Pulchriphyllium schultzei Giglio-Tos, 1912: 56, herein designated.

This genus has a combination of features which link it to several genera, thus making a higher-level taxonomic placement difficult and requiring molecular confirmation in the future. The thorax and tegmina venation in the females are reminiscent of some Pulchriphyllium sensu stricto species and the profemoral and protibial lobes are reminiscent of some Phyllium Illiger, 1798 and Comptaphyllium Cumming et al., 2019 species, whereas the male thorax is similar to Trolicaphyllium Cumming et al., 2021. At present within Phylliidae there are two recognized tribes, the Nanophylliini Zompro and Größer 2003 (which contains only the Nanophyllium Redtenbacher, 1906) and the Phylliini Brunner von Wattenwyl, 1893 (which contains all other genera). Therefore, at this time this genus is placed within the tribe Phylliini as notable features such as a two lobed posteromedial tubercle and a prescutum which is wider than long (features which help to define the Nanophyllium) are absent, suggesting a closer relationship to other genera instead.

The selected type species for this new genus is Pulchriphyllium schultzei Giglio-Tos, 1912 (= Rakaphyllium schultzei (Giglio-Tos, 1912), comb. nov.) as this was the first species described within this new genus, the holotype is from an exact collection locality, and this species appears to be the more commonly encountered of the two species within this new genus.

This new genus has been recognized as unique in the past, as this clade was designated as the schultzei species group within Hennemann et al. (2009) based upon the shorter tegmina and the two lobes on the exterior protibiae. While these features are helpful to differentiate this genus from others, the below noted autapomorphic features allow differentiation from all phylliid genera.

Within each sex there is an easily observed morphological feature which supports their monophyly and readily separates them from other phylliid genera. For females, the gonapophyses VIII are exceptionally long, with approximately half of their length projecting from under the terminal abdominal segment (Figs 7G, 9D), a feature not seen in any other phylliid as typically gonapophyses VIII only reach to the apex of the terminal abdominal segment or at most only exceed the tip slightly (Fig. 10E). For males the alae venation is unique as the radius vein is simple (Fig. 3A), not bifurcate as is seen in all other phylliid genera (Fig. 3B). These autapomorphic features help to define the new genus Rakaphyllium gen. nov. within the phylliids as well as differentiate them from the Pulchriphyllium sensu stricto.

The Rakaphyllium gen. nov. are average sized phylliids, with females ranging from ca. 80 mm to 90 mm long and males ca. 60 mm long. Typical coloration appears to be green, but with so few specimens known and color variation a common occurrence in phylliids it is possible that this genus may also exhibit color forms (such as possibly Rakaphyllium exsectum comb. nov. but this brown coloration may simply be due to the age and preservation technique (Fig. 9)).

Antennae. Females have antennae with nine segments with the terminal antennomere not notably long (only as long as the previous one to two segments combined) and segments IV through VIII all of a similar length (Figs 6D, 7F, 9E). Males have antennae which range from 20 to 23 segments with most segments covered densely in setae, and overall antennomere shape somewhat flattened.

Head capsule. Males have well-developed ocelli (Fig. 8B), females do not have ocelli (Figs 6D, 7H). Males have compound eyes which are strongly protruding and occupy ca. 2/5 of the head capsule lateral margins (Fig. 8B) versus females which have compound eyes which are notably smaller, only occupying less than 1/3 of the head capsule lateral margins and which do not strongly protrude from the capsule (Fig. 7H). Both sexes have head capsules which are marked by irregularly sized and space granulation.

Figure 8. 

Rakaphyllium schultzei comb. nov. male from the MNHU collection A habitus, dorsal B details of front legs, base of antennae, head, thorax, and tegmina, dorsal C genitalia, ventral D details of front legs, base of antennae, head, thorax, and tegmina, dorsolateral. Scale bar only associated with A. Photographs by Frank Hennemann (Germany).

Thorax. The thorax is similar in both sexes with mesopleurae that are narrowly diverging from the anterior to the posterior (evenly so in females, almost parallel in males for the anterior half and then more prominently on the posterior half) are marked on the anterior half with three to four small tubercles with granulation interspersed with the posterior half relatively smooth or with only minimal granulation (Figs 6D, 7H). In both sexes the prescutum is about the same length as the width of the anterior margin, with a posterior margin that is slightly narrower giving the prescutum a slight isosceles trapezoid appearance. The margins of the prescutum are marked with granulation and the prescutum surface is covered with granulation with those along the sagittal plane slightly larger and in males a weak sagittal crest is present (Figs 6D, 8B, 9C). When viewed laterally, both sexes have a weakly formed prescutum anterior rim with a granular surface (Figs 7B, 8B).

Legs. Both sexes have interior protibial lobes which do not span the full length of the shaft, instead they are only situated on the proximal half (Figs 6B, 7C) and exterior protibiae which are marked by two lobes, one on the proximal and one on the distal end (Figs 6B, 7C). The exterior meso- and metatibiae are notably reduced, but if lobes are present its just as small spurs (sometimes just a distal spur or sometimes one on each end of the shaft) never as prominent lobes. Profemoral exterior lobes are rather variable as within Rakaphyllium schultzei comb. nov. females they are simply arcing smoothly from end to end without a strong angle (Fig. 6B), but in Rakaphyllium exsectum comb. nov. females the exterior lobe is distinctly boxy with a right angle (Fig. 9B). Males are only known for Rakaphyllium schultzei comb. nov. in which the femoral morphology is similar to the female, simply arcing tightly along the profemoral shaft (Fig. 8B).

Wings. Female tegmina are average in length, only reaching onto abdominal segments V or VI and male tegmina are moderate in length, only reaching onto abdominal segment II or III. Females have rudimentary alae. Male alae are fully developed in an oval-fan configuration and reach onto abdominal segments VIII to X. Female tegmina have a subcoastal vein; radial vein which runs parallel with the media and splits into the first radial about halfway through its length and terminates in a radial sector which bends distinctly away from the media and arcs to the wing margin; a bifurcate medial vein; a bifurcate cubitus vein; and a first anal vein which fuses with the cubitus early on (Fig. 4A). Male tegmina (Fig. 3A) have a simple subcoastal vein; radial vein which runs parallel/subparallel with the media almost throughout the full length of the wing and branches into the first radial about one third of the way through the wing length and terminates as the radial sector; the media runs parallel/subparallel with the radius and has the media posterior split near the middle of the wing and terminates as the media anterior; the cubitus is unbranched; and there is a first anal which fuses with the cubitus early on. Male alae (Fig. 3A) have a costal vein running along the anterior margin; a subcostal vein which runs parallel with the costal vein for the full length; the radial vein is the most unique feature of the alae as it is simple, running parallel to the subcostal vein; the media splits early into the media anterior and posterior which run parallel until the media posterior fuses with the media anterior near the wing margin and they run fused to the apex of the wing; the cubitus is fused with the first anterior anal for ca. half of the length until the first anterior anal splits and runs to the wing margin; the anal veins are split into two groups, the anterior anals and the posterior anals (with seven anterior anals and four or five posterior anals).

