Cryptophyllium bankoi 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 bankoi gen. et sp. nov. Figures 5G , 13 , 14 , 15

Material examined.

Holotype ♂: "VIETNAM: Quang Nam, Tay Giang, Axan Mt, 1,300 meters: June 2017, Coll RC 17-228". Deposited in the Montreal Insectarium (IMQC).

Paratypes: (4 ♂♂, 1 ♂ nymph, 1 ♀ nymph) • 1 ♂; "Vietnam; Daknong, June, 2017, Coll RC 17-338" (Coll RC) • 1 ♂; "Vietnam; Quang Ngai Province, Bato Mt. 900 m. elv: May 2015, Coll RC 16-141" (Coll RC) • 1 ♂: "Coll. I.R.Sc.N.B., Da Nang prov., Ba Na-Nui Chua Nat. Res., 18°09'N 105°55'E, 16-19.vii.2017, GTI Project, Leg. J. Constant and J. Bresseel, I.G.: 33.498" [vomer dissected] (RBINS) • 1 ♂: "Ngoc Linh, Kon Tum prov., Vietnam, 1700 m, VI.2016, leg. Luong coll. TB-05-134" (Coll TB) • 1 subadult ♀: "C. Vietnam, Bach Ma N.P., 16°12'N 107°52'E, 12-17.vii.2011, Leg J. Constant and J. Bresseel, I.G.: 31.933" [RBINS- Phyllium -DNA sample 0006] (RBINS) • 1 subadult ♂: "Vietnam, Gia Lai prov. Kon Chu Rang N.R., 600-1200 m, 13-20.vii.2018, GTI project, 14°28 ’28” N 108°32 ’27” E, Leg. J. Constant, J. Bresseel and X. Vermeersch, I.G.:33.769" [RBINS- Phyllium -DNA sample 0008] (RBINS).

Remarks.

This species, in true "cryptic leaf" fashion, was only recognized as unique when the molecular results were reviewed as morphologically it was hidden within the Cryptophyllium rarum comb. nov. males. Their morphological resemblance is uncanny and even with a series of males it is difficult to adequately differentiate these two species based on male morphology. Only the male Cryptophyllium bankoi sp. nov. is known at present and therefore it is unknown if the female also morphologically resembles Cryptophyllium rarum comb. nov. or is morphologically more unique. Despite several expeditions to southern Vietnam by the RBINS expedition members, only one nymph female Cryptophyllium bankoi sp. nov. has been collected to date, unfortunately she did not survive to adulthood (Fig. 13 ). With this species recorded from three Vietnamese provinces, we hope that as a widely ranging species the female can one day be identified and morphologically described.

Differentiation.

Females unknown. Males are morphologically very similar to Cryptophyllium rarum comb. nov. in their antennae length, profemoral interior lobe serration, protibial interior lobe shape, tegmina length, and their thorax shape and spination. The molecular results revealed however that this species was in fact not identical to Cryptophyllium rarum comb. nov. with which it shares a sympatric geographic range in central Vietnam and a striking morphological resemblance. Thankfully with the molecular analysis we were able to separate out these two species to allow us to see the subtle differences between them, which we originally thought to be simple morphological variation. The only consistent morphological differences we were able to locate between these species is that Cryptophyllium rarum comb. nov. tends to have an abdomen which is slightly more rounded (Fig. 59A ), vs. Cryptophyllium bankoi sp. nov. which has the abdomen ever so slightly spade-shaped (Fig. 14 ). Additionally, it appears as though Cryptophyllium rarum comb. nov. tend to be larger (79.8-89.0 mm) vs. Cryptophyllium bankoi sp. nov. (61.9-69.5 mm), and the profemoral exterior lobes have slightly differing serration, with Cryptophyllium bankoi sp. nov. having no teeth or up to five small teeth, vs. Cryptophyllium rarum comb. nov., which tend to have seven or eight teeth.

Distribution.

Central and Southern Vietnam. Known from Quang Ngai, Thua Thien Hue, Da Nang, Gia Lai, Quang Nam, and Dak Nong Provinces.

