Pulchriphyllium bhaskarai sp. nov.

Cumming, Royce T., Le Tirant, Ste ́ phane, Linde, Jackson B., Solan, Megan E., Foley, Evelyn Marie, Eulin, Norman Enrico C., Lavado, Ramon, Whiting, Michael F., Bradler, Sven & Bank, Sarah, 2023, On seven undescribed leaf insect species revealed within the recent " Tree of Leaves " (Phasmatodea, Phylliidae), ZooKeys 1173, pp. 145-229 : 145

Pulchriphyllium bhaskarai sp. nov.

Figs 8H , 21E , 25 , 26 , 27 , 28 , 29 , 30

Material examined.

Holotype ♀: " Indonesia, West Java, Sukabumi, Kabandungan Village, Mt. Halimun, found 19th August 2020, first started laying eggs 16th September 2020, died 8th December 2020; laid a total of 78 eggs " (Fig. 25 ). Deposited in the Montreal Insectarium , Quebec, Canada (IMQC) . Paratypes: (19♂♂, 36♀♀, 2 eggs) See Suppl. material 1 for details about paratype specimens, their collection data, and depositories.

Differentiation.

Adult males and females have proven difficult to differentiate morphologically from Pulchriphyllium giganteum . Within both sexes of both species there is significant overlap in overall size and due to intraspecific variation, few to no features have allowed confident differentiation. Instead, the only reliable and easily observed morphological differences are the freshly hatched nymphs and egg morphology.

Females are very similar to Pulchriphyllium giganteum but can be differentiated by the mesopleurae. In Pulchriphyllium giganteum the posterior two or three spines are grouped together as a broad set projecting away from the mesopleuron margin, while in Pulchriphyllium bhaskarai sp. nov. these posterior-most spines are less prominent, only slightly projecting away from the mesopleuron margin (Fig. 26C ).

Males (Fig. 27 ) have proven impossible to differentiate from Pulchriphyllium giganteum with any certainty from morphology alone. Upon review of material from multiple locations and from both fresh and antique specimens, no single feature was consistently different enough to be useful ( Cumming et al. 2020b).

Freshly hatched nymphs (Figs 8H , 29 ) can easily be differentiated from congeners as the abdomen is notably thinner than any other Pulchriphyllium species, with a maximum width only ca ⅗ of the abdomen length (where all other Pulchriphyllium species have a wider and longer abdominal shape, with a width nearly equal to the abdomen length).

The eggs of Pulchriphyllium bhaskarai sp. nov. are rather unique and can easily differentiate this species from Pulchriphyllium giganteum by the presence of five laterally running semi-hollow tubes which are open on the anterior end (two tubes are located on each of the lateral surfaces and one tube is located on the ventral surface: Fig. 28K ). Additionally, the operculum is notably wider than long in Pulchriphyllium bhaskarai sp. nov. (Fig. 28H ) but approximately as long as wide and slightly tapered in Pulchriphyllium giganteum (Fig. 28B ).

Description.

Female. Coloration. The coloration description is based upon the holotype living individual when it was alive (Fig. 25 ). Interestingly the coloration altered throughout its development after it was brought into captivity. When first collected from the wild the overall base coloration was primarily yellow with muddled, variable patches of lime green to brown/gray coloration (Fig. 25A ). After several months in captivity the individual shifted the yellow base coloration to lime green, with variable patches of yellow and brown/gray coloration (Fig. 25B, C ). The areas that were brown/gray altered little during this period, only the base coloration appears to have changed.

