Pulchriphyllium anangu 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 anangu sp. nov.

Figs 8E , 31 , 32 , 33

Material examined.

Holotype ♂: "WS 415; India: Karnataka, Agumbe Ghats, Canopy Light Trap, N 13 29.386' E 75 04.537' 10-X-2004 " (Fig. 31 ). Deposited in Brigham Young University (BYU), Monte L. Bean Life Science Museum, Provo, Utah, USA . Paratypes: (3♂♂, 3♀♀, 2♀♀ nymphs, 1♂ nymph). See Suppl. material 1 for details about paratype specimens, their collection data, and depositories.

Differentiation.

Pulchriphyllium anangu sp. nov. are morphologically most similar to Pulchriphyllium bioculatum and Pulchriphyllium agathyrsus , of which the adult morphology is difficult to differentiate, but the freshly hatched nymphs (Fig. 32B, C ) allow reliable differentiation. Presently eggs of Pulchriphyllium anangu sp. nov. are not known to us.

Female Pulchriphyllium anangu sp. nov. (Fig. 32A ) are difficult to differentiate from Pulchriphyllium bioculatum and the only feature which appears to be somewhat reliable is the intensity of the serration on the proximal margin of the profemoral exterior lobe with Pulchriphyllium anangu sp. nov. tending to have slightly more prominent serration. For adult females Pulchriphyllium agathyrsus we have yet to identify a reliable morphological feature for differentiation.

Male Pulchriphyllium anangu sp. nov. has proven impossible to differentiate morphologically from Pulchriphyllium agathyrsus due to notable coloration and abdominal shape variation and therefore no consistent differences have been identified (Fig. 33A, B ). Pulchriphyllium anangu sp. nov. males can be differentiated from Pulchriphyllium bioculatum by subtle differences in the profemoral exterior lobe shape (slightly more obtuse and thinner in Pulchriphyllium anangu sp. nov.) and the abdominal shape in Pulchriphyllium bioculatum tends to be slightly more ovoid vs Pulchriphyllium anangu sp. nov. which have abdominal segments V and VI typically with more parallel margins (Fig. 33A, B ).

Freshly hatched nymphs of Pulchriphyllium anangu sp. nov. are very similar to Pulchriphyllium bioculatum and only subtle differences have been identified in their coloration. Typically, the overall coloration of Pulchriphyllium bioculatum tends to be a bit darker (Fig. 8D ), with the profemoral lobes the most reliable difference between the species as Pulchriphyllium anangu sp. nov. (Fig. 8E ) tend to have lighter colored profemoral lobes than Pulchriphyllium bioculatum (Fig. 8D ). Pulchriphyllium anangu sp. nov. can easily be differentiated from its sister species Pulchriphyllium agathyrsus by the coloration of the femoral and tibial lobes vs the head and thorax coloration. In Pulchriphyllium agathyrsus the femoral and tibial lobes contrast significantly with the body coloration as the lobes are dark brown/black while the head and thorax are bright red (Fig. 8F ) vs Pulchriphyllium anangu sp. nov. which has lobes on the legs with a similar color to the head and thorax (Fig. 8E ).

Description.

Female. Coloration. Coloration description is based upon numerous images of live individuals from iNaturalist (Fig. 32A ). To date we have only seen records of females where their base coloration was pale green with some areas highlighted in yellow/tan (such as the antennae and venation of the tegmina). In a few individuals we have seen some areas marked with spotting of tan/brown coloration on the abdomen, tegmina, and femoral lobes, but these areas tend to be rather minimal. We have yet to observe extreme color variations in this species.

