Cryptophyllium westwoodii (Wood-Mason, 1875) Cumming & Bank & Bresseel & Constant & Tirant & Dong & Sonet & Bradler, 2021
publication ID |
https://dx.doi.org/10.3897/zookeys.1018.61033 |
publication LSID |
lsid:zoobank.org:pub:7E9360A5-A359-437A-91C0-04C74B1FE9D6 |
persistent identifier |
https://treatment.plazi.org/id/602EFA0F-6D16-5D66-9074-356F7C502A98 |
treatment provided by |
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scientific name |
Cryptophyllium westwoodii (Wood-Mason, 1875) |
status |
comb. nov. |
Cryptophyllium westwoodii (Wood-Mason, 1875) View in CoL comb. nov. Figures 5A View Figure 5 , 6G View Figure 6 , 6H View Figure 6 , 8A View Figure 8 , 8B View Figure 8 , 9F View Figure 9 , 68 View Figure 68 , 69 View Figure 69 , 70 View Figure 70 , 71 View Figure 71 , 72 View Figure 72
Material examined.
Neotype ♂: "THAILAND: Chiang Mai Province: October 2010. Coll RC 16-148". Deposited within the Montreal Insectarium (IMQC). Molecular sample 16-148 within this study.
Additional material examined.
(15 ♀♀, 21 ♂♂, 4 eggs): 3 ♀♀: "Thailand: Chiang Mai, July 2017." (Coll RC 18-145, 18-146, 18-147); 1 ♀: "Chiang Mai, Fang: February, 2011" (Coll RC 16-211); 1 ♀: "Thailand, Fang, II-2011" (Coll RC 16-212); 1 ♀: "Lamphun Province, Maetha: September, 2011" (Coll RC 16-080); 1 ♀: "Thailand, Lamphun Province, 2009 November" (Coll RC 16-078); 1 ♀: "Northern Thailand, Chiang Mai Province, 2010, October" (Coll RC 16-079); 1 ♀: "Laos: Luang Prabang Province, Kiew Mak Nao Village, 900m.: June, 2014" (Coll RC 16-077); 1 ♂: "Thailand: Chiangmai, Doi Pui, 25 May 1985" (Coll RC 16-082); 1 ♂: "Thailand: Lampon, Mae Tha, 09/2011" (Coll RC 16-083); 1 ♂: "Thailand: Chiangmai, Doi Pui, 19 May 1985" (Coll RC 16-214); 1 ♂: "Thailand: Chiangmai, Doi Pui, 28 May 1985" (Coll RC 16-215); 1 ♂: "Thailand: Chiangmai, Doi Pui, 24 May 1985" (Coll RC 16-216); 1 ♂: "Thailand: Chiangmai, Doi Pui, 25 May 1985" (Coll RC 16-217); 1 ♂: "Thailand: Lampang, May 2001" (Coll RC 16-218); 1 ♂: "North Laos: Kiew Mak Nao, VII.2015, 900m. S. Collard leg" (Coll RC 18-030); 2 ♂♂: "Thailand: Lampoon, Mae Tha, 09-2011" (Coll RC 16-147, 16-213); 1 ♂: "Burma: 4km E. Karathuri, Top of Hill, 350 to 400m., VI. 2011., Coll. A. Banko/ Collected by beating tree in Forest" (Coll RC 18-029); 1 ♂: "Coll. I.R.Sc.N.B., Thailande (Loei), Na Haeo (bio station), 05-12.V.2001, Light trap, Leg. J. Constant & P. Grootaert" [vomer dissected] (RBINS); 2 ♂♂: "Coll. I.R.Sc.N.B., Thailande (Loei), Na Haeo (field res stat)), 15-19.V.2003, Light trap, Leg. J. Constant, K. Smets & P. Grootaert" (RBINS); 1 ♂: "Coll. I.R.Sc.N.B., Thailande (Loei), Na Haeo, light trap, 15-19.V.2003, Light trap, Leg. J. Constant & K. Smets" (RBINS); 1 ♂: "Coll. I.R.Sc.N.B., Thailande (Loei), Na Haeo, light trap, 15-19.V.2003, Light trap, Leg. J. Constant & K. Smets, RBINS-PHYLLIUM DNA sample 0002" (RBINS); 1 ♂: "Coll. I.R.Sc.N.B., Thailande (Loei), Na Haeo, forest clearing, light trap, 16.V.2003, Light trap, Leg. J. Constant & K. Smets" (RBINS); 1 ♂: "Coll. I.R.Sc.N.B., Thailand, Loei, Na Haeo, 22.V.2000, Station 20007, Leg P. Grootaert" (RBINS); 1 ♂: "Coll. I.R.Sc.N.B., Laos, Bokeo prov., Ban Muang Kan, 1-15.vi.2012, local collectors, I.G.: 32.213, RBINS-PHYLLIUM DNA sample 0012" (RBINS); 1 ♀: "Coll. I.R.Sc.N.B., Laos, Bokeo prov., Ban Muang Kan, 1-15.vi.2012, local collectors, I.G.: 32.213, RBINS-PHYLLIUM DNA sample 0011" (RBINS); 2 ♀♀: "Coll. I.R.Sc.N.B., Laos, Bokeo prov., Ban Muang Kan, 1-15.vi.2012, local collectors, I.G.: 32.213" (RBINS); 1 ♀: "Coll. I.R.Sc.N.B., Thailand, SE Chiang Mai, Salok, Wang Chin, Near Lamphang, ex breeding A. & C. Bauduin, 2015" (RBINS); 1 ♀, 1 ♂: "Coll. I.R.Sc.N.B., Ex breeding Bruno Kneubühler, 2017, Thailand, Lamphun prov., Tha Pla Duk" (RBINS); 1 ♀, 1 ♂: "Thailand, Ex Culture Kristien Rabaey (RBINS); 4 eggs: N-Thailand, Cultured F.Hennemann 1995-2001. Ex. Coll. Frank Hennemann (Germany)" (Coll RC 18-242-18-245).
