Cryptophyllium khmer 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 khmer gen. et sp. nov. Figures 5B , 5D , 6D , 9C , 37 , 38 , 39 , 40 , 41

Material examined.

Holotype ♂: "Coll. I.R.Sc.N.B., collected as nymph, Coll. I.R.Sc.N.B., Cambodia, Koh Kong prov., Tatai, 11°35 ’13” N 103°05 ’50” E, 9-19.x.2016, day collecting, GTI Project, Leg. J. Constant and J. Bresseel, I.G.: 33.345 (RBINS-PHYLLIUM DNA sample 0002)" [vomer dissected], deposited in RBINS.

Paratypes (9 ♀♀, 2 ♂♂): • "Coll. I.R.Sc.N.B., collected as nymph, Coll. I.R.Sc.N.B., Cambodia, Koh Kong prov., Tatai, 11°35 ’13” N 103°05 ’50” E, 9-19.x.2016, day collecting, GTI Project, Leg. J. Constant and J. Bresseel, I.G.: 33.345, RBINS-PHYLLIUM DNA sample 0001" (RBINS) • 3 ♀♀, 1 ♂: "Coll. I.R.Sc.N.B., Ex breeding Tim Bollens, 2018, Coll. I.R.Sc.N.B., Cambodia, Koh Kong prov., Tatai, 11°35 ’13” N 103°05 ’50” E, 9-19.x.2016, day collecting, GTI Project, Leg. J. Constant and J. Bresseel, I.G.: 33.345" • 2 ♂♂: "Coll. I.R.Sc.N.B., Ex breeding Tim Bollens, 2018, Coll. I.R.Sc.N.B., Cambodia, Koh Kong prov., Tatai, 11°35 ’13” N 103°05 ’50” E, 9-19.x.2016, day collecting, GTI Project, Leg. J. Constant and J. Bresseel, I.G.: 33.345; ex breeding Tim Bollens, 2018" • 2 ♀♀: "Coll. I.R.Sc.N.B., CAMBODIA, Siem Reap Prov., Phnom Kulen N.P., Forest near Preah Thom, 26-27-VII-2006, Leg K. Smets, Y. Oul and D. Jump." (1 ♀: RBINS; 1 ♀: RUPP) • 1 ♂: "Cambodia, Siem Reap; Kbal Spean, 13°40.858'N 104°01.111'E, 122 m, 6-jul-2015, Hap, Sour, Phauk, Khearn, Chhum, Ly, Lom, Heang, Hok, CA0028, Lighttrap in the forest with canopy cover." (RUPP) • 3 ♀, 1 ♂: "Coll. I.R.Sc.N.B., Ex breeding Tim Bollens, 2019, Coll. I.R.Sc.N.B., Cambodia, Koh Kong prov., Tatai, 11°35 ’13” N 103°05 ’50” E, 9-19.x.2016, day collecting, GTI Project, Leg. J. Constant and J. Bresseel, I.G.: 33.345; ex breeding Tim Bollens, 2019" (RBINS) • 1 ♀, 1 ♂: "Coll. I.R.Sc.N.B., Ex breeding Tim Bollens, 2019, Coll. I.R.Sc.N.B., Cambodia, Koh Kong prov., Tatai, 11°35 ’13” N 103°05 ’50” E, 9-19.x.2016, day collecting, GTI Project, Leg. J. Constant and J. Bresseel, I.G.: 33.345; ex breeding Tim Bollens, 2018" (Coll RC).

Remarks.

When this species was first reviewed morphologically, it was assumed to be an additional distribution record of Cryptophyllium westwoodii comb. nov., which is known from a relatively expansive range (Fig. 2 ). However, our molecular analysis revealed that the Tatai and Siem Reap (Cambodia) populations instead represented an undescribed species distinct from Cryptophyllium westwoodii comb. nov. The recognition of this cryptic species from Cambodia, leaves many observational records> from Laos, Thailand, and Cambodia without confirmed identification (represented by the bi-colored circles in our distribution map noting a record which, due to the lack of molecular confirmation, could represent either species; Fig. 2 ).

