apex which is larger and one lateral to it which is slightly smaller (Fig. 5H ). Legs. Profemoral exterior lobe a rounded arc without a distinct angle, slightly thinner than the interior lobe (at its widest slightly> 2 × the greatest width of the profemoral shaft), and with the distal half marked by three or four small but sharp anteriorly pointing teeth (Fig. 35 ). Profemoral interior lobe roundly triangular, at its widest ca. 2½× as wide as the profemoral shaft at its widest. The profemoral interior lobe is generally marked with five, serrate, anteriorly pointing teeth arranged in a two-one-two pattern, with shallow looping gaps between them, and occasionally marked with an extra tooth within the set (like as can be seen in Fig. 35C ). Mesofemoral exterior lobe arcs end to end but with a more prominent bend near the distal ⅓ of the lobe which is marked with two or three small serrate teeth on the distal ⅓ only, with the proximal portion of the lobe smooth. The mesofemoral interior lobe at its widest is approximately the same width as the exterior lobe, but the proximal half is slightly wider than the proximal half of the exterior lobe. The distal half of the mesofemoral interior lobe is marked with six or seven small serrate teeth and the proximal half is rather smooth. Metafemoral exterior lobe lacks dentation and has a straight margin along the metafemoral shaft. Metafemoral interior lobe smoothly arcs end to end with six or seven small serrate teeth on the distal ½ only. Protibiae lacking exterior lobe, interior lobe reaching end to end in a smooth triangle ca. 3 × as wide as the protibial shaft, with the widest point just distal to the midline, and all margins notably marked with short setae throughout (Fig. 35C ). Mesotibiae and metatibiae simple, lacking lobes completely.

Measurements of holotype male [mm]. Length of body (including cerci and head, excluding antennae) 58.6, length/width of head 3.7/3.3, antennae 32.5, pronotum 2.8, mesonotum 4.5, length of tegmina 16.1, length of alae 42.0, greatest width of abdomen 15.8, profemora 12.0, mesofemora 9.5, metafemora 11.4, protibiae 7.5, mesotibiae 6.3, metatibiae 8.2.

Measurements of paratype males [mm]. Length of body (including cerci and head, excluding antennae) 56.0-69.4, length/width of head 2.9-4.6/2.9-4.0, antennae 34.5-43.9, pronotum 2.5-3.0, mesonotum 3.8-4.8, length of tegmina 16.4-19.8, length of alae 40.5-49.9, greatest width of abdomen 14.4-18.3, profemora 11.7-14.8, mesofemora 9.5-12.3, metafemora 11.0-13.6, protibiae 7.7-9.9, mesotibiae 6.5-8.0, metatibiae 8.5-10.4.

Description of egg (Fig. 36 ). The lateral surfaces are flat but with the posterior half slightly wider than the anterior half. The dorsal surface is slightly convex, which gives the margin a slight undulating appearance when viewed from the lateral aspect as the middle is thinner than either end of the egg. When viewed from the lateral aspect; the ventral margin is also not straight but is instead with the posterior slightly protruding more than the anterior. All surfaces have numerous small sized pits throughout with short moss-like pinnae interspersed throughout the capsules surfaces with those on the margins and those on the dorsal surface slightly more prominent. Dorsal surface with irregular medium sized pitting and moss-like pinnae around the micropylar plate. Micropylar plate long, ca. 7/8 of the overall dorsal surface length, with the widest portion around the micropylar cup. Micropylar plate nearly symmetrical with the anterior and posterior thin and the area around the micropylar cup the widest point. Micropylar cup of moderate size and placed just slightly posterior to the micropylar plate midline. Operculum slightly ovular, with the outer margin with a distinct row of moss-like pinnae surrounding the operculum. Operculum is roundly raised with the height ca. ½ the operculum width. The overall color is light brown, with the moss-like pinnae sometimes slightly lighter in color.

Measurements including the extended pinnae

[mm]. Length (including operculum) 2.2, maximum width of capsule when viewed from lateral aspect 1.3 mm, length of micropylar plate 1.1 mm.

Newly hatched nymphs. (Fig. 9I ). The body is made up of two general blocks of color, the legs, head, pronotum, and mesonotum are primarily chocolate brown and the mesonotum and abdomen are burnt red in color. Basitarsi are white and the remaining tarsal segments are burnt reddish. The tibiae only have interior lobes which smoothly span the full length and lack exterior lobes. The tibiae interior lobes are brown with two white patches on the proximal half only, the proximal most is notably larger. Interior and exterior femoral lobes are all about the same width, are smoothly arcing, and all have minimal serration. On the profemoral interior lobe there is a notable whitish patch, the exterior lobe and profemoral shaft itself are devoid of prominent white markings. The meso- and metafemoral interior lobes are similar in that they have a small white patch right at the proximal end, and then another white marking ca. ⅖ of the way through the length. The exterior meso- and metafemoral lobes only have one white patch located on the proximal ⅓, but notably wider than the interior lobe white patch. The meso- and metafemoral shafts lack white coloration. The abdomen is slender with segment II through the anterior ½ of IV diverging and the posterior ½ of IV-X converging.