Morphology. Head. Head capsule is slightly longer than wide, with a vertex that is marked throughout by moderately formed granulation (that is typically a paler green than the head capsule base color) and a posteromedial tubercle which is notably larger than the other granulation (and is often brown in color). Frontal convexity finely pointed, with several short setae on the apex. Compound eyes slightly protruding from the head capsule, not overly large, taking up ca ¼ of the head capsule lateral margins. Ocelli absent. Antennal fields slightly wider than the width of the first antennomere. Antennae. Antennae consist of nine segments (including scapus and pedicellus), with the terminal segment approximately the same length as the preceding 2⅓ segments’ lengths combined. Antennomeres I-VIII are marked with slight granulation and sparse short setae. Thorax. Pronotum with slightly convex anterior margin and gently converging lateral margins, which converge to the posterior margin that is ⅔ the width of the anterior margin. The pronotum surface is slightly lumpy and marked with a distinct sagittal furrow and slight central perpendicular furrow. The pronotum has distinct, smooth anterior and lateral rims. Prosternum, mesosternum, and metanotum are covered sparsely with nodes that are irregularly spaced throughout the surface, but all relatively small. Prescutum approximately as long as wide, with lateral rims marked by 10-12 small tubercles relatively evenly sized and spaced, giving the lateral rims a rough appearance. The prescutum anterior rim is distinct but not strongly protruding and the rim surface is relatively smooth. Prescutum surface slightly raised along the sagittal plane and the surface is only slightly marked with granulation. The mesopleura start to diverge slightly posterior to the prescutum anterior rim and diverge uniformly with straight margins. The mesopleuron margin is marked throughout with five or six larger tubercles and six or seven smaller tubercles interspersed throughout. Mesopleuron surface slightly granular. Wings. Tegmina long, typically reaching significantly onto abdominal segments VII or VIII. Tegmen venation; the subcosta (Sc) is the first vein in the forewing and runs parallel with the margin for the anterior ½, and then bends slightly and runs towards the margin where it terminates ca ¼ of the way through the tegmen length. The radius (R) spans the central portion of the forewing with two subparallel branched veins; the first radius (R1) branches ca ¼ of the way through the tegmen length and terminates slightly proximal to the midline, and the radial sector (Rs) branches ca ⅓ of the way through the tegmen length and terminates ca ⅗ of the way through the length. The media (M) is bifurcate with media anterior (MA) splitting near the middle of the tegmen length and the media posterior (MP) splits ca ⅗ of the way through the length. The media anterior and media posterior terminate near to the posterior ¼ of the tegmen after arcing gently and running parallel with each other. The cubitus (Cu) is bifurcate, branching near the posterior ¼ of the wing into the cubitus anterior (CuA) and cubitus posterior (CuP) which both terminate near the tegmen apex. The first anal vein (1A) is simple and fuses with the cubitus ca ¼ of the way through the tegmen length. Ala rudimentary, no more than a nub. Abdomen. Abdominal segments II through the anterior ⅔ of segment IV diverge strongly, the remainder of IV diverges only slightly to segment V which is the widest segment, followed by segments VI and VII which converge slightly with straight margins to VIII through X which converge more significantly to the broadly rounded apex. Genitalia. Subgenital plate starts on the anterior margin of tergum VIII, is moderately broad, and extends to the anterior margin of tergum X, ending in a blunt apex. Gonapophyses VIII are moderate in length, extending halfway onto tergum X, and are moderately broad; gonapophyses IX are smaller and hidden under gonapophyses VIII. The cerci are flat, not strongly cupped, with a granular surface and numerous small setae. Legs. Profemoral exterior lobe broad, and somewhat recurved with a slightly acute angle and a greatest width ca 5 ½ to 6 × the width of the profemoral shaft. Profemoral exterior lobe proximal margin is marked with four or five distinct serrate teeth, and the distal margin is smooth, lacking serration. Profemoral interior lobe is only situated on the distal ⅔ of the profemora, ca 3 × as wide as the greatest width of the profemoral shaft and is roundly arcing. The proximal margin of the profemoral interior lobe is relatively smooth and the distal margin is marked with three or four dulled, weakly formed teeth with looping gaps between them. The mesofemoral exterior lobe is roundly triangular and arcs from end to end. The mesofemoral exterior lobe proximal margin is straight and smooth, while the distal margin is slightly rounded and marked with four or five serrate teeth. The mesofemoral interior lobe is slightly wider than the mesofemoral shaft, and the exterior lobe is ca 1½× wider than the mesofemoral shaft. The mesofemoral interior lobe arcs unevenly end to end with seven or eight dulled teeth throughout the length, and the distal end slightly wider than the proximal end. The metafemoral exterior lobe is thin and smooth, hugging the metafemoral shaft and lacks dentition. The metafemoral exterior lobe is thin on the proximal ⅓ and then a gently arcing lobe on the distal ⅔ approximately as wide as the metafemoral shaft width. The metafemoral exterior lobe has moderate serration throughout with those on the distal ½ more prominent. Protibial exterior lobe unevenly arcing with a smooth margin from end to end with a steady increase in width from the proximal to the distal ends. Protibial interior lobe also spans the entire length in a smooth, rounded triangle, with the widest point on the distal ⅓. Mesotibiae and metatibiae lacking interior lobes, with exterior lobes having a similar rounded triangular shape weighted towards the distal end and ca 2 × as wide as the shafts on which they are found.