During GTI expeditions several nymphs ranging from L1 to subadult were collected on multiple closely situated guava trees behind a house near the start of the trail leading to Tatai falls. Nymphs were successfully reared to adulthood by Tim Bollens (Belgium). Strangely locals had never noticed the insects before due to their excellent camouflage (Fig. 37 ).

Interestingly, in 2006, an attempt was made to describe a Cryptophyllium westwoodii comb. nov. like species from Rayong, Thailand as ' Phyllium rayongii ' ( Sorpongpaisal and Thanasinchayakul 2006). Cumming and Le Tirant (2020) note however that this name is a nomen nudum and therefore unavailable according to ICZN Article 16.4.1. ( ICZN 1999). With this population rather geographically close to the type locality of Cryptophyllium khmer sp. nov. it is entirely possible that ' Phyllium rayongii ' may have been intended to represent a valid population, but with the lack of a holotype specimen to define this species it was never confirmed and is now a moot point.

Due to the cryptic nature of this new species, we hope that efforts will be undertaken in the future to molecularly sample from throughout Thailand, Myanmar, Laos, and Cambodia to determine with more clarity the geographic distributions where Cryptophyllium khmer sp. nov. and Cryptophyllium westwoodii comb. nov. occur.

Differentiation.

Morphological differentiation of this species has proven to be difficult, with the only clear and consistent differences being ascertained through molecular analysis (Fig. 4 ).

Females are most morphologically similar to Cryptophyllium westwoodii comb. nov., Cryptophyllium bollensi sp. nov., Cryptophyllium phami sp. nov., and Cryptophyllium nuichuaense sp. nov. females based on the general shape of the abdomen, lobes of the legs, and the thorax. The later three species can be differentiated by their shorter alae reaching to abdominal segments II or III vs. Cryptophyllium khmer sp. nov. which has long alae reaching onto abdominal segment VI. We have yet to identify a reliable morphological feature between Cryptophyllium khmer sp. nov. and Cryptophyllium westwoodii comb. nov. as both species have long alae and at least for Cryptophyllium westwoodii comb. nov. there is significant intraspecific variation which often encompasses the range of Cryptophyllium khmer sp. nov. female variation.

Males are most morphologically similar to Cryptophyllium westwoodii comb. nov., Cryptophyllium chrisangi comb. nov., Cryptophyllium bollensi sp. nov., and Cryptophyllium phami sp. nov. based on features of the thorax, tegmina, and lobes of the legs. Cryptophyllium khmer sp. nov. males can be differentiated from the first two species by the general shape of the abdomen as it is thinly elliptical with a maximum width only 30-34% the abdominal length in Cryptophyllium westwoodii comb. nov. and Cryptophyllium chrisangi comb. nov. vs. broadly elliptical or broadly spade-shaped with a maximum width ca. 38-45% the abdominal length in Cryptophyllium khmer sp. nov., Cryptophyllium bollensi sp. nov., and Cryptophyllium phami sp. nov. males. Unfortunately, due to intraspecific variation within Cryptophyllium khmer sp. nov., Cryptophyllium bollensi sp. nov., and Cryptophyllium phami sp. nov. we could not identify a reliable morphological feature for differentiation males based on morphology alone.

This difficulty for differentiating a single sex alone emphasizes the importance of captive rearing of specimens to reveal the informative set of female, male, and egg morphology, and of course the importance of molecular comparison.

Distribution.

At present only confirmed from two Cambodian provinces, Koh Kong Province (Tatai) and Siem Reap Province (Kbai Spean and Phnom Kulen N.P., Forest Near Prean Thom). It is likely that other nearby localities may also represent this species, but due to a lack of molecular data we cannot at this time confirm them.

Description.

