Phyllium ortizi 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

Phyllium ortizi sp. nov.

Figs 8A , 10 , 11 , 12 , 13 , 14

Material examined.

Holotype ♀: " Philippines, Mindanao, Lanao Del Sur, Wao, February 2019; Collection SLT; DNA no. 35". Deposited in the Montreal Insectarium , Quebec, Canada (IMQC; Fig. 11D ) . Paratypes: (5 ♀♀, 14 ♂♂, 2 ♀♀ nymphs, 99 eggs). See Suppl. material 1 for details about paratype specimens, their collection data, and depositories.

Differentiation.

Females are morphologically most similar to Phyllium bilobatum due to their small size, strongly lobed abdomen, broad profemoral exterior lobe, large triangular teeth of the profemoral interior lobe, and similar thorax spination. The antennae morphology can differentiate these species as Phyllium ortizi sp. nov. has the terminal antennal segment notably longer (approximately the same length as the preceding two and a half or three segments; Fig. 11E ) vs Phyllium bilobatum which has a stout terminal segment which is only approximately as long as the preceding two segments. Additionally, the tegmina venation can differentiate these species as Phyllium ortizi sp. nov. has tegmina with the distance between the first radial (R1) split and the wing base ca 1½× greater than the distance between the first radial (R1) split and the radius to media crossvein (R-M) vs in Phyllium bilobatum where the distance between the first radial (R1) split and the wing base is notably greater than the distance between the first radial (R1) split and the radius to media crossvein (R-M) (ca 2½× longer). Phyllium ortizi sp. nov. occurs within the range of Phyllium mabantai and due to the extreme morphological variability of Phyllium mabantai , certain strongly lobed forms of Phyllium mabantai females can look similar to Phyllium ortizi sp. nov. females. However, size appears to be able to consistently differentiate these two species as Phyllium ortizi sp. nov. are smaller (66.8-76.4 mm long) vs Phyllium mabantai which are larger (91.2-99.4 mm long; Hennemann et al. 2009). Besides size, it can be very difficult to differentiate these two species when only dealing with female specimens; however, it appears as though the spination of the prescutum sagittal plane might allow differentiation as there are typically only three tubercles in Phyllium ortizi sp. nov. females which are systematically decreasing in size and are distinctly raised above the prescutum surface (Fig. 11B ) vs Phyllium mabantai which can have three to five blunted nodes which follow the anterior spine, and these are not as prominent.

For male Phyllium ortizi sp. nov. they are morphologically most similar to Phyllium mabantai and Phyllium iyadaon sp. nov. based upon their size, wing venison, and general lobes of the legs. The abdomen of Phyllium ortizi sp. nov. appears to be relatively stable in shape based upon reared and wild collected specimens and within Phyllium mabantai male abdominal shape is also rather stable despite female Phyllium mabantai having drastically different forms. Between these two species the abdominal general shape and the size of the eye sports appears to be a reliable feature for differentiation as the Phyllium mabantai abdomen has smooth abdominal edges (either giving the abdomen a spade shaped or slightly ovoid appearance) and abdominal segment V has small eye spots vs Phyllium ortizi sp. nov. which has somewhat of a rectangular abdomen (with segments V and VI approximately even in width) and the margins of VI, VII, and VIII gently rounded giving the abdomen a slightly scalloped edge and the eye spots of abdominal segment V are always large taking up at least ½ of the segment length (Fig. 10D ). This scalloped edge abdominal shape makes Phyllium ortizi sp. nov. superficially appear similar to Phyllium iyadaon sp. nov. but these two species can be differentiated by the width of the interior protibial lobe, interior profemoral lobe, and exterior mesofemoral lobe. In Phyllium ortizi sp. nov. the protibial interior lobe is at least 2 × wider than the width of the protibial shaft (Fig. 12A ) vs Phyllium iyadaon sp. nov. which is much narrower, at most only slightly wider than the protibial shaft width (Fig. 9C ). In Phyllium ortizi sp. nov. the protibial interior lobe is much more pronounced with a width greater than 2 × the profemoral shaft width and an angle ca 90 degrees (Fig. 12A ) vs Phyllium iyadaon sp. nov. which has a notably more slender profemoral interior lobe with a distinctly obtuse angle and a width only ca 1½× as great as the profemoral shaft width (Fig. 9C ). Additionally, the exterior mesofemoral lobe allows differentiation as the lobe in Phyllium ortizi sp. nov. is distinctly marked by serrate teeth (at least two) but in Phyllium iyadaon sp. nov. the exterior lobe is bare, lacking teeth.