Measurements of paratype females [mm]. Length of body (including cerci and head, excluding antennae) 76.6-78.7, length/width of head 8.9-9.4/6.9-7.1, antennae 4.0-4.1, pronotum 5.2-5.5, mesonotum 4.5-4.7, length of tegmina 44.7-47.6, greatest width of abdomen 34.9-35.4, profemora 19.5-19.9, mesofemora 14.0-14.4, metafemora 15.8-16.1, protibiae 9.8-10.0, mesotibiae 9.0-9.4, metatibiae 11.9-12.2.

Male. Coloration. Coloration description based upon numerous images of live individuals recorded on iNaturalist (Fig. 33A, B ). Base coloration variable, ranging from yellow to pale green. Throughout the body are variable patches of tan/brown, primarily found on the abdomen, venation of the wings, and lobes of the legs. Typically, the protibiae and the profemoral interior lobe are brown as well as the antennae. Abdominal segment V has a set of large, dark eye spots.

Morphology. Head. Head capsule approximately as long as wide, with a vertex that is weakly granular; posteromedial tubercle small but distinctly raised from the capsule surface (Fig. 31D ). Frontal convexity finely pointed, with two or three short setae near the apex. Compound eyes notably bulbous occupying ca ⅖ of the head capsule lateral margins. Three ocelli are well-developed and located between and slightly posterior to the compound eyes. Antennae. Antennae (including the scapus and pedicellus) consist of 22 or 23 segments. The scapus and pedicellus are bare, all other segments are covered in dense, thin setae that are as long as or longer than the antennal segment is wide. The terminal three segments have shorter and denser setae than the other segments. Each antennomere on the distal end projects ventrally slightly, so the overall antenna has a serrate appearance. Thorax. Pronotum with anterior margin gently concave and lateral margins that are nearly straight and converge to a straight posterior margin that is approximately ½ the width of the anterior rim (Fig. 31C ). Anterior and lateral margins of the pronotum with distinct rims and the posterior margin lacks a distinct rim (Fig. 31C ). Face of the pronotum is marked by a slightly wrinkled surface, a distinct sagittal furrow, and a small perpendicular furrow near the center. The prosternum, mesosternum, and metasternum surfaces are nearly smooth, with slightly wrinkled surfaces and sparse granulation. The prescutum is slightly longer than wide, with lateral margins converging to the posterior which is slightly narrower than the anterior rim width (Fig. 31C ). The lateral rims of the prescutum are marked with nodes throughout, giving the margins a rough textured appearance. The surface of the prescutum is somewhat raised along the sagittal plane and this crest has a weakly granular texture while the remainder of the surface is relatively smooth. The prescutum anterior rim is distinct, but not prominent, with a surface that is relatively smooth. The mesopleura are narrow on the anterior then diverge more prominently from the anterior to the posterior. Lateral margins of the mesopleurae with granulation throughout, giving the margin a rough textured appearance. Mesopleuron surface relatively smooth. Wings. Tegmina short, extending ca ½ onto abdominal segment II. Tegmen wing venation: the subcosta (Sc) is the first vein and runs relatively straight for ca ½ of the tegmen length before terminating on the wing margin. The radius (R) spans the entire length of the tegmen, running as the radial sector (Rs) straight through the center to the apex after the first radius (R1) branches near the center of the tegmen and runs to the margin on the posterior ⅓ where it terminates. The media (M) spans the entire length of the tegmen, running parallel with the radius and radial sector and terminates at the tegmen apex as the media anterior (MA) after the branching of a weakly formed media posterior (MP) near the middle of the tegmen and runs at an angle towards the apex. The cubitus (Cu) runs through the tegmen surface angled away from the media (M) for ca ⅓ of the length to the tegmen margin and then runs along the margin where it fades before meeting the apex. The first anal (1A) vein runs subparallel to the cubitus until they meet ca ¼ of the way through the tegmen length. The alae are 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 wing 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 through the first ½ of their lengths, then run parallel until fusing with the cubitus at different locations on the posterior ⅕ of the wing. 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. The media anterior fuses with the cubitus (Cu) near the posterior ¼ of the ala length. The media posterior fades before fully fusing with the cubitus. The cubitus runs unbranched and terminates at the wing apex after the media anterior, 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 slightly distal to the point where the media branches into the media anterior and media posterior and then the first anterior anal branches from the cubitus ¾ 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. The general abdominal shape is ovular. Abdominal segments II through the anterior ⅔ of IV are gradually diverging, the posterior ⅓ of segment IV diverges less prominently, V through VII are subparallel or slightly converging, VIII through X converge rapidly to the apex. Genitalia. Poculum broad and rounded, ending in a straight margined apex that passes beyond the anterior margin of segment X (Fig. 31E ). The cerci are slightly cupped, covered in a partially granulose surface with sparse short setae throughout. The vomer is broad and slightly longer than wide, with relatively straight sides evenly converging to the apex, which is armed with a single thick upwards turning hook. Legs. Profemoral exterior lobe broad and slightly obtuse angled, ca 1½× widener than the interior lobe. Profemoral exterior lobe proximal margin slightly granular, and distal margin with small serration throughout the length (Fig. 31C ). Profemoral interior lobe begins approximately ⅓ of the way through the profemoral length, and arcs gently with a maximum width of ca 2½× the profemoral shaft width. Profemoral interior lobe is marked by three to four teeth on the distal ½ of the lobe, the proximal most is small, followed by two or three broadly triangular teeth. Mesofemoral exterior lobe is roundly triangular, relatively evenly weighted from end to end, and broad, with the proximal margin straight and smooth, and the distal margin slightly arcing and marked by four or five prominent serrate teeth. The greatest width of the mesofemoral exterior lobe is ca 2 × the width of the mesofemoral shaft. The mesofemoral interior lobe is slightly thinner than the exterior lobe and is unevenly weighted from end to end with the widest portion near the distal ⅓. The proximal ⅓ of the mesofemoral interior lobe is narrow, followed by a rounded arc on the distal ⅔. There is unevenly spaced and variable sized serration throughout the full margin of the mesofemoral interior lobe (typically 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. Metafemoral interior lobe is narrow and straight for the proximal ½ and then on the distal ½ there is a slightly expanding lobe that is slightly wider than the shaft’s width. There is serration of variable size and spacing throughout the length (eight or nine teeth). Protibial exterior lobe is a thin scalene triangle weighted towards the distal end which is 1- 1½× as wide as the protibial shaft width. The protibial interior lobe is also a scalene triangle but it is slightly wider than the exterior lobe, ca 2½× as wide as the protibial shaft width, and the widest portion is weighted near the distal ⅖. Mesotibial exterior lobe is roughly a thin, smooth triangle which is weighted towards the distal ⅓ and has a greatest width ca 1½× the width of the mesotibial shaft. Metatibial exterior lobe roughly a thin smooth triangle which is weighted towards the distal ⅓ and has a greatest width ca 3 × the width of the metatibial shaft. Mesotibiae and metatibiae lack interior lobes.

