Cryptophyllium echidna gen. et, 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 echidna gen. et sp. nov. Figure 30

Material examined.

Holotype ♀: INDONESIA: Wangi-wangi Island. Collected prior to 2020, but no exact date given. Deposited in the Montreal Insectarium (IMQC).

Remarks.

This is the first phylliid record we have seen from the small island of Wangi-wangi in the Wakatobi Regency in Southeast Sulawesi Province. This small island appears to be rather unique biogeographically and unexplored as a yet to be described bird species presently known as the "Wangi-wangi White-eye" ( Zosterops sp. nov.) has recently been identified as well, suggesting this island may hold many endemic undescribed species ( O’Connell et al. 2019).

Differentiation.

Presently we only know of a single female specimen of this new species which we here designate as the holotype. Morphologically and molecularly this species is closely related to Cryptophyllium celebicum comb. nov. which has a much wider distribution to the north on the islands of Buton, Sulawesi, and Peleng (Fig. 2 ).

Female Cryptophyllium echidna sp. nov. can be differentiated from Cryptophyllium celebicum comb. nov. by only subtle differences in the thorax and profemoral exterior lobes. In Cryptophyllium echidna sp. nov. the prescutum is slightly broader and with a weaker sagittal crest (Fig. 30D ) than in Cryptophyllium celebicum comb. nov. (Fig. 22E ). Additionally the profemoral exterior lobe of Cryptophyllium echidna sp. nov. has a right exterior angle (Fig. 30A ), not acute like in Cryptophyllium celebicum comb. nov. (Fig. 22D ).

Males are presently unknown, but as the sister species to Cryptophyllium celebicum comb. nov. the males likely have a similar morphology.

Distribution.

At present only known from the small Indonesian island of Wangi-wangi off the east coast of Buton Island.

Description.

Female. Coloration. At present we only have the dried holotype female to describe the color from which has a bit of rot through the legs, head, thorax, and the central area of the abdomen. The rotten areas are brown but are assumed to have been green in life. The remainder of the female is lime-green in color throughout, with no indication of natural brown patches (which even on somewhat rotten specimens can generally be identified) but this female appears to have been uniform green in life.

Morphology. Head. Head capsule slightly longer than wide, vertex with minimal granulation throughout the surface, all relatively well-spaced with no areas on the head tightly packed. The posteromedial tubercle is the most prominent feature on the vertex of the head capsule. Frontal convexity broad and stout, shorter than the length of the first antennomere, and with a lumpy surface marked by few short transparent setae. Compound eyes not particularly large, only slightly protruding from the head capsule, taking up ca. ¼ of the length of the lateral head capsule margins (Fig. 30A ). Ocelli absent. Antennal fields slightly wider than and about as long as the length of the first antennomere. Antennae. Antennae consist of nine segments, with the terminal segment approximately the same length as the preceding two segments’ lengths combined (Fig. 30B ). Antennomeres I-III sparsely marked with small transparent setae (with the longest on the first segment), the terminal two antennomeres are densely covered in stout, brown setae. Thorax. Pronotum with a gently concave anterior margin and slightly convex lateral margins, which converge to a straight posterior margin that is half the width of the anterior margin (Fig. 30D ). The pronotum surface lacks granulation but is slightly lumpy, with only a prominent pit in the center, and slight furrows anterior and lateral to the pit (Fig. 30D ). The pronotum has a prominent anterior rim and moderate lateral and posterior rims (Fig. 30D ). Prosternum and mesosternum with numerous nodes throughout the surface, all about the same size and spacing throughout. Metasternum with slightly less granulation but they are slightly larger than those on the pro- and mesosternum. Prescutum longer than wide, with a slightly broader anterior margin (Fig. 30D ). Lateral rims with 6-8 prominent tubercles with various small, lumpy granules interspersed throughout the margins (Fig. 30D ). Prescutum anterior rim prominent but not strongly protruding, surface is granular and lacks a prominent sagittal spine (Fig. 30D ). Prescutum surface without a strongly raised sagittal crest, instead the surface is only slightly raised along the sagittal plane. The prescutum surface has moderate granulation throughout ranging in size from small to medium with irregular spacing (Fig. 30D ). Mesopleura not spanning the entire length, instead with the anterior ⅓ narrow and only starting to fan out near the midline of the prescutum length. Mesopleura lateral margins with five or six larger, sharp tipped tubercles with an additional five or six smaller nodes interspersed throughout (Fig. 30D ). Face of the mesopleura lacking granulation, but instead highly wrinkled and with two notable pits, one on the anterior ⅓ and one nearer the posterior ⅓ (Fig. 30D ). Wings. Tegmina long, reaching nearly to the posterior margin of abdominal segment VII. The subcosta (Sc) is the first vein in the forewing and runs subparallel with the wing for the first half of its length, and then bends towards the wing margin for the second half. The radius (R) spans the central portion of the tegmina with two subparallel branched veins. The first radius (R1) branches ca. ⅗ of the way through the radius length and terminates ca. ⅓ of the way through the wing length. The radial sector (Rs) branches from the end of the radius and runs angled to the wing margin where it terminates just posterior to the wing midline length. There is a weak continuation of the radius following the prominent radial sector branching which continues on as a short and thin radius to media crossvein (R-M). The media (M) is simply bifurcate with both the media anterior (MA) and media posterior (MP) terminating close to the posterior ¼ of the wing. The cubitus (Cu) runs throughout the entire wing length simply, and then near the posterior ⅕ of the wing becomes bifurcate into the cubitus anterior (CuA) and cubitus posterior (CuP) which both terminate at or very near the wing posterior apex. The first anal vein (1A) is simple and fuses with the cubitus early on, near where the radial sector branches from the radial. Alae of moderate length, reaching abdominal segment IV. Abdomen. Abdominal segments II through the anterior ½ of IV diverging, with the middle of segment IV the widest segment. Segments V-VII gently converging, with segment VII ending with a distinct lobe which bends inward to a notably narrower segment VIII. Segments VIII-X converging to a broad apex. Genitalia. Subgenital plate starts at the anterior margin of segment VIII, is broad, and only extends ⅓ of the way under segment X, ending in a fine point (Fig. 30E ). Gonapophyses VIII are long (exceeding the tip of the abdomen but not as long as the tips of the cerci) and moderately broad (together side by side are about as broad as the subgenital plate; Fig. 30E ). Gonapophyses IX are smaller and shorter, and mostly covered from view by the notably larger gonapophyses VIII. Cerci flat, with a granular surface throughout and few setae on the ventral surface, the dorsal surface has thin, transparent setae throughout the surface (Fig. 30E ). Legs. Profemoral exterior lobes broad (ca. 2 × width of the interior lobe) and approximately right angled (Fig. 30A ). Proximal edge of the exterior profemoral lobe gently undulates giving this margin a slightly wavy appearance, whereas the distal margin is nearly straight, both margins have notable serration throughout their lengths (Fig. 30A ). Profemoral interior lobe with a slightly obtuse angle and marked with five large, serrate teeth with looping gaps between them. These teeth are arranged somewhat into a two-one-two pattern with the gaps between these sets larger. Mesofemoral exterior lobe arcs from end to end and is almost evenly weighted on both sides, but with the broadest point just off center to the distal side of the midline, and on the distal half only marked with three small serrate teeth. Interior mesofemoral lobe arcs end to end, is ca. ¼ narrower than the exterior lobe, not as strongly angled, and marked with six or seven teeth on the distal half. Metafemoral interior lobe arcs end to end but is notably wider on the distal ⅔ of the lobe and this wider portion is marked by 11 or 12 serrate teeth. Metafemoral exterior lobe is thin and smooth, hugging the metafemoral shaft and marked with one or two small, rounded teeth on the distal edge. Protibiae lacking an exterior lobe. Protibiae interior lobe spans the entire length of the protibiae as a rounded scalene triangle with the widest portion on the distal ⅓. Mesotibiae and metatibiae simple, lacking exterior and interior lobes.