The eggs of Phyllium ortizi sp. nov. are rather unique as they have two general types of pinnae on their surfaces (of which all surfaces have the same types) some that are the more typical feather-like pinnae and a second type which is thinner and more filament-like, lacking projections like is seen in the feather-like pinnae. These thinner filament-like pinnae are not seen in other phylliids eggs, and the closest types are those seen in Phyllium tobeloense bhaskarai Cumming et al. 2019b; however, that species typically has the thinner pinnae ending in a fork or a hooked tip vs in Phyllium ortizi sp. nov. where the tips taper off and are thin (Fig. 13 ). Phyllium ortizi sp. nov. eggs also lack any "bald patches", a feature which many species have such as Phyllium mabantai .

Freshly hatched nymphs are only known for a handful of Phyllium species at present, but from what is known Phyllium ortizi sp. nov. can be differentiated from congenerics reliably. Phyllium ortizi sp. nov. (Figs 8A , 14 ) is most similar to Phyllium mabantai (Fig. 8B ) and Phyllium samarense sp. nov. (Fig. 8C ) due to each abdominal segment lateral lobe being split approximately in half by two colors (with the anterior ½ mint/lime green and the posterior 1/2 brown; Fig. 8A-C ). All other Phyllium species have the abdomen rather uniform in color (brown/black) and at most only have the margins slightly marked with white or yellow. Phyllium ortizi sp. nov. can be differentiated from Phyllium mabantai and Phyllium samarense sp. nov. based on the coloration of the meso- and metafemoral lobes. In Phyllium ortizi sp. nov. the white stripe is broken on both the meso- and metafemora therefore giving them a somewhat large spotted appearance, vs, the other two species which have these white stripes unbroken reaching from one end of the femora to the other. Additionally, the green color on the abdomen of Phyllium ortizi sp. nov. appears to be a bit darker and more richly green than the others which are paler and more mint green but depending on the lighting they are observed under the coloration can vary slightly.

Description.

Female. Coloration. Coloration description is based upon living individuals in culture (Fig. 10A, B, E ). Overall coloration is lime green throughout, with certain areas slightly variable in coloration with tan, orange, or brown. To date the majority of females in captive culture have mostly been solidly green with minimal additional colored area, but consistently the mesonotum has been tan and brown in color. Within captive cultures a few individuals have expressed significant coloration variations, such as fully brown/tan (Fig. 10E ). The margins of the femoral lobes and the protibial interior lobe are the most variable with darker colored individuals having most of the lobe colored (Fig. 11A ) but often these features in paler individuals are only lightly highlighted in brown/orange. Compound eyes are of a tan coloration. The ventral coloration of the meso- and metacoxae are bright orange (Fig. 10B ).

