Cryptophyllium, the hidden leaf insects - descriptions of a new leaf insect genus and thirteen species from the former celebicum species group (Phasmatodea, Phylliidae) Author Cumming, Royce T. https://orcid.org/0000-0001-7930-1292 Montreal Insectarium, 4581 rue Sherbrooke est, Montreal, Quebec, Canada, H 1 X 2 B 2 & Richard Gilder Graduate School, American Museum of Natural History, New York, NY 10024, USA & Biology, Graduate Center, City University of New York, NY, USA roycecumming@gmail.com Author Bank, Sarah https://orcid.org/0000-0001-6952-1590 Department of Animal Evolution and Biodiversity, Johann-Friedrich-Blumenbach Institute for Zoology and Anthropology, University of Go ̈ ttingen, Untere Karspu ̈ le 2, 37073, Go ̈ ttingen, Germany sarah.bank@uni-goettingen.de Author Bresseel, Joachim Royal Belgian Institute of Natural Sciences, O. D. Taxonomy and Phylogeny and JEMU, rue Vautier 29, B- 1000, Brussels, Belgium Author Constant, Je ́ ro ̂ me Royal Belgian Institute of Natural Sciences, O. D. Taxonomy and Phylogeny and JEMU, rue Vautier 29, B- 1000, Brussels, Belgium Author Tirant, Stephane Le Montreal Insectarium, 4581 rue Sherbrooke est, Montreal, Quebec, Canada, H 1 X 2 B 2 Author Dong, Zhiwei State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences, Kunming, Yunnan, 650223, China Author Sonet, Gontran Royal Belgian Institute of Natural Sciences, O. D. Taxonomy and Phylogeny and JEMU, rue Vautier 29, B- 1000, Brussels, Belgium Author Bradler, Sven https://orcid.org/0000-0001-9307-1032 Department of Animal Evolution and Biodiversity, Johann-Friedrich-Blumenbach Institute for Zoology and Anthropology, University of Go ̈ ttingen, Untere Karspu ̈ le 2, 37073, Go ̈ ttingen, Germany text ZooKeys 2021 2021-02-18 1018 1 179 http://dx.doi.org/10.3897/zookeys.1018.61033 journal article http://dx.doi.org/10.3897/zookeys.1018.61033 1313-2970-1018-1 7E9360A5A359437A91C004C74B1FE9D6 84B0D9BEE71D5171B80C3F4CBFDC7366 Cryptophyllium tibetense (Liu, 1993) comb. nov. Figures 8C , 8D , 9D , 63 , 64 , 65 , 66 Material examined. (7 ♀♀, 4 ♂♂, 10 eggs): 1 ♂: "Tibet, Nyingchi Area, De'Ergong Village, Motutown 2020" (Coll SLT); 1 ♂: "China, Tibet, Motuo, Beibeng. 2019. 6-8." (Coll RC 20-001); 1 ♀ nymph: "Crowley Bequest. 1901-78. Sikkim" (NHMUK); 1 ♀ nymph: "Arunachal Pradesh from the Mishmi Hills. Delei River. 1,700ft. 28.i.1935, M.Steele." (NHMUK); 1 ♂ nymph: "Arunachal Pradesh from the Mishmi Hills. Lohit River. 22.iii.1935, M.Steele." (NHMUK); 1 ♂, 1 ♀: "bred from material collected in: Tibet China: Beibeng Township,Medog County, IX-2016, Jin Chen." (Coll ZD); 10 eggs: "China: Tibet: Medong Region: Bred by Bruno Kneubühler,2018" (Coll RC 18-396-18-405). Photographic records :1 ♀: "Kalimpong, West Bengal, India, September, 2019, photographed by Vandana Wadwa Sood (West Bengal, India)"; 1 ♀nymph: "Digboi, Assam 786171, India, iNaturalist user @rajib, by Rajib Rudra Tariang" (https://www.inaturalist.org/observations/61945900); 1 ♀: "Samthar, iNaturalist user @ripbumlepcha" (https://www.inaturalist.org/observations/35911522); 1 ♀: "Pasighat in the East Siang district, observed and photographed by Oken Tayeng" Remarks. Cryptophyllium tibetense comb. nov. is the highest latitude phylliid species known at present, and interestingly it is also one of the largest known species of Cryptophyllium gen. nov. (although the unknown female for Cryptophyllium animatum sp. nov. may be larger as the holotype male is 89.4 mm long suggesting a significantly sized female). The region where Cryptophyllium tibetense comb. nov. is found is more temperate (Fig. 66B ) than the tropics where phylliids are more typically known, and we hope that future ecological studies will reveal more about their generation time and general ecology in this region. This species was described from a female holotype, and it has entered the phasmid breeding community, however, as a parthenogenetic culture (Fig. 63 ), therefore records> of males are absent from culture and rarely collected. We have examined few male specimens, but all have matched morphologically to the male we illustrate from Beibengxiang, Medog County, Nyingchi, Tibet (Fig. 64B ). The first image we are aware of for this species is a beautiful color image from Inglis (1930) which was meant to be illustrating ' Phyllium scythe ' taken by Mr. T. A. Baldry (a mammologist active in the Darjeeling area, West Bengal in the 1920's and 30's). Inglis (1930) although confused as to which species he was illustrating (although to be fair we are aware of Phyllium (Pulchriphyllium) bioculatum scythe Gray, 1843 is also known from this area so he may have at one time seen the species he was intending to mention), he did note that leaf insects found in this area (although rare) were found feeding on Castanopsis hystrix , the Common Chestnut. This is the first record of phylliids feeding on Castanopsis , the second being of Cryptophyllium oyae comb. nov. from northern Laos (Cumming and Le Tirant 2020). Figure 63. Cryptophyllium