Morphology. Head. Head capsule is slightly longer than wide, with a vertex that is marked throughout by moderately formed granulation and a singularly pointed posteromedial tubercle which is notably larger than the other granulation on the head capsule (Fig. 26C ). Frontal convexity triangular with a blunt point, which has a slightly granular surface and three to five setae near the apex. The compound eyes slightly protrude from the head capsule, not overly large, with a width taking up ca ¼ of the head capsule lateral margins (Fig. 26C ). Ocelli appear to be either very weakly developed, or absent. Antennal fields slightly wider than the width of the first antennomere. Antennae. Antennae consist of nine segments including the scapus and pedicellus, with the terminal segment approximately the same length as the preceding three segments’ lengths combined (Fig. 26A ). Antennomeres I-VIII sparsely marked with few transparent setae. Terminal antennomere covered densely in short, brown setae (Fig. 26A ). Thorax. Pronotum with slight concave anterior margin and lateral margins that are straight and subparallel for the anterior ⅔ and the posterior ⅓ converges slightly to the posterior margin that is ⅔ the width of the anterior margin (Fig. 26C ). The pronotum surface is marked with prominent granulation throughout, a distinct sagittal furrow, and a smaller perpendicular furrow near the center (Fig. 26C ). The pronotum has distinctly formed anterior and lateral rims and a weakly formed posterior rim (Fig. 26C ). Prosternum, mesosternum, and metanotum are marked by nodes of various size and spacing, with those along the margins more prominent. Prescutum approximately as long as the greatest width, with lateral rims marked by a granular surface with those on the anterior third larger than the others (Fig. 26C ). Prescutum anterior rim not strongly protruding, rim surface is granular and lacks a sagittal spine (Fig. 26C ). Prescutum surface slightly raised along the sagittal plane and the entire surface is marked throughout with granulation. Mesopleura are narrow for the anterior ¼, then diverge strongly for the middle ½, bend prominently and then run only slightly diverging for the remainder of their lengths. On the mesopleuron near the distinct bend from prominently diverging to slightly diverging, there is a cluster of two or three tubercles which are grouped together and project slightly away from the mesopleuron margin (Fig. 26C ). Mesopleuron lateral margin with eight or nine various sized tubercles, with those on the anterior typically smaller (Fig. 26C ). Mesopleuron surface granular and marked with a distinct divot near the middle (Fig. 26C ). Wings. Tegmina long, reaching at least ½ onto abdominal segment VII or slightly onto VIII. Tegmen venation; the subcosta (Sc) is the first vein in the forewing, runs parallel with the margin for the first ½, and then bends slightly and runs towards the margin where it terminates ca ⅕ of the way through the wing length. The radius (R) spans the central portion of the tegmen with two slightly diverging veins; the first radius (R1) branches ca ⅕ of the way through the tegmen length and terminates approximately ⅓ of the way through; the radial sector (Rs) branches ca ⅓ of the way through the tegmen length and terminates near the distal ⅖. The media (M) is bifurcate with the media anterior (MA) branching near the middle of the tegmen length and terminating near the distal ¼ and the media posterior (MP) branches near the distal ¼ of the tegmen and terminates near the apex. The cubitus (Cu) is also bifurcate, branching near the posterior ⅕ of the wing into the cubitus anterior (CuA) and cubitus posterior (CuP) which both terminate at or near the wing apex. The first anal vein (1A) is simple and fuses with the cubitus ca ⅕ of the way through the tegmen length. Alae rudimentary. Abdomen. Abdominal segments II through the anterior ½ of IV gradually diverging. The posterior ½ of segment IV through the anterior ½ of segment V weakly diverging. The posterior ½ of abdominal segment V through VII with straight, slightly converging margins. Abdominal segment VIII arcing inward to form a small lobe, followed by segment IX and X converging to the apex. Genitalia. Subgenital plate starts at the anterior margin of tergum VIII, is not particularly broad, only extending at most to the anterior margin of tergum X and terminating in a fine point (Fig. 26E ). Gonapophyses VIII are long and moderately broad, nearly reaching the apex of abdominal tergum X; gonapophyses IX are shorter and broader (Fig. 26E ). Cerci flat, with a relatively smooth surface and few detectable setae (Fig. 26E ). Legs. Profemoral exterior lobe broad, slightly recurved with an acute angle, with a width ca 2 × the width of the interior lobe (Fig. 26D ). Edge of the profemoral exterior lobe primarily smooth or slightly lumpy with at most three dulled, small teeth on the recurved angle (Fig. 26D ). Profemoral interior lobe only located on the distal ⅔ as an arcing lobe ca 2½× as wide as the greatest width of the profemoral shaft, with the distal ½ with four or five dulled triangular teeth of irregular size and spacing with looping gaps between them (Fig. 26D ). The mesofemoral exterior lobe arcs from end to end in a rounded triangular shape that is weighted towards the distal ⅖ and is at most slightly wider than the mesofemoral shaft width. The proximal margin is smooth but the distal margin following the bend is marked with three or four serrate teeth. The mesofemoral interior lobe is ca 1½× as wide as the mesofemoral shaft, and also shaped as a rounded triangle, but the interior lobe is slightly weighted towards the proximal ⅓. The mesofemoral interior lobe proximal margin is smooth and the distal margin following the bend is marked by six or seven dulled, serrate teeth. The metafemoral exterior lobe arcs end to end hugging the metafemoral shaft and lacks teeth. The metafemoral interior lobe arcs gently from end to end but is slightly thicker on the distal ⅔ (which is at most as wide as the metafemoral shaft width). The distal ½ of the metafemoral interior lobe has five or six distinct serrate teeth. The protibial exterior lobe is a thin triangle weighted to the distal end with a maximum width of ca 1½× the width of the protibial shaft. The protibial interior lobe spans the entire length as a rounded triangle weighted slightly towards the distal ½ with a maximum width ca 2 × the width of the protibiae shaft itself. Mesotibiae and metatibiae lacking interior lobes.