Abdomen. Both sexes have variable abdominal shapes, but all forms are broad in the middle with the widest segments V or VI; in both sexes the anterior halves are uniformly broadening to the middle segments, and then the posterior half of the abdomen is variable either with smooth margins giving the abdomen an ovoid appearance, or with the posterior segments gently or strongly undulating giving the abdomen a lobed appearance. Female subgenital plate is short and relatively narrow with the apex slightly reaching onto the anterior margin of the terminal abdominal segment and ending in a fine point; gonapophyses VIII are exceptionally long (with ca. half of their length projecting out from underneath the abdomen) with a uniform width through most of their length; the cerci are relatively flat, marked sparsely with a granular surface, and end in blunt points (Figs 7G, 9D). Males have a long, relatively narrow, triangular vomer which is singularly pronged, hooking up into the paraproct (Fig. 8C).

Figure 9. 

Rakaphyllium exsectum comb. nov. holotype female (NHMUK) A habitus, dorsal B profemora, dorsal C details of head through thorax, dorsal D genitalia, ventral E details of antenna, dorsal F associated data labels G details of head through thorax, dorsolateral. Scale bar only associated with A. Photographs A, F, G taken by Paul Brock (United Kingdom), other photographs taken by first author.

Egg. Egg morphology is not yet known for this rare genus, but with such unusually long gonapophyses to hold the eggs before they are flung away, the eggs must have a unique shape to require such ungainly gonapophyses.

Rakaphyllium meaning “walking leaf”. This generic epithet is a compound of the Latinized name Phyllium the type genus for the family (from Greek φυλλον, -ου (phyllon, -oy) + -um; Poitout 2007), coupled with the prefix raka from the Hiri Motu language of New Guinea which means “to walk” (Chatterton 1975; Dutton and Voorhoeve 1975). We wish to honor the original inhabitants of this area by using the traditional language of Hiri Motu which is one of the official languages of Papua New Guinea and as Papua New Guinea pushed towards sovereignty in 1975, Hiri Motu was seen as a unifying force which was instrumental in the awakening of national pride (Dutton and Voorhoeve 1975). We chose this name because of the amazing camouflage these insects possess, allowing them to appear miraculously as a leaf that simply stands up and walks away when disturbed. This new genus is neuter in gender, following Phyllium.

At present our knowledge of the Rakaphyllium gen. nov. is rather limited due to its rarity, however, interestingly despite it being rarely collected, this genus appears to be somewhat widespread, with records from throughout New Guinea, and even a record from the Aru Islands (Fig. 1A) off the western coast of Papua Province, Indonesia (Fig. 5).

(3 ♀♀, 3 ♂♂, 3 ♂♂ nymphs, 2 ♀♀ nymphs): Holotype (♀): “New Guinea (No 131) VIII. 1910 L. Schultze L.J.; Pulchriphyllium schultzei Giglio-Tos, E. Giglio-Tos, det.; Mossu südl von Germainhuk; Holotypus; DEI Hemimetabola #100122” (SDEI; Fig. 6). See Suppl. material 1 for additional specimens reviewed, their collection data, and depositories.

This rarely encountered species was described from a female from northern New Guinea (Fig. 6), and since its description over a century ago few additional specimens have been collected. At present due to the rarity of material and lack of fresh material for molecular comparison, the male specimens associated with this species are only assumed based on shared morphology (Fig. 8) but have not been confirmed yet.

For female Rakaphyllium schultzei comb. nov., the abdominal shape and profemoral lobes easily differentiate it from Rakaphyllium exsectum comb. nov. Within Rakaphyllium schultzei comb. nov. the profemoral exterior lobe arcs smoothly from end to end without a strong angle (Fig. 6B), but in Rakaphyllium exsectum comb. nov. the exterior lobe is distinctly boxy with a right angle (Fig. 9B). Additionally, abdominal shape appears to be reliable for differentiation, but some abdominal variation has been observed in a few Rakaphyllium schultzei comb. nov. females to include perfectly smooth margins and some with slight undulations on the terminal abdominal segments (although none have been observed to be as extreme as in Rakaphyllium exsectum comb. nov.) therefore abdominal shape may also be useful for differentiation.

At present we are aware of records from several locations throughout New Guinea (both on the Indonesian and Papuan sides) and records from the Aru Islands which visually appear to be this species despite the geographic disconnect from mainland New Guinea (see Fig. 5 and Suppl. material 1 for details of these records). The distribution of this species is vague as a thorough knowledge of morphological variation (due to limited material) and molecular analyses are lacking at present, and therefore our identification assumptions are only based upon general morphology. Unfortunately, without positive confirmation of the males we assume are this species, this distribution may be over expansive (possibly representing several species instead of one), and especially for the observational record from the Aru Islands (Fig. 1A), the specimen could not be examined and although the habitus appears to match the morphology of the mainland New Guinea male the fine details may suggest otherwise if a specimen could be examined.

Holotype (♀): “HOLO-TYPE; Rawlinson Mts. inland Huon Gulf. (Keysser).; Rothschild Bequest B. M. 1939-I.; Rawlinson Mt; Phyllium exsectum ♀ Zompro n. sp. HOLOTYPUS det. O. Zompro X.2000; BMNH(E) #845235” (NHMUK; Fig. 9).

This remarkable looking species is unfortunately only known from the singular holotype female from the little accessed Rawlinson Mountains of the Huon Peninsula (Fig. 5). Even in other phylliid species with prominent lobes on the abdomen, they pale in comparison to the unique habitus of this species, which likely evolved to mimic some strongly lobed host plant. Unfortunately, as this is the only specimen known and it lacks ecological data, nothing is known about a possible plant that the unique shape of this species may have evolved to resemble through advergent evolution.

At present only the female is known for this species, no tentative males have been found which could represent the opposite sex. This species can be differentiated from its single other congenic by the elaborate lobed abdomen (although some schultzei-like females have been observed with slight lobes suggesting morphological plasticity within that species) or more reliably by the profemoral exterior lobe, which in Rakaphyllium exsectum comb. nov. is right angled (Fig. 9B), versus Rakaphyllium schultzei which always has the profemoral exterior lobe roundly arcing from end to end without a distinct angle present (Fig. 6B).