Type material and discussion.
Unfortunately, the male/female pair of syntypes which were originally deposited within the NZSI are considered lost ( Hennemann et al. 2009; Brock et al. 2020). Philip E. Bragg, well-known phasmid researcher from the United Kingdom, several years ago was able to obtain photographs of what was assumed to be the female syntype within the NZSI collection (the possible male type could not be located). However, the female labeled as such within the collection is the wrong size, does not have an original Wood-Mason collection label, and when compared with the illustration in Wood-Mason (1875) instead clearly represents a different species (a Phyllium (Pulchriphyllium) female). While it is possible that the original syntypes are still in the NZSI collection, if their original data labels were accidentally removed, or moved to other specimens, then there would be no way to positively identify either the male or the female. Besides their lost status, with our understanding of Cryptophyllium gen. nov. morphology now consolidated and extensively reviewed, we believe that Wood-Mason’s (1875) original syntype pair actually represented two different species, one from the mainland, and a second species from the Andaman Islands. Additionally, with the identification of a cryptic, nearly morphologically indiscernible Cryptophyllium westwoodii -like species from Cambodia ( Cryptophyllium khmer sp. nov.), a clear identification of the population which corresponds to this original name is necessary. A male specimen could not be located from the exact type locality "Pahpoon (Hpapun, Papun), 150 miles north of Moulmein (Mawlamyine, formerly Moulmein) in Salween Country (Salween River, officially Thanlwin River)" therefore we instead chose a rather morphologically average male from Chiang Mai, Thailand. Although this location is across an international border, it is only 175 kilometers away, a distance which is negligible when considering the range over which we have found Cryptophyllium westwoodii comb. nov.; we have specimens genetically confirmed as the same species across a distance of over 1,000 kilometers (from northern Laos to southern Myanmar; Fig. 2 View Figure 2 ). Additionally, this neotype location is the closest to the syntype locality for which we have seen specimens recorded, and Chiang Mai is a well-known breeding site for Cryptophyllium westwoodii comb. nov., therefore many museums and collections around the world have ample specimens of this particular population. Also, obtaining specimens from Myanmar is exceedingly difficult and no museum specimens could be located during our review. Therefore, due to the above complex taxonomic problems involved with the original name (Article 75.1, ICZN 1999), we here establish a neotype male which matches the morphological description in Wood-Mason (1875) and the illustration of the syntype presented in Wood-Mason (1877). Consequently, due to our above reasonings and the fact that historically a lectotype was never designated from the syntype pair, the qualifying conditions for designating a neotype are satisfied in accordance with Article 75.3 of the ICZN (1999).