Measurements of holotype female [mm]. Length of body (including cerci and head, excluding antennae) 97.0, length/width of head 8.3/7.1, antennae 5.2, pronotum 6.2, mesonotum 4.9, length of tegmina 55.5, greatest width of abdomen 50.7, profemora 21.3, mesofemora 15.3, metafemora 18.9, protibiae 10.6, mesotibiae 11.1, metatibiae 15.6.

Measurements of paratype females [mm]. Length of body (including cerci and head, excluding antennae) 95.6-114.5, length/width of head 8.2-10.8/7.0-8.3, antennae 5.1-6.2, pronotum 6.1-6.8, mesonotum 4.8-5.5, length of tegmina 55.3-67.0, greatest width of abdomen 50.5-56.1, profemora 21.0-27.6, mesofemora 15.2-17.5, metafemora 18.7-21.3, protibiae 10.5-12.6, mesotibiae 11.0-13.0, metatibiae 15.4-17.8.

Male. Coloration. Coloration description based upon images of live specimens. Overall coloration is pale green throughout with somewhat variable patches of tan, brown, or black (Fig. 27A ). The antennae, head, thorax, protibiae, profemoral interior lobe, mesotibiae, mesofemora, and metatibiae are brown. The sclerotized area of the alae has variable tan portions around the veins. Abdominal segments V and VI have variable patches of brown/tan, and segment V has a pair of large brown/black eye spots.

