Cryptophyllium rarum (Liu, 1993) 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 rarum (Liu, 1993) comb. nov. Figures 5I , 59 , 60 , 61 , 62

Material examined.

(5 ♀♀, 7 ♂♂): 1 ♀: "China: Guangxi Province, Nanning city, Shuangding town （ 双定镇）N22°59’41.39” E108°6 ’53.59”, VI-2016, Xiao-Yu Zhu. Molecular sample: DZW08" (Coll ZD); 1 ♀: "China, Guangxi, Liuzhou, Dayaoshan Mts, IX.2019" (IMQC); 1 ♀: "China, Guangxi, Jinxiu County, Dayao Mountain, July-Oct 2020" (Coll SLT); 1 ♀: "China, Guangxi, Jinxiu County, Dayao Mountain, Oct 2020" (Coll SLT); 1 ♀: "Museum Paris, Tonkin, reg. de Hoa-Binh, A. De Cooman 1927" (MNHN); 1 ♂: "Kon Tum Province, Ngoc Linh Mt. 1,700 m. elv. May 2015" (Coll RC 16-116); 1 ♂: "Vietnam: Da Nang Prov., Ba Na Mt.: May,2015 1,450 m." (Coll RC 16-115); 1 ♂ "Da Nang Province, Ba Na Mt. 1,450 m. elv. May 2015" (Coll RC 16-119); 1 ♂: "Coll. I.R.Sc.N.B., Vietnam, Ninh Binh prov., Cuc Phuong Nat. Park, 20°20 ’53” N 105°35 ’52” E, 31.vii-3.viii.2016, GTI Project, Leg. J. Constant & J. Bresseel, I.G.: 33.282, RBINS-PHYLLIUM DNA sample 0004" [vomer dissected] (RBINS); 1 ♂: "Coll. I.R.Sc.N.B., Vietnam, Ninh Binh prov., Cuc Phuong Nat. Park, 20°20 ’53” N 105°35 ’52” E, 2-8.vii.2019, Leg. J. Constant, I.G.: 34.032" (RBINS); 1 ♂: "Coll. I.R.Sc.N.B., Vietnam, Tay Yen Tu Nat. Res., 21°11 ’10’’ N 106°43 ’25” E, 7-11.vii.2013, night collecting, Leg J. Constant & J. Bresseel, I.G. 32.454" (VNMN); 1 ♂: "Coll. I.R.Sc.N.B., Vietnam, Tay Yen Tu Nat. Res., 21°11 ’10’’ N 106°43 ’25” E, 7-11.vii.2013, night collecting, Leg J. Constant & J. Bresseel, I.G. 32.454, RBINS-PHYLLIUM DNA sample 0005" (RBINS).

Remarks.

The holotype female Cryptophyllium rarum comb. nov. is from Hexian, Baise prefecture level division, Guangxi Province, we were able to obtain a tissue sample from a female specimen from Guangxi Province from the adjoining division of Nanning City (Fig. 59C ). With the species until now only known from a female, we were excited to uncover that large unknown males from Vietnam were molecularly in close relation to the female, allowing this species to be illustrated from both sexes (Fig. 60 ). During a joint GTI expedition, two males in Tay Yen Tu Nature Reserve and one male from Cuc Phuong N.P. were collected at night using a light trap. They were found while checking the surroundings of the light trap, having been attracted to the light but not landing on the white collection sheet. Additionally, a second male in Cuc Phuong N.P. was collected when it was attracted by the streetlights. Interestingly, inclusion of molecular data from six males from throughout north and central Vietnam did not reveal clearly delineated clades (which could represent distinct species) or a tightly formed clade with little genetic diversity (indicating a close kinship/homogenizing gene flow throughout this wide range), but instead somewhat revealed a nestedness of our samples from north to south with significant molecular distances between most samples (Fig. 4 ). Similar molecular distances were revealed between the geographically close Cryptophyllium bollensi sp. nov., Cryptophyllium nuichuaense sp. nov., and Cryptophyllium phami sp. nov. but our set of Cryptophyllium rarum comb. nov. samples come from a geographically wide range of several hundred kilometers. This trio of molecularly distinct and morphologically discernable species from within a limited geographic area contrasts greatly with our sampling of Cryptophyllium rarum comb. nov. males from a wide geographic area but no morphological feature was found to allow differentiation of these notably molecularly distinct samples. Due to the extreme geographic distance likely allowing a certain degree of intraspecific molecular variation and our lack of continuous samples of Cryptophyllium rarum comb. nov. from throughout the full range, we at this time treat this clade as a singular species but expect that future analysis may reveal it to be several once the larger geographic gaps in our sampling are filled.