The type locality of Rawlinson Mts. on the Huon Gulf, Morobe Province, Papua New Guinea is the only record known at present (Fig. 5).

Phyllium groesseri Zompro, 1998: 159, herein designated.

Several features link this genus to the Phylliini genera and not the Nanophylliini, such as males with the alae radius split near the wing base (not splitting on the distal half of the wing as in Nanophyllium and Acentetaphyllium gen. nov.), females with tegmina venation similar to Chitoniscus due to the diverging radius and media veins and distinct cubitus anterior and cubitus posterior (cubitus only weakly bifurcate or not bifurcate in Nanophyllium), and both sexes have a singular posteromedial head tubercle (not bilobed as in Nanophyllium or Acentetaphyllium gen. nov.). Due to these notable morphological features, we place this herein described genus within the tribe Phylliini Brunner von Wattenwyl, 1893.

The selected type species for this new genus is Phyllium groesseri Zompro, 1998 (= Vaabonbonphyllium groesseri (Zompro, 1998), comb. nov.) as this was the first species described from within this new genus and this species is well represented by multiple specimens from the geographically isolated Solomon Islands (Fig. 4). Unfortunately, fresh specimens of either species within this genus have not yet been sequenced so hopefully this species can be collected one day and sequenced to add clarity to the higher-level taxonomy.

Within females, the first radial vein (R1) of the tegmina splits from the radius early on, ca. one third of the way between the wing base and the radius to media crossvein (R–M)/the bend in the radial sector (Fig. 4B) versus all other phylliids which have the first radius splitting from the radius midway or more than midway from the wing base to the radius to media cross vein (R–M)/the bend in the radial sector or even after this point.

For males, the media (M) vein of the tegmina runs parallel with the radial sector (Rs) several vein widths away and the media posterior (MP) splits on the distal half of the tegmina, bends immediately and runs parallel with the media anterior (MA) to the wing apex (Fig. 3B) versus in other genera typically the media and radial veins run side by side touching or nearly so, and the posterior branch(es) of the media split on the proximal half or near the middle of the length and run diverging (not parallel) from the media posterior and run to the wing margin, not the apex (Fig. 3A).

The Vaabonbonphyllium gen. nov. are small to medium, with females ca. 70.0 mm in length (in Vaabonbonphyllium groesseri comb. nov.) and with males from ca. 43.0 mm (Vaabonbonphyllium groesseri comb. nov.) to ca. 53.0 mm (in Vaabonbonphyllium rafidahae gen. et sp. nov.).

Antennae. Females have antennae with nine segments with segments I, III, and IV notably broader than the other segments and the terminal antennae segment is as long as the preceding three and a half segments (Fig. 10D). Males have antennae with around 22 segments (in Vaabonbonphyllium groesseri comb. nov. both known male specimens have broken antennae that are missing segments, therefore they only have 19 or 20 segments, but in Vaabonbonphyllium rafidahae gen. et sp. nov. there are 22 segments that are not broken; Fig. 11F) and most segments covered in setae which are longer than the segment is wide.

Head capsule. Males have well-developed ocelli (Figs 11C, 12G) and female lack ocelli (Fig. 10C). Males have compound eyes which are strongly protruding and occupy ca. 2/5 of the head capsule lateral margins (Fig. 11C) versus females which have compound eyes which are notably smaller, only occupying ca. ¼ of the head capsule lateral margins and which to not strongly protrude from the capsule (Fig. 10C). In both sexes the head capsules are marked throughout by weak (Fig. 10C) to distinct granulation (Fig. 12G) which is relatively evenly spaced. The posteromedial head tubercle is singular and distinctly raised from the head capsule (Figs 11E, 12H).

Thorax. The thorax is similar in both sexes with the pronotum notably longer than wide (Figs 10C, 11B, 12G). The mesopleurae are gradually diverging from the anterior to the posterior and are marked with four to six tubercles and some minor granulation throughout (Figs 10C, 11B, 12G). In both sexes the prescutum is ca. 2X wider on the anterior than long with lateral margins marked by three to five nodes/tubercles, and a prescutum surface which can be relatively smooth or slightly granular. When viewed laterally, both sexes have the prescutum anterior rim marked with a raised sagittal spine (Fig. 12H).

Figure 10. 

Vaabonbonphyllium groesseri comb. nov. females A data labels associated with the holotype B holotype, habitus, dorsal C details of head through thorax, dorsal D details of antennae, dorsal E details of genitalia, ventral F front legs, head, and thorax, anterodorsal. Scale bar associated with A and B. A and B photographs by Christian Schmidt (SMTD), other images taken by the first author of specimens from the NHMUK.

Legs. Both sexes have interior tibial lobes on the protibiae which do not span the full length of the shaft (only occupying the proximal ½ to 4/5, but not fully reaching), have two lobes on the protibial exterior (one on the distal end which is smaller and one near the middle which is larger; Figs 10F, 11B, 12C), and the meso-, metatibiae are mostly bare except for the exterior distal ends which can be variable, by either being slightly raised or with small to medium sized lobes present (Fig. 12D, 12E). In both sexes the profemoral interior lobe is broader than the exterior lobe and marked with three or four broad teeth which can be more sharply pointed (Fig. 11B) or dulled (Fig. 12C). Both sexes have the interior mesofemoral lobes slightly broader or about even in width to the exterior lobes, and both lobes can have serration on the distal ½ to 1/3 of the lobe, but the interior lobe serration is more prominent than the exterior lobe. Both sexes have the interior metafemoral lobes several times broader than the exterior lobes and the interior lobes are prominently marked by serration.

Figure 11. 

Vaabonbonphyllium groesseri comb. nov. males A–C, E, F paratype male, (ANIC (CSIRO)), photographs by Bonnie Koopmans (CSIRO) D, G non-type male (NHMUK), photographs by first author A habitus, dorsal (left front leg is physically detached, photoshopped onto the specimen for this figure) B left front leg, dorsal C details of head through thorax, dorsal D habitus, dorsal E details of head through thorax, lateral F associated data labels G genitalia, ventral. Scale bar only associated with A.