Morphology. Head. Head capsule slightly longer than wide, vertex moderately wrinkled with evenly spaced, small nodes across most of the surface (most distinct on the posterior ½), with the posteromedial tubercle the most prominent feature on the head capsule, many times larger and broader than any of the nodes (Fig. 11C ). Frontal convexities broad and moderately pointed, with a slightly granular surface and sparse short setae throughout. Compound eyes slightly protruding from the head capsule, not overly bulbous, taking up ca ⅖ of the head capsule lateral margins (Fig. 11C ). Ocelli absent. Antennal fields ca ⅓ wider than the width of the first antennomere. Antennae. Antenna consist of nine segments, with the terminal segment approximately the same length as the preceding 2½ or 3 segments’ lengths combined (Fig. 11E ). Antennomeres I-VII sparsely marked with short and thin transparent setae, the terminal two antennomeres are covered in stout, dark brown setae which are irregularly spaced (Fig. 11E ). Thorax. Pronotum with gently concave anterior margin and straight lateral and posterior margins. The overall shape is an isosceles trapezoid with the anterior margin the longer side and the narrower posterior margin ca ⅗ of the anterior width (Fig. 11C ). The pronotum surface is slightly wrinkled, with only a prominent pit in the center, and a distinct furrow anterior to the central pit and weakly formed posterior and lateral furrows to the pit (Fig. 11C ). The pronotum has moderately formed anterior and lateral rims and a weakly formed posterior rim, all of which are smooth, lacking distinct granulation or setae (Fig. 11C ). Prosternum and the anterior ⅓ of the mesosternum have a slightly wrinkled surface marked with numerous broad nodes, with the remainder of the mesosternum and the metasternum relatively smooth without notable features. Prescutum approximately as long as wide, with lateral rims marked by seven or eight medium to large tubercles with somewhat irregular spacing (Fig. 11C ). Prescutum anterior rim prominent, strongly protruding as a large sagittal spine, with the rim surface slightly lumpy (Fig. 11C ). Prescutum sagittal crest with two additional distinct tubercles, reducing in size steadily from the prominent anterior tubercle to a smaller posterior tubercle (Fig. 11B ). Prescutum surface irregularly lumpy and slightly raised along the sagittal crest (Fig. 11B ). Mesopleura begin at the anterior of the prescutum and diverge slowly on the anterior margin but immediately angle outwards prominently and then evenly diverge throughout the rest of their lengths (Fig. 11C ). Mesopleuron lateral margin with six or seven medium to large tubercles with fine points and three or four smaller nodes interspersed, all situated on the anterior ⅗ with the remainder of the margin relatively smooth (Fig. 11C ). Face of the mesopleuron heavily wrinkled, with two notable divots, one on the anterior margin and one near the middle (Fig. 11C ). Wings. Tegmina long, reaching onto abdominal segment VIII. Tegmen venation; the subcosta (Sc) is the first vein and runs relatively straight from the base with just a slight arcing in the middle as it angles towards the margin, terminating on the margin ca ⅕ of the way through the tegmen length. The radius (R) spans the central portion of the tegmen and branches into two diverging veins; the first radius (R1) branches ca ⅕ of the way through the wing length, runs straight, and terminates on the margin slightly <⅓ of the way through the tegmen length, and the radial sector (Rs) branches ca 2/7 of the way through the tegmen length, arcs gently as it spans through the majority of the tegmen central surface, and terminates on the margin of the distal ⅓ of the tegmen length. There is a weak continuation of the radius following the prominent Rs branching which continues on as a short and thin radius to media crossvein (R-M) that weakly connects the two veins. The media (M) is bifurcate with the media anterior (MA) branching near the middle of the tegmen length and media posterior (MP) branches near the distal ⅓ of the tegmen length with both veins arcing gently and running parallel/subparallel towards the tegmen margin with the media anterior terminating near the distal ¼ of the tegmen and the media posterior terminating near the distal ⅕ of the tegmen length. Following the branching of the media posterior there is a thin media to cubitus crossvein (M-Cu). The cubitus (Cu) is also bifurcate, branching near the posterior ¼ of the tegmen into the cubitus anterior (CuA) which runs arcing parallel with the media posterior to the tegmen apex and the cubitus posterior (CuP) which runs away from the cubitus anterior as it follows the tegmen margin and it fades near the apex. The first anal vein (1A) is simple and fuses with the cubitus early on, at the length approximately midway between the splitting of the first radial and the radial sector. Alae rudimentary, only just a nub. Abdomen. Abdominal segments II through the anterior ⅔ of segment IV uniformly diverging with straight margins. The posterior ⅓ of segment IV through segment VI are subparallel, giving the abdomen a boxy appearance as these segments only slightly converge. The posterior margin of segments V and VI are slightly broader than the anterior margin of their following segment and this difference is notably more significant in segments VII and VIII which have distinct lobes projecting slightly beyond the anterior margin of the following segment as these segments narrow notably and converge towards the abdomen apex. Segments IX and X are notably narrower than the previous segments, and have converging, straight margins to the broad rounded apex. Genitalia. Subgenital plate starts at the anterior margin of tergum VIII, is moderately broad, and the apex extends ca ⅓- ½ of the way onto tergum X with margins that arc smoothly at first and then run slightly converging to the bluntly pointed apex (Fig. 11F ). Gonapophyses VIII are long and moderately broad, with their tips slightly exceeding the apex of the abdominal segment X; gonapophyses IX are shorter and narrower, mostly hidden below the gonapophyses VIII (Fig. 11F ). Cerci are mostly flat with only the interior margin near the middle slightly cupped, a surface that is relatively smooth with little granulation, and only the very slightly serrate terminal margins are marked with sparse, short setae (Fig. 11F ). Legs. Profemoral exterior lobe reaching from end to end in a broad, rounded, and obtusely angled lobe which is only slightly wider than the interior lobe. Edge of the profemoral exterior lobe granular with two or three small teeth present on the distal margin. Profemoral interior lobe ca 2 × as wide as the greatest width of the profemoral shaft, with the lobe in the shape of a right angle with the distal margin ornamented with four or five serrate or triangular teeth irregularly arranged in a two-two or three-two pattern with looping gaps between them, and typically the largest teeth are the ones in the middle. Mesofemoral exterior lobe arcs from end to end with the greatest width on the distal ⅓ and distal to this widest point are two small serrate teeth. Mesofemoral exterior lobe is slightly wider than the mesofemoral shaft width, and the interior lobe is slightly <2 × the width of the mesofemoral shaft. Mesofemoral interior lobe arcs from end to end with the greatest width on the distal ⅓ and from this greatest width to the distal end there are five serrate teeth while the thinner proximal portion of the lobe lacks dentition. Metafemoral interior lobe arcs end to end, with the distal half slightly wider than the proximal half and this distal half is marked with six to seven serrate teeth. Metafemoral exterior lobe is thin and smooth, hugging the metafemoral shaft and lacks dentition. The protibial interior lobe spans the entire length and is ca 2 × the width of the protibial shaft. The lobe is roundly triangular with the widest portion slightly distal to the midline. Protibiae lacks exterior lobes, mesotibiae and metatibiae lack both exterior and interior lobes.

