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Live adult female leaf insects showing intergeneric and intraspecific color variability A Pulchriphyllium anangu sp. nov. photographed near Agumbe, India on 17 May 2022 by iNaturalist user vishwanathgowda. Used with permission of Vishwanath Gowda (India) https://www.inaturalist.org/observations/118478029 B, C Phyllium ortizi sp. nov. reared and photographed by Maxime Ortiz (France) in captivity B brown form female C green form female.
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Phylogenetic tree of the Phylliidae based on six nuclear and mitochondrial loci with a focus on Phyllium and Pulchriphyllium (please refer to Suppl. material
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Live Phyllium samarense sp. nov. observed in the wild A female nymph, dorsal habitus, found feeding on Mallotus floribundus (Euphorbiaceae) B adult female, dorsal habitus, found feeding on Mallotus floribundus (Euphorbiaceae) C adult male, dorsal habitus D detail of the coxae, ventral, female E detail of the coxae, ventral, male F Phanera sp. (Fabaceae) G Mallotus floribundus (Euphorbiaceae).
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Phyllium samarense sp. nov. female type material A, C–G paratype (IMQC) B holotype (IMQC), photographs by René Limoges (IMQC) A details of antennae, head capsule, and thorax, dorsal B habitus, dorsal C profemoral and protibial lobes, dorsal D habitus, ventral E genitalia, ventral F thorax details, lateral. Scale bars: 25 mm (B, D).
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Phyllium samarense sp. nov. male paratype, Coll RC #16-259 A details of thorax, ventral B habitus, dorsal C details of head and thorax, dorsal D genitalia, ventral E details of front leg, dorsal F thorax details, lateral. Scale bar: 25 mm (B).
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Phyllium samarense sp. nov. eggs, collected from the female in Fig.
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Phyllium samarense sp. nov. freshly hatched nymph reared in captivity A dorsal habitus B dorsal habitus C details of the abdomen and back legs.
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Illustrations of the freshly hatched nymph coloration for herein referenced species A Phyllium ortizi sp. nov. (Mindanao) B Phyllium mabantai (Mindanao) C Phyllium samarense sp. nov. (Samar) D Pulchriphyllium bioculatum (West Malaysia) E Pulchriphyllium anangu sp. nov. (Southwest India) F Pulchriphyllium agathyrsus (Sri Lanka) G Pulchriphyllium giganteum (West Malaysia) H Pulchriphyllium bhaskarai sp. nov. (Java). Illustrations by scientific illustrator Liz Sisk (Washington D.C., USA).
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Phyllium iyadaon sp. nov. type specimens A–E holotype (IMQC) Coll RC 17-221 F paratype (Coll RC) Coll RC 17-222 A habitus, dorsal B details of the head through thorax, dorsal C details of the front leg, dorsal D genitalia, ventral E details of the head through thorax, lateral F details of the head through thorax, lateral. Scale bar: 20 mm (A).
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Phyllium ortizi sp. nov. live individuals from captive culture highlighting aspects of their coloration. Photographs by Maxime Ortiz (France) A adult male and female pair, dorsal habitus B adult female, postero-ventral view, showing the bright orange coxae coloration C adult male, habitus, dorsal D adult male, habitus, ventral E brown form nymph, same individual as in Fig.
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Phyllium ortizi sp. nov. female holotype (IMQC) D, paratype (Coll RC) A–C, E, F photographs by René Limoges (IMQC). Scale bars associated with A, D respectively A dorsal habitus, paratype B lateral view of the thorax showing spination C details of the head and thorax, dorsal D dorsal habitus, holotype E details of the antennae, dorsal F genitalia details, ventral.
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Phyllium ortizi sp. nov. male paratypes A–D (Coll RC 21-001). Photographs by René Limoges (IMQC) E, F (Coll RC 21-002) A details of front leg B dorsal habitus C details of the head and thorax, dorsal D antenna, dorsal E genitalia details, ventral F lateral view of the thorax showing spination. Scale bar: 15 mm (B).
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Phyllium ortizi sp. nov. egg capsule, IMQC collection, photographs by René Limoges (IMQC), scale bar associated with all images A lateral view B dorso-lateral view C dorsal view D opercular (anterior) view E posterior view F ventral view.
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Phyllium ortizi sp. nov. freshly hatched nymphs reared and photographed by Maxime Ortiz (France) A habitus, dorsolateral B habitus, dorsal C two nymphs drinking moisture off of bramble (Rubus sp.), one of the few times freshly hatched nymphs hold still.