Remarks.
The female and egg morphology of Cryptophyllium westwoodii comb. nov. were well-described by Hennemann et al. (2009) and this species is common within the phasmid breeding community (Fig. 68 View Figure 68 ) and therefore we only describe the neotype male morphology herein. This species appears to be one of the most widespread and most commonly encountered Cryptophyllium gen. nov. species as the range from north to south is over 1,000 km and from our review of museum collections and citizen scientist records> this is by far the most commonly observed species.
Differentiation.
Female Cryptophyllium westwoodii comb. nov. are morphologically inseparable from Cryptophyllium khmer sp. nov. due to the wide range of morphological forms observed within Cryptophyllium westwoodii comb. nov. (both in shape and coloration; Fig. 69 View Figure 69 ) which do not allow a reliable morphological feature to be identified for differentiation. When only comparing females, solely through molecular comparison can these two species be differentiated with confidence (see Fig. 4 View Figure 4 ). Thankfully, males of these two species can be morphologically differentiated, discussed further below. Additionally, Cryptophyllium westwoodii comb. nov. is also morphologically similar to the three southern Vietnam species: Cryptophyllium bollensi sp. nov., Cryptophyllium phami sp. nov., and Cryptophyllium nuichuaense sp. nov. females. All of these species share similar femoral and tibial lobe shape and serration (Fig. 70D View Figure 70 ), narrow anterior margin of the mesopleura and serration (Fig. 70E View Figure 70 ), and an abdominal shape which is boxy with a rounded lobe VII (Fig. 70A View Figure 70 ). From all of these southern Vietnam species however Cryptophyllium westwoodii comb. nov. can be differentiated by the length of the alae which are long, reaching onto abdominal segment VI (Fig. 68D View Figure 68 ), vs. all these others which have shorter alae, reaching segments II or III only.
Male. Cryptophyllium westwoodii comb. nov. are morphologically similar to Cryptophyllium athanysus comb. nov. and Cryptophyllium chrisangi comb. nov. due to their similar femoral lobe shape and serration, their shorter tegmina length (only reaching abdominal segment IV), and their general abdominal shape (Fig. 71A View Figure 71 ). Cryptophyllium athanysus comb. nov. can immediately be differentiated by the presence of fully spanning metatibial exterior lobes, as Cryptophyllium westwoodii comb. nov. lacks exterior lobes on all tibiae. Cryptophyllium westwoodii comb. nov. and Cryptophyllium chrisangi comb. nov. are very similar in morphology, and the only consistent feature we have seen between these to differentiate them is the size, with Cryptophyllium chrisangi comb. nov. slightly larger (73-74 mm long; Seow-Choen 2017) and Cryptophyllium westwoodii comb. nov. smaller (63-69 mm long; Hennemann et al. 2009). Despite female Cryptophyllium westwoodii comb. nov. being inseparable morphologically from Cryptophyllium khmer sp. nov., the males do consistently differ in the width of their abdomen, with Cryptophyllium westwoodii comb. nov. having an abdominal shape that is thinly elliptical, with a maximum width only 30-34% of the abdominal length (Fig. 71A View Figure 71 ), vs. Cryptophyllium khmer sp. nov. which has an abdominal shape broadly elliptical or broadly spade-shaped with a maximum width ca. 38-45% of the abdominal length (Fig. 40A View Figure 40 ).
