Cryptophyllium bollensi 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 bollensi gen. et sp. nov. Figures 5E , 6B , 8E , 8F , 9H , 16 , 17 , 18 , 19 , 20 , 21

Material examined.

Holotype ♂: "Coll. I.R.Sc.N.B., Ex breeding Tim Bollens, 2018, Coll. I.R.Sc.N.B., Vietnam, Ninh Thuan prov., Phuoc Binh N.P., 12°04'N, 108°45'E, 26.vii.2014, night coll., Leg.: J. Constant & J. Bresseel, GTI project, IG: 32.779". Deposited in the Royal Belgian Institute of Natural Sciences (RBINS).

Paratypes: (16 ♀♀, 25 ♂♂, 46 eggs) • 1 ♀: "Coll. I.R.Sc.N.B., Vietnam, Ninh Thuan prov., Phuoc Binh N.P., 12°04'N, 108°45'E, 26.vii.2014, night coll., Leg.: J. Constant and J. Bresseel, GTI project, IG: 32.779" (RBINS) • 6 ♂♂: same data as HT [3 with vomer dissected] (RBINS) • 2 ♂♂, 3 ♀♀, 1 subadult ♂: "Coll. I.R.Sc.N.B., Ex breeding Tim Bollens, 2019, Vietnam, Ninh Thuan prov., Phuoc Binh N.P., 12°04'N, 108°45'E, 26.vii.2014, night coll., Leg.: J. Constant and J. Bresseel, GTI project, IG: 32.779" (RBINS) • 1 ♂: "Coll. I.R.Sc.N.B., Ex breeding B. Kneubuhler, Vietnam, Ninh Thuan prov., Phuoc Binh N.P., 12°04'N, 108°45'E, 26.vii.2014, night coll., Leg.: J. Constant and J. Bresseel, GTI project, IG: 32.779" (RBINS) • 1 ♂, 1 ♀: "Coll. I.R.Sc.N.B., Ex breeding Bruno Kneubühler, 2017" "Coll. I.R.Sc.N.B., Vietnam, Ninh Thuan prov., Phuoc Binh N.P., 12°04'N, 108°45'E, 26.vii.2014, night coll., Leg.: J. Constant and J. Bresseel, GTI project, IG: 32.779" (VNMN) • 1 ♀; "VIETNAM: Ninh Thuan prov. Phuoc Binh N.P., 12°04'N 108°45'E, Bred from eggs supplied by Bruno Kneubühler (Switzerland), May, 2018, Coll RC 18-414" (Coll RC) • 2 subadult ♀♀; "VIETNAM: Ninh Thuan prov. Phuoc Binh N.P., 12°04'N 108°45'E, Bred from eggs supplied by Bruno Kneubühler (Switzerland), May, 2018" Coll RC 18-412 and 18-413 (Coll RC) • 1 ♂; "VIETNAM: Ninh Thuan prov. Phuoc Binh N.P., 12°04'N 108°45'E, Bred from eggs supplied by Bruno Kneubühler (Switzerland), April, 2018, Coll RC 18-2017" (Coll RC) • 19 eggs; "VIETNAM: Ninh Thuan prov. Phuoc Binh N.P., 12°04'N 108°45'E, eggs supplied by Bruno Kneubühler (Switzerland), 2018", Coll RC 17-375, 17-376, 18-282-18-298 (Coll RC) • 1 ♀, 3 ♂♂, 27 eggs; "ex Zucht T. Bollens 2018, Herkunft: Vietnam, Prov. Nin Thuan, Bác Ái Distr., Phuoc Binh N.P., leg. Bresseel and Constant 2014" [coll. FH, No’s 1061-1 to 4, E], (Coll FH) • 1 ♀, 1 ♂ (nymph n3); "ex Zucht F. Hennemann 2019, Herkunft: Vietnam, Prov. Nin Thuan, Bác Ái Distr., Phuoc Binh NP, leg. Bresseel and Constant 2014" [coll. FH, No’s 1061-5 and 6], (Coll FH) • 2 ♀♀, 3 ♂♂; "VIETNAM: Ninh Thuan prov. Phuoc Binh N.P., 12°04'N 108°45'E, Bred by Maxime Ortiz, France, circa 2020" (Coll MO) • 4 ♀♀, 6 ♂♂; "VIETNAM: Ninh Thuan prov. Phuoc Binh N.P., bred by Bruno Kneubühler (Switzerland), circa 2017-2018" (Coll OC).