Female. Coloration. Coloration description is based upon photographs of living individuals (Fig. 38A, B ) reared by Tim Bollens (Belgium). Overall coloration pale mint green with variable slight highlighting of orange or tan coloration throughout. Compound eyes are slightly more yellow with tan highlights. Antennae are tan. The prescutum and mesopleura are reddish tan with pale cream granulation throughout. Throughout the head, legs, and body there is slight speckling as granulation is slightly paler in color than the surface it is found on. In lighter individuals, the venation of the tegmina is pale yellow to pale mint green (Fig. 38A ) and in darker individuals the venation is yellow with highlights of orange interspersed throughout (Fig. 38B ). Darker individuals also have variable reddish patches throughout the lobes of the legs and slightly darker coloration on the abdomen.

Morphology. Head. Head capsule slightly longer than wide, vertex with granulation throughout the surface, none as prominent as the posteromedial tubercle which is not notably wide but is distinctly taller than any other nodes on the head (Fig. 39E ). Frontal convexity stout, marked throughout with slight granulation and several short setae. Compound eyes slightly protruding from the head capsule, but are significantly large, taking up slightly <⅓ of the head capsule margins (Fig. 39E ). Ocelli absent. Antennal fields slightly wider than the first antennomere. Antennae. Antennae consisting of nine segments, with the terminal segment slightly longer than the preceding two segments’ lengths combined (Fig. 39C ). Antennomeres I-VIII sparsely marked with small transparent setae, the terminal antennomere is covered densely in slightly shorter setae. Thorax. Pronotum slightly wider than long, with gently concave anterior margin and slightly convex lateral margins, which converge to a slightly convex posterior margin that is half the width of the anterior margin (Fig. 39E ). The pronotum surface is marked with granulation throughout, a prominent pit in the center, and slight furrows anterior and lateral to the pit (Fig. 39E ). The pronotum has a prominent anterior rim and weakly formed lateral and posterior rims (Fig. 39E ). Prosternum and the anterior half of the mesosternum are marked with stout and numerous nodes, with the remainder of the mesosternum and the metasternum lacking prominent nodes (Fig. 39B ). Prescutum about as long as wide with lateral rims with 11 or 12 lumpy tubercles ranging in size from small to medium with granulation present throughout the length giving the margins a tough textured appearance (Fig. 39E ). Prescutum anterior rim not strongly protruding and marked with a granular surface (Fig. 39F ). Prescutum surface with 14 or 15 distinct nodes predominantly along the sagittal plane, with those on the anterior half slightly larger than the rest (Fig. 39E ). Mesopleura are narrow and parallel on the anterior ⅓, and then bend distinctly and diverge uniformly throughout their length; lateral margin with 13-16 small to medium lumpy tubercles, of which three or four are slightly larger than the rest, but most are small and variable in shape, giving the margin a rough textured appearance (Fig. 39E ). Face of the mesopleura with granulation along the margin, with the remainder of the surface relatively smooth or with slight wrinkles. The surface of the mesopleura also has two distinct pits, one near the anterior ⅓ where the mesopleura bend, and one near the posterior ⅓ (Fig. 39E ). Wings. Tegmina long, reaching onto abdominal segment VIII. The subcosta (Sc) is the first vein in the forewing and runs parallel with the wing for the first half of its length, and then bends towards the wing margin for the second half, terminating ca. ⅓ of the way through the wing length. The radius (R) spans the central portion of the tegmina with two subparallel branched veins. The first radius (R1) branches ca. ½ through the radius length and terminates ca. ⅖ of the way through the wing length. The radial sector (Rs) branches from the end of the radius and runs angled to the wing margin where it terminates near the posterior ⅓ of the wing length. There is a weak continuation of the radius following the prominent radial sector branching which continues on as a short and thin radius to media crossvein (R-M). The media (M) is simply bifurcate with both the media anterior (MA) and media posterior (MP) terminating close to the posterior ⅕ of the wing. The cubitus (Cu) runs throughout the entire wing length simply, and then near the posterior ⅕ of the wing splits 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, at around the midline between the first radial branching and the radial sector branching. Alae well-developed, reaching abdominal segment VI. Abdomen. Abdominal segments II through the anterior half of IV diverging, the posterior half of IV through the anterior half of VII parallel-sided (giving the abdomen a boxy appearance), the remainder of VII smoothly rounded and converging to the apex with segments VIII-X. Genitalia. Subgenital plate starts at the anterior margin of segment VIII, is moderately broad, and extends ½ to ¾ of the way onto segment X, ending in a fine point (Fig. 39H ). Gonapophyses VIII are long and moderately broad, exceeding the apex of the abdomen with the tips slightly longer than the cerci, gonapophyses IX are thinner and shorter, hidden below gonapophyses VIII (Fig. 39H ). Cerci flat, not strongly cupped, with a finely granular surface and moderately marked with a few short setae. Legs. Profemoral exterior lobe broad and smoothly rounded, ca. 1½ to ca. 2 × wider than the interior lobe (Fig. 39D ). Margin of the profemoral exterior lobe with 10-12 small weakly formed teeth throughout the length (Fig. 39D ). Profemoral interior lobe obtusely angled and typically marked with five teeth arranged in a two-one-two pattern with looping gaps between them, but occasionally individuals can have doubly serrate teeth or an extra small tooth between sets (Fig. 39D ). Mesofemoral exterior lobe arcs from end to end but is weighted towards the distal half with a detectable bend and marked with four or five rounded teeth distributed on the distal half only. Interior and exterior lobes of a similar width. Mesofemoral interior lobe arcs end to end smoothly with five or six small serrate teeth only on the distal half of the arc which is slightly wider than the proximal half of the arc. Metafemoral interior lobe arcs end to end and has five or six serrate teeth on the distal half of the lobe which is slightly wider than the proximal half. Metafemoral exterior lobe is thin and smooth, hugging the metafemoral shaft and lacks notable teeth but the distal ⅓ can be slightly granular. Protibial interior lobe spans the entire length of the protibiae and is ca. 2½ the width of the protibiae shaft itself. The lobe is roundly triangular and is slightly wider on the distal half. Protibiae lacking a distinct exterior lobe. Mesotibiae and metatibiae lacking exterior and interior lobes.