Wings. Female tegmina are always long, reaching onto abdominal segments VIII or IX and male tegmina are moderate in length, reaching onto abdominal segment III. Female alae are highly reduced to no more than just a nub and male alae are fully developed in an oval-fan configuration reaching slightly past the apex of the terminal abdominal segment. Female tegmina have a subcostal (Sc) vein which is simple and runs parallel with the wing margin for ca. ⅕ of the wing length before fading; radial vein which diverges steadily away from the media and splits into the first radial vein (R1) which diverges from the radius (R) early on, ca. ⅓ of the way between the wing base and the radius to media crossvein (R–M)/the bend in the radial sector, the primary vein terminates as the radial sector (Rs) on the wing margin just past the middle of the wing length, and as a small radial to medial crossvein (R–M) which does fully connect; the media vein (M) runs side by side the cubitus vein (Cu) and splits into the media anterior (MA) ca. halfway through the wing length and the media posterior (MP) diverges from the cubitus near the distal third of the wing length; the cubitus vein runs along the wing margin until it splits into the cubitus anterior (CuA) and cubitus posterior (CuP) near the distal quarter of the wing; and a first anal vein fuses with the cubitus early on (Fig. 4B). Male tegmina have a simple subcoastal vein; a radial vein (R) which runs parallel with the media (M) throughout the full length of the wing and branches into the first radial (R1) ca. 2/5 of the way through the wing length and the primary vein terminates as the radial sector (Rs) at the wing apex; the media runs parallel with the radius and splits into the media posterior (MP) near the distal 2/5 of the wing, bends immediately and runs parallel with the media anterior (MA) until reaching the wing margin; the cubitus is unbranched and runs along the wing margin; and there is a first anal which fuses with the cubitus ca. ⅓ of the way through the wing length (Fig. 3B). Male alae (Fig. 3B) have a costal vein (C) running along the anterior margin and a subcostal vein (Sc) which runs for nearly the same length parallel with the costal vein; the radial vein (R) is bifurcate when it splits slightly less than 1/3 of the way through the wing length where they gradually diverge, run parallel/subparallel, then converge gradually to the apex where they terminating very near each other but not touching; the media (M) is also bifurcate, splitting about 1/8 of the way through the alae length and these veins run parallel for most of their length until the media posterior (MP) fades near the wing apex and the media anterior (MA) fuses with the radial sector near the wing apex; the cubitus (Cu) is fused with the first anterior anal (1AA) for ca. ¾ of the length until the first anterior anal splits and runs to the wing margin and the cubitus runs singularly to the wing apex; the anal veins are split into two groups, seven anterior anals (1AA–7AA) and five posterior anals (1PA–5PA).

Figure 12. 

Holotype male Vaabonbonphyllium rafidahae gen. et sp. nov., (IMQC) (Coll RC 19-106) photographs by the first author A genitalia, ventral B habitus, dorsal C details of the protibial and profemoral lobes D details of the mesotibial and mesofemoral lobes E details of the metatibial and metafemoral lobes F antenna, dorsal G details of the head and thorax, dorsal H head–thorax, lateral. Scale bar only associated with B.

Abdomen. Abdominal shapes are variable within this genus; females are only known for Vaabonbonphyllium groesseri (Zompro, 1998), comb. nov. and have a boxy abdomen (with segments IV through VI parallel sided) and segment VII strongly lobed and tapering strongly to a narrower segment VIII which is weakly lobed (Fig. 10B). For the males, Vaabonbonphyllium groesseri comb. nov. have segment II converging to the narrowest point of the abdomen, III diverging (giving the abdomen a pinched waist appearance), IV weakly projecting and converging, V diverging, VI strongly diverging and converging giving it a notable lobe, VII appears variable as one male known has this segment converging (Fig. 11A) and the other male has a notable lobe on this segment as well (Fig. 11D), the remainder of the segments converge to the apex. In Vaabonbonphyllium rafidahae gen. et sp. nov. the abdominal shape is simple with all segments similar in width and margins all nearly parallel sided giving it a narrow, blade-like appearance (Fig. 12B).

Female subgenital plate is moderate in length and width, projecting slightly less than halfway under the terminal abdominal segment; gonapophyses VIII are long, broad, and parallel sided for most of their length, with an apex which projects from under the terminal abdominal segment about as much as the cerci do; the cerci are flat and marked sparsely with a granular surface (Fig. 10E). Males have an approximately equilateral triangular vomer which is singularly pronged, hooking up into the paraproct (Figs 11G, 12A).

Egg. Egg morphology is not yet known for this rare genus.

Vaabonbonphyllium meaning “leaf that waits for the night to come”. This generic epithet is a compound of the Latinized name Phyllium the type genus for the family (from Greek φυλλον, -ου (phyllon, -oy) + -um; Poitout 2007), coupled with the prefix vaabonbon from the Teop (Tiop) language phrase which means “wait for the night to come” (Mosel 2019). We wished to honor the original inhabitants of this area by using a traditional language from Bougainville Island, the type locality of the genus. The Teop language is little known and considered a threatened language with as few as 6,000 speakers left in the autonomous region of Bougainville Island (Mosel 2014). We chose this name as these insects are exceptionally camouflaged and nocturnal, holding still during the day, thus hiding from view and only venturing out at night, making them exceptionally rare in collections and little known. This new genus is neuter in gender, following Phyllium.

At present Vaabonbonphyllium gen. nov. is primarily known from the Solomon Islands (Vaabonbonphyllium groesseri comb. nov.) and a single record from Western Highlands Province, Papua New Guinea (Vaabonbonphyllium rafidahae gen. et sp. nov.; Fig. 5).

(15 ♀♀, 2 ♂♂, 3 ♀♀ nymph, 2 ♂♂ nymph, 2 unsexed nymph): Holotype (♀): “Buin I. Bougainville Arch. Salomons; 1941 4; Chitoniscus lobiventris Blanch. K. Gunther det.; lobiventris Blanch.; Chitoniscus Stal; Phyllium groesseri Zompro 1997 HOLOTYPUS det. O. Zompro X. 1996; HOLOTYPUS; Phyllium groesseri ♀ Zompro n. sp. HOLOTYPUS det. O.Zompro V.1998” (SMTD; Fig. 10A). See Suppl. material 1 for additional specimens reviewed, their collection data, and depositories.

At present this is the only known phylliid species from the Solomon Islands, although the phylliid knowledge of these islands is rather limited as even this species is rarely collected so if additional species were present but not yet formally described it would not be surprising.

Only two males are known to us at the present (from within the NHMUK and CSIRO collections) both of which are notably damaged (Fig. 11). These males have not yet been positively confirmed as the male Vaabonbonphyllium groesseri comb. nov. via molecular comparison or from captive breeding but based upon shared morphology between the sexes and the current knowledge that only one species of phylliid is present on the Solomon Islands it is safe to assume these are the male for this species.

This species was originally described simply as a Phyllium species (not placed within a subgenus) and was placed within the Pulchriphyllium subgenus by Größer (2001) due to the presence of exterior lobes of the tibiae. During the intrageneric revision of Hennemann et al. (2009) this species was placed within the frondosum species group to which this species has a strong general resemblance (when reviewing the size, abdominal shape, and female wing venation). In 2020 the frondosum species group was found to represent the unknown female Nanophyllium; however, this species was not transferred to the Nanophyllium but was instead moved to the Pulchriphyllium subgenus schultzei species group based upon the bilobed protibiae (Cumming et al. 2020a). Now thanks to a full review of morphology and review of multiple specimens, the fine details which differentiate this clade from others allows its formal recognition as a unique genus.

