Cryptophyllium limogesi 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 limogesi gen. et sp. nov. Figures 5J , 8Q , 8R , 42 , 43 , 44 , 45

Material examined.

Holotype ♀: "VIETNAM, Lam Dong Province, Bao Lam, Dambri, V.2018". Deposited within the Montreal Insectarium (IMQC).

Paratypes: (13 ♀♀, 2 ♂, 7 eggs) • 1 ♂: "Coll. I.R.Sc.N.B., VIETNAM, Dak Nong prov., Ta Dung N.P., 11°52 ’22” N 107°58 ’40” E, 5-8.viii.2019, GTI Project, Leg. J. Constant and J. Bresseel, I.G.: 34.048 (Coll. I.R.Sc.N.B.)", vomer dissected (RBINS) (SB0531 molecular sample within our analysis) • 1 ♂: "Vietnam: Dak Lak Province, Local collector, September 2020" (IMQC) • 5 ♀♀; "Vietnam: Dak Lak Province, Local collector, September 2020" (Coll RC 20-127, 20-128, 20-129) • 5 ♀♀; "Vietnam: Dak Lak Province, Local collector, September 2020" (Coll SLT) • 3 ♀♀; "Vietnam: Dak Lak Province, Local collector, September 2020" (IMQC) • 1 egg: "Vietnam: Dak Lak Province, Local collector, September 2020" (Coll SLT) • 1 egg: "Vietnam: Dak Lak Province, Local collector, September 2020" (Coll FH) • 3 egg: "Vietnam: Dak Lak Province, Local collector, September 2020" (IMQC) • 2 egg: "Vietnam: Dak Lak Province, Local collector, September 2020" (Coll RC).

Remarks.

This large and morphologically unique species has been located in several provinces of southern Vietnam over the last few years and molecularly we found it to be sister species to Cryptophyllium icarus sp. nov. (Fig. 4 ) a species also only at the present known from southern Vietnam (Fig. 2 ). Interestingly, these molecular sister species are morphologically drastically different, perhaps due to their geographic sympatry. During a GTI joint expeditions two small nymphs were collected in Ta Dung N.P. on the forest edge adjacent to a coffee plantation. Tim Bollens (Belgium) reared one nymph to adulthood, thus revealing the male morphology (Fig. 42 ).

Differentiation.

Female Cryptophyllium limogesi sp. nov. are most morphologically similar to Cryptophyllium celebicum comb. nov. and Cryptophyllium tibetense comb. nov. based on the wide profemoral exterior lobe with an acute angle and broad boxy abdomen. From Cryptophyllium celebicum comb. nov., the easiest observed difference is the structure of the thorax, with the mesopleura of Cryptophyllium celebicum comb. nov. notably narrower on the anterior half vs. the mesopleura of Cryptophyllium limogesi sp. nov. reaching nearly to the anterior margin of the prescutum with straight margins (Fig. 42E ). From Cryptophyllium tibetense comb. nov. the female subgenital plates easily differentiate these species as Cryptophyllium tibetense comb. nov. has a long subgenital plate with a point which exceeds the apex of the terminal abdominal segment (Fig. 63B ) vs. Cryptophyllium limogesi sp. nov. where the subgenital plate is short, reaching no more than ½ through the terminal abdominal segment (Fig. 42H ).

Male Cryptophyllium limogesi sp. nov. are similar morphologically to Cryptophyllium oyae comb. nov. due to the broad spade-shaped abdomen and the profemoral lobe shapes. These species can be differentiated by the length of their tegmina (only reaching onto abdominal segment III in Cryptophyllium limogesi sp. nov. but reaching halfway onto segment IV in Cryptophyllium oyae comb. nov.); and they can be differentiated by the shape of the mesopleura as they are broad and nearly straight margined in Cryptophyllium oyae comb. nov. but slightly narrower on the anterior in Cryptophyllium limogesi sp. nov. males (Fig. 44C ).

Distribution.

