Research Article |
Corresponding author: Royce T. Cumming ( roycecumming@gmail.com ) Academic editor: Sven Bradler
© 2021 Royce T. Cumming, Stephane Le Tirant, Thies H. Büscher.
This is an open access article distributed under the terms of the CC0 Public Domain Dedication.
Citation:
Cumming RT, Tirant SL, Büscher TH (2021) Resolving a century-old case of generic mistaken identity: polyphyly of Chitoniscus sensu lato resolved with the description of the endemic New Caledonia Trolicaphyllium gen. nov. (Phasmatodea, Phylliidae). ZooKeys 1055: 1-41. https://doi.org/10.3897/zookeys.1055.66796
|
With every molecular review involving Chitoniscus Stål, 1875 sensu lato samples from Fiji and New Caledonia revealing polyphyly, the morphology from these two distinct clades was extensively reviewed. Morphological results agree with all previously published molecular studies and therefore Trolicaphyllium gen. nov. is erected to accommodate the former Chitoniscus sensu lato species restricted to New Caledonia, leaving the type species Chitoniscus lobiventris (Blanchard, 1853) and all other Fijian species within Chitoniscus sensu stricto. Erection of this new genus for the New Caledonian species warrants the following new combinations: Trolicaphyllium brachysoma (Sharp, 1898), comb. nov., Trolicaphyllium erosus (Redtenbachher, 1906), comb. nov., and Trolicaphyllium sarrameaense (Größer, 2008a), comb. nov. Morphological details of the female, male, freshly hatched nymph, and egg are illustrated and discussed alongside the Chitoniscus sensu stricto in order to differentiate these two clades which have been mistaken as one for decades.
Camouflage, Drehu, Grande Terre, Ile de Bélep, L’Île-des-Pins, Lifou, Lifu, Maré, mimicry, new combination, Phasmida, Tiga, walking leaf
Phasmatodea, the stick and leaf insects, are a group of rather large herbivorous insects that is well known for their diversity of morphological adaptations facilitating imitation of parts of plants (
The following collection acronyms are used.
Coll DG Private collection of Detlef Größer, Berlin, Germany;
Coll RC Private collection of Royce T. Cumming, California, USA;
Coll SLT Private collection of Stéphane Le Tirant, Québec, Canada;
IAC Institut Agronomique néo-Calédonien, La Foa, Nouvelle-Calédonie;
IMQC Insectarium de Montréal, Montréal, Québec, Canada;
MHN Muséum d’Histoire Naturelle, Geneva, Switzerland;
Photographs of specimens deposited within the IMQC collection and Coll SLT were taken by René Limoges using a Nikon D850 DSLR camera (Nikon Corporation, Tokyo, Japan) with Nikon Micro-Nikkor 200mm f/4 lens on Manfrotto 454 micrometric positioning sliding plate (Manfrotto, Casolla, Italy). Lighting was provided by two Nikon SB-25 flash units with a Cameron Digital diffusion photo box (Henry’s, Vancouver, Canada).
Photographs from
Photographs of specimens within the first authors collection (Coll RC) were taken by RC using a Canon 5D Mark II and a MP-E 65mm macro lens and stacked using Zerene Stacker (Zerene Systems LLC, Richland, USA). The eggs of Trolicaphyllium sarrameaense comb. nov. were photographed using a Nikon D3500 camera (Nikon Corporation, Tokyo, Japan) mounted on a Wild M3C stereomicroscope (Wild, Heerbrugg, Switzerland) and manually stacked. Adobe Photoshop Elements 13 (Adobe Inc., San Jose, USA) was used as post processing software.
Egg orientation terminology follows
Micrographs were obtained from dried samples, which were sputter coated with 10 nm gold–palladium. Overview images were obtained with a Hitachi TM3000 SEM (Hitachi High-technologies Corp., Tokyo, Japan) at 15 kV acceleration voltage using a rotatable specimen holder (Pohl, 2010). Detailed micrographs were obtained in the SEM Hitachi S4800 (Hitachi High-technologies Corp., Tokyo, Japan) at an acceleration voltage of 5 kV. Adobe Photoshop Elements 13 (Adobe Inc., San Jose, USA) was used as post processing software.