Female Vaabonbonphyllium groesseri comb. nov. are the only females known for this genus, so differentiation from congenerics is not possible until the unknown female Vaabonbonphyllium rafidahae gen. et sp. nov. is located and described. Male Vaabonbonphyllium groesseri comb. nov. can be differentiated from Vaabonbonphyllium rafidahae gen. et sp. nov. based upon the abdominal shape as Vaabonbonphyllium groesseri comb. nov. has notably differing abdominal segment widths, with the middle segments many times wider than the anterior and distal segments (Fig. 11A, D) versus Vaabonbonphyllium rafidahae gen. et sp. nov. which has an abdomen with all segments relatively even in width giving it a smooth margined appearance (Fig. 12B). Also, the lobes of the pro- and mesofemora allow differentiation as Vaabonbonphyllium groesseri comb. nov. has the profemoral exterior lobe highly reduced (Fig. 11B) and the interior mesofemoral lobe is notably broader than the mesofemoral exterior lobe versus Vaabonbonphyllium rafidahae gen. et sp. nov. which has a profemoral exterior lobe which is slightly wider than the profemoral shaft width (Fig. 12C) and the mesofemoral interior and exterior lobes which are approximately the same width (Fig. 12E). Additionally, the prosternum allows for differentiation as in Vaabonbonphyllium groesseri comb. nov. the prosternum is relatively smooth, lacking a distinct protrusion (Fig. 11E) versus Vaabonbonphyllium rafidahae gen. et sp. nov. which has a distinct protrusion with a granular surface (Fig. 12H).

This species was originally only recorded from Bougainville Island, but since then additional records have been recovered from numerous collections representing previously unknown localities (see Suppl. material 1 for a full list of known specimens and their data). Additional islands recorded are (north to south): Kolombangara, Nggela, Guadalcanal, and Small Malaita (Fig. 5). With how rarely this species is collected, and from the wide range that it can be found from throughout the Solomon Islands, it is likely that this species can be found on several of the other larger islands as well, but just has not been formally recorded yet.

Holotype ♂: “Papua New Guinea: Western Highlands Province (Highlands Region). Mt. Hagen Dist. (New Guinea Island) Mt. Hagen, along Highlands Hwy. (5 51'46.20"S, 144 13'30.75"E) Elev. 1730 m 23-VI-1989 Coll. L D. Munsey. (Coll RC 19-106)”. Deposited in the Montreal Insectarium (IMQC).

This mainland New Guinea species is presently only known from the singular holotype male (Fig. 12). This species is tentatively placed within Vaabonbonphyllium gen. nov. due to the protibial exterior being bilobed (a feature only known from Vaabonbonphyllium gen. nov. and Rakaphyllium gen. nov.) but the wing venation of this male differs notably from Rakaphyllium gen. nov. and therefore, this species appears to be related to Vaabonbonphyllium groesseri comb. nov. based upon our poor knowledge of male morphology within that species. Unfortunately, for the Vaabonbonphyllium gen. nov. only three males are known (two Vaabonbonphyllium groesseri comb. nov. and this herein described species) so our knowledge on the intrageneric variability is severely limited and until specimens of both species can be sequenced a confident placement cannot be made beyond this initial morphology-based review.

Females unknown. Male Vaabonbonphyllium rafidahae gen. et sp. nov. can be differentiated from Vaabonbonphyllium groesseri comb. nov. based upon the abdominal shape as Vaabonbonphyllium rafidahae gen. et sp. nov. has an abdomen with all segments relatively even in width giving it a smooth margined appearance (Fig. 12B), versus Vaabonbonphyllium groesseri comb. nov. which have notably differing abdominal segment widths, with the middle segments many times wider than the anterior and distal segments (Fig. 11A, D). The lobes of the femora allow differentiation as Vaabonbonphyllium rafidahae gen. et sp. nov. has a profemoral exterior lobe which is slightly wider than the profemoral shaft width (Fig. 12C) and mesofemoral interior and exterior lobes which are approximately the same width (Fig. 12D) versus Vaabonbonphyllium groesseri comb. nov. which has the profemoral exterior lobe highly reduced (Fig. 11B) and the interior mesofemoral lobe notably broader than the mesofemoral exterior lobe. Additionally, the prosternum allows for differentiation as in Vaabonbonphyllium rafidahae gen. et sp. nov. there is a distinct protrusion (Fig. 12H), and in Vaabonbonphyllium groesseri comb. nov. the prosternum is relatively smooth, lacking a distinct protrusion (Fig. 11E).

Currently only known from the type locality of Mt. Hagen, Western Highlands Province, Papua New Guinea (Fig. 5).

Male. Coloration. The coloration description is based on the single dried holotype specimen which appears to have been collected in alcohol which turns phylliids yellow (Fig. 12). In life the specimen likely was green, but color variation is known within phylliids so a definitive coloration description cannot be given at this time for possible living coloration. The overall coloration of the holotype specimen is straw yellow throughout with the only feature which is distinctly different in color being the compound eyes which are a dark red (Fig. 12G). Abdominal segment V has a set of eye spots which are ovular and slightly darker in color than the remainder of the abdomen.

Morphology. Head. Head capsule approximately as long as wide, with a vertex that is marked throughout by nodes which are relatively evenly spaced and of uniform size (Fig. 12G). The posteromedial tubercle is singularly pointed, large, and notably raised above the head capsule, many times larger than any of the capsule granules (Fig. 12H). Frontal convexity stout and bluntly pointed with several short setae on the apex. Compound eyes large and bulbous, occupying ca. ⅖ of the head capsule lateral margins (Fig. 12G). Between and slightly posterior to the compound eyes are three ocelli that are well-developed (Fig. 12G). Antennal fields are slightly wider than and approximately as long as the scapus.

Antennae. Antennae (including the scapus and pedicellus) consist of 22 segments. The scapus and pedicellus are bare, segments III through XIX have dark setae which are mostly around two times longer than the segments are wide and are not densely packed, the terminal three segments are covered by fine, densely packed, transparent setae (Fig. 12F).