Southern Vietnam: presently known from three provinces: the type locality of Lam Dong Province, Bao Lam, Dambri; the male paratype record from Dak Nong Province, Ta Dung N.P.; an observational record from Dak Lak Province, Chu Yang Sin N.P.; and paratype records> from Dak Lak Province.

Description.

Female. Coloration. Coloration description is based on photos of the holotype female shortly after being preserved. Nearly the entire body was of a uniform lime-green without differing colored markings (all legs and even wing venation a similar color to the rest of the body). Only the compound eyes were slightly yellow and not the same shade of green as the rest of the body.

Morphology. Head. Head capsule longer than wide, vertex with a moderately granular surface, and the posteromedial tubercle which is three or four times larger than the most prominent granules on the capsule. Frontal convexity broad and stout, notably shorter than the length of the first antennomere, and with several long, thin, clear setae across the surface. Compound eyes are not large, only slightly protruding from the head capsule and with a width of ca. ¼ the head capsule length (Fig. 42C ). Ocelli absent. Antennal fields wider than the first antennomere but not protruding back farther than the frontal suture. Antennae. Antennae consist of nine segments, with the terminal segment approximately the same length as the previously three segments combined (Fig. 42D ). The eighth antennal segment has a distinct furrow around the middle which makes the segment appear to be two separate segments (giving the antennae a ten segmented appearance), but this furrow appears to only be superficial (Fig. 42D ). Antennomeres I-III sparsely marked with thin transparent setae, similar to those found on the frontal convexity, but slightly shorter in length. Antennomere IV is short, disk-like, and wider than the following segments, and interestingly has a base which is narrow and somewhat longer than other congeneric antennal segments IV, giving it a raised appearance (Fig. 42D ). The terminal antennomere and the distal half of segment VIII (distal to the midline furrow) are covered in dense, stout, setae. Thorax. Pronotum with anterior margin slightly concave and lateral margins that are relatively straight, converging to a narrow, straight posterior margin that is ca. ½ the width of the anterior rim (Fig. 42E ). The pronotum surface is smooth, with only a prominent pit in the center, and slight furrows anterior and lateral to the pit, no prominent wrinkles or granulation. The pronotum has a prominent anterior rim and moderate lateral rims, the posterior is lacking a rim. Prosternum with notable nodes throughout the surface, relatively evenly spaced. Mesosternum with prominent nodes on the anterior margin, followed by moderate nodes on the anterior ⅓ of the surface, with the remainder with dispersed, weak granulation which continues onto the metasternum. Prescutum notably longer than wide, with lateral margins running parallel to the posterior margin giving it a distinctly rectangular appearance. Lateral rims with five or six medium sized tubercles situated on the anterior ⅔, with only small granulation on the remainder. Prescutum anterior rim prominent but not strongly protruding, with the surface granular and lacking a prominent sagittal tubercle. Prescutum surface with granulation throughout with those along the sagittal plane slightly larger (Fig. 42E ). Mesopleura start ca. ⅓ down the prescutum and evenly diverge with straight lateral margins. Lateral margin with six or seven major and distinctly pointed tubercles and six or seven smaller tubercles intermixed amongst them. This mix of tubercles is only prominent along the anterior ⅔ of the length with the remaining ⅓ lacking notable tubercles and instead marked with consistent granulation (Fig. 42E ). Face of the mesopleura with slight wrinkles throughout most of the surface and a few irregular nodes along the lateral margin, as well as two faint divots, one on the anterior margin and one closer to the center. Wings. Tegmina reaching slightly past the anterior margin of abdominal segment VII. Tegmina venation is rather typical for the Cryptophyllium gen. nov. 