Morphology. Head. Head capsule approximately as wide as long, with a vertex that is slightly wrinkled/rough textured. The posteromedial tubercle is small but distinctly raised above the head capsule. Frontal convexity stout and ending with a blunt point and marked with only a few short setae. Compound eyes bulbous, occupying slightly> ⅖ of the head capsule lateral margins (Fig. 27G ). There are three well-developed ocelli located between and slightly posterior to the compound eyes. Antennal fields are approximately as wide as the scapus. Antennae. Antennae (including the scapus and pedicellus) consist of 22 segments (Fig. 27D ). The scapus and pedicellus are bare, the following 17 segments are covered with thin, short setae, and the terminal three segments have notably shorter and denser setae. Each antennomere on the distal end projects ventrally slightly, so the overall antenna has a serrate appearance. Thorax. Pronotum with anterior margin gently concave, with a well-defined margin. Lateral margins have moderately formed rims that converge gently to a straight posterior margin that lacks a distinct margin and is approximately ½ the width of the anterior rim (Fig. 27G ). Face of the pronotum is smooth or slightly wrinkled and marked with a distinct sagittal furrow, and a slight perpendicular furrow near the center (Fig. 27G ). The prosternum surface is slightly granular, and the mesosternum and metasternum surfaces are smooth centrally, but have slightly granular margins. The prescutum is approximately as long as wide, with lateral margins slightly converging to the posterior margin which is ca ¾ as wide as the anterior margin. Prescutum lateral rims roughly textured, with the nodes on the anterior more prominent than the posterior. The surface of the prescutum is heavily granulose and moderately wrinkled, with the surface slightly raised along the sagittal plane. Prescutum anterior rim weakly formed, surface only slightly granular, and lacking a distinct central tubercle (Fig. 27G ). Mesopleura narrow for the anterior ⅓, then gradually diverging steadily throughout their lengths (Fig. 27G ). Mesopleuron lateral margin with four or five small tubercles throughout the length, with the posterior two slightly more prominent (Fig. 27G ). Face of the mesopleuron with a slightly wrinkled texture. Wings. Tegmina short, extending ¾ of the way onto abdominal segment II. Tegmen wing venation: the subcosta (Sc) is the first vein, runs relatively straight for ca ½ of the wing length and terminates on the margin. The radius (R) spans the entire length of the tegmen, running as the radial sector (Rs) straight through the center of the tegmen to the apex after the first radius (R1) branches near the distal ⅓ of the wing and terminates near the posterior margin. The media (M) spans the entire length of the tegmen, running parallel with the radius (R) and radial sector (Rs) and terminates at the wing apex as the media anterior (MA) after the branching of a weakly formed media posterior (MP) near the proximal ⅓ of the tegmen. The cubitus (Cu) runs through the tegmen surface angled away from the media (M) for slightly < ½ of the tegmen length to the margin and then runs along the margin where it fades before reaching the apex. The first anal (1A) vein runs subparallel to the cubitus until they meet ca ⅓ of the way through the tegmen length. Alae well-developed in an oval fan configuration, reaching onto abdominal segment VIII or IX. Ala wing venation: the costa (C) is present along the entire foremargin giving stability to the wing. The subcosta (Sc) is fused with the radius for the anterior ⅗ of the ala length, then splits and runs parallel with the ala margin until it fades before reaching the apex. The radius (R) branches ca ⅔ of the way through the ala length into the first radius (R1) and radial sector (Rs) which run gradually diverging until they fuse with the cubitus at different locations. The media (M) branches early, near the anterior ⅛ into the media anterior (MA) and the media posterior (MP) which run diverging for ⅓ of their lengths, then parallel for ⅓ of their lengths, and converging for the final ⅓. Both the media anterior and the media posterior weakly fuse with the cubitus at different locations. The cubitus runs unbranched and terminates at the wing apex after the media anterior, media posterior, first radius, and radial sector fuse with it at different locations. Of the anterior anal veins, the first anterior anal (1AA) fuses with the cubitus ca ⅕ of the way through the wing length and then the first anterior anal branches from the cubitus on the distal ⅓ 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 segments II through the anterior ½ of segment IV gradually diverge. The posterior ½ of segment IV through the anterior ½ of segment V diverge slightly less than the previous segments. The posterior ½ of segment V through segment VII have parallel margins, giving the abdomen a rectangular appearance. The posterior end of abdominal segment VIII converges significantly and forms a slight posterior pointing lobe. Abdominal segment IX has margins converging directly to abdominal segment X which is small and converges to a blunt apex. Genitalia. Poculum broad and rounded (with a maximum width broader than abdominal segment X) and ending in a blunt apex that passes beyond the anterior margin of segment X (Fig. 27F ). Cerci slender, with ca ½ of their length extending out from under abdominal segment X. The cerci are slightly cupped, have a granulose surface, and numerous long setae along the margins. Vomer broad with straight margins evenly converging to the apex, which is armed with a singular, broad upwards turning hook. Legs. Profemoral exterior lobe broad, nearly right angled, ca 1½× wider than the interior lobe, with the proximal margin smooth and the distal margin marked with three to five teeth near the bend (Fig. 27B ). Profemoral interior lobe only on the distal ⅔ of the profemoral shaft, roundly arcing, with the distal ½ ornamented with four teeth arranged in a two-two pattern, with a large looping gap between the sets (Fig. 27B ). Mesofemoral exterior lobe is roundly triangular, slightly weighted towards the distal end, and broad (ca 1½× as wide as the mesofemoral shaft width), with the proximal margin straight and smooth, and the distal margin slightly arcing and marked by four distinct serrate teeth. The mesofemoral interior lobe is slightly thinner than the exterior lobe and it is relatively evenly weighted from end to end. The proximal ½ of the mesofemoral interior lobe is straight and smooth, while the distal ½ is armed with six or seven serrate teeth. Metafemoral exterior lobe arcs thinly along the metafemoral shaft, with a maximum width slightly thinner than the metafemoral shaft width. Metafemoral exterior lobe lacks dentition throughout most of its length, with only the distal ⅓ marked by three or four small serrate teeth. Metafemoral interior lobe is narrow and straight for the proximal ½ and then on the distal ½ it widens out slightly to a maximum width approximately as wide as the metafemoral shaft width. The distal ½ of the metafemoral interior lobe is armed with six or seven serrate teeth. The protibial exterior lobe is only present on the distal ½ as a thin arcing lobe at most 2 × the width of the protibial shaft. Protibial interior lobe is a rounded, thin scalene triangle slightly weighted just distal to the midlength, with a maximum width ca 3 × the width of the protibial shaft. Mesotibial exterior lobe is a small, triangular spur, only slightly wider than the width of the mesotibial shaft, situated just distal to the midlength, and occupying slightly <⅓ of the overall length. The metatibial exterior lobe is ca 2 × the size of the mesotibial exterior lobe but slightly more rounded in shape, is situated only on the distal ⅔ of the length and has a maximum width of 1½× the metatibial shaft width. Mesotibiae and metatibiae lack interior lobes.