Remarks.

This species was first collected by Jérôme Constant (RBINS) and Joachim Bresseel (RBINS) during a GTI research expedition. Only a single adult female was found (Fig. 16 ) while collecting at night in Phuoc Binh N.P.. The female was found on a shrub in a field that had been cleared of trees and planted with crops. The clearing of the forest had been done fairly recently as tree stumps were still present. In this area there was no adjacent old growth forest, only secondary forest nearby. The female was kept alive long enough to lay a series of fertilized eggs which were shared with expert phylliid breeder Tim Bollens (Belgium) who was able to rear a nice series of specimens and eventually share this species with other leaf insect breeder enthusiasts (Fig. 17 ).

Differentiation.

Females are morphologically most similar to Cryptophyllium phami sp. nov., Cryptophyllium chrisangi comb. nov., and Cryptophyllium nuichuaense sp. nov. based on the general femoral lobe shapes, the broad rounded exterior profemoral lobe, and the thorax shape and spination. Cryptophyllium bollensi sp. nov. have moderately long alae reaching onto abdominal segment IV that can be differentiated from Cryptophyllium phami sp. nov., which have shorter alae only reaching the anterior margin of abdominal segment III. The ventral surface of the antennae differentiates Cryptophyllium bollensi sp. nov. from the other similar species as Cryptophyllium bollensi sp. nov. have the ventral surface of segments VI, VII, and VIII flush (Fig. 6B ) vs. Cryptophyllium chrisangi comb. nov. and Cryptophyllium nuichuaense sp. nov. which have the ventral surface of antennal segments VI and VII projecting beyond segment VIII, giving the antennae a slight lamellate appearance (Fig. 6E ).

Males are morphologically most similar to Cryptophyllium westwoodii comb. nov., Cryptophyllium chrisangi comb. nov., Cryptophyllium phami sp. nov., and Cryptophyllium khmer sp. nov. due to the femoral shape and spination, the length of antennae and alae, and thorax shape and spination. Cryptophyllium westwoodii comb. nov. and Cryptophyllium chrisangi comb. nov. can be differentiated by their narrower abdominal shape with a maximum width only 30-34% of the abdominal length, vs. the others which have an abdominal shape that is broadly elliptical or broadly spade-shaped with a maximum width ca. 38-45% of the abdominal length. Due to similarities in morphology and the intraspecific variation within Cryptophyllium bollensi sp. nov., Cryptophyllium phami sp. nov., and Cryptophyllium khmer sp. nov., we could not identify a reliable morphological feature for differentiation within the males. The female morphology does allow differentiation of these species, and of course molecular analysis (Fig. 4 ) allows reliable differentiation even between these variable and difficult to distinguish species.

Distribution.

At present only known from the type locality of Phuoc Binh N.P., Ninh Thuan Province, southern Vietnam.

Description.

Female. Coloration. Coloration description is based upon living individuals (Figs 16 , 17A ). Overall coloration is pale green throughout, with variable areas highlighted with burnt red or brown coloration. These areas tend to be the margins on the lobes of the legs, some striping on the lobes of the legs, the thorax, abdominal margins, and the venation in the tegmina (Fig. 16 ).