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. 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. ⅓ 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 wing midline 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 ¼ of the wing. The cubitus (Cu) runs throughout the entire wing length simply, and then near the posterior ⅕ of the wing becomes bifurcate 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 short, only 21.8 mm long. Abdomen. Abdominal segments II through the anterior half of IV diverging, posterior half of IV through the anterior ⅓ of VII parallel. Abdominal segment VII with a distinct looping lobe which meets abdominal segment VIII which is notably narrower. Segments VIII-X uniformly converge to a broad rounded apex. Genitalia. Projecting portion of the subgenital plate stout, beginning at the anterior margin of abdominal segment IX and projecting with nearly straight sides to just under the anterior margin of the terminal abdominal segment (Fig. 42H ). Gonapophyses VIII are long and broad with a dagger-like shape (parallel-sided at first and then after ca. ½ of the length uniformly converging to the point) with the points just projecting from under the terminal abdominal segment (Fig. 42H ). Gonapophyses IX are smaller and not visible from under the large gonapophyses VIII. Cerci strongly pointed and relatively flat (not strongly cupped) with weakly crenate margins, and the dorsal surface heavily granular and marked by thin transparent setae throughout (Fig. 42G, H ). Legs. Profemoral exterior lobes broad, at its broadest ca. 2 × wider than the interior lobe, and distinctly serrate throughout the entire length (with 13-17 small, pointed teeth). The proximal edge is slightly concave and the distal edge is smoothly convex, therefore giving the lobe a distinct recurved appearance and an acute exterior angle (Fig. 42F ). Interior profemoral lobe ca. 2½× the width of the profemoral shaft at its widest and with doubly serrate, large, triangular teeth. The largest teeth are grouped into a two-one-two pattern with large looping gaps between these groupings, with these large gaps also finely serrate, not smooth (Fig. 42F ). Interior mesofemoral lobe arcs evenly weighted from end to end, and at its widest is approximately the same width as the mesofemoral shaft itself. The interior mesofemoral lobe is finely serrate for ca. ¾ of its distal length with seven or eight teeth. Mesofemoral exterior lobe is also approximately at its widest as wide as the mesofemoral shaft, but the exterior lobe is distinctly bent in the center with straight margins, not smoothly arcing from end to end. On the distal half of the lobe only there are six or seven small serrate teeth. Metafemoral interior lobe arcs end to end and has five or six dull teeth pointing distally. Metafemoral exterior lobe is thin and smooth, hugging the metafemoral shaft and lacks dentation. Protibiae with a thin but notable exterior lobe on the distal ⅕ only. Protibial interior lobe spans the entire length as a broad scalene triangle with the broad end on the distal half of the protibiae. Mesotibiae simple, completely lacking lobes. Metatibiae lacking interior lobes, exterior is marked by a very slender lobe which only occupies the distal ¼ of the shaft.