Morphological abbreviations (listed morphologically anterior to posterior)
a1–a9> antennomeres 1–9
st stridulatory file
sr stridulatory ridge
ar arolium
cl claw
eu1–5 euplantula 1–5
ta1–5 tarsomere 1–5
ri median ridgelike expansion
C costa
Sc subcosta
R radius
R1 radius 1
R2 radius 2
Rs radial sector
R–M radius to media crossvein
M media
MA media anterior
MP media posterior
MP1 first media posterior
MP2 second media posterior
Cu+MA+MP fused cubitus, media anterior, and media posterior
Cu cubitus
CuA cubitus anterior
CuP cubitus posterior
Cu+1AA cubitus and first anterior anal
1A first anal
1AA–7AA first–seventh anterior anal
1PA–5PA first–fifth posterior anal
The results of our morphological and biogeographic examination of Chitoniscus sensu lato strongly support the presence of at least two independent lineages, thereby necessitating the recognition of a new genus, Trolicaphyllium gen. nov., to accommodate the endemic New Caledonia taxa. The erection of this novel genus is supported by the identification of autapomorphic morphological features (discussed below) as well as years of molecular results recovering Chitoniscus sensu lato as polyphyletic (e.g.,
Phyllium brachysoma Sharp, 1898.
Due to the general phylliid morphological features, and the consistent recovery of this clade nested within the greater phylliids in molecular studies, we herein place this genus within the tribe Phylliini Brunner von Wattenwyl, 1893.
The selected type species for this new genus is Phyllium brachysoma Sharp, 1898 (= Trolicaphyllium brachysoma (Sharp, 1898), comb. nov.) which was the first species described and is represented by two female syntype specimens collected on Lifou Island (Fig.
This new genus has been confused for decades with the similarly sized Chitoniscus Stål, 1875 sensu stricto from nearby Fiji due to their superficial similarities. All molecular phylogenies which have included both Fijian and New Caledonian samples have recovered these as polyphyletic (e.g.,
Little is presently known about the Trolicaphyllium gen. nov. ecology at the moment, as the only host plant records we have seen to date are from a Ficus sp. (recorded by Thierry Salesne; New Caledonia) and Syzygium cumini (recorded by Sylvie Cazeres (IAC); Fig.
Features which liken these two genera together are their short length (ca. 40 to 60 mm) and broad bodies. Both genera have species which have smooth/tapered abdominal shapes or can be strongly lobed (within both males and females). The eggs of both species are small and lack pinnae therefore they superficially resemble each other.
However, when the finer details of these two genera are reviewed, the differences between them are significant (Table
Summary of morphological features for differentiating Chitoniscus Stål, 1875 sensu stricto from Trolicaphyllium gen. nov. as these two have been mistakenly associated for more than a century. Listed morphologically from the anterior to posterior.