Etymology.

Noun, from Greek mythology. Named for the tragic story of Icarus, son of Daedalus. During their escape from the island of Crete, Icarus flew too close to the sun and melted the wax wings his father built. We felt it was fitting that this mythological name is shared with this species that lacks the hindwings within the former Cryptophyllium celebicum species group (characterized by females with well-developed alae).

Figures

Figure 5. Details of the Cryptophyllium gen. nov. male vomer. All from the RBINS collection, prepared and photographed by Jerome Constant (RBINS). Views 1 ventral 2 right lateral of the ventrally oriented vomer 3 left lateral of the ventrally oriented vomer A Cryptophyllium westwoodii comb. nov., Thailand, Na Haeo B Cryptophyllium khmer gen. et sp. nov. Cambodia, Kbal Spean C Cryptophyllium phami gen. et sp. nov., Vietnam, Cat Tien D Cryptophyllium khmer gen. et sp. nov., Cambodia, Tatai E Cryptophyllium bollensi gen. et sp. nov., Vietnam, Phuoc Binh F Cryptophyllium oyae comb. nov., Laos, Mt. Phu Phan G Cryptophyllium bankoi gen. et sp. nov., Vietnam, Ba Na-Nui Chua H Cryptophyllium icarus gen. et sp. nov., Vietnam, Bidoup-Nui Ba I Cryptophyllium rarum comb. nov., Vietnam, Cuc Phuong J Cryptophyllium limogesi gen. et sp. nov., Vietnam, Ta Dung. Scale bars, 1.0 mm.

Figure 8. Known eggs for the Cryptophyllium gen. nov. set in couplets of lateral and then dorsal views. All images except for K and L were taken by Bruno Kneubu ̈ hler (Switzerland), K and L images taken by the first author, Q and R images taken by Rene ́ Limoges (IMQC) A, B Cryptophyllium westwoodii comb. nov. C, D Cryptophyllium tibetense comb. nov. E, F Cryptophyllium bollensi gen. et sp. nov. G, H Cryptophyllium oyae comb. nov. I, J Cryptophyllium icarus gen. et sp. nov. K, L Cryptophyllium liyananae gen. et sp. nov. M, N Cryptophyllium chrisangi comb. nov. O, P Cryptophyllium celebicum comb. nov. Q, R Cryptophyllium limogesi gen. et sp. nov. Scale bars 2.0 mm long.

Figure 9. Freshly hatched nymphs known for the Cryptophyllium gen. nov., all except for C bred and photographed by Bruno Kneubu ̈ hler (Switzerland) A Cryptophyllium celebicum comb. nov. B Cryptophyllium chrisangi comb. nov. C Cryptophyllium khmer gen. et sp. nov. bred and photographed by Tim Bollens (Belgium) D Cryptophyllium tibetense comb. nov. E Cryptophyllium oyae comb. nov. F Cryptophyllium westwoodii comb. nov. G Cryptophyllium phami gen. et sp. nov. H Cryptophyllium bollensi gen. et sp. nov. I Cryptophyllium icarus gen. et sp. nov.

Figure 32. Live Cryptophyllium icarus found in July 2014 by Joachim Bresseel (RBINS) and Jerome Constant (RBINS) in Bidoup Nui Ba N. P., Vietnam. Photographs by Jerome Constant (RBINS) A paratype female, dorsal closeup B male nymph, paratype C same female as in A but showing the host tree she was found on D disturbed area along the road where the original stock was found with arrows pointing out host trees.

Figure 33. Live adult Cryptophyllium icarus bred and photographed by Bruno Kneubu ̈ hler (Switzerland) A female B male C lateral view of female with tegmina held up to expose the dwarfed alae underneath.

Figure 34. Cryptophyllium icarus gen. et sp. nov. female paratype, photographs by Jerome Constant (RBINS) A habitus, dorsal B habitus, ventral C antennae and anterior of head capsule D profemoral and protibial lobes E head and thorax details dorsal F head and thorax details lateral G genitalia dorsal H genitalia ventral.

Figure 35. Cryptophyllium icarus gen. et sp. nov. male holotype, photographs by Jerome Constant (RBINS) A habitus, dorsal B habitus, ventral C profemoral and protibial lobes D head and thorax details dorsal E head and thorax details lateral F genitalia dorsal G genitalia ventral.

Figure 36. Cryptophyllium icarus gen. et sp. nov. egg, RBINS collection, photographs by Jerome Constant A lateral view B dorso-lateral view C dorsal view D opercular (anterior) view E posterior view F ventral view.