Thorax. Pronotum is slightly longer than the greatest width, with anterior margin concave and lateral margins that are straight and converge to a gently convex posterior margin that is ca. ⅗ the width of the anterior rim (Fig. 12G). Anterior and lateral margins of the pronotum have distinct rims and the posterior margin lacks a rim (Fig. 12G). Face of the pronotum is notably lumpy, marked with a distinct sagittal furrow on the anterior half, and a distinct pit in the center (Fig. 12G). Prosternum has a notably granulose surface with a distinctly projecting area in the center and an additional smaller projection near the posterior margin, which is smaller than the central projection, both projections are marked with granulation (Fig. 12H). Mesosternum has a notably granulose surface similarly marked as the prosternum with uniform spacing between granules which are all nearly even in size. Metasternum not as heavily marked with granulation, but instead has slightly wrinkled anterior areas and a nearly smooth central and posterior area. Prescutum on the anterior is wider than long, with lateral margins converging uniformly to the posterior which is ca. ⅗ as wide as the anterior rim width (Fig. 12G). Lateral rims with a lumpy texture due to irregularly sized tubercles/nodes (ca. nine distinct but some weakly formed) with the largest near the anterior margin (Fig. 12G). The surface of the prescutum is marked with a few nodes but it is mostly smooth as there is no distinct sagittal crest, instead the anterior margin is the only distinct feature as the rim is prominently formed with a granular surface and raised into a distinct sagittal point (Fig. 12G, H). Mesopleura narrow, only slightly diverging from the anterior to the posterior giving the mesosternum a rectangular appearance when viewed ventrally. Lateral margin with one notably larger tubercle on the anterior margin with three setae protruding from it followed by four or five moderately sized tubercles which in some cases have a seta or two protruding from them, with these tubercles evenly spaced throughout the length (Fig. 12G). Face of the mesopleura with a weakly lumpy surface (most areas are relatively smooth) and marked with two weakly formed pits near the middle of the length (Fig. 12G, H).

Wings. Tegmina moderate length, extending ca. ⅓ of the way onto abdominal segment III (Fig. 3B). Tegmina wing venation: the subcosta (Sc) is the first vein, it runs relatively straight and terminates slightly less than ½ of the way through the overall tegmina length. The radius (R) spans the entire length of the tegmina with the first radius (R1) branching slightly greater than ⅓ of the way through the wing length and running to the wing margin where it terminates ca. ⅗ of the way through the wing length, and then the radial sector (Rs) runs straight from the branching point to the wing apex. The media (M) spans the entire length of the tegmina, terminating at the wing apex as the media anterior (MA). The media posterior (MP) branches ca. ⅔ of the way through the wing length, immediately bends and runs parallel with the media anterior until it terminates near the wing apex. The cubitus (Cu) runs through the wing surface angled towards the margin which it meets ca. ⅓ of the way through the tegmina length at the same location where the first anal (1A) vein fuses with it and the cubitus runs along the wing margin until it fades near the point where the media posterior terminates near the wing margin. Alae well-developed in an oval fan configuration, reaching beyond the apex of the abdominal segments, instead reaching to the apex of the cerci. Alae wing venation (Fig. 3B): the costa (C) is present along the entire forewing margin. The subcosta (Sc) runs along the costa throughout almost its entire length. The radius (R) branches ca. ¼ of the way through the wing length into the first radius (R1) and radial sector (Rs) which run for half of their length slightly diverging, then parallel/subparallel until the wing apex where thy converge and terminate very near each other at the wing apex. The media (M) branches early, ca. ⅛ of the way through the wing length into the media anterior (MA) and the media posterior (MP) which run parallel with each other until the media anterior bends towards the radial sector and fuses with it near the wing apex, and the media posterior bends towards the media anterior but fades before fusing with it. The cubitus (Cu) runs unbranched and terminates at the wing apex. Of the anterior anal veins, the first anterior anal (1AA) fuses with the cubitus near the point where the media branches into the media anterior and media posterior and then the first anterior anal branches from the cubitus ca. ¼ 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–5PA) share a common origin separate from the anterior anal veins and run unbranched to the wing margin.

Abdomen. The general shape of the abdomen is long and slender with margins that are relatively parallel and segments which are nearly even in width, varying little from one to the next (Fig. 12B). Abdominal segments II through IV diverge only slightly, segments V and VI are parallel sided, and VII through the apex converge to the apex.

Genitalia. Poculum roundly rectangular in shape with anterior and lateral margins relatively straight and the posterior margin only slightly passing onto the terminal abdominal segment as it gently arcs towards the abdomen apex. Cerci long and slender (approximately ⅖ as wide as long), with slightly more than ½ of their length extending out from under the anal abdominal segment. The cerci are relatively flat and covered in a granulose surface with numerous short setae evenly spaced throughout (Fig. 12A). The vomer general shape is an equilateral triangle with straight sides evenly converging to the apex, which is armed with a singular upward turning hook that is broad and notably darker than the rest of the vomer (Fig. 12A).

Legs. Profemoral exterior lobe arcing gently with a width only slightly greater than the greatest width of the profemoral shaft with the margin marked slightly with granulation (Fig. 12D). Profemoral interior lobe roundly right angled with a greatest width ca. three times that of the exterior lobe and the distal margin is marked with three or four stout and blunted teeth with varying spacing between them (Fig. 12D). Mesofemoral exterior lobe arcs unevenly end to end with the widest point on the distal ⅓, and the greatest width slightly wider than the mesofemoral shaft or mesofemoral exterior lobe widths (Fig. 12D). The mesofemoral interior lobe is marked by four dulled teeth on the distal ⅓ of the lobe (Fig. 12D). The mesofemoral exterior lobe is very similar to the interior lobe in terms of shape, size, and dentition, with the only notable difference being that at its greatest width it is slightly thinner than the interior lobe (Fig. 12D). Metafemoral exterior lobe has a slightly granular margin but lacks dentition, and the margin is straight running along the metafemoral shaft (Fig. 12E). Metafemoral interior lobe is unevenly weighted on the shaft with the majority occupying the distal ⅔ of the length with the distal ⅓ marked with three or four teeth, with the proximal most rather dulled and the distal most teeth more finely pointed (Fig. 12E). Protibiae with two exterior lobes, one small lobe on the distal end and a larger lobe (ca. three times as large as the small distal lobe) slightly proximal to halfway through the length (Fig. 12D). Protibiae with a single interior lobe which is the shape of a rounded scalene triangle which occupies the proximal ¾ of the protibial length and at its maximum width is ca. two and a half times the width of the protibial shaft width (Fig. 12D). Mesotibiae and metatibiae lack interior lobes but both have singular small exterior lobes situated on the distal end, with the mesotibial exterior lobe only weakly formed (not as wide as the shaft width: Fig. 12D) and the metatibial exterior lobe more distinctly formed (ca. as wide as the metatibial shaft width: Fig. 12E).