Female | Trolicaphyllium gen. nov. | Chitoniscus Stål, 1875 sensu stricto |
---|---|---|
Antennae | Antennomeres III, VIII, and IX widened, broader than the antennomeres between; Fig. |
Antennomeres III, VIII, and IX not widened, with similar diameter as the antennomeres between; Fig. |
Antennae: third antennomere | Broadened, sr not shifted anteriorly, sf meeting sr in anterior third; Fig. |
Not broadened, sr shifted anteriorly, sf meets sr at half of its length; Fig. |
Antennae: third antennomere (sf teeth) | Teeth with a smooth apex; Fig. |
Teeth with a weakly bilobed apex; Fig. |
Antennae: first antennomere | Dorsal surface without notable expansion, flush with anterior of the segment; Fig. |
Dorsal surface with expansion projecting anteriorly alongside the 2nd antennomere, projecting beyond the anterior end of the segment; Fig. |
Protibial interior lobe | Always spanning the full length of the protibial shaft | Either absent or even in the most well-developed forms only on the proximal half, never fully spanning |
Prescutum anterior rim sagittal spine | Spine and rim distinct, but not large; Fig. |
Spine and rim prominent; Fig. |
Ventral coxae color | Green, similar shade as the remainder of the insect; Fig. |
Sky blue in color; Fig. |
Tegmina: R and M | R runs parallel with M until the split of Rs, at which point Rs bends away distinctly; Fig. |
R diverges steadily from M for the full length, therefore the split of the Rs is not a significant bend; Fig. |
Tegmina: R–M crossvein | R–M crossvein does not fade, but fully reaches to and connects with M; Fig. |
R–M crossvein thins and fades before reaching M; Fig. |
Terminal abdominal segment | Broad; almost two times as wide as long; Fig. |
Narrow; approximately as long as the greatest width; Fig. |
Cerci texture | Weakly granular/smooth; Fig. |
Heavily granular; rough textured; Fig. |
Tarsus | Euplantula 2 and 3 with ridgelike expansion along the entire tarsomere; Fig. |
Euplantula 2 and 3 without ridgelike expansion; Fig. |
Male | ||
Ocelli | Well developed; Fig. |
Absent; Fig. |
Protibial interior lobe | Always spanning the full length of the protibial shaft | Typically, absent or in well-developed forms only on the proximal half, rarely fully spanning and if so only as a thin lobe |
Prescutum | Anterior margin more typical of a phylliid with the margin not strongly curved, making the prescutum appear less compacted; Fig. |
Anterior margin angled posteriorly, making the prescutum appear very stout; Fig. |
Alae: R split into R1 and Rs | Split is approximately ⅖ of the way through the wing; Fig. |
Split is approximately halfway through the wing |
Alae: MA and MP | Media anterior (MA) and media posterior (MP) veins fuse with the cubitus (Cu) at different locations along the cubitus and run fused to the wing margin; Fig. |
Media anterior (MA) runs unfused to the wing margin; media posterior (MP) fades without fusing or reaching the wing margin |
Eggs | ||
Operculum | Raised on the ventral end, not centrally raised, no pit, minimal granulation throughout; Fig. |
Centrally raised and with a pit in the center; Fig. |
General chorionic texture | Small spherical surface structures; Fig. |
Tuberculate chorionic surface, rough; Fig. |
Microstructures | Mushroom-like smooth granula; Fig. |
Small pinnae arranged in ridges; Fig. |
Freshly hatched nymph | ||
Meso-, metafemoral coloration | Prominent white patch on the center of the exterior lobe and onto the femoral shaft; Fig. |
Mostly black in color, no prominent white patches; Fig. |
Mesonotum | Slender, posterior width similar to length; Fig. |
Stout, posterior width greater than length; Fig. |
Abdominal coloration | Abdomen black with the margins of segment II–IV and VI–IX green; Fig. |
Abdomen uniformly black, no green margins; Fig. |
Distribution | New Caledonia; Fig. |
Fiji |
Several morphological features unite the New Caledonian species and support monophyly of this clade within the phylliids. Within females, the euplantula 2 and 3 on the tarsus has the unique feature of a ridge-like expansion running along the entire tarsomere (Fig.
The Trolicaphyllium gen. nov. are small to medium, with females ranging from 42.0 mm (in the smallest recorded Trolicaphyllium erosus comb. nov.;
Legs. Both sexes have interior tibial lobes on the protibiae which span the full length, 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 broader than the exterior lobe (distinctly so in males with a width almost two times that of the exterior lobe, sometimes in females the interior and exterior are almost even in width). In both sexes the profemoral interior lobe is generally only marked with three or four broadly spaced teeth (quite dulled in females; slightly more serrate in males). Both sexes have the interior meso-, and metafemoral lobes slightly broader or about even in width to the exterior lobes, but the interior lobes are always more prominently marked by serration.