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Distribution map for the herein discussed Pulchriphyllium species with known records plotted. For data on the distribution points used for this map, see Suppl. material
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Pulchriphyllium scythe (Gray, 1843) stat. rev., comb. nov. herein designated lectotype female, photographs by Robert Douglas (OUMNH) A habitus, dorsal B habitus, ventral C detail of the front leg, dorsal D data label E detail of the genitalia, ventral F detail of the antenna, head, and anterior of the thorax, dorsal.
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Live females of Pulchriphyllium scythe (Gray, 1843) stat. rev., comb. nov. A adult female, dorsal habitus, found near Makunda Christian Hospital, Karimganj District, Assam, 20 April 2016, photographed by Vijay Anand Ismavel (India), (https://www.inaturalist.org/observations/4311333) B adult female, dorsal habitus, found in Assam 788727, India, 27 December 2017, photographed by iNaturalist user @lovelymonlamin (https://www.inaturalist.org/observations/30552832) C adult female, dorsal habitus, found in Assam 788727, India, 27 August 2019, photographed by Dwithun Moshahary, (https://www.inaturalist.org/observations/51076352) D female nymph, dorsal habitus, found in Vaisam, Damchhara R. F., Tripura, India, 12 August 2020, photographed by Michael Lalruatfela (India) E female nymph, dorsal habitus, found near Makunda Christian Hospital, Karimganj District, Assam, 9 June 2019, photographed by Vijay Anand Ismavel (India), (https://www.inaturalist.org/observations/29119549) F same individual as in D but dorso-anterior view. Photographs used were uploaded to iNaturalist and are here used under license (CC BY-NC 4.0) or with explicit permission by the photographer.
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Live males and nymphs of Pulchriphyllium scythe (Gray, 1843) stat. rev., comb. nov. A male habitus, dorsal, found in Vaisam, Damchhara R. F., Tripura, India, 30 June 2019 and photographed by Michael Lalruatfela (India) B male habitus, dorsal, found in Badshahi Forest, Assam, India, 18 September 2020 and photographed by Rejoice Gassah (India) (https://www.inaturalist.org/observations/64092851) C nymph, dorso lateral view, found in Lawachara National Park, 31 July 2019, photographed by Hassan-al Razi Chayan (Bangladesh) (https://www.inaturalist.org/observations/103359437) D nymph, dorso-anterior view, found in Dosdewa Khasi Village, Katamoni, Assam, India, 24 April 2021 and photographed by Rejoice Gassah (India) (https://www.inaturalist.org/observations/79254856). Photographs used were uploaded to iNaturalist and here used under license (CC BY-NC 4.0) or with explicit permission by the photographer.
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Pulchriphyllium crurifolium stat. rev., comb. nov. specimens which, based upon their labels, appear to be the syntype specimens designated by Audinet-Serville in 1838 A paralectotype, male, dorsal habitus, associated data labels to the right B lectotype, female, dorsal habitus, associated data labels to the right C paralectotype, female, dorsal habitus, associated data labels to the left and below D paralectotype, male, dorsal habitus, associated data labels to the left and below. Photographs by Robert Douglas, courtesy of Oxford University Museum of Natural History.
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Phyllium dardanus Westwood, 1859 holotype male (= Pulchriphyllium crurifolium stat. rev., comb. nov.) A dorsal habitus B holotype data labels C details of the antennae, front legs, head, and thorax, dorsal A, B photographs by Robert Douglas, courtesy of Oxford University Museum of Natural History C photograph by Paul Brock (
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Adult female profemoral lobes from various Pulchriphyllium species for comparison A Pulchriphyllium crurifolium stat. rev., comb. nov. (
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Live Pulchriphyllium crurifolium stat. rev., comb. nov. A adult female, dorsal habitus, observed September 2019 at Sans Souci, Mahé, Seychelles, by Juan Jose Areso uploaded by iNaturalist user @liahg (Amalia Herrera Grau) (https://www.inaturalist.org/observations/76082181) B adult male, dorsal habitus, observed December 2006 at Grand’ Anse, Mahé, Seychelles by iNaturalist user @thierrycordenos (https://www.inaturalist.org/observations/22992534). Both images uploaded to iNaturalist and used under license (CC BY-NC 4.0).
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Eggs from Pulchriphyllium bioculatum-like species, where the eggs are known A–C Pulchriphyllium bioculatum from West Malaysia (Coll RC #18-042) D–F Pulchriphyllium crurifolium stat. rev., comb. nov. from the Seychelles (Coll RC #18-233) G–I Pulchriphyllium agathyrsus from Sri Lanka (Coll RC #18-232). Top row, dorsal view. Middle row, lateral view. Bottom row, anterior view.
