Cryptophyllium chrisangi (Seow-Choen, 2017) 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 chrisangi (Seow-Choen, 2017) comb. nov. Figures 8M , 8N , 9B , 24 , 25 , 26

Material examined.

(7 ♀♀, 5 ♂♂, 10 eggs): 2 ♂♂, 2 ♀♀: "Singapore, ex breeding" (RBINS); 1 ♂: "Singapore (Pulau Ubin), Collected at light, by Francis Seow-Choen, January 5th, 2018 (0-75 m elv)" (Coll RC 18-279); 1 ♂: "Thailand: Satun Province, Tarutao Island" (UCR); 1 ♀: "Sarawak: Kuching, 12.12, J.M. Bryan., B.M.1931-150." (NHMUK); 1 ♀: "P. Phyllium siccifolium , 4.93 Tapah, Dr. Yeh" (LKCNHM); 10 eggs: “Singapore” (Coll RC 18-332-18-341); 1 ♀: "Presented by Dr. Brooke, St. John’s I. 26.4.09, Coll Freie Universitat Berlin Sammlung Exotische Insekten, DEI Hemimetabola #100117".

Photographic records: 2 ♀♀: Thailand: Nakhon Si Thammarat Province, Thung Song District, August 2020 (photographed by Tatsatorn Dharithai, Thailand);

1 ♂: Indonesia, Pulau Weh Island off the coast of Sumatra (photographed by Mathieu MJP Van Goethem, South Africa).

Remarks.

This species is one which was brought into the breeding community (Fig. 24A ) and has therefore allowed thorough observation of the adults, nymph, and egg morphology. Molecularly, Cryptophyllium chrisangi comb. nov. is sister species to Cryptophyllium westwoodii comb. nov. with these two species biogeographically separated by the Isthmus of Kra. The Isthmus of Kra is a notable line of biogeography for several organisms (e.g., Li and Li 2018; Parnell 2013) which separates the Ranong and Chumphon Provinces of Thailand (Fig. 2 ). This isthmus appears to be significant for the Phyllium (Phyllium) as no species are presently known north of this line, but insignificant for the Phyllium (Pulchriphyllium) as this group is found on both sides of this line.

With our herein designation of a neotype for Cryptophyllium westwoodii comb. nov. we can help to clear up possible significant confusion which surrounds this species. When Wood-Mason described this species in 1875 he did so with a male and female pair of syntypes from two different localities. The female was from "South Andaman" and the male was from "near Pahpoon, ca. 150 miles north of Moulmein, in the Salween country" ( Wood-Mason 1875). It is almost certain biogeographically that these syntypes represent two different phylliid species, not a singular Cryptophyllium westwoodii comb. nov. as he intended. Morphologically, Wood-Mason’s description and wonderful illustration of the female syntype interestingly appears very similar morphologically to Cryptophyllium chrisangi comb. nov. with the correct femoral and abdominal shapes, and alae length (Fig. 25 ). With our designation of a neotype Cryptophyllium westwoodii comb. nov. from a male from the mainland, this leaves the Andaman female somewhat mysterious as few records> of phylliids have been noted from the Andaman Islands, all of which are Phyllium (Pulchriphyllium) bioculatum -like species, none representing a Cryptophyllium gen. nov. species. Hopefully one day a fresh specimen of this Cryptophyllium gen. nov. species from the Andamans can be located and molecularly compared with congenerics to identify if it is an additional Cryptophyllium chrisangi comb. nov. range expansion or an undescribed species.

Differentiation.

Females can be differentiated by the following combination of features: spade-shaped abdomen (as segment VII lacks lobes), mesopleura which are distinctly narrower on the anterior half, and alae which are only ca. ½ as long as the tegmina. Morphologically, this species is similar to Cryptophyllium westwoodii comb. nov. due to the femoral and mesopleura shape, but can be differentiated by the shorter alae (only half of the tegmina length) as Cryptophyllium westwoodii comb. nov. has alae which are nearly the same length as the tegmina (Fig. 68D ). Additionally, Cryptophyllium athanysus comb. nov. is morphologically similar due to the femoral, mesopleura, and abdominal shape, but can immediately be differentiated by the metatibial exterior lobes which are lacking in Cryptophyllium chrisangi comb. nov.

Males are morphologically very similar to Cryptophyllium westwoodii comb. nov. and we have yet to find a reliable morphological feature to differentiate these two species. Both are morphologically variable and can even have a wide range of sizes which does not allow for confident differentiation when molecular markers and locality are unknown. One of the more consistent features however is the abdominal shape as Cryptophyllium westwoodii comb. nov. tends to have a slightly more spade-shaped abdomen with segments V-IX converging, and Cryptophyllium chrisangi comb. nov. having a slightly more ovoid abdomen with segments V-VI parallel or subparallel, but we have seen morphological intermediates which do not allow this as a diagnostic feature.

Distribution.

The type locality for Cryptophyllium chrisangi comb. nov. is mainland Singapore and it has additionally been recorded from St. John’s Island from a record in the SDEI collections. Additionally, we have observed specimens and photographs from several areas, including several from Thailand: Nakhon Si Thammarat Province, Thung Song District (Fig. 24D ) and Satun Province, Tarutao Island (UCR coll.); one tentative old record of a large female from Malaysia, Sarawak, Kuching, from the NHMUK collection (Fig. 26 ); an adult female from Tapah, Malaysia (LKCNHM); and a record from Indonesia, Pulau Weh Island off the coast of Sumatra (Fig. 24C ). Hopefully molecular samples from these far-reaching areas can one day be obtained to confirm the identification as Cryptophyllium chrisangi comb. nov. but for now morphologically this is what these records> appear to represent.

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 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 24. Live Cryptophyllium chrisangi comb. nov. A female bred and photographed by Bruno Kneubu ̈ hler (Switzerland) B male genitalia, ventral view, bred by Bruno Kneubu ̈ hler (Switzerland) C male observed and photographed by Mathieu MJP Van Goethem (South Africa) on Weh Island off the north shore of Sumatra D female observed and photographed in Thailand: Nakhon Si Thammarat Province, Thung Song District, by Tatsatorn Dharithai (Thailand) in August 2020.

Figure 25. Wood-Mason's 1875 plate XVII with the beautifully illustrated ' Phyllium westwoodii ' female syntype from South Andaman. Note the morphological similarity to Cryptophyllium chrisangi comb. nov. and given the wide range of Cryptophyllium chrisangi comb. nov. outside of the type locality of Singapore, this record may represent a range expansion. Public domain work downloaded from the Biodiversity Heritage Library (https: // www. biodiversitylibrary. org / item / 114410 page / 227 / mode / 1 up)

Figure 26. Tentative female Cryptophyllium chrisangi comb. nov. from Kuching, Sarawak, Malaysia which represents a range expansion for this species, photographs by Jerome Constant (RBINS) A habitus, dorsal B details of the anterior of the head, dorsal C habitus, ventral D front leg details, dorsal E details of the head and thorax, dorsal F details of the head and thorax, lateral G genitalia details, dorsal H genitalia details, ventral I data label.

Figure 68. Live adult Cryptophyllium westwoodii comb. nov. bred and photographed by Bruno Kneubu ̈ hler (Switzerland) from stock collected in Tha Pla Duk subdistrict, Mae Tha district, Lamphun, Thailand, collected in May / June 2014 by Suttah Ek-Amnuay (Thailand) A female habitus B male habitus C female details of the thorax, dorsal D lateral view female with tegmina lifted to show long alae E female genitalia details, ventral.