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 C3C5916A-9C50-5F26-9B98-9AA2B8BFBEF9 Trolicaphyllium sarrameaense (Größer, 2008)comb. nov.Figures 1, 3, 4A, 4C, 5A, C, 6A, C, 7A, 8A, 9A, 10A, 11A, 12B, 13A, B, 15A–C, 16A–C, 17A–D, 18A, 19, 20, 29Material examined.

(8 ♀♀, 9 ♂♂, 3 eggs): Holotype and paratypes examined: 1 ♀, 1 ♂, 3 eggs: “Chitoniscus, sarrameaensis, Neu Kaledonien, Sarramea, Sep. 2006, det.Größer” (SDEI: HT♀, DEI Hemimetabola #100215; PT♂, DEI Hemimetabola #100214; PT eggs, DEI Hemimetabola #100216); (SDEI; Figs 7A, 8A, 13A, B, 29).

See Suppl. material 1 for additional specimens reviewed, their collection data, and depositories.

Remarks.

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 (Größer 2008a). Within the original description this species was not explicitly compared with the sympatric and morphologically very similar Trolicaphylliumerosus comb. nov. but was instead only differentiated from Chitoniscuslobipes Redtenbacher, 1906, where most features given for differentiation were simply the features we discuss above as significant for differentiating the two genera.

Other lobed specimens have been recovered from throughout New Caledonia, but unfortunately most have been nymphs (such as several from within the QM collection) and therefore they could not be confidently identified as Trolicaphylliumsarrameaense comb. nov. or as Trolicaphylliumerosus comb. nov. nymphs. Unfortunately, in Größer (2008b) the key to species of Chitoniscus sensu lato tried to use the female tegmina radial and media venation pattern to differentiate species, but mixed up the species. Within the key Trolicaphylliumerosus comb. nov. and Trolicaphylliumbrachysoma comb. nov. (from New Caledonia) instead key out as the Fijian population and Chitoniscuslobiventris (Blanchard, 1853) and Chitoniscuslobipes Redtenbacher, 1906 (from Fiji) key out as the New Caledonian population. We have reviewed the type specimen photos available on the Phasmid Species Files (Brock et al. 2021; http://phasmida.speciesfile.org) as well as numerous museum specimens, and always the female tegmina venation allowed accurate distinction of these two genera. Even if you look past this inaccuracy within the key, unfortunately no additional features can be gleaned from the further couplets to allow differentiation of Trolicaphylliumsarrameaense comb. nov. from Trolicaphylliumerosus comb. nov. (as the further couplets discuss abdominal shape, which in these two species is identical/variable). At this moment in time, we still lack significant details about the population of Trolicaphyllium gen. nov. on Grande Terre as material is limited and molecular data has yet to be compared across a wide sampling on the island. With phylliid abdominal shapes sometimes rather variable within a single species, this makes us wonder if Trolicaphylliumsarrameaense comb. nov. is in fact a valid species, or simply a synonym of Trolicaphylliumerosus comb. nov. which was described more than 100 years previously from the same island. Our examination of all type specimens which could be traced has not yet revealed additional features for morphological differentiation besides the overall size of these two species. Hopefully, future molecular analyses from across New Caledonia will reveal if there are several species present on Grande Terre or if it is simply a single species which can range in size from smaller (ca. 40 mm; Trolicaphylliumerosus comb. nov.) to larger (ca. 60 mm; Trolicaphylliumsarrameaense comb. nov.). It is due to this lack of sound molecularly based evidence and the propensity for phylliids to be morphologically variable that we refrain from synonymizing Trolicaphylliumsarrameaense comb. nov. with Trolicaphylliumerosus comb. nov. as we feel a significant decision such as this should be based upon a solid foundation. If future molecular analyses reveal that there is only a single morphologically variable species of Trolicaphyllium gen. nov. on Grande Terre based upon a sampling throughout the island, then we feel a synonymization will be necessary, but not until that time.

The etymology given in the original description is that this name is a toponym, named after the type locality, Sarramea, New Caledonia (Größer 2008a). The original combination was with the masculine genus (Chitoniscus) and therefore in order for the species name to agree in gender with our newly erected genus, the spelling of “sarrameaensis” is changed to the neuter gender “sarrameaense”.

Differentiation.

Females can be differentiated from Trolicaphylliumbrachysoma comb. nov. based upon abdominal shape, as Trolicaphylliumbrachysoma comb. nov. are considered to have a spade-shaped abdomen, with smooth margins, versus Trolicaphylliumsarrameaense comb. nov. which has a broad abdominal shape with parallel sides, ending in lobed segments VII and VIII. From Trolicaphylliumerosus comb. nov. the only feature we have been able to identify as useful is the overall length, with Trolicaphylliumerosus comb. nov. ca. 40 mm long versus Trolicaphylliumsarrameaense comb. nov. ca. 60 mm long.