Morphology. Head. Head capsule about as long as wide, vertex relatively smooth with the only notable feature being the posteromedial tubercle which is finely pointed (Fig. 18E ). Frontal convexity broad and blunt, with a slightly granular surface. Compound eyes slightly protruding from the head capsule, and are not particularly large, taking up ca. ¼ of the head capsule lateral margins (Fig. 18E ). Ocelli absent. Antennal fields slightly wider than the width of the first antennomere. Antennae. Antennae consist of nine segments, with the terminal segment about the same length as the preceding 2½ segments’ lengths combined (Fig. 6B ). Antennomeres I-VIII sparsely marked with small transparent setae, the terminal antennomere and the anterior margin of antennomere VIII are covered in stout, brown setae (Fig. 18C ). Thorax. Pronotum with gently 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. 18E ). The pronotum surface is smooth, with only a prominent pit in the center, and slight furrows anterior and lateral to the pit (Fig. 18E ). The pronotum has moderately formed anterior and lateral rims and a weakly formed posterior rim, all of which are relatively smooth (Fig. 18E ). Prosternum and the mesosternum are covered with numerous broad nodes, but the metasternum has a somewhat wrinkled surface. Prescutum longer than wide, lateral rims with 9-11 small to medium tubercles, similar in size giving the margin a rough appearance (Fig. 18E ). Prescutum anterior rim prominent but not strongly protruding, rim surface is granular, lacking a large sagittal spine (Fig. 18F ). Prescutum surface heavily granular, with those along the sagittal plane slightly larger than the rest (Fig. 18E ). Mesopleura begin ca. ¼ of the way through the prescutum length and evenly diverge; lateral margin with nine or ten small tubercles with about half of those slightly larger than the rest, with the smaller ones interspersed throughout (Fig. 18E ). Face of the mesopleura smooth or slightly wrinkled, with two notable divots, one on the anterior margin and one near the middle (Fig. 18F ). Wings. Tegmina long, reaching ½ through abdominal segment VII. Tegmina venation; the subcosta (Sc) is the first vein in the forewing, running parallel with the margin for the first half, and then bending and running towards the margin. The radius (R) spans the central portion of the forewing with two subparallel branched veins; the first radius (R1) branches ca. ¼ of the way through the wing length and terminates slightly proximal to the midline, and the radial sector (Rs) branches ca. ⅖ of the way through the wing length and terminates near the distal ⅓ of the wing length. There is a weak continuation of the radius following the prominent Rs branching which continues on as a short and thin R-M crossvein that weakly connects the two veins. The media (M) is simply bifurcate with both the media anterior (MA) and media posterior (MP) terminating near to 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, at the length about midway between the splitting of the R1 and Rs. Alae short, with their apex only just passing the anterior margin of abdominal segment IV. Abdomen. Abdominal segments II through the anterior half of IV uniformly diverging. The posterior half of segment IV through the anterior of segment VII are parallel, giving the abdomen a boxy appearance. The posterior half of segment VII ends in a slightly rounded lobe. Segments VIII-X are notably narrower than the previous segments, and have converging margins to the broad rounded apex (Fig. 18G ). Genitalia. Subgenital plate starts at the anterior margin of tergum VIII, is moderately broad, and extends halfway onto tergum X with straight margins ending in a fine point (Fig. 18H ). Gonapophyses VIII are long and moderately broad, slightly exceeding the apex of abdominal tergum X; gonapophyses IX are shorter and narrower, hidden below (Figs 17C , 18H ). Cerci flat, not strongly cupped, with a granular surface and few detectable setae (Fig. 18H ). Legs. Profemoral exterior lobe broad, rounded, and obtusely angled, smoothly arcing from end to end, ca. ⅓ wider than the width of the interior lobe (Fig. 18D ). Edge of the profemoral exterior lobe granular, or with a slightly serrate surface of eight or nine small teeth (Fig. 18D ). Profemoral interior lobe ca. 2 × as wide as the greatest width of the profemoral shaft, obtusely angled, and marked with five teeth arranged in a two-one-two pattern with looping gaps between them (Fig. 18D ). Mesofemoral exterior lobe arcs from end to end but is slightly bent in the center, weighted towards the distal half, and marked with three or four small serrate teeth distributed on the distal half only. Interior lobe is about the same width as the mesofemoral shaft, and the exterior lobe is slightly wider. Mesofemoral interior lobe arcs smoothly end to end with 6-8 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, with the distal half slightly wider than the proximal half and marked with 7-10 serrate teeth on the distal half of the lobe. Metafemoral exterior lobe is thin and smooth, hugging the metafemoral shaft and lacks dentation. Protibiae lacking an exterior lobe (Fig. 18D ). Protibiae interior lobe spans the entire length of the protibiae and is ca. 2 × the width of the protibiae shaft itself. The lobe is roundly triangular with the widest portion on the distal half. Mesotibiae and metatibiae lacking exterior and interior lobes.