Measurements of paratype females [mm] (from Tatai, Cambodia). Length of body (including cerci and head, excluding antennae) 83.3-90.0, length/width of head 8.4-8.7/6.6-7.1, antennae 4.1-4.6, pronotum 5.6-6.0, mesonotum 7.6-7.8, length of tegmina 52.8-53.6, length of alae 42.6 (only measured on one specimen, the others have the alae covered by the tegmina), greatest width of abdomen 31.3-36.2, profemora 19.1-21.4, mesofemora 15.1-15.4, metafemora 18.7-19.6, protibiae 12.5-12.6, mesotibiae 11.4-11.6, metatibiae 14.7-15.0.

Male. Coloration. Coloration description based on images of live males bred by Tim Bollens (Belgium). Overall coloration pale mint green throughout with highlighting of tan to orange (Fig. 39C ). The areas most often with the orange highlighting are the tips of the antennae, margins of the lobes of the legs, the thorax, and the margins of the abdomen. Additionally, on more prominently colored individuals the base of the antennae and the posteromedial tubercle of the head capsule can also be colored. Compound eyes are a muddled tan to reddish.

Morphology. Head. Head capsule about as long as wide, with a vertex that has moderate granulation throughout and a prominent but not broad posteromedial tubercle which is larger than any of the granules on the head capsule (Fig. 40E ). Frontal convexity not particularly long but ending in a fine point and covered with sparse thin setae. Compound eyes large and bulbous, taking up ca. ⅖ of the head capsule lateral margins (Fig. 40E ). There are three moderately developed ocelli located between and slightly posterior to the compound eyes. Antennal fields about as wide as the scapus. Antennae. Antennae (including the scapus and pedicellus) consists of 25 segments, all segments except the scapus and pedicellus and terminal three segments are covered in dense setae that are as long as or longer than the antennae segment is wide. The terminal three segments are covered in dense short setae and the scapus and pedicellus are nearly completely bare. Thorax. Pronotum with anterior margin slightly concave and lateral margins that are straight or slightly convex and converging to a straight posterior margin that is half the width of the anterior rim (Fig. 40E ). Anterior margin of the pronotum has a distinct rim, lateral margins have moderate rims, and the posterior margin lacks a rim (Fig. 40E ). Face of the pronotum is marked by a distinct sagittal furrow and pit in the center, a granular surface, and a slight perpendicular furrow from the central pit. Prosternum is granulose throughout with small nodes of nearly even size. Mesosternum anterior half with nodes of a similar size to the prosternum and those on the posterior half slightly less prominent. The metasternum has a slightly wrinkled surface and sparse granulation. Prescutum longer than wide, with lateral margins slightly converging to the posterior (Fig. 40E ). Lateral rims with small granulation throughout giving them a rough textured appearance, only three or four are slightly larger than the rest. Prescutum surface with granulation throughout with those along the sagittal plane slightly larger than the others. Prescutum anterior rim weakly formed but marked with a surface which is granular. Mesopleura narrow, almost parallel for the anterior quarter, and then only gradually diverge for the remainder of the length (Fig. 40E ). Lateral margin lacking prominent tubercles, instead marked with sharp granulation throughout with only two or three slightly larger than the rest, giving the margins a rough textured appearance. Face of the mesopleura slightly wrinkled and with two faint divots, one near the anterior margin and one half-way through the length (Fig. 40D ). Wings. Tegmina moderate length, reaching ⅓ to ½ onto abdominal segment III. Tegmina wing venation: the subcosta (Sc) is the first vein, is simple, and terminates