Research Article |
Corresponding author: Royce T. Cumming ( roycecumming@gmail.com ) Corresponding author: Sarah Bank ( sarah.bank@uni-goettingen.de ) Academic editor: Fedor Konstantinov
© 2023 Royce T. Cumming, Stéphane Le Tirant, Jackson B. Linde, Megan E. Solan, Evelyn Marie Foley, Norman Enrico C. Eulin, Ramon Lavado, Michael F. Whiting, Sven Bradler, Sarah Bank.
This is an open access article distributed under the terms of the CC0 Public Domain Dedication.
Citation:
Cumming RT, Le Tirant S, Linde JB, Solan ME, Foley EM, Eulin NEC, Lavado R, Whiting MF, Bradler S, Bank S (2023) On seven undescribed leaf insect species revealed within the recent “Tree of Leaves” (Phasmatodea, Phylliidae). ZooKeys 1173: 145-229. https://doi.org/10.3897/zookeys.1173.104413
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With the recent advance in molecular phylogenetics focused on the leaf insects (Phasmatodea, Phylliidae), gaps in knowledge are beginning to be filled. Yet, shortcomings are also being highlighted, for instance, the unveiling of numerous undescribed phylliid species. Here, some of these taxa are described, including Phyllium iyadaon sp. nov. from Mindoro Island, Philippines; Phyllium samarense sp. nov. from Samar Island, Philippines; Phyllium ortizi sp. nov. from Mindanao Island, Philippines; Pulchriphyllium heracles sp. nov. from Vietnam; Pulchriphyllium delislei sp. nov. from South Kalimantan, Indonesia; and Pulchriphyllium bhaskarai sp. nov. from Java, Indonesia. Several additional specimens of these species together with a seventh species described herein, Pulchriphyllium anangu sp. nov. from southwestern India, were incorporated into a newly constructed phylogenetic tree. Additionally, two taxa that were originally described as species, but in recent decades have been treated as subspecies, are elevated back to species status to reflect their unique morphology and geographic isolation, creating the following new combinations: Pulchriphyllium scythe (Gray, 1843) stat. rev., comb. nov. from Bangladesh and northeastern India, and Pulchriphyllium crurifolium (Audinet-Serville, 1838) stat. rev., comb. nov. from the Seychelles islands. Lectotype specimens are also designated for Pulchriphyllium scythe (Gray, 1843) stat. rev., comb. nov. and Pulchriphyllium crurifolium (Audinet-Serville, 1838) stat. rev., comb. nov. from original type material.
India, Indonesia, Kalimantan, Phasmida, Philippines, Seychelles, Vietnam, walking leaf
Stick and leaf insects (Phasmatodea; colloquially “phasmids”) are a clade that has evolved remarkable morphological adaptations for botanical camouflage and mimicry. Presently Phasmatodea includes more than 3,000 known species distributed across most regions of the world, with dozens of species new to science described annually (
One clade which mimics general angiosperm leaf morphology particularly well are the Phylliidae, or true leaf insects. These masters of leaf masquerade are widely distributed across the tropical regions of Asia, Australasia, and the Pacific (
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.