Measurements of paratype female [mm] (wild caught). Length of body (including cerci and head, excluding antennae) 75.6, length/width of head 8.0/6.1, antennae 4.3, pronotum 4.8, mesonotum 7.2, length of tegmina 45.3, length of alae 23.1, greatest width of abdomen 28.6, profemora 16.7, mesofemora 13.6, metafemora 16.3, protibiae 10.8, mesotibiae 9.4, metatibiae 12.6.

Measurements of paratype females [mm] (ex culture). Length of body (including cerci and head, excluding antennae) 74.6-80.1, length/width of head 7.6-8.5/5.9-6.2, antennae 3.9-4.2, pronotum 4.5-5.4, mesonotum 6.7-6.8, length of tegmina 43.2-45.1, length of alae (not measurable as they were hidden by the tegmina), greatest width of abdomen 29.7, profemora 17.2-18.2, mesofemora 13.7-14.4, metafemora 16.5-16.6, protibiae 10.9-11.0, mesotibiae 9.1-9.6, metatibiae 12.5-12.8.

Male. Coloration. Coloration based upon live bred specimens in captivity (Fig. 17B ). Overall coloration pale green throughout with variable patches of tan to reddish coloration (Fig. 17B ). These tan to reddish areas are primarily around the margins of the lobes of the legs, the margins of the thorax, the tips of the antennae, and the margins of the abdomen. In darker colored specimens the mesofemoral lobes can also have coloration, not just along the margins. Abdominal segment V has a pair of slightly transparent eye spots.

Morphology. Head. Head capsule about as long as wide, with a vertex that is relatively smooth with only light granulation throughout. Frontal convexity stout with sparse thin setae. The posteromedial tubercle is not broad but is distinctly raised from the head capsule. Compound eyes large and bulbous, taking up ca. ⅖ of the head capsule lateral margins (Fig. 19E ). There are three well-developed ocelli located between and slightly posterior to the compound eyes. Antennae. Antennae (including the scapus and pedicellus) consist of 23-26 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 slightly convex and converging to a straight posterior margin that is ca. ½ the width of the anterior rim (Fig. 19E ). Anterior and lateral margins of the pronotum have moderately formed rims and the posterior margin lacks a rim (Fig. 19D ). Face of the pronotum is marked by a distinct furrow and pit in the center and a relatively smooth lumpy surface with weak granulation (Fig. 19D ). Prosternum surface is weakly granular with small nodes of even size and spacing. Mesosternum surface marked with slightly more prominent nodes, with the largest along the sagittal plane and denser on the anterior margin, posterior margin with less prominent and smaller nodes. Prescutum slightly longer than wide, with lateral margins that are only slightly converging to the posterior (Fig. 19E ). Lateral rims with eight or nine node-like tubercles, giving the lateral margins a rough textured appearance. Prescutum surface with minimal nodes throughout, with those along the sagittal plane slightly larger than the others. Prescutum anterior rim moderately formed but not strongly raised, with a granular surface and lacking a prominent sagittal tubercle. Mesopleura begin on the anterior prescutum margin but are narrow throughout the anterior ⅓ of their length, only diverging gently for the posterior ⅔ (Fig. 19E ). Lateral margin with eight or nine minor tubercles throughout the length except for the posterior ⅓ which is relatively smooth. Face of the mesopleura mostly smooth, with slight wrinkling throughout. Wings. Tegmina moderate length, extending ⅓ of the way onto abdominal segment III. Tegmina wing venation: the subcosta (Sc) is the first vein, is simple, and terminates the earliest ca. ⅓ of the way through the overall tegmina length. The radius (R) spans the entire length of the tegmina with the first radius (R1) branching just proximal to the midline 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 slightly more than ⅓ of the way through the wing length, and then the second media posterior (MP2) branches just distal to 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 ca. ⅓ of the way through the wing length. Alae well developed in an oval fan configuration, long, reaching to the middle or posterior of abdominal segments IX. Alae wing venation: the costa (C) is present along the entire foremargin giving stability to the wing. The subcosta (Sc) is long, spanning 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 slightly proximal to the midline 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 ⅕ of the wing where the media posterior fuses with the media anterior which then run fused 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. Lateral margins of abdominal segment II parallel, III through the anterior ⅔ of segment IV gradually diverging, the remainder of IV and segment V are parallel-sided, segment VI starts parallel-sided but then gently starts to converge and the remaining segments converge uniformly to the rounded apex of the abdomen. Genitalia. Poculum broad and ends in a rounded apex that slightly passes the anterior margin of segment X (Fig. 19G ). Cerci long and slender, extending from under the anal abdominal segment, nearly flat, not strongly cupped, covered in a granulose surface and numerous short setae (Fig. 19G ). Vomer broad and stout with straight sides evenly converging and ending in a thick apical hook with a smaller second hook adjacent to it (Fig. 5E ). Interestingly while examining the vomers of type material we found several aberrant vomers with some bearing only a singular hook (Fig. 20A ) or even three hooks (Fig. 20B ), based on other specimens examined and the trend within the Cryptophyllium gen. nov. males we expect that a typical male of this species has a two hooked vomer. Legs. Profemoral exterior lobe slightly broader than the interior lobe, ca. 2½× the greatest width of the profemoral shaft, roundly arcing end to end in a broad obtuse angle that is not distinctly bent, with the proximal margin slightly granulose, and the distal margin with four or five small serrate teeth (Fig. 20C ). Profemoral interior lobe roundly triangular and marked with five sharp teeth arranged in a two-one-two pattern with looping gaps between them, and the central tooth slightly larger than the others (Fig. 20C ). Mesofemoral exterior lobe arcs end to end but is slightly wider on the distal ⅓ which is marked with three or four serrate teeth, and a proximal half that is rather thin. Mesofemoral interior lobe is about the same width as the exterior, is broader on the distal end and is marked with 6-8 small serrate teeth. 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 small serrate teeth on the distal ⅔, which is slightly wider than the proximal ⅓. Protibiae lacking exterior lobe, interior lobe reaching end to end in a smooth triangle which is slightly weighted to the distal half and at its widest is ca. 2½ as wide as the protibial shaft (Fig. 20C ). Meso- and metatibiae simple, lacking lobes completely.