Measurements of holotype female [mm]. Length of body (including cerci and head, excluding antennae) 66.8, length/width of head 6.0/5.4, antennae (broken in the holotype), pronotum 5.9, mesonotum 6.2, length of tegmina 43.5, greatest width of abdomen 31.6, profemora 12.3, mesofemora 11.0, metafemora 13.8, protibiae 7.3, mesotibiae 8.5, metatibiae 12.1.

Measurements of paratype females [mm] (ex culture). Length of body (including cerci and head, excluding antennae) 74.2-76.4, length/width of head 6.2-6.4/5.9- 6.1, antennae 4.1-4.4, pronotum 3.9-4.2, mesonotum 6.4-6.6, length of tegmina 47.8-48.2, length of alae 2.4-2.6, greatest width of abdomen 35.5-36.8, profemora 14.0-14.4, mesofemora 12.8-13.2, metafemora 15.7-16.1, protibiae 7.5-8.2, mesotibiae 8.4-8.9, metatibiae 13.7-14.1.

Male. Coloration. Coloration description is based upon living specimens in captivity (Fig. 10A, C, D ). Overall coloration lime green throughout with variable patches of brown to ruddy brown coloration. These variable brown to ruddy brown areas are primarily around the margins on the lobes of the legs, the thorax, the tips of the antennae, a set of eye spots on abdominal segment V, and the margins of the abdomen. The areas that are most variable are the lobes of the pro- and meso- legs which can be fully or almost fully colored or only partially with just the margins colored. Ventral coloration of the meso- and metacoxae can be pale cream to slightly yellow, not identical to the green of their bodies (Fig. 10D ).

Morphology. Head. Head capsule approximately as long as wide, with a vertex that is slightly lumpy and marked by a few irregularly spaced nodes (Fig. 12C ). Frontal convexities stout, notably short, and marked by only a few short setae. The posteromedial tubercle is not large (but it is larger than any of the nodes on the head) and it is only slightly raised from the head capsule. Compound eyes large and bulbous, occupying ca ⅖ of the head capsule lateral margins (Fig. 12C ). There are three well-developed ocelli that are notably raised and located between the compound eyes (Fig. 12C ). Antennae. Antennae (including the scapus and pedicellus) each consist of 23 segments, all segments except the scapus and pedicellus and terminal five segments are covered in varyingly spaced setae that vary in length (many approximately as wide as the segment or slightly shorter, others longer than the segment is wide; Fig. 12D ). The terminal five segments are covered in dense, short, darkly colored setae and the scapus and pedicellus are nearly completely bare as they only have a few thin and short setae (Fig. 12D ). Thorax. Pronotum with anterior margin notably concave and lateral margins slightly convex and converging to a straight posterior margin that is slightly> ½ the width of the anterior rim (Fig. 12C ). Anterior and lateral margins of the pronotum have well-formed rims and the posterior margin lacks a rim (Fig. 12C ). Face of the pronotum is marked by a distinct furrow and pit in the center and a relatively smooth surface with sparse, irregularly spaced granulation (Fig. 12C ). Prosternum surface is heavily marked with large nodes with those on the posterior half slightly larger than those on the anterior. Almost the entire mesosternum surface is distinctly wrinkled and marked with granulation mostly along the sagittal plane. Metasternum surface with only weak wrinkles and minimal or no granulation. Prescutum approximately as long as the greatest anterior width, with lateral margins that are converging to the notably narrower posterior (Fig. 12C ). Lateral rims with eight or nine tubercles, of these around five are slightly more prominent than the others (which are smaller and can vary in placement/size). Prescutum surface smooth and rises up to meet the sagittal crest which is marked by four tubercles which decrease in size from the anterior to the posterior. Prescutum anterior rim is well-formed and prominently raised into a distinct sagittal tubercle, and the surface of the rim is smooth (Fig. 12F ). The mesopleura begin on the anterior prescutum margin and diverge at an increasing degree throughout their length, at first only gradually but increasingly diverging to the posterior (Fig. 12C ). Mesopleuron lateral margin with five larger tubercles throughout the length with seven or eight smaller nodes spaced throughout (sometimes one or two between major tubercles) and almost all tubercles end with a seta or two. Face of the mesopleuron notably wrinkled and marked with two weak divots (one near the middle and the other near the anterior margin). Wings. Tegmina moderate length, extending ½ of the way onto abdominal segment III. Tegmen wing venation: the subcosta (Sc) is the first vein, is simple, runs through the tegmen for the first half of its length and then on the distal half it bends and runs angled towards the tegmen margin where it terminates ca ½ of the way through the overall tegmen length. The