Unfortunately, males of Trolicaphylliumbrachysoma comb. nov. and Trolicaphylliumerosus comb. nov. have never been confidently confirmed through breeding or molecular comparison. Based upon the confidently confirmed male/female Trolicaphylliumsarrameaense comb. nov. however, we expect that the male morphology should mirror the female morphology. Most likely the male Trolicaphylliumbrachysoma comb. nov. will lack prominent abdominal lobes and the male Trolicaphylliumerosus comb. nov. will have distinct lobes to match with their female counterparts. Based upon the female Trolicaphylliumerosus comb. nov. smaller size, we expect that the male must also be rather small, which could likely be used as a feature for differentiation.

Distribution.

To date we have only confirmed adult specimens and observations which are the correct morphology and size of Trolicaphylliumsarrameaense comb. nov. from central Grande Terre (Fig. 21). We have however seen nymph specimens which had characteristically lobed abdomens which may represent this species from other locations on Grande Terre, so we expect this species may be widespread throughout the island.

Within the MZPW collection there is a pair of Trolicaphylliumsarrameaense comb. nov. specimens with the data of simply “Lifu”, which if true could lend credibility to the hypothesis that perhaps these species are all variable in their abdominal shape (if there is only one species present on Lifou island), but as these are antique and give no other data, we do not take these as highly credible, and therefore exclude this record from further discussion and they are not included within the distribution map (Fig. 21). Or it is possible Lifou island holds several morphologically different species.

10.3897/zookeys.1055.66796.figure1F4CDCD52-3C65-50BC-BF6A-F5CB34F451A5

Dorsal habitus of a live adult female Trolicaphylliumsarrameaense comb. nov. photographed by Thierry Salesne (New Caledonia) in March 2011, in Vallée Pierrat, La Foa, Grand Terre.

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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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Scanning electron micrographs of female antennae A, CTrolicaphylliumsarrameaense comb. nov. B, DChitoniscus 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).

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Scanning electron micrographs of female stridulatory organs A, CTrolicaphylliumsarrameaense comb. nov. B, DChitoniscus sensu stricto A, B overview of stridulatory ridge C, D teeth of stridulatory file. Scale bars: 100 µm (A, B), 20 µm (C, D).

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Scanning electron micrographs of female scapus (1st antennomere) A, CTrolicaphylliumsarrameaense comb. nov. B, DChitoniscus sensu stricto A, B overview of scapus, lateral view C, D medial view. Scale bars: 200 µm.

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Details of the head through thorax, dorsal ATrolicaphylliumsarrameaense comb. nov. (holotype) (DEI Hemimetabola, #100215) BChitoniscus sp. “Suva” (RC Coll 18-176).

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Lateral details of the head and thorax ATrolicaphylliumsarrameaense comb. nov. (holotype) (DEI Hemimetabola, #100215) BChitoniscus sp. “Suva” (RC Coll 18-176).

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Ventro-posterior view of live females showing their exposed coxae coloration ATrolicaphylliumsarrameaense comb. nov. taken by Thierry Salesne (New Caledonia) March 2011, in Vallée Pierrat, La Foa, Grand Terre BChitoniscus sp. “Suva” (RC Coll 18-176) live photograph taken by Thierry Heitzmann (Philippines).

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Female tegmina venation ATrolicaphylliumsarrameaense comb. nov. (Coll SLT) BChitoniscusfeejeeanus from SDEI (#100213). Abbreviations: Sc (subcosta); R (radius); R1 (radius 1); Rs (radial sector); R–M (radius to media crossvein); M (media); MA (media anterior); MP (media posterior); Cu (cubitus); CuA (cubitus anterior); CuP (cubitus posterior); 1A (first anal).

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Female genitalia details, ventral view ATrolicaphylliumsarrameaense comb. nov. (Coll SLT) BChitoniscus sp. “Suva” (RC Coll 18-176).

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Scanning electron micrographs of tarsi A, CChitoniscus sensu stricto BTrolicaphylliumsarrameaense 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.

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Details of male head through thorax A, BTrolicaphylliumsarrameaense comb. nov., paratype male (#100214); photographs by Arne Köhler (SDEI) C, DChitoniscus sensu stricto A, C, D head through thorax, dorsal B head through thorax, lateral CChitoniscus sp. from the NHMUK, photograph by RC D live Chitoniscus sp. “Suva”, photograph by Thierry Heitzemann (Philippines).

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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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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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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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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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Dorsal habitus Trolicaphylliumsarrameaense comb. nov. A holotype female (#100215) B paratype male (#100214). Photographs by Arne Köhler (SDEI). Scale bar: 10 mm.

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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.figure218B981238-C20B-5D83-B466-B893EE78E9AB

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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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.GrößerD (2008a) First description of a new species of Chitoniscus Stal, 1875, since 100 years (Phasmatodea: Phylliidae). Arthropoda 16: e32.BrockPDBüscherTHBakerE (2021) Phasmida SF: Phasmida Species File Version 5.0/5.0. In: Roskov Y et al. (Eds) Species 2000 and ITIS Catalogue of Life. Species 2000: Naturalis, Leiden. https://www.catalogueoflife.org/col