ca. ½ through the overall tegmina length. The radius (R) spans the entire length of the tegmina with the first radius (R1) branching <ca. ½ through the wing length and terminating just distal to the midline, followed by the branching and termination of the second radius (R2) near the distal ⅓ of the wing, and then the radial sector runs to the wing apex. The media (M) also spans the entire length of the tegmina with the first media posterior (MP1) branching off ca. ⅖ of the way through the wing length, and then the second media posterior (MP2) branches near the midline, and the media anterior (MA) runs to the wing apex. The cubitus (Cu) runs along the edge of the wing as the two media posterior veins fuse with it and as the cubitus reaches the apex it fades. The first anal (1A) vein terminates upon reaching the cubitus slightly <⅓ of the way through the wing length. Alae well-developed in an oval fan configuration, long, reaching onto abdominal segments IX or X. Alae wing venation: the costa (C) is present along the entire foremargin giving stability to the wing. The subcosta (Sc) is long, spanning slightly> 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. ⅓ of the way through into the first radius (R1) and radial sector (Rs) which run gently diverging for most of their length and then converge at the apex of the wing where they terminate 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 throughout the wing until the distal ⅙ of the wing where the media posterior fuses with the media anterior which then run fused together to the wing apex where they terminate near the radial sector. 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-6PA) 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. Margins of abdominal segment II either slightly converging or parallel-sided. Abdominal segments III through the anterior ⅔ of IV diverging. Segment V with parallel margins and VI-X converging slowly at first then more prominently for the terminal three segments, giving the abdomen a spade-shaped appearance. Genitalia. Poculum broad, posteriorly rounded and with a shallow notch medioapically; slightly passes the anterior margin of segment X (Fig. 40G ). Cerci long and slender, extending from under the anal abdominal segment, slightly cupped with a granular surface and numerous short setae throughout (Fig. 40F ). Vomer broad and stout with sides evenly converging and terminating in an upward hooking apical spine with a smaller hook next to the base of the primary spine (Fig. 5D ). Legs. Profemoral exterior lobe about the same width as the interior lobe or slightly wider, smoothly arcing from end to end and marked with a granular margin and five or six small serrate teeth on the distal half only (Fig. 40C ). Profemoral interior lobe roundly triangular and marked with five teeth arranged in a two-one-two pattern with prominent looping gaps between the sets and the middle tooth larger than the others (Fig. 40C ). Mesofemoral exterior lobe arcs end to end, but is slightly more bent than the interior lobe and is broader on the distal half which can either be lacking dentation or have three or four dulled teeth, and the proximal half that is rather thin and lacking teeth. Mesofemoral interior lobe of a similar width to the exterior lobe, is broader on the distal end and is marked with five or six serrate teeth mostly situated on the distal ⅓ to ½ of the lobe. Metafemoral exterior lobe lacks dentation, and has a straight margin along the metafemoral shaft. Metafemoral interior lobe smoothly arcs end to end with eight or nine serrate teeth on the slightly wider distal half. Protibiae lacking exterior lobe, interior lobe reaching end to end in a smoothly rounded triangle with the widest portion ca. 3- 3½× as wide as the protibial shaft and situated just distal to the midline. Meso- and metatibiae simple, lacking lobes completely.