Measurements of holotype male [mm]. Length of body (including cerci and head, excluding antennae) 63.1, length/width of head 3.7/3.9, antennae (missing from holotype specimen), pronotum 3.1, mesonotum 2.7, length of tegmina 13.2, length of alae 49.1, greatest width of abdomen 22.5, profemora 14.2, mesofemora 11.6, metafemora 9.2, protibiae 6.9, mesotibiae 6.8, metatibiae 9.1.

Newly hatched nymphs.

(Fig. 32B, C ). The general color throughout the body ranges from orange to bright vermillion with muddled lighter and darker areas. The basitarsi are white/yellow, with the remaining tarsal segments similar to the body base color. All tibiae have well-developed exterior lobes of similar shapes, all a rounded arc that is widest approximately halfway through the tibial length. The protibial interior lobe is prominent, a similar size to the exterior lobe, but not as strongly angled. The protibial lobes have widths that are ca 2 × as width of the protibia shaft width. Profemoral interior lobe occupies the distal ⅔ and is slightly <2 × the profemoral shaft width. The profemoral interior lobe has two weakly formed teeth and is slightly angled. Profemoral exterior lobe has a clearly defined obtuse angle with a distal margin that is marked by weak dentition. The profemoral exterior lobe is ca 3 × the width of the profemoral shaft. All femoral lobes have mild serration (with the exterior lobes marked by slightly more prominent serration). 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 have a similar shape to their interior lobe counterparts but are less angled than the interior lobe. The meso- and metafemoral interior and exterior lobes have two or three small teeth on the distal end, with the teeth of the mesofemoral interior lobe less prominent than the mesofemoral exterior lobe, and the metafemoral interior lobe more prominent than the metafemoral exterior lobe. The head and thorax base color are variable, ranging from vermillion to light orange, often muddled in color. The thorax is of a similar color pattern to the head, but with margins marked in a darker color, ranging from reddish to brown. The abdomen base color is of a similar muddled coloration to the head/thorax, but typically with darker markings. Segments II through VIII are marked near the middle with an undulating darker pattern, with the inner area a lighter, uniform color, and the remainder of the segment of a darker muddled coloration. Segments IX and X are mostly muddled brown without a distinct pattern. The abdomen is notably wide, with a maximum width only slightly less than the abdomen length. The widest point of the abdomen is abdominal segment V.