Measurements of paratype males [mm]. Length of body (including cerci and head, excluding antennae) 73.4-74.0, length/width of head 4.5-4.6/2.1-2.2, antennae 25.7-25.9, pronotum 3.2-3.3, mesonotum 2.9-3.0, length of tegmina 14.3-14.5, length of alae 56.8-57.0, greatest width of abdomen 29.0-29.2, profemora 14.8-14.9, mesofemora 10.5-10.6, metafemora 11.2-11.3, protibiae 7.4-7.6, mesotibiae 6.9-7.0, metatibiae 9.0-9.2.

Eggs. (Fig. 28G-L ). The overall color is dark brown, all surfaces are roughly textured, with some of the rough textured areas lighter brown in color. The cross-section shape is approximately pentagonal, with the dorsal surface the widest surface. Each lateral surface has two longitudinally running semi hollow tubes running the full length of the capsule with their openings on the anterior end (Fig. 28K ). When viewed laterally, the dorsal surface is slightly arcing, and the ventral surface is straight (Fig. 28G ). The dorsal surface has the micropylar plate which spans the entire length of the capsule in a roughly, rounded, diamond shape with the micropylar cup centrally located (Fig. 28I ). The area around the micropylar plate lacks distinct features, and is relatively simple, only roughly textured (Fig. 28I ). The operculum is slightly ovular, wider than long, and is semi hollow, ending in a distinct opening (Fig. 28K ). The ventral surface has a fully hollow tube running the full length, with openings on each end (Fig. 28K-L ). On the posterior (Fig. 28L ) only the ventral tube has an opening, the rest of the surface is rounded from the closed bases of the lateral, longitudinally running tubes.

Measurements [mm]. Length (including operculum): 7.9-8.0; maximum width of capsule when viewed from lateral aspect 4.2-4.3; length of micropylar plate 5.0-5.1.

Newly hatched nymphs.

(Fig. 29 ). The general color throughout the body is vermillion with muddled lighter and darker areas. The basitarsi are white, with the remaining tarsal segments a similar red to the body base color. All tibiae have well-developed exterior lobes of a similar shape, a narrow triangle occupying the central ⅓ of the tibiae. The protibial interior lobe is prominent, gently arcing the full length of the tibia. The protibial lobe has a width that is ca 2 × as wide as the protibia shaft width. The profemoral interior lobe occupies the distal ⅔, is broad and obtusely angled, with a width that is slightly larger than 2½× the profemoral shaft width. The profemoral interior lobe has two weakly formed teeth on each end of the distal margin. The profemoral exterior lobe is of a similar shape as the interior lobe, but differs in being slightly wider, lacking distinct serration, and the proximal end reaches closer to the base of the profemora. All femoral lobes have mild serration, typically only two or three dulled teeth on the distal halves of the lobes. The meso- and metafemoral interior lobes span end to end, but are more heavily weighted towards the distal end, with the proximal end thin. The meso- and metafemoral exterior lobes are slightly more evenly spread across their lengths and slightly thinner, giving them a more arcing appearance. The head and lobes of the legs are of a similar burnt red coloration while the thorax is brighter red. The abdomen base color is vermillion with darker red to black markings along the margins and looping along the central abdominal cavity, giving the appearance of spots. The abdomen is notably thinner than any other Pulchriphyllium species, with a maximum width ca ⅗ of the abdomen length. The widest point of the abdomen is abdominal segment V.