Despite the stunning appearance of leaf insects, they are exceedingly rare within collections and in many cases solely known from the holotype (
The relationship of the Phylliidae to the greater Phasmatodea has been debated for decades, with various authors placing them in dramatically different locations in the phasmid family tree (examples of past relationships summarized in
Despite the uncertainty of the higher-level phylogenetic relationship of the phylliids, the application of molecular data has been found to be a useful tool to clarify phylogenetic relationships within the Phylliidae (
Specimens and observation records for this study come from permitted fieldwork conducted by the authors, records from reviewed museum collections, records from several private collections, and observations from citizen science-based platforms such as iNaturalist and biodiversity enthusiast groups on Facebook. For each species discussed herein, we explicitly state within Suppl. material
Despite the usefulness of internal genitalia for differentiation of many insects, including phasmids (
Measurements of specimens were made to the nearest 0.1 mm using digital calipers and are given for individual specimens where applicable (such as for holotype specimens) or for series recording a minimum to maximum range (such as for paratype specimens). Images taken using: a Canon EOS 5DS R camera (Canon, Tokyo, Japan) with a 100-mm lens. Zerene Stacker (v. 1.04) was used to generate images from stacked photos (Zerene Systems LLC, Richland, WA, USA). Holotype and paratype specimens herein designated are deposited within several different institution collections, which are explicitly listed within the type material information of the new species descriptions and/or within Suppl. material
Coll EF Private collection of Evelyn Marie Foley, New Jersey, USA;
Coll FH Private collection of Frank H. Hennemann, Germany;
Coll HMY Private collection of H. M. Yeshwanth, Bangalore, India;
Coll MO Private collection of Maxime Ortiz, France;
Coll MS Private collection of Megan Solan, Texas, USA;
Coll NE Private collection of Norman Enrico C. Eulin, Samar, Philippines;
Coll PEB Private collection of Phil E. Bragg, Derbyshire, United Kingdom;
Coll RC Private collection of Royce T. Cumming, California, USA;
Coll SLT Private collection of Stéphane Le Tirant, Québec, Canada;
IEBR Entomological Collection of the Institute of Ecology and Biological Resources Hanoi, Vietnam;
IMQC Insectarium de Montréal, Montréal, Québec, Canada;
OUMNH University Museum of Natural History, Oxford, United Kingdom;
SASRD School of Agricultural Sciences and Rural Development, Nagaland, India;
Photographs of specimens deposited within the IMQC collection were taken 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). Two Nikon SB-25 flash units provided lighting with a Cameron Digital diffusion photo box (Henry’s, Vancouver, Canada). Adobe Photoshop Elements 13 (Adobe Inc., San Jose, USA) was used as post-processing software. For details that could not be seen with the naked eye, such as the stridulatory file of the third antennomere in females, scanning electron microscopy (SEM) was used. Samples were sputter-coated with a 10 nm gold-palladium layer. SEM images were then taken with a SEM Hitachi TM3000 (Hitachi High-technologies Corp., Tokyo, Japan) at 15 kV acceleration voltage as outlined in
The phylogenetic relationships of 169 taxa with 15 newly added specimens resulted in an almost identical phylogeny to that inferred by
Phylogenetic tree of the Phylliidae based on six nuclear and mitochondrial loci with a focus on Phyllium and Pulchriphyllium (please refer to Suppl. material
Phylliidae, Phylliinae, Phylliini
Phyllium siccifolium (Linnaeus, 1758), type locality: “Indies”.
Phyllium historically encompassed most leaf insect species, with numerous internal subdivisions and species groups, some recognized as unique for more than a century (e.g., Pulchriphyllium Griffini, 1898) and other divisions more recent (e.g., those found in
Holotype
♀: “Philippines: Eastern Visayas, Northern Samar, Lope De Vega: April, 2017; Coll RC 17-206; DNA ID #W9”. Deposited in the Montreal Insectarium, Quebec, Canada (IMQC). Paratypes: (9 ♀♀, 6 ♂♂, 1 ♂ nymph, 2 ♀♀ nymphs, 6 eggs). See Suppl. material
Female Phyllium samarense sp. nov. (Figs
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).
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).
Male Phyllium samarense sp. nov. (Figs
Eggs of Phyllium samarense sp. nov. (Fig.
Freshly hatched nymph Phyllium samarense sp. nov. (Figs
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).
Female. Coloration. Coloration description is based upon living individuals in which some variation has been observed. Overall coloration has been observed to range from fully pale green (Fig.
Morphology. Head. Head capsule slightly longer than wide; vertex is irregularly lumpy due to moderately sized nodes throughout the surface and a base texture which is slightly wrinkled (Fig.
Measurements of holotype female [mm]. Length of body (including cerci and head, excluding antennae) 78.7, length/width of head 7.3/6.3, antennae 4.5, pronotum 5.4, mesonotum 7.1, length of tegmina 48.6, length of alae 4.3, greatest width of abdomen 29.3, profemora 15.2, mesofemora 12.9, metafemora 16.0, protibiae 9.2, mesotibiae 10.0, metatibiae 13.5.