Differentiation.

Females can be differentiated by the unique combination of exterior profemoral lobes which are right angled and distinctly serrate, alae which are long (reaching abdominal segment VI), and an abdomen which is distinctly tapering from the sixth segment to the tip of the abdomen (not parallel-sided or with a lobe). Species which are morphologically similar are Cryptophyllium athanysus comb. nov. (which has right angled exterior lobes and a spade-shaped abdomen) and Cryptophyllium icarus sp. nov. which as the slender form female can have a similar abdominal shape (Fig. 34A ). Cryptophyllium athanysus comb. nov. can immediately be differentiated however as this species has prominent exterior lobes on the meso- and metatibiae whereas Cryptophyllium rarum comb. nov. has simple tibiae, lacking lobes. Cryptophyllium icarus sp. nov. can be differentiated by the profemoral exterior lobes which are notably more obtuse angled, not right angled, and the lack of well-developed alae which are small, only reaching the second abdominal segment (Fig. 33C ).

Males are morphologically similar to Cryptophyllium bankoi sp. nov. due to their similarly shaped lobes on all legs, similar wing lengths, shape and spination of the thorax, and general abdominal shape. These two species occur sympatrically in central Vietnam and therefore we were only able to confidently differentiate them through molecular analyses which allowed us to observe the morphological variation in these species to more confidently separate them morphologically. The only two features which we consistently saw between these species were of the exterior profemoral lobe and the abdominal shape. Cryptophyllium rarum comb. nov. has a slightly more ovoid abdomen ( Cryptophyllium bankoi sp. nov. is slightly more tapered on the abdominal segments VI and VII) and the profemoral exterior lobe on Cryptophyllium rarum comb. nov. has 6-8 distinct serrate teeth (Fig. 61E ) vs. Cryptophyllium bankoi sp. nov. which at most have one to four small minor teeth (Figs 14A , 15C ), never a full set of distinctly serrate margins.

Distribution.

China, Guangxi Province (recorded from Hexian, Shuangding town, and Dayaoshan Mountains) and distributed south through Vietnam. At present we have records> for five Vietnamese provinces: Vinh Phuc, Quang Ninh, Ninh Binh, Da Nang, and Kon Tum Provinces.

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 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 33. Live adult Cryptophyllium icarus bred and photographed by Bruno Kneubu ̈ hler (Switzerland) A female B male C lateral view of female with tegmina held up to expose the dwarfed alae underneath.

Figure 34. Cryptophyllium icarus gen. et sp. nov. female paratype, photographs by Jerome Constant (RBINS) A habitus, dorsal B habitus, ventral C antennae and anterior of head capsule D profemoral and protibial lobes E head and thorax details dorsal F head and thorax details lateral G genitalia dorsal H 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).

Figure 60. Cryptophyllium rarum comb. nov. male collected by Jerome Constant (RBINS) and Joachim Bresseel (RBINS) in Cuc Phuong N. P., Vietnam in August 2016. Photographs by Jerome Constant (RBINS). This male is molecular sample RBINS 04 in our analysis.

Figure 61. Cryptophyllium rarum comb. nov. male from Cuc Phuong N. P., Ninh Binh Province, Vietnam collected by Jerome Constant and Joachim Bresseel (RBINS), 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 pro- femoral and tibial lobes, dorsal F terminalia, dorsal G genitalia, ventral.

Figure 62. Presumed Cryptophyllium rarum comb. nov. a historic female from Hoa Binh, Vietnam, loaned to the RBINS from the MNHN collection, photographs by Jerome Constant (RBINS) A habitus, dorsal B habitus, ventral C details of antennae and anterior of head capsule D profemoral lobe, dorsal E details of the head and thorax, dorsal F details of the head-thorax, lateral G terminalia, dorsal H genitalia, ventral I data label.