Etymology.

Patronym. Named to honor Edy Bhaskara who supplied the specimens which solved the mystery surrounding this species. Previously only males were known, which are morphologically indistinguishable from Pulchriphyllium giganteum , until Edy Bhaskara supplied the authors with fresh specimens (which allowed DNA analyses) as well as adult females, freshly hatched nymphs, and eggs, all of which allowed a full understanding to be generated for this species and allowed differentiation from congenerics.

Distribution.

At present only known from Java, Indonesia, with records from throughout the island (Suppl. material 1).

Remarks.

Originally, it was thought that the " Pulchriphyllium giganteum "-like population on Java, Indonesia was simply a range expansion for Pulchriphyllium giganteum ( Cumming et al. 2020b). This erroneous assumption was based solely upon male morphology as no significant morphological features could be identified at the time. It was not until fresh specimens could be sequenced and the adult female, freshly hatched nymph, and the egg morphology were known that it became apparent that this " Pulchriphyllium giganteum "-like species from Java was unique and undescribed.

Within the DNA-based analyses of Bank et al. (2021), this population was included as " Pulchriphyllium sp. 2" and not recovered as close relative to Pulchriphyllium giganteum from Malaysia, but instead was recovered with weak support to be sister to a clade comprising Pulchriphyllium rimiae (Seow-Choen, 2017), Pulchriphyllium shurei (Cumming & Le Tirant, 2018), and Pulchriphyllium mannani (Seow-Choen, 2017). Herein, the Javan taxon was instead recovered with high support as sister to all other included Pulchriphyllium species (Fig. 2 ). Following DNA analyses, we were sent images of adult female specimens (Fig. 25 ) by Edy Bhaskara (Java, Indonesia) which, although appearing very similar to Pulchriphyllium giganteum from Malaysia, the Javan taxon has less prominent mesopleurae (Fig. 26C ). Additionally, Edy Bhaskara was able to share images of the eggs (Fig. 28G-L ) and freshly hatched nymphs (Fig. 29 ), both of which added to our understanding of the uniqueness of this species. Upon review of all available evidence for this taxon, this population can now confidently be identified as an undescribed species and described.

Thanks to DNA analyses, the geographic distribution of Pulchriphyllium giganteum has been clarified to encompass West Malaysia, West Borneo, and North Borneo as a single species. Unfortunately, Sumatra and Nias island leave an area of significant uncertainty as no DNA samples are yet available, and to date only male specimens have been observed/collected (which look identical to both Pulchriphyllium giganteum and Pulchriphyllium bhaskarai sp. nov.). Interestingly, despite Pulchriphyllium bhaskarai sp. nov. being presently understood as endemic to Java, Pulchriphyllium giganteum has a wide range from South Thailand, West Malaysia, and throughout Borneo ( Cumming et al. 2020b, Jiaranaisakul and Pawangkhanant 2022). Hopefully, future collecting efforts on the islands of Sumatra and Nias will produce DNA data as well as the adult female, egg, and freshly hatched nymph morphology, and thus allow these populations to be identified.