Measurements of holotype female [mm]. Length of body (including cerci and head, excluding antennae) 99.0, length/width of head 8.9/6.8, antennae 5.1, pronotum 5.9, mesonotum 6.5, length of tegmina 55.7, length of alae 21.8, greatest width of abdomen 41.5, profemora 22.8, mesofemora 16.0, metafemora 19.1, protibiae 11.2, mesotibiae 11.8, metatibiae 15.0.

Male. Coloration. Coloration description based on the captive reared paratype male when it was alive (Fig. 43 ). Overall coloration mint-green throughout with highlights of tan coloration on the distal half of the protibiae, along all femoral lobe margins, the frontal convexity, the margins of the thorax, the base of the antennae and the distal tips of each longer antennomere, and intermittently along the tegmina and alae sclerotized veins. Abdominal segment V has a slightly darker patch and a transparent eye spot on each side of the midline. Compound eyes are pale yellow with slight orange marbling throughout.

Morphology. Head. Head capsule approximately as long as wide, with a vertex that is only slightly granular with no discernable pattern (Fig. 44C ). The posteromedial tubercle is not broad and is only weakly raised from the head capsule. Compound eyes are large and bulbous, taking up ca. ⅖ of the head capsule lateral margins (Fig. 44C ). There are three well-developed ocelli between and slightly posterior to the compound eyes (Fig. 44C ). Antennal fields are about as wide as the scapus. Antennae. Antennae (including the scapus and pedicellus) consist of 29 segments. The scapus and pedicellus are nearly completely bare, lacking long setae. The following segments except the terminal three are covered in dense setae that are as long as or longer than the antennae segment is wide, and the terminal three segments are covered in dense short setae. Thorax. Pronotum with anterior margin slightly concave and lateral margins that are straight and converging to a slightly curved posterior margin that is ca. ½ the width of the anterior rim (Fig. 44C ). Anterior and lateral margins have moderate rims and the posterior margin lacks a rim (Fig. 44C ). Face of the pronotum is marked by a distinct pit in the center with a furrow anterior to the pit along the sagittal plane, weakly formed furrows lateral to the central pit, and a smooth surface with only slight granulation in no detectable pattern (Fig. 44C ). The prosternum is slightly granulose throughout and the anterior ⅓ of the mesosternum surface is marked with more prominent nodes, with the remainder of the surface with fewer and smaller nodes (Fig. 44B ). Prescutum longer than wide, with lateral margins slightly converging to the posterior (Fig. 44C ). Lateral rims with eight or nine small tubercles (Fig. 44C ). Prescutum surface slightly raised along the sagittal plane with six or seven nodes of varying size, with the remainder of the surface with only slight granulation throughout. Prescutum anterior margin weakly formed and with a granular surface, lacking a prominent central tubercle. Mesopleura narrow on the anterior quarter of the length but then gently diverging with nearly straight margins to the posterior (Fig. 44C ). Mesopleura lateral margin with three large conical tubercles, three or five small tubercles, and four or five nodes throughout the length (Fig. 44C ). Face of the mesopleura slightly wrinkled, with slight granulation throughout, and with two faint divots, one on the anterior margin and one near the midline. Wings. Tegmina moderate length, extending halfway through abdominal segment III. Tegmina wing venation: the subcosta (Sc) is the first vein, is simple, and terminates the earliest slightly < ½ 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 near the midline, followed by the branching and termination of the second radius (R2) slightly distal to the midline, 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. ⅓ of the way through the wing length, then the second media posterior (MP2) branching 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 ca. ⅓ of the way through the wing length. Alae well-developed in an oval fan configuration, long, reaching the middle of abdominal segment VIII. Abdomen. Abdominal segment II gently diverging, III through the anterior half of segment IV diverging to the widest portion of the abdomen. The posterior of IV-V parallel-sided giving the abdomen a broad spade-shaped appearance. Segments VI-X uniformly converging with slightly undulating margins (Fig. 44B ). Genitalia. Poculum broad, and ends in a broadly rounded apex that slightly passes the anterior margin of segment X (Fig. 44G ). Cerci long and slender, with> ½ their length extending from underneath the terminal abdominal segment (Fig. 44F ), relatively flat, covered in a heavily granulose surface and with equally granular margins, and surface with numerous short setae (Fig. 44G ). Vomer stout with slightly rounded sides converging to an apex with two side by side thick apical hooks which are the same size and hook upwards into the paraproct (Fig. 43C ). Legs. Profemoral exterior lobe broader than the interior lobe, arcing end to end with a distinct rounded bend in the center, with the proximal half margin with a distinctly granular surface, and the distal half with six small serrate teeth (Fig. 44E ). Profemoral interior lobe roundly triangular, at least 3 × wider than the profemoral shaft, and marked with five large, serrate teeth arranged in a two-one-two pattern with looping gaps between them (Fig. 44E ). Mesofemoral exterior lobe arcs end to end, is slightly wider than the mesofemoral shaft, but with the widest portion on the distal ⅓ which is marked with granulation or with one or two weakly formed teeth. Mesofemoral interior lobe is slightly thinner than the exterior lobe and is slightly broader on the distal end which is marked with six 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 the distal half wider than the proximal half, and the distal half is marked with seven or eight serrate teeth on the distal half. Protibiae with a small but notable exterior lobe on the anterior ⅕ which is no wider than the width of the protibial shaft (Fig. 44E ). Protibial interior lobe reaching end to end in a smoothly rounded triangle with the widest portion on the distal third ca. 3 × as wide as the protibial shaft (Fig. 44E ). Meso- and metatibiae simple, lacking lobes.