Antennae. Females have antennae with nine segments with segments I, III, VIII, and IX notably broader than the other segments (Fig.
Scanning electron micrographs of female antennae A, C Trolicaphyllium sarrameaense comb. nov. B, D Chitoniscus sensu stricto A, B overview of the antenna, medial view A right antenna B left antenna C, D third antennomere. Abbreviations: a1–a9, antennomeres 1–9 st stridulatory file, sr stridulatory ridge. Scale bars: 300 µm (A, B), 200 µm (C, D).
Head capsule. Males have well-developed ocelli (Fig.
Thorax. The thorax is similar in both sexes with mesopleurae that are narrowly diverging from the anterior to the posterior and are marked with five to seven tubercles, occasionally with sparse setae interspersed (Figs
Wings. Female tegmina are always long, reaching onto abdominal segments VII or VIII and male tegmina are moderate in length, reaching onto abdominal segment III. Females always have highly reduced alae, no more than just a nub (Fig.
Ventro-posterior view of live females showing their exposed coxae coloration A Trolicaphyllium sarrameaense comb. nov. taken by Thierry Salesne (New Caledonia) March 2011, in Vallée Pierrat, La Foa, Grand Terre B Chitoniscus sp. “Suva” (RC Coll 18-176) live photograph taken by Thierry Heitzmann (Philippines).
Female tegmina venation A Trolicaphyllium sarrameaense comb. nov. (Coll SLT) B Chitoniscus feejeeanus from
Abdomen. Both sexes have variable abdominal shapes; females can range from spade-shaped to broad and boxy with prominently projecting abdominal lobes VII and VIII; males can be narrowly-ovoid and lack lobes to broadening until segment VII and converging with lobes. Female subgenital plate is short and stout with the apex reaching the anterior margin of the terminal abdominal segment and ending in a fine point; the gonapophyses VIII are long and slender, slightly exceeding the apex of the terminal abdominal segment; the cerci are relatively flat, marked sparsely with a granular surface with margins slightly marked with setae (Fig.
Egg.
Egg morphology is only known at present from Trolicaphyllium sarrameaense, comb. nov. (Fig.
Scanning electron micrographs of tarsi A, C Chitoniscus sensu stricto B Trolicaphyllium sarrameaense comb. nov. A male, overview B, C females, tarsomeres 1–5. Abbreviations: ar, arolium, cl, claw, eu1–5, euplantula 1–5. Ta1–5, tarsomere 1–5, ri, median ridgelike expansion. Scale bars: 500 µm.
Nymphs. Freshly hatched nymphs are known at present for Trolicaphyllium sarrameaense comb. nov. (Fig.
Trolicaphyllium brachysoma (Sharp, 1898), comb. nov.
Trolicaphyllium erosus (Redtenbachher, 1906), comb. nov.
Trolicaphyllium sarrameaense (Größer er, 2008a), comb. nov.
Etymology. Trolicaphyllium meaning “leaf that walks noiselessly”. This generic epithet is a compound of the Latinized name Phyllium the type genus for the family (from Greek φυλλον, -ου (phyllon, -oy) + -um;
Details of male head through thorax A, B Trolicaphyllium sarrameaense comb. nov., paratype male (#100214); photographs by Arne Köhler (
Distribution. At present Trolicaphyllium gen. nov. specimens are only known from the country of New Caledonia, with records from Grande Terre, Lifou, Tiga, Maré, Ile de Bélep, and L’Île-des-Pins islands (Fig.