Distribution.
Cryptophyllium westwoodii comb. nov. has only been confirmed through genetic analysis from northern Thailand, northern Laos, and southern Myanmar. With the description of Cryptophyllium khmer sp. nov. which morphologically cannot be differentiated from photographs of nymphs or females, we are unsure where these two species biogeographically are separated, but at this time we only know of Cryptophyllium khmer sp. nov. from Cambodia and are unsure if Cryptophyllium westwoodii comb. nov. also occurs in this country. Until additional Cryptophyllium westwoodii comb. nov. samples from throughout the range are also sequenced, the true distribution must remain somewhat vague at this point (as indicated by the bi-colored symbols in our distribution map; Fig. 2 View Figure 2 ).
Neotype male.
Coloration. Coloration description is based upon the dried neotype specimen (Fig. 72A View Figure 72 ), living individuals are more vibrant. Overall coloration pale green with variable patches of straw yellow throughout due to the drying process (primarily around the center of the body and the antennae). Compound eyes burnt red in color and basitarsi are orange.
Morphology. Head. Head capsule about as long as wide, with a vertex that is smooth except for the posteromedial tubercle which is not broad but is distinctly raised from the head capsule (Fig. 72B View Figure 72 ). Frontal convexity stout with a narrow point and marked with sparse thin setae. Compound eyes large and bulbous, taking up ca. ⅖ of the head capsule lateral margins (Fig. 72B View Figure 72 ). There are three moderately developed ocelli located between and slightly posterior to the compound eyes. Antennal fields as wide and as long as the scapus. Antennae. Antennae (including the scapus and pedicellus) consists of 27 segments, all segments except the scapus and pedicellus and terminal three segments are covered in dense pale setae that are as long as or longer than the antennae segment is wide. The terminal four segments are covered in dense, dark, short setae and the scapus and pedicellus are nearly completely bare. Thorax. Pronotum with anterior margin slightly concave and lateral margins that are nearly straight and converging to a straight posterior margin that is ½ the width of the anterior rim (Fig. 72B View Figure 72 ). Anterior and lateral margins have moderate rims, and the posterior margin lacks a rim (Fig. 72B View Figure 72 ). Face of the pronotum is marked by a distinct furrow in the center, short furrows lateral to this central sagittal furrow, and a smooth but slightly lumpy surface (Fig. 72B View Figure 72 ). Prosternum is moderately granular throughout with small nodes of even size. Mesosternum surface marked with slightly more prominent nodes on the anterior half and the posterior half has similar small nodes as those on the prosternum. Metasternum with a slightly wrinkled surface and small sparse nodes. Prescutum slightly longer than wide, with lateral margins slightly converging to the posterior (Fig. 72B View Figure 72 ). Lateral rims with eight or nine nodes giving the surface a rough textured appearance, not very large or prominent (Fig. 72B View Figure 72 ). Prescutum surface rather smooth except for along the sagittal plane which has seven or eight small nodes of about even size (Fig. 72B View Figure 72 ). Prescutum anterior margin prominent but not strongly raised above the surface, margin slightly granular and lacking a prominent central tubercle (Fig. 72D View Figure 72 ). Mesopleura rather narrow, gently diverging throughout the length, lateral margin with only slight granulation throughout, no prominent tubercles, at most three or four nodes and slight interspersed granulation throughout (Fig. 72B View Figure 72 ). Face of the mesopleura slightly wrinkled and with two distinct divots, one on the anterior ⅓ and one near the middle. Wings. Tegmina of moderate length, extending ½ through abdominal segment III. Tegmina wing venation: the subcosta (Sc) is the first vein, is simple, and terminates slightly < ½ through the overall tegmina length. The radius (R) spans the entire length of the tegmina with the first radius (R1) branching ca. ⅖ of the way through the wing length and terminating ca. ⅗ of the way through the wing length, followed by the branching and termination of the second radius (R2) near the distal ⅓ of the wing, 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. ⅖ of