radius (R) spans the entire length of the tegmen with the first radius (R1) branching ca ⅖ of the way through the tegmen length and terminates near the distal ⅓ of the tegmen, and then the radial sector (Rs) runs straight to the wing apex. The media (M) also spans the entire length of the tegmen and runs side by side along the radius/radial sector with the media posterior (MP) branching off near the middle of the tegmen and running angled towards the apex/cubitus, and the media anterior (MA) runs straight to the tegmen apex. The cubitus (Cu) cuts across the tegmen to the margin ca ⅓ of the way through the length and runs along the edge of the tegmen. The media posterior vein fuses with the cubitus and as the cubitus reaches the apex it fades. The first anal (1A) vein terminates upon reaching the cubitus ca ⅓ of the way through the tegmen length. Alae well-developed in an oval fan configuration, long, reaching onto abdominal segments IX. Ala wing venation: the costa (C) is present along the entire foremargin giving stability to the ala. The subcosta (Sc) is long, spanning ca ½ of the ala length and is mostly fused with the radius near the base of the wing but terminates when it meets the costa. The radius (R) spans the entire wing and branches slightly <⅓ of the way through the ala length into the first radius (R1) and radial sector (Rs) which run gently diverging for ca ½ of their length, then run parallel until they near the apex of the ala where the radial sector begins to arc towards the first radial and either weakly joins it or fades before fully fusing with it, and the first radial either weakly reaches the apex or fades before fully reaching the apex. The media (M) branches early, ca ⅛ of the way through the ala length into the media anterior (MA) and the media posterior (MP) which run gently diverging at first, then parallel for a short while, then gently converging for ca ½ their length until the distal ¼ of the wing where the media posterior fuses with the media anterior which runs fused to the apex where it terminates. 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 slightly> ⅕ of the way through the ala length and then the first anterior anal branches from the cubitus ⅗ of the way through the ala length where it uniformly diverges away 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 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. Lateral margins of abdominal segment II parallel, III slightly diverging, IV diverging more prominently for the anterior ⅔ then run parallel, V and VI parallel or nearly so with the posterior end slightly wider on each segment so the abdomen has a slight undulating appearance, VII has a rounded margin narrowing to the notably narrower VIII, VIII through X converging relatively uniformly to the rounded apex of the abdomen. Genitalia. Poculum broad and ends in a blunt apex that slightly passes the anterior margin of segment X (Fig. 12E ). Cerci long and slender, extending from under the anal abdominal segment, nearly flat, surface slightly granular, and notably pubescent. Vomer broad and stout with straight sides evenly converging and ending in a singular thick apical hook (Fig. 12E ). Legs. Profemoral exterior lobe arcs from end to end, hugging the profemoral shaft, with a maximum width as wide as the profemoral shaft width, and the margin on the distal half is marked with five small, finely pointed serrate teeth, with the proximal half marked with minimal granulation, and sparsely ornamented with short, fine setae throughout the lobe length (Fig. 12A ). Profemoral interior lobe roundly triangular (approximately right angled) and typically marked with five sharp teeth arranged in a three-two pattern with looping gaps between them, with the central two teeth often larger than the others (Fig. 12A ). Mesofemoral exterior lobe is almost entirely situated on the distal half with the proximal half thin to almost fully reduced. Mesofemoral exterior lobe has its greatest width on the distal end and is approximately the same width as the greatest mesofemoral shaft width. The distal ⅓ of the mesofemoral exterior lobe is marked with two or three sharply pointed serrate teeth. Mesofemoral interior lobe has a similar shape as the exterior lobe (very thin on the proximal ½ and widest on the distal ½) but the distal ⅖ is marked with 6-8 sharply pointed serrate teeth. Metafemoral exterior lobe lacks dentition and has a straight margin hugging the metafemoral shaft. Metafemoral interior lobe arcs end to end with the distal ½ wider than the proximal ½, and the distal ½ margin is marked with seven or eight strongly angled serrate teeth. Protibiae lacking exterior lobe, interior lobe reaching end to end in a broad arc with the proximal and distal ends notably tapered and the center the widest point which at its maximum is slightly> 2 × the width of the protibial shaft (Fig. 12A ). Meso- and metatibiae simple, lacking lobes completely.