Measurements of holotype male [mm]. Length of body (including cerci and head, excluding antennae) 57.6, length/width of head 4.⅓.4, antennae 41.4, pronotum 3.0, mesonotum 4.3, length of tegmina 18.3, length of alae 43.4, greatest width of abdomen 16.0, profemora 12.8, mesofemora 11.2, metafemora 13.0, protibiae 8.9, mesotibiae 7.7, metatibiae 9.5.

Measurements of paratype males [mm] (ex culture). Length of body (including cerci and head, excluding antennae) 55.8-65.5, length/width of head 4.1-4.9/3.2-3.5, antennae 39.9-41.5, pronotum 3.0-3.4, mesonotum 4.0-4.8, length of tegmina 18.2-20.1, length of alae 41.5-47.2, greatest width of abdomen 13.6-16.3, profemora 12.4-13.6, mesofemora 10.4-12.3, metafemora 12.7-14.4, protibiae 8.7-10.2, mesotibiae 7.5-8.4, metatibiae 9.0-10.2.

Eggs. (Fig. 21 ). The overall color is muted dark brown, with the moss-like pinnae lighter in color, generally tan or light brown. The lateral surfaces are flat or slightly convex, with eggs either the same width anterior to posterior or with the posterior of the egg slightly wider. The lateral surfaces are marked with 40-50 small to medium sized pits, unevenly spaced in no detectable pattern, with sparse tufts of moss-like pinnae between these pits (Fig. 21A ). The dorsal surface has the micropylar plate spanning a majority of the length but not quite reaching end to end. On either side of the micropylar plate is variable pitting (generally eight or so pits) with those one the anterior and posterior ends slightly larger than the central pits (Fig. 21C ). The micropylar plate is symmetrical with the anterior and posterior thin and the middle the widest point. The micropylar cup is not located at this widest midpoint but is instead located on the posterior ⅓ of the micropylar plate. The micropylar plate margin is lined with short moss-like pinnae. Operculum slightly ovular, outer margin with a distinct row of short moss-like pinnae and in from the outer margin is a singular semi-circle of small to medium pits on the dorsal and lateral aspects (not fully surrounding the apex of the operculum as the ventral portion lacks these pits; Fig. 21D ). Operculum is roundly raised with the height ca. ½ the operculum width and the apex of the raised operculum has a tuft of moss-like pinnae. The ventral surface of the egg capsule has a slightly raised sagittal crest marked sparsely with short moss-like pinnae on the anterior ⅔, and the posterior ⅓ has longer moss-like pinnae. On either side of this raised sagittal crest is pitting, near the posterior ⅓ on each side of the longer moss-like pinnae of the sagittal crest is a large circular pit, and anterior to the lowest large pit are around ten small to medium pits arranged in no detectable pattern (Fig. 21F ).

