AnimaliaPhasmatodeaPhasmatodeaCummingRoyce T.TirantStéphane LeBüscherThies H.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)Zookeys0582021105514110.3897/zookeys.1055.66796 E9E27569-68C8-57E5-93DD-2094FFFD0480 Trolicaphyllium brachysoma (Sharp, 1898)comb. nov.Figures 14, 22, 24, 25Material examined.

(35 ♀♀, 11 ♂♂, 2 unsexed nymphs): Syntypes (2 ♀♀): “Phyllium (Chitoniscus) brachysoma. Type D.S. Lifu. Dr. Willey. 1897” and “Phylliumbrachysoma. Type ex parte. D. Lifu. Willey. 1897” (CUMZ; Fig. 22). See Suppl. material 1 for additional specimens reviewed, their collection data, and depositories.

Remarks.

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.

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Male tegmina and alae venation Trolicaphylliumcf.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); 1AA7AA (first–seventh anterior anal); 1PA5PA (first–fifth posterior anal).

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10.3897/zookeys.1055.66796.figure15C4A543EC-7C4D-5CCE-BA1D-BAE56FE99DB0

Comparison of Chitoniscus sensu stricto and Trolicaphyllium gen. nov. eggs A–CTrolicaphylliumsarrameaense comb. nov., imaged by TB of eggs from Coll DGA dorsal B lateral C opercular (anterior) D–FChitoniscus sp. “Suva” (RC Coll 18-272) D dorsal E lateral F opercular (anterior).

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10.3897/zookeys.1055.66796.figure163D777D22-5656-544C-BB3D-967503FB072C

Scanning electron micrographs of specialized chorionic structures of the eggs A–CTrolicaphylliumsarrameaense comb. nov. D–FChitoniscus 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).

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Scanning electron micrographs of chorionic microstructures on the eggs A–DTrolicaphylliumsarrameaense comb. nov. E–HChitoniscus 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).

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10.3897/zookeys.1055.66796.figure187F73E96F-560F-57AB-8182-337E2FD1F16D

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 ATrolicaphylliumsarrameaense comb. nov.; overall nymph length from head to tip of abdomen approximately 7 mm (Größer 2008b); illustration based upon photographs from Detlef Größer (Germany) BChitoniscus sp. “Suva’’ based upon images supplied by Mayk de Haan (Belgium).

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Captive bred Trolicaphylliumsarrameaense comb. nov. dorsal, habitus, female reared and photographed by Detlef Größer (Germany).

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The syntype females were collected by Dr. Arthur Willey in 1897 while he was living on Lifou Island (Fig. 23; Sharp 1898). Dr. Willey was traveling and living in New Britain, New Hanover, eastern New Guinea, and Lifou Island between 1895 and 1897 in search of living Pearly Nautilus colonies which he could capture, collect eggs from, and rear through development in order to study their embryology (Wiley 1899; Kerr 1943). Although his years of expedition yielded many great discoveries of which he published prolifically, he was unfortunately unsuccessful in his primary goal of rearing eggs to maturity (Willey 1899). Willey lived on the west coast of Lifou Island on “Sandal Bay” (modern Santal Bay) from July 1896 to March 1897; and while no exact date was given with the syntype set of females, they are noted as being collected in 1897. Therefore, they are from the beginning of the year (January through March), and most likely from late January when a severe gale passed through the area (wreaking havoc on his Nautili traps; Kerr 1943) which likely knocked the phylliids from the canopy enabling them to be found by Dr. Willey. While males occasionally will fly to lights at night, females and nymphs are most often only found on the ground after storms when they are knocked from their typical canopy habitat and found lower (Brock and Hasenpusch 2003, 2015).

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Trolicaphylliumsarrameaense comb. nov. eggs showing the variation in coloration, photographs by Sylvie Cazeres (IAC), eggs laid by females from Vallée Pierrat, Commune de La Foa A lighter colored eggs B first five eggs laid by the female from Figure 3.

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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).

Differentiation.

For female Trolicaphylliumbrachysoma 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 (Cumming et al. 2020b), and even in the syntype set of two females, one female is notably more tapered (Fig. 22B) than the other (Fig. 22A). Trolicaphylliumbrachysoma, comb. nov. is about the same size as Trolicaphylliumsarrameaense comb. nov. (ca. 60 mm) which can differentiate them from Trolicaphylliumerosus comb. nov. which are notably smaller (ca. 40 mm).

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 (Cumming et al. 2020c; Cumming et al. 2021). Unfortunately, we have yet to confidently confirm a male Trolicaphylliumbrachysoma comb. nov. and therefore, at this time can only illustrate presumed males (Fig. 24) which follow the morphology of the female by having a smooth tapered abdomen and falling within an appropriate size range for a potential male (38 to 43 mm). Trolicaphylliumerosus comb. nov. has no presumed male records we are aware of (as no possibilities have been located in collections which are small enough to represent a male of this species) but based upon female size the male Trolicaphylliumerosus comb. nov. is likely rather small.