Male.

Coloration. Living individuals are a more vibrant green, and our descriptions are only based on preserved specimens. Overall coloration pale green throughout with variable patches of yellow and tan coloration (Figs 14 , 15 ). Compound eyes are slightly reddish to pink (Figs 14B , 15D ). The antennae are yellow to orange in color. In all specimens examined there were no variable brown patches on the lobes of the legs as is sometimes common in male phylliids.

Morphology. Head. Head capsule approximately as long as wide, with a vertex that is irregularly lumpy and with small granulation throughout, and a posteromedial tubercle which is larger than any of the nodes on the head (Fig. 14B ). Frontal convexity stout with sparse thin setae throughout the surface. Compound eyes large and bulbous, occupying ca. ⅖ of the head capsule lateral margins (Fig. 14B ). Between the compound eyes are three well-developed ocelli (Fig. 14B ). Antennal fields approximately as wide and as long as the scapus. Antennae. Antennae (including the scapus and pedicellus) consists of 25 or 26 segments, all segments except the scapus and pedicellus and terminal four segments are covered in dense setae that are as long as or longer than the antennae segment is wide. The terminal four segments are covered in dense short setae and the scapus and pedicellus are nearly bare, lacking setae. Thorax. Pronotum with anterior margin slightly concave and lateral margins that are straight and converging to a straight posterior margin that is ca. ½ as wide as the anterior margin (Fig. 14B ). Anterior and lateral margins of the pronotum have distinct rims, but the posterior margin has a weakly formed rim (Fig. 14B ). Face of the pronotum is marked by a distinct pit in the center with prominent anterior and lateral furrows from this central pit, and the pronotum surface is marked with wrinkles throughout, but not prominent nodes (Fig. 14B ). Prosternum surface is moderately granulose throughout, the mesosternum surface is mostly smooth with only small granulation along the sagittal plane which then continues onto the metasternum surface which has moderate granulation throughout (Fig. 15B ). Prescutum ca. ⅓ longer than the anterior margin width, with lateral margins slightly converging to the posterior (Fig. 14B ). Lateral margins lacking distinct tubercles, instead with granulation throughout giving them a rough appearance (Fig. 14B ). Prescutum surface throughout with granulation, with smaller and tighter packed granulation along the lateral margins, and slightly larger nodes along the sagittal plane (Fig. 14B ). Prescutum anterior margin distinct but not significantly raised, with a surface that is marked with minimal granulation. Mesopleura not notably wide, narrow near the anterior margin and then only gradually diverge for the remainder of their length (Fig. 14B ). Lateral margin with five or six small tubercles throughout the length and several smaller minor tubercles interspersed throughout. Face of the mesopleura slightly wrinkled and with two faint divots, one near the anterior margin and one on the posterior ⅓ (Fig. 15E ). Wings. Tegmina moderate length, extending ⅓ to ½ through 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 ca. ⅖ of the way through the wing length and terminating near the midline, followed by the branching and termination of the second radius (R2) near the distal ⅓ of the wing, and 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. ⅓ of the way through the wing length, the second media posterior (MP2) branching near the midline, and the media anterior (MA) running 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 of abdominal segments VIII. 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 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 ⅕ of the wing when 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. Abdominal segment II with parallel margins, III through the anterior ⅓ of segment IV gradually diverging to the widest portion. The posterior of IV-V are parallel or slightly subparallel. Segments VI-X uniformly converging, giving the abdomen a spade-shaped appearance (Fig. 14C ). Abdominal segment X distinctly longer than wide (Fig. 15F ). Genitalia. Poculum broad, and ending in a rounded apex that slightly passes the anterior margin of segment X (Fig. 14D ). Cerci long and slender, extending from under the anal abdominal segment by slightly> half of their length, surface slightly cupped and with a granulose surface with numerous short setae throughout (Fig. 14D ). Vomer broad and stout with straight sides evenly converging to the apex, which is marked by two apical hooks, one slightly larger than the other (Fig. 5G ). Legs. Profemoral exterior lobe slightly thinner than the interior lobe, arcing end to end in a smoothly bending lobe which has a granular margin throughout and can range from lacking dentation to having four or five small serrate teeth on the distal half of the lobe (Figs 14A , 15C ). Profemoral interior lobe roundly triangular and marked with five teeth arranged in a two-one-two pattern with large looping gaps between these groups (Fig. 15A ). Mesofemoral exterior lobe arcs end to end, but is more heavily weighted toward the distal half which is slightly broader than the mesofemoral shaft width, and which is marked with 1-3 serrate teeth on the distal half only, and the proximal half is rather thin and lacks dentation. Mesofemoral interior lobe is about the same width as the mesofemoral shaft and arcs end to end, is slightly broader on the distal end and is marked with four or five small serrate teeth. Metafemoral exterior lobe lacks dentation, and has a straight margin hugging the metafemoral shaft. Metafemoral interior lobe smoothly arcs end to end with seven or eight serrate teeth on the distal half only. Protibiae lacking exterior lobe, interior lobe reaching end to end in a smoothly rounded triangle which is slightly weighted to the distal half and at the widest point slightly <2 × the protibial shaft width (Fig. 15A ). Meso- and metatibiae simple, lacking lobes completely.