Measurements of reared paratype male [mm]. Length of body (including cerci and head, excluding antennae) 79.7, length/width of head 4.9/3.9, antennae 41.8, pronotum 3.7, mesonotum 5.7, length of tegmina 24.4, length of alae 57.5, greatest width of abdomen 26.5, profemora 16.4, mesofemora 11.4, metafemora 13.7, protibiae 9.1, mesotibiae 7.7, metatibiae 10.9.

Eggs. (Fig. 45 ). When viewed from the anterior the egg capsule cross section is rounded pentagonal, therefore the lateral surfaces are raised into dorsolateral and ventrolateral surfaces. All surfaces and margins slightly undulate giving the egg an overall lumpy appearance. The ventrolateral surface is marked by four large, evenly spaced pits from the anterior to the posterior in a singular line. The dorsolateral surface has five large pits, arranged with one on the anterior, one on the posterior, and three in the middle spaced out in a two-one pattern. Both lateral surfaces are primarily bare but do have sparse and small moss-like pinnae between the pits. The dorsal surface is marked with seven medium sized pits total (one on the anterior end along the sagittal plane at the apex of the micropylar plate, followed by three on each slide of the plate with broad spacing between them ending with the posterior most near the base of the capsule). The dorsal surface is also marked throughout with short moss-like pinnae around the micropylar plate, with the area immediately around each pit bare. The micropylar plate is long, ca. 5/7 of the overall dorsal surface length with the micropylar cup situated on the posterior ⅓ of the length. The micropylar plate is thin with the widest portion the area around the micropylar cup. The operculum is slightly ovular with a surface that is roundly raised and a height slightly < ½ the operculum width. The operculum is marked intermittently with moss-like pinnae similar in shape but slightly smaller than those found on the rest of the capsule, as well as marked with two medium sized pits (one on each side of the sagittal plane). The overall egg color is dark brown, with the moss-like pinnae light brown in color so they stand out clearly on the surface.

Measurements including the extended pinnae [mm]. Length (including operculum): 5.2; maximum width of capsule when viewed from lateral aspect 3.8; length of micropylar plate 3.0.

Etymology.

Patronym. Named after René Limoges (Canada) from the Montreal Insectarium to thank him for his many years of assisting Team Phyllies with countless publication worthy photographs.

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 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 42. Holotype female Cryptophyllium limogesi, photographs by Rene Limoges (IMQC) A habitus, dorsal B habitus, ventral C head and antennae dorsal D antennae detail dorsal E thorax dorsal F front leg, dorsal G terminalia, dorsal H genitalia, ventral.

Figure 43. Paratype male Cryptophyllium limogesi gen. et sp. nov. from Ta Dung N. P. reared to adulthood and photographed by Tim Bollens (Belgium) A habitus, dorsal B detail of the front legs and head C genitalia details, ventral.

Figure 44. Paratype male Cryptophyllium limogesi gen. et sp. nov. same individual as in Figure 43 photographs by Jerome Constant (RBINS) A habitus, dorsal B habitus, ventral C details of the base of antennae, head, and thorax, dorsal D details of the base of antennae, head, and thorax, lateral E front leg details, dorsal F genitalia details, dorsal G genitalia details, ventral.

Figure 45. Paratype Cryptophyllium limogesi gen. et sp. nov. egg, laid by paratype females from Dak Lak Province, Vietnam. Photographs by Rene Limoges (IMQC) A lateral B dorsal C ventral D opercular (anterior) view.

Figure 63. Cryptophyllium tibetense comb. nov. live female bred and photographed by Bruno Kneubu ̈ hler (Switzerland) A dorsal, habitus B ventral genitalia, note the subgenital plate which reaches all the way to the apex of the abdomen C dorsal head, thorax, and front legs D lateral view with tegmina raised to expose the alae length.