Measurements of paratype males [mm] (ex culture). Length of body (including cerci and head, excluding antennae) 53.5-55.8, length/width of head 2.9-3.0/2.8- 2.9, antennae 2.6.1-27.9, pronotum 2.3-2.4, mesonotum 2.8-2.9, length of tegmina 17.6-19.0, length of alae 39.7-41.2, greatest width of abdomen 12.3-12.9, profemora 8.7-9.4, mesofemora 8.9-9.3, metafemora 10.4-10.8, protibiae 5.8-6.2, mesotibiae 5.3-6.0, metatibiae 7.8-7.9.

Eggs. (Fig. 13 ). The overall color is brown, with the capsule surface lighter in color and the pinnae darker. The lateral surfaces are nearly flat (only slightly convex), with the posterior of the egg slightly wider than the anterior. There are two types of pinnae on the capsule surfaces; moderate to long feather-like pinnae which are found along the operculum rim and along the capsule margins (these feather-like pinnae are not as well formed as in other species with large feather-like pinnae and instead these appear more like a "wet feather" with the lateral frills less defined) and an addition pinna type which are short to medium length “filament-like” pinnae which lack lateral frills and are instead a simple stock without elaborate expansions. The lateral surfaces of the capsule are densely marked throughout with short to medium length filament-like pinnae emerging from a roughly textured surface (Fig. 13A ). The dorsal surface has the micropylar plate (which is marked with a distinct margin) and the plate spans approximately the central ⅗ of the capsule length with a shape that is distinctly tear-shaped with the wide, rounded end around the micropylar cup which is located on the posterior ⅖ of the capsule (Fig. 13C ). On either side of the micropylar plate is a similar surface texture/pinnae arrangement to the lateral surfaces. The operculum is slightly ovular, the surface is flat and marked moderately by filament-like pinnae, and the outer margin is rimmed with medium length feather-like pinnae (Fig. 13D ). The ventral surface of the egg capsule has a surface texture and pinnae similar to the lateral surfaces (Fig. 13F ).

Measurements including the extended pinnae [mm]. Length (including operculum): 4.5-4.6; maximum width of capsule when viewed from lateral aspect 2.9-3.0; length of micropylar plate 2.2-2.3.

Newly hatched nymphs.

(Fig. 14 ). The general color throughout the body is a rich dark brown. The basitarsi are cream colored and the remaining tarsal segments are dark brown. All tibiae lack exterior lobes but the protibiae do have a smoothly arcing interior lobe which is approximately as wide as the protibial shaft width and the meso- and metatibiae have very thin interior lobes only approximately as wide as the shaft they are on. All tibiae have a similar color pattern with the proximal ¼ -⅓ pure white and the remainder brown with sparse white/tan partial striping (less so on the metatibiae and the most prominent on the protibiae). All femoral lobes are similar in color pattern with their primary color brown and near the middle (of the profemora) or the proximal ⅖ (of the meso- and metafemora) there are distinct white spots on each side of the femoral shaft that are crisply colored and not connected across the shaft. Additionally, on the meso- and metafemoral interior lobes at the base are small single white patches. All femoral interior lobes have mild serration and the meso- and metafemoral exterior lobes have dulled dentition on their distal ends. The distal end of the metafemoral exterior lobe also has a faint white edge. The head and thorax base color are brown, and the mesothorax and metathorax margins are marked with lime green. The abdomen base color is brown, with the centerline of the abdomen where the internal organs are fully brown but the thin lobes on each side of this center are bicolored. Each segment is marked on the anterior ½ -⅓ with green and the remainder of the segment is brown, except for segments VIII-X that are mostly brown and only marked minimally on the lateral margin with green. The abdomen’s general shape is long and narrow, with a maximum width < ½ of the abdomen length. The widest point of the abdomen is abdominal segment IV.

Etymology.

Patronym. Named after Maxime Ortiz (France) who is a well-known phasmid breeder who established this new species in culture and who shared type material with the authors for this study. Thanks to his efforts we were able to understand the morphological diversity within the males, females, freshly hatched nymphs, and eggs.

Distribution.

At present this species is known from four provinces in Mindanao; Lanao Del Sur Province (with the exact type locality of Wao Municipality), South Cotabato Province, Davao del Sur Province, and from the inexact locality of Bukidnon Province from captive reared specimens.

Remarks.