Measurements of holotype male [mm]. Length of body (including cerci and head, excluding antennae) 64.3, length/width of head 3.8/3.6, antennae 34.5, pronotum 2.9, mesonotum 3.7, length of tegmina 17.4, length of alae 47.0, greatest width of abdomen 20.6, profemora 13.7, mesofemora 10.6, metafemora 12.5, protibiae 8.2, mesotibiae 7.0, metatibiae 9.4.

Measurements of paratype males [mm]. Length of body (including cerci and head, excluding antennae) 61.9-69.5, length/width of head 3.2-3.6/3.1-3.7, antennae 32.8-41.2, pronotum 2.7-3.6, mesonotum 4.0-5.1, length of tegmina 19.7-19.9, length of alae 48.3-50.1, greatest width of abdomen 18.4-20.6, profemora 12.5-15.7, mesofemora 9.9-12.1, metafemora 10.8-14.2, protibiae 7.6-9.2, mesotibiae 7.8-8.2, metatibiae 8.0-10.4.

Etymology.

Patronym. This species is dedicated to our friend and colleague Alexandre Banko for his extensive efforts to discover new species and his long collaboration with Team Phyllies to present us with fresh material to sequence and study.

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 13. Cryptophyllium bankoi gen. et sp. nov. paratype female nymph collected in Bach Ma N. P. on Rubus sp. (Rosaceae), Vietnam, photographed by Jerome Constant (RBINS) on 11 th July 2011. Molecular sample RBINS 06 within this study A dorsal view with abdomen curled B dorsal view with abdomen held flat.

Figure 14. Cryptophyllium bankoi gen. et sp. nov. holotype male, photographs by RTC A front leg details, dorsal B details of the base of antennae, head, thorax C habitus, dorsal D details of genitalia, ventral.

Figure 15. Cryptophyllium bankoi gen. et sp. nov. paratype male from the Ba Na-Nui Chua Nature Reserve (RBINS), photographs by Jerome Constant (RBINS) A habitus, dorsal B habitus, ventral C pro- femoral and tibial, dorsal D details of the base of antennae, head, and thorax, dorsal E details of the base of antennae, head, and thorax, lateral F terminalia, dorsal G genitalia, ventral.

Figure 59. Cryptophyllium rarum comb. nov. male and female A male, habitus, dorsal, molecular sample within this study Coll RC 16 - 115 from Vietnam, Da Nang Province, photograph by RTC B details of the head and thorax, dorsal, Coll RC 16 - 115, photograph by RTC C female, habitus, dorsal, from China: Guangxi Province, Nanning city, Shuangding town, molecular sample DZW 08 within this study, Photographs by Zhiwei Dong (KIZ) D female, genitalia, ventral, photographs by Zhiwei Dong (KIZ).