Cryptophyllium oyae (Cumming & Le Tirant, 2020) Cumming & Bank & Bresseel & Constant & Tirant & Dong & Sonet & Bradler, 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 oyae (Cumming & Le Tirant, 2020) comb. nov. Figures 5F , 8G , 8H , 9E , 50 , 51 , 52

Material examined.

The extensive paratype series within Coll RC and Coll SLT were examined to review the intraspecific variation within this species. See Cumming and Le Tirant (2020) for a detailed list of material examined. Additionally, 1 ♂: "Coll. I.R.Sc.N.B., Laos, NE, Mt Phu Phan, vi.2019, local collectors, I.G.: 34.159" [vomer dissected] (paratype male from original description) (RBINS).

Photographic records: 1 ♂, 1 ♀: "China, Yunnan Province, Maguan County, Gulinqing town ( 古林箐乡)” (observed by Xiang-Jing Liu, China); 1 ♂: "Vietnam, Ha Giang Province, Dung Ba Commune" (observed by Chien C. Lee, Malaysia); 1 ♂: "Thailand, Nan Province, Bo Kluea Tai District" (observed by Tatsatorn Dharithai, Thailand); 1 ♂: "Thailand: Phetchabun Province, Phetchabun Research Station" (Photograph shared by Tatsatorn Dharithai, Thailand); 1 ♀: "San Ku Ruins, Chiang Mai, Thailand" (observed by Rob Thacker, United Kingdom).

Remarks.

This species was only recently described and has entered the phylliid enthusiast breeding community in the last year or so (Fig. 50 ). Despite the relatively recent description, as a large prominent species it has been easy to identify from numerous observational records> from throughout mainland Asia, thus appearing to be a somewhat widespread species than was originally known (Fig. 2 ).

Differentiation.

Females are morphologically similar to Cryptophyllium tibetense comb. nov. and Cryptophyllium yunnanense comb. nov. due to the mesopleura which are prominent and reach uniformly from the anterior to the posterior of the prescutum and are marked with large prominent tubercles; the general femoral lobe shapes; the general abdominal shape with a boxy abdomen with a gently lobed abdominal segment VII; and due to the distal portion of the tibial exterior areas having small lobes. Both of these similar species can however be differentiated by the female subgenital plate as it is notably long in Cryptophyllium tibetense comb. nov. and Cryptophyllium yunnanense comb. nov. with the apex reaching the tip of the abdomen (Figs 63B , 77E ), vs. Cryptophyllium oyae comb. nov. which has a subgenital plate which is shorter, only reaching ca. ½ through abdominal segment X (Fig. 50D ).

Males are morphologically similar to Cryptophyllium limogesi sp. nov. and Cryptophyllium yunnanense comb. nov. due to their large size and spade-shaped abdomens. Cryptophyllium oyae comb. nov. shares the following morphological similarities to Cryptophyllium limogesi sp. nov.: similar femoral and tibial lobe shapes (particularly the profemoral exterior lobe which is broad, distinctly bent, and marked with prominent serration); and the occasional presence of small distal lobes of the tibial exteriors. Cryptophyllium oyae comb. nov. can be differentiated by the length of the tegmina as they reach significantly onto abdominal segment IV when folded, and in Cryptophyllium limogesi sp. nov. the tegmina only reach ½ onto abdominal segment III, and the mesopleura of Cryptophyllium limogesi sp. nov. are not as broad on the anterior as those of Cryptophyllium oyae comb. nov. which has mesopleura which are nearly straight margined, diverging evenly not gradually. Male Cryptophyllium oyae comb. nov. can be differentiated from Cryptophyllium yunnanense comb. nov. by the profemoral exterior lobe shape as it is slightly thinner than the interior lobe and smoothly arcing from end to end in Cryptophyllium yunnanense comb. nov. (Fig. 75B ) but notably broad and clearly bent in the center like in Cryptophyllium oyae comb. nov.

Distribution.

Widely ranging through several countries with only inland records> to date, no coastal records> yet known. This species has been located in Vietnam, Ha Giang Province, Dung Ba Commune (observed by Chien C. Lee, Malaysia; Fig. 51A ); the type locality of Laos, Houaphanh Province, Xam Neua District, Mount Phu Phan; China, Yunnan Province, Maguan County, Gulinqing town (古林箐乡) (observed by Xiang-Jing Liu, China; Fig. 52 ); and three localities from Thailand, one from Nan Province, Bo Kluea Tai District (observed by Tatsatorn Dharithai, Thailand, Fig. 51B ), from San Ku Ruins, Chiang Mai (observed by Rob Thacker, United Kingdom; Fig. 51C ), and from Phetchabun Province, Phetchabun Research Station.

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

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

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

Figure 50. Live Cryptophyllium oyae comb. nov. bred and photographed by Bruno Kneubu ̈ hler (Switzerland) A female B male C lateral view of female with tegmina held up to expose the alae D female genitalia, ventral E male genitalia, ventral.

Figure 51. Live Cryptophyllium oyae males from additional, non-type localities A from Vietnam: Ha Giang, Dung Ba Commune, observed and photographed by Chien C. Lee (Malaysia) in January 2020 B from Thailand: Nan Province, Bo Kluea Tai District observed and photographed by Tatsatorn Dharithai (Thailand) in September 2019 C from Thailand: Chiang Mai Province, San Ku Ruins, observed and photographed by Rob Thacker (United Kingdom) in January 2021.

Figure 52. Live Cryptophyllium oyae pair from China, Yunnan Province, Maguan County, Gulinqing town (古林箐乡) collected by Xiang-Jing Liu (China) in October 2020, photographed by Zhiwei Dong (KIZ) A adult female dorsal B adult male dorsal.

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.

Figure 75. Male Cryptophyllium yunnanense comb. nov. (Coll RC 16 - 120), photographed by RTC A habitus, dorsal B left front leg details C head-thorax details D genitalia details, ventral E head-thorax details, lateral.

Figure 77. Female Cryptophyllium yunnanense comb. nov. A, C, D, F are photographs of the female used in our molecular analysis (DZW 04) (the same female in Fig. 76) collected by Zhiwei Dong in China, Yunnan, Daweishan, Pinbian County, Honghe Prefecture, photographs by Zhiwei Dong B, E, G, H of a female from Vietnam, Lao Cai Province, Mt. Fan-si-pan (Coll RC 17 - 270), photographs by RTC A habitus, dorsal B habitus, dorsal C details of antennae, dorsal D details of front leg, dorsal E genitalia, ventral F details of thorax, dorsal G details of thorax, dorsal H details of the thorax, lateral.