Measurements of holotype male [mm]. Length of body (including cerci and head, excluding antennae) 61.9, length/width of head 4.5/3.9, antennae 37.31

, pronotum 3.6, mesonotum 4.3, length of tegmina 19.0, length of alae 49.3, greatest width of abdomen 17.3, profemora 13.5, mesofemora 11.3, metafemora 13.4, protibiae 9.7, mesotibiae 7.8, metatibiae 10.1.

Measurements of paratype males [mm]. Length of body (including cerci and head, excluding antennae) 63.8-70.2, length/width of head 5.0-5.5/4.1-4.3, antennae 38.8-39.5, pronotum 3.6-4.1, mesonotum 5.0-6.2, length of tegmina 20.0-20.4, length of alae 50.0-52.1, greatest width of abdomen 17.1-17.9, profemora 15.72

, mesofemora 12.1, metafemora 13.7-14.1, protibiae 9.8**, mesotibiae 8.7-8.9, metatibiae 11.2-11.4.

Eggs. (Fig. 41 ). The lateral surfaces are flat with a length ca. 1½× the width with parallel margins, giving the capsule a rectangular appearance. All surfaces have numerous small to medium sized pits throughout, the lateral surface has around 35 pits (mostly on the smaller end of the spectrum) arranged in no detectable order, some more closely spaced than others. In addition, between the pits the surfaces are covered with short moss-like pinnae with the pinnae along the margins slightly longer than the pinnae on the other surfaces. The dorsal surface is marked with six or seven slightly irregular medium sized pits on each half running the length of the capsule with short moss-like pinnae around the micropylar plate and between the pits. The micropylar plate is not overly long, occupying ca. ½ of the dorsal surface length but not perfectly centered, with ca. ⅓ of the unoccupied space below and ⅔ above the micropylar plate. The micropylar cup is the widest portion of the micropylar plate and is located ca. ⅓ of the dorsal surface length from the posterior. The micropylar plate is approximately teardrop-shaped with the anterior portion longer and thinner than the posterior after the micropylar cup. Operculum slightly ovular, with the outer margin encircled with short moss-like pinnae surrounding the operculum and four or five medium pits surrounding the dorsal and lateral margins. The operculum is roundly raised with a height slightly> ½ operculum width. This rounded raised cap is marked with a sagittal raised row of pinnae similar in length to those along the capsule margins. The rounded raised cap is not perfectly centered and instead the rounded projection is shifted slightly towards the ventral surface. The overall egg color is tan to light brown, with the moss-like pinnae sometimes slightly lighter in color.

Measurements including the extended pinnae [mm]. Length (including operculum): 5.6; maximum width of capsule when viewed from lateral aspect 3.2; length of micropylar plate 3.0.

Etymology.

Noun. The species epithet is the Hindi word Cryptophyllium khmer , meaning Cambodia, referring to the country of origin for this species.