Length of body (including cerci and head, excluding antennae) 52.8, length/width of head 3.1/3.0, antennae 21.5, pronotum 2.5, mesonotum 2.3, length of tegmina 17.2, length of alae 45.6, greatest width of abdomen 10.5, profemora 7.7, mesofemora 7.4, metafemora 7.4, protibiae 4.8, mesotibiae 4.8, metatibiae 6.8.

The specific epithet of the new species is selected to honor the wife of Larry D. Munsey, the individual who collected the specimen upon which the new species description is based. Rafidah, a native of West Malaysia, and Larry, a native Californian, live in Borneo where they jointly have been studying the megadiverse cerambycid fauna of the island for the past 17 years.

Phyllium brevipennis Größer, 1992: 164, herein designated.

This clade, based upon molecular analyses, was recovered as sister to the Nanophyllium a placement which is supported by the linked morphological features of the wider than long prescutum, two lobed posteromedial tubercle, and similar antennae morphology (Bank et al. 2021). Therefore, this new genus is included within the tribe Nanophylliini Zompro & Größer, 2003 along with its sister genus Nanophyllium.

The selected type species for this new genus is Phyllium brevipennis Größer, 1992 (= Acentetaphyllium brevipenne (Größer, 1992), comb. nov.) as this was the first species described from this group and is represented by a good condition holotype female with detailed collection data so little confusion surrounds the type species. Additionally, this species has been sequenced and included in molecular phylogenies (Bank et al. 2021) while the other species are still lacking.

Again, the uniqueness of this clade was recognized by Hennemann et al. (2009), with their designation of the brevipenne species group, and they even recognized this clade as closely related to the Nanophyllium (which as the time were still considered the Phyllium (Pulchriphyllium) frondosum species group, not the opposite sex of the Nanophyllium yet which would not be until a decade later in Cumming et al. 2020a).

Within the phylliid-wide phylogeny of Bank et al. (2021) Acentetaphyllium brevipenne comb. nov. was recovered as sister to all sampled Nanophyllium species with high support. Despite the drastic morphological differences between this new genus (Fig. 13A) and the Nanophyllium sensu stricto (Fig. 13B), there are still several morphological features which help to link these two genera. Most notably the two lobed posteromedial tubercle of the head capsule and the similar mesothorax morphology, which is notably wider than long, immediately link these two genera.

Figure 13. 

Dorsal habitus of adult Acentetaphyllium gen. nov. and Nanophyllium to help illustrate the morphological differences/similarities between these two genera. Images scaled to relative size; scale bar associated with all images A Acentetaphyllium brevipenne female (Coll TM) B Nanophyllium chitoniscoides (Größer, 1992) female (Coll TM) C Acentetaphyllium stellae comb. nov. holotype male (SDNHM) D Nanophyllium rentzi holotype male (NHMUK) A, B photographs by Rene Limoges (IMQC) C photograph by first author D photograph by Paul Brock (NHMUK).

Little is known about the Acentetaphyllium gen. nov. ecology except for labels included with two females from within the BPBM collection noting their host plant as “Eucalyptus” (Fig. 14). Several other genera/species are known to accept Eucalyptus spp. as a host within captivity (pers. comm. Tomas Stijnts (Belgium; Flanders)). Additionally, these BPBM specimens also note an elevation of collection at 750 meters, whereas a specimen in the Coll SLT from Wau, Morobe Province was collected at 1,200 meters, suggesting this species is not restricted to a limited elevation range.

Figure 14. 

Examples of abdominal variation observed within Acentetaphyllium brevipenne comb. nov. adult females from within the BPBM collection, dorsal habitus, with their associated data labels to the right of each specimen. Photographs by Miho Maeda, Jerilynn Chun, and James Boone, 2020 (BPBM) A brown form with distinct eye spots on three of the abdominal segments, and a notably lobed abdomen B green form female with eye spots on a single abdominal segment and an abdominal shape which has smoother margins.

Within the Nanophyllium the association of opposite sexes has always been problematic due to limited material and notable age of many specimens not allowing molecular comparison. Only recently was the confusion surrounding the “missing” Nanophyllium females for the most part been resolved (Cumming et al. 2020a). However, even within that work it was recognized that there were two distinct clades known for the males (noted within Cumming et al. 2020a as the pygmaeum species group and the stellae species group, see therein figures and side-by-side morphological comparisons) but females were only confirmed for the pygmaeum species group. The stellae species group was recognized as having numerous morphological similarities and differences to the pygmaeum species group but no tentative female had been linked with clarity. Now with the molecular results of Bank et al. (2021) recovering brevipenne as sister to the Nanophyllium, the males of the stellae species group can with higher confidence be associated with these morphologically unique females (see Fig. 13 for a side by side of these two clades). Although not yet confirmed by breeding or from molecular comparison, the numerous shared morphological features of the males of the stellae species group and brevipenne strongly suggest these are opposite sexes of the same clade (Fig. 13A, C).

For detailed discussion and figures of the non “brevipenne” Acentetaphyllium gen. nov. species which were originally placed within Nanophyllium, see the recent work of Cumming et al. (2020a) as these species (Acentetaphyllium larssoni (Cumming, 2017), comb. nov.; Acentetaphyllium miyashitai (Cumming et al., 2020), comb. nov.; and Acentetaphyllium stellae (Cumming, 2016), comb. nov.) are not discussed below as no additional information can be added at this time to supplement the recent publication and their original descriptions.

Features which link these two genera are their antennae morphology (in females), thorax shape and spination (both sexes), and genitalia (both sexes). There are several notable differences between the genera, for female Acentetaphyllium gen. nov. the tegmina are highly sclerotized and rudimentary (Fig. 15D), at most reaching onto the anterior of abdominal segment II leaving the dorsal of the abdomen fully exposed versus tegmina which are fully developed and covering most of the abdomen in Nanophyllium. For male Acentetaphyllium gen. nov. the shape of the femoral lobes differentiates them as the profemoral interior lobes are rounded without a sharp angle and the mesofemoral interior lobes have a large rounded triangular lobe, reaching from end to end and lacking prominent spination (versus Nanophyllium which have profemoral interior lobes which are angular and mesofemoral interior lobes which are heavily weighted towards the distal end and are marked by distinct serrate teeth on the distal half). Also, the alae of the male Acentetaphyllium gen. nov. allow differentiation as the media anterior and the media posterior do not fuse, instead they both run to the wing margin, and the cubitus after splitting from the first anterior anal fuses with the media posterior near the wing margin and they run fused to the wing apex (versus Nanophyllium which have the media posterior fusing to the media anterior and then they run fused to the wing margin without fusing with other veins (see Cumming et al. 2020a: figs. 10–13 for side by side comparisons)).