tibetense comb. nov. live female bred and photographed by Bruno Kneubühler (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 64. Cryptophyllium tibetense comb. nov. live pair from Tibet and bred and photographed by Zhiwei Dong A female, habitus, dorsal B male, habitus, dorsal C male, details of the antennae, front legs, head, thorax, and tegmina, dorsal D male, details of the antennae, front legs, head, thorax, and tegmina, lateral view. Differentiation. Females are morphologically similar to Cryptophyllium drunganum comb. nov. and Cryptophyllium liyananae sp. nov. based on the general abdominal and femoral lobe shapes, the shape and spination of the thorax (Fig. 63C ), long alae at least extending abdominal segment VI (Fig. 63D ) and the presence of small exterior lobes on the tibiae (Fig. 63C ). Cryptophyllium tibetense comb. nov. can be differentiated from both however by the shape of the subgenital plate which is long and projecting beyond the tip of the abdomen in Cryptophyllium tibetense comb. nov. (Fig. 63B ) and shorter, only ca. 1/2 the length of abdominal segment X in the other two species (Fig. 47E ). Male Cryptophyllium tibetense comb. nov. are morphologically similar to Cryptophyllium rarum comb. nov. and Cryptophyllium bankoi sp. nov. due to the thorax shape and fine granular mesopleura margins (Fig. 64C ), tegmina lengths (Fig. 64D ), femoral lobe shapes (Fig. 64C ), and general abdominal shape. Both of these species can be differentiated from Cryptophyllium tibetense comb. nov. by the absence of small but distinct exterior, anteriorly situated protibial and metatibial lobes in the former two species. Distribution. The type locality for Cryptophyllium tibetense comb. nov. stated by Liu (1993) is Tibet (Xizang Autonomous Region), Medog County (Motuo). There do appear to be additional distribution records> for this high elevation species however. Unfortunately, we have only been presented with photographs of individuals or antique subadults/nymphs, and no fresh adult specimens to examine or include in our molecular phylogeny at this time. Therefore, these additional distribution records> are only presumed to be Cryptophyllium tibetense comb. nov. due to their morphological similarity to bred nymphs of this species and are here presented to give as thorough a view into the Cryptophyllium gen. nov. distribution as possible. From Southeast Tibet, Tenga Valley, we have been presented with an image of an adult male which looks to be the right size and has morphological features which suggest it could represent the Cryptophyllium tibetense comb. nov. male. Additionally, there is a pair of nymphs in the NHMUK from "Arunachal Pradesh from the Mishmi Hills" from 1935, a female from Anjaw District "Delei River. 1,700ft. 28.i.1935, M.Steele." (Fig. 65C ) and a male from the Lohit District "Lohit River. 22.iii.1935, M.Steele." (Fig. 65D ). Recently we also were sent photos of a female from Pasighat in the East Siang district, which matches well with the Cryptophyllium tibetense comb. nov. morphology (Fig. 66A ). Figure 65. Additional Cryptophyllium tibetense comb. nov. distribution records> A live adult female from Kalimpong, West Bengal, India, observed and photographed by Vandana Wadwa Sood (West Bengal, India) in September 2019 B female nymph, NHMUK from Sikkim, India, photographs by RTC C female nymph, NHMUK: "Burma: Mishmi Hills. Delei River. 1,700ft. 28.i.1935, M.Steele.", photographs by RTC D male nymph, NHMUK: "Burma: Mishmi Hills. Lohit River. 22.iii.1935, M.Steele.", photographs by RTC E female nymph observed and photographed in Digboi, Assam by Rajib Rudra Tariang (India). Figure 66. A Cryptophyllium tibetense comb. nov. from Pasighat in the East Siang district, observed and photographed by Oken Tayeng (India) next to a visually similar leaf from the area the female was found B habitat near where the female was found. From Nepal we are aware of two records>, both female subadults, one found in Gandaki Pradesh, Tanahun District and the other found in Province No. 1, Ilam. Both of these subadult females appear to have Cryptophyllium tibetense comb. nov. shaped abdomens, profemoral lobes, and importantly small exterior tibial lobes which help to characterize Cryptophyllium tibetense comb. nov. females. From India we have located a nymph from within the NHMUK collection from Sikkim (Fig. 65B ), and we have been lucky enough to be presented with photographs of an adult female from West Bengal, Kalimpong (Fig. 65A ) both of which have Cryptophyllium tibetense comb. nov. like features. From the state of Assam we have been sent an image of a female nymph from the town of Digboi which can clearly be seen as having distinct but small exterior tibial lobes (Fig. 65E ). Interestingly, we have yet to be presented with records> from Bhutan, but as it lies between areas where Cryptophyllium gen. nov. species have been confirmed we expect that there likely is at least one species present, just not yet officially recorded.