(35 ♀♀, 11 ♂♂, 2 unsexed nymphs): Syntypes (2 ♀♀): “Phyllium (Chitoniscus) brachysoma. Type D.S. Lifu. Dr. Willey. 1897” and “Phyllium brachysoma. Type ex parte. D. Lifu. Willey. 1897” (
This was the first phylliid species recorded from New Caledonia and was therefore the first described Trolicaphyllium gen. nov. species, consequently, we here designate it as the type species for the new genus. Additionally, it was chosen as it has acceptably accurate collection data (Lifou Island; a rather small island instead of a general locality from the larger main island, which possibly contains several species) thereby removing some degree of possible confusion which could surround such old and difficult to distinguish specimens. This precise locality will allow future reviewers with adequate material sampled from numerous islands to identify species boundaries and determine if this species ranges across New Caledonia or if it is restricted to Lifou Island.
Male tegmina and alae venation Trolicaphyllium cf. brachysoma comb. nov. (Coll RC 16-094). Abbreviations: C (costa); Sc (subcosta); R (radius); R1 (radius 1); R2 (radius 2); Rs (radial sector); M (media); MA (media anterior); MP (media posterior); MP1 (first media posterior); MP2 (second media posterior); Cu+MA+MP (fused cubitus, media anterior, and media posterior); Cu (cubitus); Cu+1AA (cubitus and first anterior anal); 1A (first anal); 1AA–7AA (first–seventh anterior anal); 1PA–5PA (first–fifth posterior anal).
Scanning electron micrographs of specialized chorionic structures of the eggs A–C Trolicaphyllium sarrameaense comb. nov. D–F Chitoniscus sensu stricto A, D overview of micropylar plate B, E detail of micropylar plate C, F micropylar cap. Scale bars: 300 µm (A, D), 100 µm (B, E, F), 50 µm (C).
Scanning electron micrographs of chorionic microstructures on the eggs A–D Trolicaphyllium sarrameaense comb. nov. E–H Chitoniscus sensu stricto A, B mushroom-like granula C, D, G, H surface microsculpture C surface of the granula D exochorionic surface microstructures E, F pinnae. Scale bars: 100 µm (A, E), 20 µm (B, F), 10 µm (G), 5 µm (D), 3 µm (H), 1 µm (C).
Illustrations of freshly hatched nymphs for comparison, dorsal habitus. Illustrations by Liz Sisk (USA). Nymph size is approximated to be relative to each other based upon the few photographs available but is only an estimate A Trolicaphyllium sarrameaense comb. nov.; overall nymph length from head to tip of abdomen approximately 7 mm (
The syntype females were collected by Dr. Arthur Willey in 1897 while he was living on Lifou Island (Fig.
No etymology was given by Sharp, but it can be assumed that he chose brachysoma to denote the size of the species, from the Greek words brachy- (short) and -soma (body).
For female Trolicaphyllium brachysoma comb. nov., one feature which appears to differentiate this species from the other two is the abdominal shape, which is lobeless, and tapered, giving them a spade-shaped appearance. It is worth noting however that in many phylliids abdominal shape is often a poor feature for differentiation as it is often variable within a single species (
Correctly matching up male and female phylliids is frequently a significant challenge due to their elusiveness in nature and sexual dimorphism and therefore opposite sexes can only be confirmed through molecular comparison or captive rearing (
The type locality for this species is Lifou island, but brachysoma-like specimens with the tapered, lobeless abdomen have been found on Grande Terre (Fig.