Figure 15. 

Acentetaphyllium brevipenne comb. nov. adult female details A, B, E from Coll SLT C, D from Coll TM Photographs by Rene Limoges (IMQC) A details of head, dorsal B details of antennae, dorsal C details of protibiae and profemora, dorsal D details of head through thorax, dorsal E details of genitalia, ventral.

For females the tegmina are highly sclerotized and rudimentary, no longer than the anterior of abdominal segment II (Fig. 14), whereas all other phylliids have well-developed tegmina which at least reach halfway onto the abdomen and in most cases cover most of the abdomen. For males the alae venation is autapomorphic as the media anterior and the media posterior do not fuse, instead the media anterior runs alone to the wing margin and the cubitus (after splitting from the first anterior anal) fuses with the media posterior near the wing margin and then they run fused to the wing apex as one (see Cumming et al. 2020a: fig. 13B for an illustration of this venation pattern).

The Acentetaphyllium gen. nov. have females that for phylliids are average length (ca. 80 mm) and males which are on the smaller end (ca. 40 mm), which gives a male to female length ratio of ca. 1.0:2.0 between the sexes for this genus which is the most extreme known in the family (Fig. 13A, C) as typically a male to female ratio of 1.0:1.2 to 1.0:1.6 is common. Interestingly for Acentetaphyllium gen. nov. coloration, males are only known to have brown forms but within females brown, green, and yellow are all known to exist. Hopefully as more males are located it will become apparent if males are exclusively brown or also come in different color forms.

Antennae. Females have antennae with nine segments with segments IV through VIII which are uniform in length, disk-like, marked with granulation, and the terminal segment is stout and densely covered in setae (Fig. 15B). Males have antennae with 21 to 22 segments which are relatively flat, and most segments are covered by setae which are longer than the segment is wide.

Head capsule. Males have well-developed ocelli and females lack ocelli. Males have compound eyes which are strongly protruding and occupy ca. 2/5 of the head capsule lateral margins versus females which have compound eyes which are notably smaller, only occupying ca. ¼ of the head capsule lateral margins and which to not strongly protrude from the capsule (Fig. 15A). Both sexes have head capsules which are marked throughout by distinct but somewhat irregularly sized granulation which is relatively evenly spaced (Fig. 15A).

Thorax. In both sexes the prescutum can be two to three times wider on the anterior than long with lateral margins that are granular, and a prescutum surface which is only slightly granular, lacking a prominent anterior rim or sagittal crest (Fig. 15D). In both sexes the mesopleurae begin notably wider than the prescutum anterior rim and weakly diverge gradually and can be marked with a single anterior tubercle or by as many as nine warty tubercles (Fig. 15D).

Legs. Both sexes have interior protibial lobes which span the full length in a broadly rounded triangle, lack lobes on the protibial exterior, and the meso-, metatibiae are simple, lacking both interior and exterior lobes. In both sexes the profemoral interior lobe is notably smaller than the exterior lobe and is rounded without a strong angle and marked by two to four dulled weakly formed teeth (Fig. 15C). The profemoral exterior lobe is somewhat variable in males (being roundly obtuse or with a slight recurve giving the lobe a boxier appearance) but in females it appears to be more stable as it is notably recurving posteriorly and marked with a strongly serrate posterior margin and a smoothly arcing anterior margin (Fig. 15C). Both sexes have the interior meso-, and metafemoral lobes ca. 1.5 to 2.0 times as broad as the exterior lobes, with both arcing smoothly, giving the meso-, and metafemora smooth, pointed leaflet-like appearances which tuck in nicely to the anterior of the abdomen helping to give the habitus a full ovoid appearance (Fig. 14).

Wings. In both sexes the tegmina are highly reduced and heavily sclerotized, with a maximum length of reaching the anterior margin of abdominal segment II, but mostly shorter, only reaching the metanotum (Fig. 15D). Females lack alae but in males the alae are fully developed, reaching onto abdominal segment IX or X. Male alae have a costal vein running along the anterior margin; a subcostal vein which runs alongside the costal vein for most of its length until it fuses with the costal vein near the wing margin; the radial vein is bifurcate, splitting into the first radial and radial sector near the distal third of the alae and these run to the wing margin without fusing to other veins or each other; the media splits early on into the media anterior and media posterior which run parallel to the wing apex, with the media anterior arriving alone, but the media posterior has the cubitus fuse to it near the distal 1/8 of the wing apex; the cubitus is fused with the first anterior anal for ca. half of the wing length until the first anterior anal splits and runs to the wing margin and the cubitus fuses with the media posterior; the anal veins are split into two groups, the anterior anals and the posterior anals (with seven anterior anals and six posterior anals).

Abdomen. Abdominal shapes are somewhat variable; females are often perfectly spade-shaped (Fig. 14B) but in some forms the abdominal segments can have slightly undulating margins, giving them a wavy appearance (Fig. 14A); males have similar appearances as they can be smoothly spade-shaped (Fig. 14C) or can have abdomen which are strongly undulating (see Acentetaphyllium larssoni comb. nov., for an example of a strongly undulating abdomen in males). Female subgenital plate is short and stout with the apex only reaching halfway onto abdominal segment IX and ending in a fine point; gonapophyses VIII are long and slender, but not quite reaching to the apex of the terminal abdominal segment; gonapophyses IX are located to either side of the gonapophyses VIII and have broad bases for ca. ½ their length which taper quickly to slender points; the cerci are relatively flat and rounded, with a slightly wrinkled surface (Fig. 15E). Males have a narrow, triangular vomer which is ca. two times longer than the greatest width and terminates in a single hook. Male cerci are notably marked with granulation and distinct setae along the margins.

Egg. Egg morphology is not yet known for this rare genus.

Acentetaphyllium meaning “flawless leaf”. This generic epithet is a compound of the Latinized name Phyllium the type genus for the family (from Greek φυλλον, -ου (phyllon, -oy) + -um; Poitout 2007), coupled with the Latin prefix acenteta meaning “flawless”. One of the most remarkable features of this genus is how the abdomen and legs are shaped so that when the legs are at rest the habitus is a perfect oval, giving this genus a flawless leaf-like shape (Fig. 14B). This new genus is neuter in gender, following Phyllium.

At present only known from a handful of records as this appears to be a rarely collected genus. However, it appears as though this genus is found throughout the island of New Guinea as an undescribed Acentetaphyllium gen. nov. is known to us from Fak Fak, Indonesia, there are multiple records from northern New Guinea from the Cyclops Mountains (Indonesia side) and Bewani Mountains (Papua New Guinea side), and multiple records from Morobe Province, Papua New Guinea. See Suppl. material 1 for details on known records.