Measurements of holotype female [mm]. Length of body (including cerci and head, excluding antennae) 94.5, length/width of head 7.8/7.5, antennae 5.0, pronotum 6.6, mesonotum 7.9, length of tegmina 59.1, length of alae 31.6, greatest width of abdomen 41.3, profemora 20.7, mesofemora 16.5, metafemora 21.2, protibiae 14.0, mesotibiae 12.1, metatibiae 16.4.

Etymology.

Noun, Greek in origin. Relating to the tenth labor of Heracles (apparently a favorite story of Gray (1843) as four of his therein described species came from this myth) in which Heracles was tasked with capturing the red cattle from the monster Geryon (for which Gray named Phyllium geryon Gray, 1843). Before Heracles sailed out to the island Erythia where Geryon and his red cattle lived, he rested on the mainland and forgot to tie up his horses. When he eventually found them, they were in a cave with an Echidna (a monster which is described as women from the waist up and snake on her lower half). She refused to give back his horses unless he lay with her, which being a classic Greek hero, he did. Before he left the Echidna, she told him that she was pregnant with three of his sons and asked him which of the three should rule her lands one day. Heracles then left her with his bow and a girdle, and told her that whichever of his three sons could draw the bow and wear the girdle best would inherit her land and the other two should be banished. Those three sons were Agathyrsus, Gelonus, and Scythes (for which Gray named Phyllium agathyrsus Gray, 1843; Phyllium gelonus Gray, 1843; and Phyllium scythe Gray, 1843).

We felt that this species could help to finish telling the story which Gray was so fond of. With Phyllium geryon being a species from the Philippines, one biogeographical bridge for species to the Philippines is from Sulawesi through the Sangihe Islands ( Evans et al. 2003), therefore as a steppingstone to the Philippines we felt there should be an Echidna along the route to the Philippines where Phyllium geryon can be found.

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 22. Holotype, Cryptophyllium celebicum (de Haan, 1842), comb. nov. the type species for the Cryptophyllium gen. nov. Photographs by Luc Willemse, Naturalis Biodiversity Center (RMNH) A dorsal, habitus B genitalia, ventral C ventral, habitus, and original collection label inset to left D details of the front legs, head, and thorax, dorsal E details of the antennae and thorax, dorsal.

Figure 30. Cryptophyllium echidna gen. et sp. nov. holotype female, photographs by Rene ́ Limoges (IMQC) A habitus, dorsal B dorsal details of the antennae C habitus, ventral D dorsal details of the thorax E ventral details of the genitalia.