Distribution map noting all presently known Trolicaphyllium gen. nov. records which could be traced and accurately noted. See Suppl. material
A Arthur Willey (1867–1942), collector of the Trolicaphyllium brachysoma comb. nov. females in 1897 from Lifou Island while searching for the Pearly Nautilus; image used from the public domain due to expired copyright; from
(10 ♀♀): Syntypes (2 ♀♀): “Syntype;
See Suppl. material
This was the second species described from New Caledonia and was described by
Trolicaphyllium erosus comb. nov. syntype female originally from the collection of Brunner von Wattenwyl, then passed on to Redtenbacher, now deposited in
Within the original description by Redtenbacher the number of syntypes was vague, but at least three are explicitly stated as “New Caledonia (Coll. M., Mus. Paris); New Guinea (Mus. Budapest)” (
An additional syntype was explicitly noted as a nymph within the Budapest Museum (with the wording suggesting that there is only one syntype within that collection). Unfortunately, a fire during the Hungarian Revolution of 1956 destroyed the Budapest syntype along with many of the museum’s important type specimens (
Within the
Philippe François the collector of one of the syntype Trolicaphyllium erosus comb. nov. females from within the
Interestingly, Redtenbacher only appears to have measured one specimen of the type series, the one within his own collection (now within the
No etymology was given in the original description, but it can be assumed that Redtenbacher was referencing the lobed abdominal shape with erosus, from the Latin e- (out/away from) and -rosus (gnaw/peck), noting that the leaf-like body appears gnawed on, giving it the undulating appearance.
To date this appears to be the least known species on New Caledonia as we have not been successful in tracing any additional females which fall within this small size range of ca. 40 mm beyond one of the syntypes. All specimens examined so far appear to belong to the other species with their larger size (ca. 50–60 mm). In fact, it is only this smaller size which we have found as useful for differentiation as the other features match up with the prominently lobed Trolicaphyllium sarrameaense comb. nov. females. The additional specimens identified as Trolicaphyllium erosus comb. nov. within the
Additionally, we have yet to locate any possible male Trolicaphyllium erosus comb. nov. as all males located were much too large and appear to belong to the other two species.
The only specimen we have located with detailed locality information is the syntype female from the
(8 ♀♀, 9 ♂♂, 3 eggs): Holotype and paratypes examined: 1 ♀, 1 ♂, 3 eggs: “Chitoniscus, sarrameaensis, Neu Kaledonien, Sarramea, Sep. 2006, det.Größer” (
See Suppl. material
As it was only described in 2008, this was the most recently described species from New Caledonia with type material originally collected by Sigetake Suzuki in 2004 from Sarramea (
Other lobed specimens have been recovered from throughout New Caledonia, but unfortunately most have been nymphs (such as several from within the
The etymology given in the original description is that this name is a toponym, named after the type locality, Sarramea, New Caledonia (
Females can be differentiated from Trolicaphyllium brachysoma comb. nov. based upon abdominal shape, as Trolicaphyllium brachysoma comb. nov. are considered to have a spade-shaped abdomen, with smooth margins, versus Trolicaphyllium sarrameaense comb. nov. which has a broad abdominal shape with parallel sides, ending in lobed segments VII and VIII. From Trolicaphyllium erosus comb. nov. the only feature we have been able to identify as useful is the overall length, with Trolicaphyllium erosus comb. nov. ca. 40 mm long versus Trolicaphyllium sarrameaense comb. nov. ca. 60 mm long.
Unfortunately, males of Trolicaphyllium brachysoma comb. nov. and Trolicaphyllium erosus comb. nov. have never been confidently confirmed through breeding or molecular comparison. Based upon the confidently confirmed male/female Trolicaphyllium sarrameaense comb. nov. however, we expect that the male morphology should mirror the female morphology. Most likely the male Trolicaphyllium brachysoma comb. nov. will lack prominent abdominal lobes and the male Trolicaphyllium erosus comb. nov. will have distinct lobes to match with their female counterparts. Based upon the female Trolicaphyllium erosus comb. nov. smaller size, we expect that the male must also be rather small, which could likely be used as a feature for differentiation.
To date we have only confirmed adult specimens and observations which are the correct morphology and size of Trolicaphyllium sarrameaense comb. nov. from central Grande Terre (Fig.