Etymology.

Proper noun, named after Anangu, one of the Yakshi, or tree nymphs of Tamil mythology from southern India ( Pai-Dhungat 2020). This name was chosen to reference the range of this species in southern India and because leaf insects are mysterious, seldom seem creatures, likening them to spirits of the trees. Additionally, the Yakshi are often associated with mango trees (a favorite host plant of leaf insects).

Distribution.

At present known from multiple states in southwestern India (Fig. 15 ). States that this species have been recorded from are Goa, Karnataka, Kerala, and Tamil Nadu. See Suppl. material 1 for more details.

Remarks.

Despite being an undescribed species, leaf insect observations from India are commonly recorded on the citizen science platform iNaturalist. These submitted observations allowed a significant look into adult male (Fig. 33A, B ) and female (Fig. 32A ) morphology, as well as the coloration of the freshly hatched nymphs (Fig. 32B, C ). Freshly hatched nymphs are quite active after they hatch, brightly colored, and travel from the forest floor up into the canopy, thus allowing them to be readily observed by citizen scientists. Unfortunately, the eggs of this species are presently unknown, as the eggs are lost within leaf litter and never observed/recorded. Hopefully a captive colony in India one day will reveal the unknown egg morphology of this species and allow differentiation from congenerics. Phylogenetic analysis and morphological comparison with congenerics have now allowed this commonly encountered species to be recognized as undescribed and endemic to southwestern India.

Within the southwestern Indian state of Kerala, the Malayali people call this species "Pera Rani" (meaning "Queen of Guava plant" in Malayalam) a fitting common name as phylliids are frequently observed by farmers of guava (pers. comm. Gavas Ragesh, Kerala Agricultural University). Local farmers additionally report this species as being observed on cashew and mango trees, but never in significant enough populations to be considered a pest. Of these three common agricultural plants, only mango is native to Asia (with the other two native to the tropics of the western hemisphere; Morton 1987), but little is presently known about other potential native host plants for leaf insects.

Figures

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 15. Distribution map for the herein discussed Pulchriphyllium species with known records plotted. For data on the distribution points used for this map, see Suppl. material 1 A distribution of the " Pulchriphyllium bioculatum - like species " B distribution of the Pulchriphyllium species from mainland Asia with a boxy abdominal shape (inset outline in A). Abbreviations: KH, Cambodia; LA, Laos; TH, Thailand; VN, Vietnam.

Figure 31. Male holotype Pulchriphyllium anangu sp. nov. (BYU) A ventral, habitus B dorsal, habitus C details of the front leg, head, and thorax, dorsal D details of the head through thorax, lateral E genitalia, ventral F terminalia, dorsal. Photographs taken by JBL.

Figure 32. Live Pulchriphyllium anangu sp. nov. A adult female, dorsal habitus, observed July 2016 at Kadumane Estate, India by iNaturalist user @ ashwinv (Ashwin Viswanathan) https: // www. inaturalist. org / observations / 29374627) B freshly hatched nymph, observed October 2021 at Chikkamagaluru, Karnataka, India by iNaturalist user @ prashanthakrishnamc (Prashantha Krishna M C) https: // www. inaturalist. org / observations / 98984918) C freshly hatched nymph, observed January 2021 in Ponmudi, India by iNaturalist user @ hari _ krishnan _ (Harikrishnan S) https: // www. inaturalist. org / observations / 70254070). Uploaded to iNaturalist and used under license CC BY-NC 4.0.

Figure 33. Live Pulchriphyllium anangu sp. nov. A adult male, dorsal habitus observed October 2006 in B. R. Hills, Karnataka, India by iNaturalist user @ prashanthns (Prashanth N S) (https: // www. inaturalist. org / observations / 31929431) B adult male, dorsal habitus observed November 2019 in Pakkalakunja, Karnataka, India by iNaturalist user @ sanath _ ramesh _ manimoole (Sanath R M) (https: // www. inaturalist. org / observations / 103401996) C male nymph, dorsal habitus observed February 2020 in Shimoga, Karnataka, India by iNaturalist user @ girishgowda (Girish Gowda) (https: // www. inaturalist. org / observations / 104655374) D nymph, dorsal habitus, hiding under a leaf in the daylight observed August 2020 near Agali, India by iNaturalist user @ manavsajan (https: // www. inaturalist. org / observations / 56588953). Uploaded to iNaturalist and used under license CC BY-NC 4.0.