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) Author Cumming, Royce T. https://orcid.org/0000-0001-7930-1292 Montreal Insectarium, 4101 rue Sherbrooke est, Montre ́ al, Que ́ bec, H 1 X 2 B 2, Canada & Richard Gilder Graduate School, American Museum of Natural History, New York, NY 10024, USA & Biology, Graduate Center, City University of New York, NY, USA roycecumming@gmail.com Author Tirant, Ste ́ phane Le Montreal Insectarium, 4101 rue Sherbrooke est, Montre ́ al, Que ́ bec, H 1 X 2 B 2, Canada Author Bu ̈ scher, Thies H. Department of Functional Morphology and Biomechanics, Zoological Institute, Kiel University, Am Botanischen Garten 9, 24118, Kiel, Germany text ZooKeys 2021 2021-08-05 1055 1 41 http://dx.doi.org/10.3897/zookeys.1055.66796 journal article http://dx.doi.org/10.3897/zookeys.1055.66796 1313-2970-1055-1 DB8C0779E75744EDB9B2397EA2EF3BAE 69CACA7352355D15AE1D785BE3CA28C5 Trolicaphyllium brachysoma (Sharp, 1898) comb. nov. Figures 14 , 22 , 24 , 25 Material examined. ( 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" (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. Figure 14. 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). Figure 15. Comparison of Chitoniscus sensu stricto and Trolicaphyllium gen. nov. eggs A-C Trolicaphyllium sarrameaense comb. nov., imaged by TB of eggs from Coll DG A dorsal B lateral C opercular (anterior) D-F Chitoniscus sp. "Suva" (RC Coll 18-272) D dorsal E lateral F opercular (anterior). Figure 16. 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 ). Figure 17. 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 ). Figure 18. 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 ( Groesser 2008b ); illustration based upon photographs from Detlef Groesser (Germany) B Chitoniscus sp. "Suva'' based upon images supplied by Mayk de Haan (Belgium). Figure 19. Captive bred Trolicaphyllium sarrameaense comb. nov. dorsal, habitus, female reared and photographed by Detlef Groesser (Germany). 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 ). Figure 20. Trolicaphyllium sarrameaense comb. nov. eggs showing the variation in coloration, photographs by Sylvie Cazeres (IAC), eggs laid by females from Vallee Pierrat, Commune de La Foa A lighter colored eggs B first five eggs laid by the female from Figure 3 . No etymology was given by Sharp, but it can be assumed that he chose Trolicaphyllium brachysoma to denote the size of the species, from the Greek words brachy - (short) and - soma (body). Differentiation. 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 ( 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 ). Trolicaphyllium brachysoma , comb. nov. is about the same size as Trolicaphyllium sarrameaense comb. nov. (ca. 60 mm) which can differentiate them from Trolicaphyllium erosus 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 Trolicaphyllium brachysoma 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). Trolicaphyllium erosus 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 Trolicaphyllium erosus comb. nov. is likely rather small. Distribution. The type locality for this species is Lifou island, but Trolicaphyllium brachysoma -like specimens with the tapered, lobeless abdomen have been found on Grande Terre (Fig. 21 ) and L'Ile-des-Pins (Fig. 25 ) as well. Additionally, within the MNHN there is a female which was collected on Ile de Belep , 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. Figure 21. 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). Figure 22. Syntype females of Trolicaphyllium brachysoma comb. nov. the herein designated type species for the new genus. Photographs by Paul Brock (United Kingdom) of the set within the CUMZ A dorsal habitus with tegmina spread; note the lack of developed alae B dorsal habitus with tegmina closed. Figure 23. 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 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). Figure 24. Dorsal habitus of male Trolicaphyllium cf. brachysoma comb. nov. A Bouloupari Commune, March 2013 (Coll RC 16-095) B Sarramea Commune, February 2009 (Coll RC 16-094). Figure 25. Trolicaphyllium cf. brachysoma comb. nov. adult female observed on L'Ile-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.