Within the
With significant confusion surrounding the distribution of Trolicaphyllium gen. nov. species in New Caledonia (due to apparent intraspecific variation and sexual dimorphism), this leaves many records as unidentifiable to species (see Suppl. material
Live nymphs of Trolicaphyllium gen. nov. which could not be identified to species. Images from https://endemia.nc and used under creative commons license (CC BY-NC-SA 4.0) A Frédéric Desmoulins, Plaine des Lacs, April 2014 B Hendrik Oesterlin, Koé (Dumbéa) elevation 185 meters, June 2006 C Julien Barrault, Hienghène, November 2010 D Daniel and Irène Létocart, Tchamba, October 2009 E Bernard Suprin, Dumbéa, April 2004 F Gildas Gâteblé, Ouenghi, October 2012.
With phylliids typically rather island endemic, it is uncertain at this point if the records from Tiga (Fig.
Only now are the higher-level relationships within the phylliids being explored, with the most likely sister group to Trolicaphyllium gen. nov. appearing to be the Comptaphyllium (
Additionally, geographically, it is possible that Vanuatu to the north of New Caledonia also has suitability for phylliids and may have been part of their route of colonization to modern day New Caledonia assuming an origin of New Guinea for modern phylliids (
Morphologically it is not uncommon within the phylliids for single species to be rather variable within their abdominal shape, ranging from tapered, smooth and boxy, weakly lobed, to strongly lobed (Hennemann et al. 2009;
Some morphological features separating Chitoniscus sensu stricto and Trolicaphyllium gen. nov., namely tarsal and egg morphology, can shed further light on the evolutionary history of these groups, as the morphological difference can be a result of adaptations that accompany the process of speciation. Their functional relevance can highlight the functional constraints which led to the specific morphological traits. The overall tarsus morphology in phasmids in general and in Phylliidae in particular is quite conserved, but the differences found are probably results of adaptations to the specific environments of the species (
The eggs of both groups differ significantly in the modification of their exochorionic surface. While Trolicaphyllium gen. nov. bear distinct microscopic spherical structures on the surface of the eggs (Fig.
The presence or absence of pinnae and their respective morphology have been shown to be of taxonomic value for the eggs of Phylliidae already (
To conclude, despite more than a century of considering the Fijian and New Caledonian phylliids a single genus, we herein adjust the taxonomy of this polyphyletic clade. Our erection of the Trolicaphyllium gen. nov. reflects the unique aspects of the species of New Caledonia as distinct from their previously considered congenerics (Chitoniscus sensu lato).
Herein we have addressed the several questions presented in the introduction regarding these two groups of leaf insects. Firstly, although these clades are in general similar due to their smaller size and abdominal shapes ranging from smoothly tapered to boxy and strongly lobed, a thorough review of small and microstructures of adults, nymphs, and eggs present a plethora of morphological features for easy and reliable differentiation (Table
Therefore, based upon years of molecular results, dozens of morphological features, and the significant geographic isolation for these two clades of leaf insect, we feel that the erection of Trolicaphyllium gen. nov. properly addresses the many discrepancies which were glaringly problematic with the Chitoniscus sensu lato.
Thank you to the many photographers who shared great live images of phylliids with us: Albert Kang (Malaysia), Thierry Heitzmann (Philippines), Patrice Kaateu (New Caledonia), Thierry Salesne (New Caledonia), Detlef Größer (Germany), Thomas Buckley (New Zealand), Damien Brouste (New Caledonia), Mayk de Haan (Belgium), Sylvie Cazeres (IAC), and Christian Mille (IAC). Thank you to the many contributors who posted images of Trolicaphyllium on https://endemia.nc which allowed review of variable morphology as well as a better understanding of the distribution of species: Frédéric Desmoulins, Hendrik Oesterlin, Julien Barrault, Daniel and Irène Létocart, Bernard Suprin, Gildas Gâteblé, Christian Siroux, Jacques Dangio, Nicolas Petit, Christine Poellabauer, and Geoff Monteith. Thank you to the many people who sent us images of museum specimens for review: Jessa Thurman (Australia), Susan Wright (
Table S1. Material examined (specimen data and deposition/ observational records) for localities used within the distribution map (Fig.
Data type: specimen data