the way through the wing length, and 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 slightly <⅓ of the way through the wing length. Alae well-developed in an oval fan configuration, long, reaching onto abdominal segments IX. Alae wing venation: the costa (C) is present along the entire foremargin giving stability to the wing. The subcosta (Sc) is long, spanning ca. ⅔ of the wing length and is mostly fused with the radius in the beginning but terminates when it meets the costa. The radius (R) spans the entire wing and branches ca. ⅖ of the way through into the first radius (R1) and radial sector (Rs) which run gently diverging for most of their length and then converge at the apex of the wing where they terminate near each other but not touching. The media (M) branches early, ca. ⅙ of the way through the wing into the media anterior (MA) and the media posterior (MP) which run parallel with each other throughout the wing until the distal ⅙ of the wing where the media posterior fuses with the media anterior which then run fused together to the wing apex where they terminate near the radial sector. The cubitus (Cu) runs unbranched and terminates at the wing apex. Of the anterior anal veins, the first anterior anal (1AA) fuses with the cubitus near the point where the media branches into the media anterior and media posterior and then the first anterior anal branches from the cubitus ⅔ of the way through the wing length where it uniformly diverges from the cubitus until it terminates at the wing margin. The anterior anal veins two-seven (2AA-7AA) have a common origin and run unbranched in a folding fan pattern of relatively uniform spacing to the wing margin. The posterior anal veins (1PA-6PA) share a common origin separate from the anterior anal veins and run unbranched to the wing margin with slightly thinner spacing than the anterior anal veins. Abdomen. Abdominal segment II slightly converging, III through the anterior half of segment IV diverging to the widest portion. The posterior of IV and the anterior half of V parallel, the remainder of segments V-X gently converging to the rounded apex. Genitalia. Poculum broad and ends in a straight margined apex that slightly passes the anterior margin of segment X. Cerci long and slender, with slightly> ½ their length extending from under the anal abdominal segment, relatively flat, covered in a granulose surface and numerous short setae. Vomer broad and stout with straight sides evenly converging, and a thick single apical hook which hooks upwards into the paraproct and a notable smaller hook near the base of the primary hook, situated to the left of the primary hook when viewed ventrally. Legs. Profemoral exterior lobe slightly wider than the interior lobe (ca. 2½× as wide as the greatest width of the profemoral shaft), smoothly arcing end to end without a distinct bend and marked with a slightly granular margin and four small teeth on the distal half (Fig. 72C View Figure 72 ). Profemoral interior lobe roundly triangular and marked with five teeth arranged in a two-one-two pattern with shallow looping gaps between them (Fig. 72C View Figure 72 ). Mesofemoral exterior lobe arcs end to end but is significantly weighted more so on the distal half which is marked with three serrate teeth and the proximal half that is rather thin, lacking dentation. Mesofemoral interior lobe is slightly thinner than the exterior lobe, is broader on the distal end and is marked with six or seven small serrate teeth on the distal end. Metafemoral exterior lobe lacks dentation and has a straight margin along the metafemoral shaft. Metafemoral interior lobe smoothly arcs end to end with nine small serrate teeth on the distal half which is wider than the proximal half (Fig. 72D View Figure 72 ). Protibiae lacking exterior lobe, interior lobe reaching end to end in a rounded triangle with the widest portion on the distal half ca. 2 × the width of the protibial shaft (Fig. 72C View Figure 72 ). Meso- and metatibiae simple, lacking lobes completely.
Measurements of neotype male [mm]. Length of body (including cerci and head, excluding antennae) 70.5, length/width of head 3.9/3.8, antennae 47.8, pronotum 3.3, mesonotum 4.1, length of tegmina 20.5, length of alae 51.8, greatest width of abdomen 14.7, profemora 15.0, mesofemora 12.7, metafemora 15.4, protibiae 11.1, mesotibiae 18.5, metatibiae 11.5.
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