This species was first thought to represent a small, strongly lobed Phyllium mabantai or possibly the first record of Phyllium bilobatum Gray, 1843 in more than a century, a species with which Phyllium ortizi sp. nov. shares many general similarities. However, thanks to the rearing of additional specimens by Maxime Ortiz (France), the intraspecific variation was revealed, and the fine details instead reliably differentiate these species. Phyllium ortizi sp. nov. was sampled within Bank et al. (2021) as " Phyllium sp. 8 ‘Kitanglad’” where it was recovered as sister species to Phyllium samarense sp. nov. from Samar Island (" Phyllium sp. 3 'North Samar’” within Bank et al. (2021)). The same tree topology was recovered within our included phylogenetic analysis (Fig. 2 ).

The eggs of Phyllium ortizi sp. nov. are primarily covered with pinnae "type 5" (feather-like) as defined by Büscher et al. (2023). In addition to this type of pinnae, however, is a pinnae type not characterized within Büscher et al. (2023). Intermixed within the pinnae "type 5" are filament-like pinnae, thin tapering pinnae without significant side branches or prominent splitting at the apex (Fig. 13 ). Morphologically these filament-like pinnae appear to be simplified "type 5" pinnae, not a wholly unique type worth characterizing as an independent state, particularly in light of the addition of "type 5" pinnae on the egg which logically fits within the results of Büscher et al. (2023).

Figures

Figure 1. Live adult female leaf insects showing intergeneric and intraspecific color variability A Pulchriphyllium anangu sp. nov. photographed near Agumbe, India on 17 May 2022 by iNaturalist user vishwanathgowda. Used with permission of Vishwanath Gowda (India) https: // www. inaturalist. org / observations / 118478029 B, C Phyllium ortizi sp. nov. reared and photographed by Maxime Ortiz (France) in captivity B brown form female C green form female.

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 9. Phyllium iyadaon sp. nov. type specimens A-E holotype (IMQC) Coll RC 17 - 221 F paratype (Coll RC) Coll RC 17 - 222 A habitus, dorsal B details of the head through thorax, dorsal C details of the front leg, dorsal D genitalia, ventral E details of the head through thorax, lateral F details of the head through thorax, lateral. Scale bar: 20 mm (A).

Figure 10. Phyllium ortizi sp. nov. live individuals from captive culture highlighting aspects of their coloration. Photographs by Maxime Ortiz (France) A adult male and female pair, dorsal habitus B adult female, postero-ventral view, showing the bright orange coxae coloration C adult male, habitus, dorsal D adult male, habitus, ventral E brown form nymph, same individual as in Fig. 1 B before the final molt.

Figure 11. Phyllium ortizi sp. nov. female holotype (IMQC) D, paratype (Coll RC) A-C, E, F photographs by Rene ́ Limoges (IMQC). Scale bars associated with A, D respectively A dorsal habitus, paratype B lateral view of the thorax showing spination C details of the head and thorax, dorsal D dorsal habitus, holotype E details of the antennae, dorsal F genitalia details, ventral.

Figure 12. Phyllium ortizi sp. nov. male paratypes A-D (Coll RC 21 - 001). Photographs by Rene ́ Limoges (IMQC) E, F (Coll RC 21 - 002) A details of front leg B dorsal habitus C details of the head and thorax, dorsal D antenna, dorsal E genitalia details, ventral F lateral view of the thorax showing spination. Scale bar: 15 mm (B).

Figure 13. Phyllium ortizi sp. nov. egg capsule, IMQC collection, photographs by Rene Limoges (IMQC), scale bar associated with all images A lateral view B dorso-lateral view C dorsal view D opercular (anterior) view E posterior view F ventral view.

Figure 14. Phyllium ortizi sp. nov. freshly hatched nymphs reared and photographed by Maxime Ortiz (France) A habitus, dorsolateral B habitus, dorsal C two nymphs drinking moisture off of bramble (Rubus sp.), one of the few times freshly hatched nymphs hold still.