Figures

Figure 2. Distribution map for the 24 Cryptophyllium gen. nov. species presently known (with solid lines from their name pointing to the type locality) as well as additional Cryptophyllium gen. nov. species which we could not herein describe / differentiate (indicated by dashed lines). Note that the line for Cryptophyllium westwoodii comb. nov. is pointing to the neotype locality and the type locality for Cryptophyllium athanysus comb. nov. is simply " Ceylon " therefore the line is pointing to the present-day localities we are aware of. Inset is of southern Vietnam showing the distributions of three additional species which could not fit within the main map. The colors in this map are noted to the left of the names within the phylogenetic tree in Fig. 4. Newly herein described species have names noted in bold. Note that with Cryptophyllium khmer sp. nov. not easily distinguishable from Cryptophyllium westwoodii comb. nov. from photos alone, only the locations for these two species where they were genetically sampled are solid colored, all observational images without genetic sampling have bicolored circles and could represent either of these species. Additionally, one symbol is split three ways for Cryptophyllium gen. nov. specimens from the Thai offshore islands of Ko Phangan and Ko Samui which could not be differentiated from C. chrisangi comb. nov., C. westwoodii comb. nov., and C. khmer sp. nov. from images alone. See Suppl. material 4 for a full list of the specimens / observations utilized to make the distribution map with deposition data for specimens and links to observational records>.

Figure 4. Phylogenetic relationships among 35 phylliid species (67 specimens). The species of the Cryptophyllium gen. nov. are highlighted with colors and symbols corresponding to the distribution map (Fig. 2). Support values based on maximum likelihood (UFBoot, IQ-TREE) are indicated at each node.

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 6. Details of female phylliid antennae illustrated from specimens within the RBINS collection, illustrations by Julien Caudron (RBINS). Note that the figures start with the scapus, therefore the diagnostic fourth antennomere is the third one shown; to ensure clarity A has the segment number listed to the right of the image. Note that the fourth antennomere of Phyllium (A) is longer than the fifth antennomere in comparison to the Cryptophyllium gen. nov. representatives (B-H), which have the fourth antennomere short and disk-like A Phyllium ericoriai, Philippines B Cryptophyllium bollensi gen. et sp. nov., Vietnam, Phuoc Binh C Cryptophyllium celebicum comb. nov., Indonesia, Sulawesi D Cryptophyllium khmer gen. et sp. nov., Cambodia, Siem Reap Province E Cryptophyllium nuichuaense gen. et sp. nov., Vietnam, Nui Chua F Cryptophyllium phami gen. et sp. nov., Vietnam, Cat Tien G Cryptophyllium westwoodii comb. nov., Laos, Bokeo Province H Cryptophyllium westwoodii comb. nov., Thailand, Chiang Mai. Drawn to relative scale.

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 37. Live Cryptophyllium khmer gen. et sp. nov. type material found by the joint effort of the RBINS, RUPP, and VNMN teams in Tatai, Cambodia in October 2016, all found as nymphs. Individuals in photographs A-H were found feeding on Guava trees (Psidium guajava) and the specimen in photograph I was found feeding on an unidentified species. Photographs by Jerome Constant (RBINS) A female B Hong Thai Pham (left) and Sisonila Kangsun (right), participants of the GTI project, photographing the leaf insects, which are very popular in Asia C male nymph D female nymph E male nymph F several nymphs G nymph H nymph I female nymph on unidentified host species.

Figure 38. Live Cryptophyllium khmer gen. et sp. nov. type material bred and photographed by Tim Bolllens (Belgium), dorsal views A paler form female (paratype) B darker form female (paratype) C male (paratype).

Figure 39. Cryptophyllium khmer gen. et sp. nov. paratype female, molecular sample RBINS 01 in our analysis, from Tatai, Cambodia, photographs by Jerome Constant (RBINS) A habitus, dorsal B habitus, ventral C details of the antennae, dorsal D pro- tibial and femoral lobes, dorsal E details of the antennae, head, and thorax, dorsal F details of the antennae, head, and thorax, lateral G terminalia, dorsal H genitalia, ventral.

Figure 40. Cryptophyllium khmer gen. et sp. nov. holotype male, from Tatai, Cambodia, photographs by Jerome Constant (RBINS) A habitus, dorsal B habitus, ventral C pro- tibial and femoral lobes, dorsal D details of the antennae, head, and thorax, lateral E details of the antennae, head, and thorax, dorsal F terminalia, dorsal G genitalia, ventral.

Figure 41. Cryptophyllium khmer gen. et sp. nov. egg, RBINS collection, photographs by Jerome Constant (RBINS) A lateral view B dorso-lateral view C dorsal view D opercular (anterior) view E posterior view F ventral view.