Distribution.

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. 21) and L’Île-des-Pins (Fig. 25) as well. Additionally, within the MNHN there is a female which was collected on Ile de Bélep, which is the only phylliid record we have seen from this island, and we only tentatively note this specimen as this species as it has slight lobes on the abdomen and could not be examined in person. Hopefully future molecular analyses with material from multiple islands will reveal if these are all one species or several.

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Distribution map noting all presently known Trolicaphyllium gen. nov. records which could be traced and accurately noted. See Suppl. material 1 for full details for all records presented. Stars indicate a record based upon a specimen, circles represent a record based upon a photographic observation. Produced with SimpleMappr (Shorthouse, 2010).

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Syntype females of Trolicaphylliumbrachysoma comb. nov. the herein designated type species for the new genus. Photographs by Paul Brock (United Kingdom) of the set within the CUMZA dorsal habitus with tegmina spread; note the lack of developed alae B dorsal habitus with tegmina closed.

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A Arthur Willey (1867–1942), collector of the Trolicaphylliumbrachysoma 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 Kerr (1943)B Lifou, cliffs on the north end of the island slightly farther north of where Willey was stationed 1896–1897; photograph by user Bahnfrend in November 2007; used under Creative Commons 3.0 (CC BY-SA 3.0) (https://commons.wikimedia.org/wiki/File:Jokin,_Lifou,_2007_(4).JPG).

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Dorsal habitus of male Trolicaphylliumcf.brachysoma comb. nov. A Bouloupari Commune, March 2013 (Coll RC 16-095) B Sarramea Commune, February 2009 (Coll RC 16-094).

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Trolicaphylliumcf.brachysoma comb. nov. adult female observed on L’Île-des-Pins by Patrice Kaateu (New Caledonia) in November 2020 A dorsal, habitus B same individual as in A but zoomed out to give scale/perspective.

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10.3897/zookeys.1055.66796.suppl1DD4C43EC-0FE7-5005-9BB6-7E63546AC362

Table S1. Material examined (specimen data and deposition/ observational records) for localities used within the distribution map (Fig. 21) and to accompany the discussion of each species

specimen data

https://binary.pensoft.net/file/574826This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.Royce Cumming, Stéphane Le Tirant, Thies H. Büscher
10.3897/zookeys.1055.66796.figure3F6AA9329-25E2-5BD0-BA4F-FA06CE37D960

Adult female Trolicaphylliumcf.sarrameaense comb. nov. found feeding on Syzygiumcumini Sarramea county, near La Foa, January 2021 (recorded and photographed by Sylvie Cazeres (IAC)) A dorsoanterior, habitus B dorsal, habitus.

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GrößerD (2008b) Wandelnde Blätter. Ein Katalog aller bisher beschriebenen Phylliinae-Arten und deren Eier mit drei Neubeschreibungen (2nd Edn.).Chimaira, Frankfurt am Main, 175 pp.KerrJ (1943) Arthur Willey. 1867–1942.Obituary Notices of Fellows of the Royal Society4: 395410. https://doi.org/10.1098/rsbm.1943.0011SharpD (1898) Account of the Phasmidae, with notes on the eggs. In: WilleyA (Ed.) Zoological Results Based on Material from New Britain, New Guinea, Loyalty Islands and Elsewhere, Collected During the Years 1895, 1896, and 1897.Cambridge University Press, Cambridge, United Kingdom, 7594.WilleyA (1899) 1. General Account of a Zoological Expedition to the South Seas during the years 1894–1897.Proceedings of the Zoological Society of London67: 79. https://doi.org/10.1111/j.1469-7998.1899.tb06841.xBrockPDHasenpuschJ (2015) Australian stick and leaf insects (Insecta, Phasmida): camouflage and natural history. In: EliasAHarleyRTsoutasN (Eds) Camouflage Cultures: Beyond the Art of Disappearance.Australia, Sydney University Press, Sydney, 4562. http://www.jstor.org/stable/j.ctt1bh4b60CummingRTBakerELe TirantSMarshallJ (2020b) On the Phyllium Illiger, 1798 of Palawan (Philippines), with description of a new species (Phasmida: Phylliidae).Faunitaxys8: 19.CummingRTTirantSLTeemsmaSNHennemannFHWillemseLBüscherTH (2020c) Lost lovers linked at long last: elusive female Nanophyllium mystery solved after a century of being placed in a different genus (Phasmatodea, Phylliidae).ZooKeys969: 4384. https://doi.org/10.3897/zookeys.969.56214CummingRTBankSBresseelJConstantJLe TirantSDongZSonetGBradlerS (2021) Cryptophyllium, the hidden leaf insects – descriptions of a new leaf insect genus and thirteen species from the former celebicum species group (Phasmatodea, Phylliidae).ZooKeys1018: 1179. https://doi.org/10.3897/zookeys.1018.61033