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Pulchriphyllium heracles sp. nov. holotype male (Coll RC 16-007; IMQC) A habitus, dorsal B details of the head through thorax, dorsal C genitalia, ventral. Photographs by Steve Thurston (AMNH) using Microptics-USA/Visionary Digital photomicrographic system developed by Roy Larimer; using a Nikon D300 DSLR camera, Infiniti optics, and a Micro Nikkor 60 mm lens. Multiple layers stacked using Helicon Focus.
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Pulchriphyllium bhaskarai sp. nov. live holotype female. Photographs by Edy Bhaskara (Indonesia) A coloration shortly after being found in the wild, dorsal habitus B altered coloration after several months in captivity, dorsal habitus C habitus, dorsolateral view.
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Pulchriphyllium bhaskarai sp. nov. female holotype A details of the antennae, dorsal B dorsal habitus C details of the head and thorax, dorsal D details of the profemora E genitalia details, ventral F ventral habitus G lateral view of the thorax (anterior to the right). Scale bars: 25 mm (B, F).
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Pulchriphyllium bhaskarai sp. nov. male paratype, Coll. RC #21-032 A dorsal habitus B details of the profemora C lateral view of the thorax (anterior to the right) D details of the antennae, dorsolateral view E terminalia details, dorsal F genitalia details, ventral G details of the head and thorax, dorsal. Scale bar: 25 mm (A).
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A–F Pulchriphyllium giganteum egg capsule, IMQC collection A lateral view B dorso-lateral view C dorsal view D opercular (anterior) view E posterior view F ventral view G–L Pulchriphyllium bhaskarai sp. nov. egg capsule, IMQC collection (Coll RC 21-034) G lateral view H dorso-lateral view I dorsal view J opercular (anterior) view K posterior view L ventral view. Photographs by René Limoges (IMQC). Scale bars: 2 mm (A–L).
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Pulchriphyllium bhaskarai sp. nov. live freshly hatched nymphs A dorsal habitus B habitus, dorsolateral view C habitus, dorsolateral view. Photographs by Edy Bhaskara (Indonesia).
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Paratype historic pair of Pulchriphyllium bhaskarai sp. nov. from
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Male holotype Pulchriphyllium anangu sp. nov. (
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Live Pulchriphyllium anangu sp. nov. A adult female, dorsal habitus, observed July 2016 at Kadumane Estate, India by iNaturalist user @ashwinv (Ashwin Viswanathan) https://www.inaturalist.org/observations/29374627) B freshly hatched nymph, observed October 2021 at Chikkamagaluru, Karnataka, India by iNaturalist user @prashanthakrishnamc (Prashantha Krishna M C) https://www.inaturalist.org/observations/98984918) C freshly hatched nymph, observed January 2021 in Ponmudi, India by iNaturalist user @hari_krishnan_ (Harikrishnan S) https://www.inaturalist.org/observations/70254070). Uploaded to iNaturalist and used under license CC BY-NC 4.0.
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Live Pulchriphyllium anangu sp. nov. A adult male, dorsal habitus observed October 2006 in B.R. Hills, Karnataka, India by iNaturalist user @prashanthns (Prashanth N S) (https://www.inaturalist.org/observations/31929431) B adult male, dorsal habitus observed November 2019 in Pakkalakunja, Karnataka, India by iNaturalist user @sanath_ramesh_manimoole (Sanath R M) (https://www.inaturalist.org/observations/103401996) C male nymph, dorsal habitus observed February 2020 in Shimoga, Karnataka, India by iNaturalist user @girishgowda (Girish Gowda) (https://www.inaturalist.org/observations/104655374) D nymph, dorsal habitus, hiding under a leaf in the daylight observed August 2020 near Agali, India by iNaturalist user @manavsajan (https://www.inaturalist.org/observations/56588953). Uploaded to iNaturalist and used under license CC BY-NC 4.0.
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Female holotype Pulchriphyllium delislei sp. nov. (IMQC) A dorsal, habitus B antennae, dorsal C right profemora, dorsal D head and thorax, dorsal E thorax, lateral, anterior to the right F ventral, habitus G genitalia, ventral H terminalia, dorsal.
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Female paratype Pulchriphyllium delislei sp. nov. (IMQC) A dorsal, habitus B details of the head through thorax, dorsal C details of the front leg, dorsal D details of head and antennae, dorsal E genitalia, ventral F ventral, habitus.