The eggs of Pulchriphyllium bhaskarai sp. nov. are quite unique, as they lack the typical Pulchriphyllium dense fins, and instead have rigid, open tubes running along the egg surfaces (Fig. 28K ). No other phylliid eggs have such structures, and these rigid tubes are only similar to "type 7" eggs characterized within Büscher et al. (2023) due to texture/material, but their overall shape is wholly unique. Within our recovered phylogeny (Fig. 2 ) Pulchriphyllium bhaskarai sp. nov. was recovered as sister to all other Pulchriphyllium species, a placement which is not surprising as within Büscher et al. (2023) Pulchriphyllium giganteum with eggs that are also unlike most Pulchriphyllium species was recovered as external to the other included Pulchriphyllium species. Within our recovered phylogeny (Fig. 2 ) Pulchriphyllium bhaskarai sp. nov., Pulchriphyllium giganteum , and a clade formed by three species whose eggs are presently unknown ( Pulchriphyllium rimiae , Pulchriphyllium shurei , and Pulchriphyllium mannani ) were recovered as external to the bulk of the Pulchriphyllium species which have the dense fins. This leaves us speculating what egg morphology Pulchriphyllium rimiae , Pulchriphyllium shurei , and Pulchriphyllium mannani may have as these species (which are only known from males at the moment) are morphologically unique and not nested within the greater Pulchriphyllium . Hopefully females of these species can be located one day, and their egg morphology identified in order to better frame the evolution of phylliid egg morphology and fill in this missing knowledge.

Figures

Figure 2. Phylogenetic tree of the Phylliidae based on six nuclear and mitochondrial loci with a focus on Phyllium and Pulchriphyllium (please refer to Suppl. material 4 for the complete tree). Ultrafast bootstrap support values are depicted at each node. Specimens in green font are in addition to the already existing tree inferred by Bank et al. (2021). Highlighted in green boxes are the seven newly described species.

Figure 8. Illustrations of the freshly hatched nymph coloration for herein referenced species A Phyllium ortizi sp. nov. (Mindanao) B Phyllium mabantai (Mindanao) C Phyllium samarense sp. nov. (Samar) D Pulchriphyllium bioculatum (West Malaysia) E Pulchriphyllium anangu sp. nov. (Southwest India) F Pulchriphyllium agathyrsus (Sri Lanka) G Pulchriphyllium giganteum (West Malaysia) H Pulchriphyllium bhaskarai sp. nov. (Java). Illustrations by scientific illustrator Liz Sisk (Washington D. C., USA).

Figure 21. Adult female profemoral lobes from various Pulchriphyllium species for comparison A Pulchriphyllium crurifolium stat. rev., comb. nov. (NHMUK) B Pulchriphyllium bioculatum from West Malaysia (Coll RC 16 - 038) C Pulchriphyllium scythe stat. rev., comb. nov. from Sylhet, Bangladesh (OUMNH) D Pulchriphyllium pulcrifolum from Java, Indonesia (Coll RC 16 - 024) E Pulchriphyllium bhaskarai sp. nov. from Java, Indonesia (MZPW).

Figure 25. Pulchriphyllium bhaskarai sp. nov. live holotype female. Photographs by Edy Bhaskara (Indonesia) A coloration shortly after being found in the wild, dorsal habitus B altered coloration after several months in captivity, dorsal habitus C habitus, dorsolateral view.

Figure 26. Pulchriphyllium bhaskarai sp. nov. female holotype A details of the antennae, dorsal B dorsal habitus C details of the head and thorax, dorsal D details of the profemora E genitalia details, ventral F ventral habitus G lateral view of the thorax (anterior to the right). Scale bars: 25 mm (B, F).

Figure 27. Pulchriphyllium bhaskarai sp. nov. male paratype, Coll. RC # 21 - 032 A dorsal habitus B details of the profemora C lateral view of the thorax (anterior to the right) D details of the antennae, dorsolateral view E terminalia details, dorsal F genitalia details, ventral G details of the head and thorax, dorsal. Scale bar: 25 mm (A).

Figure 28. A-F Pulchriphyllium giganteum egg capsule, IMQC collection A lateral view B dorso-lateral view C dorsal view D opercular (anterior) view E posterior view F ventral view G-L Pulchriphyllium bhaskarai sp. nov. egg capsule, IMQC collection (Coll RC 21 - 034) G lateral view H dorso-lateral view I dorsal view J opercular (anterior) view K posterior view L ventral view. Photographs by Rene Limoges (IMQC). Scale bars: 2 mm (A-L).

Figure 29. Pulchriphyllium bhaskarai sp. nov. live freshly hatched nymphs A dorsal habitus B habitus, dorsolateral view C habitus, dorsolateral view. Photographs by Edy Bhaskara (Indonesia).

Figure 30. Paratype historic pair of Pulchriphyllium bhaskarai sp. nov. from MZPW A male with data labels inset to the lower left B female with data labels inset to the right.