Measurements including the extended pinnae [mm]. Length (including operculum): 4.1-4.4; maximum width of capsule when viewed from lateral aspect 2.6-2.8; length of micropylar plate 3.3-3.4

Newly hatched nymphs. (Fig. 9H ). The general color throughout the body is dark brown with slightly lighter brown on the legs. The basitarsi are yellow and remaining tarsal segments are dark brown. All tibiae lack exterior lobes but do have extremely thin smoothly arcing interior lobes which have several tan to brown stripes throughout their length. All femoral lobes are similar in width and have distinct serration on their distal halves. The interior profemoral lobe lacks a white spot, but the exterior lobe has a distinct white patch on the proximal ⅓ with an additional small white patch at the proximal most margin. The meso- and metafemoral interior lobes have two white patches, one on the proximal most edge, and a larger white patch ⅓ of the way through the length. The meso- and metafemoral exterior lobes also have a large white patch on the proximal ⅓, but lack a smaller white patch on the proximal most margin. The distal ends of the meso- and metafemora also have minimal white edges. The abdomen is mostly brown, but abdominal segments II and III have distinct green patches on their lateral surfaces (the centerline of the abdomen is uniform brown throughout). The terminal three abdominal segments also have a little bit of green on their margins. The widest point of the abdomen is abdominal segment IV.

Etymology.

Patronym. Named after Tim Bollens (Belgium) who has been instrumental in bringing many new phylliid species into the phasmid breeding community over the years. With his expertise in breeding these difficult phasmids he has allowed us to compare the informative sets of male, female, freshly hatched nymph, and egg morphology instead of only comparing singular dead specimens collected in the wild.

Figures

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 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 16. Live Cryptophyllium bollensi gen. et sp. nov. paratype where she was found in Phuoc Binh N. P., Vietnam in July 2014 by Joachim Bresseel (RBINS) and Jerome Constant (RBINS). Photograph by Jerome Constant (RBINS).

Figure 17. Live Cryptophyllium bollensi gen. et sp. nov. bred and photographed by Bruno Kneubu ̈ hler (Switzerland) A adult female, dorsal B adult male, dorsal C genitalia, female, ventral.

Figure 18. Cryptophyllium bollensi gen. et sp. nov. paratype female, photographs by Jerome Constant (RBINS) A habitus, dorsal B habitus, ventral C details of the antennae, dorsal D front leg details, 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 19. Cryptophyllium bollensi gen. et sp. nov. holotype male, photographs by Jerome Constant (RBINS) A habitus, dorsal B habitus, ventral C pro- tibial and femoral lobes, dorsal D details of the base of the antennae, head, and thorax, lateral E details of the antennae, head, and thorax, dorsal F terminalia, dorsal G genitalia, ventral.

Figure 20. Abnormal male vomer found in Cryptophyllium bollensi gen. et sp. nov. from Vietnam, Phuoc Binh, observed in the RBINS collection bred from the original limited stock. Prepared and photographed by Jerome Constant (RBINS). Scale bars, 1.0 mm. Views 1 ventral 2 right lateral of the ventrally oriented vomer 3 left lateral of the ventrally oriented vomer A abnormal singularly hooked vomer B abnormal three hooked vomer.

Figure 21. Cryptophyllium bollensi gen. et sp. nov. egg (RBINS), photographs by Jerome Constant A lateral view B dorso-lateral view C dorsal view D opercular (anterior) view E posterior view F ventral view.