Measurements of paratype females [mm]. Length of body (including cerci and head, excluding antennae) 76.4–80.1, length/width of head 7.2–7.4/6.3– 6.4, antennae 4.5–4.6, pronotum 5.2–5.4, mesonotum 7.0–7.4, length of tegmina 47.5–49.7, length of alae 4.3–4.4, greatest width of abdomen 28.2–30.2, profemora 15.1–15.6, mesofemora 12.7–13.2, metafemora 16.0–16.4, protibiae 9.2–9.5, mesotibiae 10.0–10.4, metatibiae 13.5–13.8.
Male. Coloration. Coloration based upon living individuals (Fig.
Morphology. Head. Head capsule slightly longer than wide, with a vertex that is slightly lumpy with sparse irregularly spaced but relatively uniformly sized nodes throughout the posterior of the capsule. The posteromedial tubercle is singularly pointed and distinctly raised above the head capsule (Fig.
Measurements of paratype males [mm]. Length of body (including cerci and head, excluding antennae) 57.6–60.2, length/width of head 3.9–4.5/3.3– 3.5, antennae 33.2–35.1, pronotum 2.9–3.1, mesonotum 4.6–4.8, length of tegmina 19.0–19.6, length of alae 42.8–43.3, greatest width of abdomen 14.2–14.5, profemora 11.6–12.0, mesofemora 10.2–10.5, metafemora 12.7–13.1, protibiae 7.6–8.0, mesotibiae 7.3–7.5, metatibiae 9.7–10.1.
Eggs. (Fig.
Measurements including the extended pinnae [mm]. Length (including operculum): 4.5–5.0; maximum width of capsule when viewed from lateral aspect 2.5–3.0; length of micropylar plate 2.7–3.0.
(Fig.
Toponym. Named after the type locality Samar Island, Philippines. As the first officially endemic species of Phyllium from the island, we wished to highlight this significant find by referencing the island.
At present only known from North Samar province, on Samar Island, Philippines in the Eastern Visayas region.
This population was previously thought to be a range expansion for Phyllium mabantai, a species which is quite variable in abdominal morphology and has a similar overall habitus (
To date, Phyllium samarense sp. nov. has been found feeding on three host plants in the wild; Phanera sp. (Fabaceae; Fig.
The Phyllium samarense sp. nov. egg morphology fits within the results of
Holotype ♂: “Philippines: Mindoro, Puerto Galera: April, 2017; Coll RC 17-221”. Deposited in the Montreal Insectarium, Quebec, Canada (IMQC). Paratype: (1♂) • “Philippines: Mindoro, Puerto Galera: April, 2017; Coll RC 17-222” (Coll RC).
Female, freshly hatched nymph, and egg unknown. Males morphologically are similar to Phyllium philippinicum and Phyllium bourquei in all features except for the undulating abdomen, the profemoral interior lobe with a notable space between the third and fourth tooth, and the sagittal crest of the mesonotum differs by having a prominent anterior spine with the remainder of the sagittal crest having weakly formed tubercles (Fig.
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).
Male. Coloration. Coloration description based on the dried type specimens. Coloration is pale green with variable areas of straw yellow throughout likely due to the drying process, particularly if alcohol was used to help preserve the specimens (Fig.
Morphology. Head. Head capsule ca ¼ longer than wide, with a vertex that is slightly lumpy, but smooth, lacking distinct granulation; posteromedial tubercle is not prominent, only slightly raised above the head capsule (Fig.
Measurements of holotype male Coll RC 17-221 [mm]. Length of body (including cerci and head, excluding antennae) 50.3, length/width of head 2.8/2.9, antennae 27.3, pronotum 2.8, mesonotum 3.8, length/width of tegmina 18.4/5.6, length of alae 35.8, greatest width of abdomen 10.7, profemora 10.4, mesofemora 8.9, metafemora 11.2, protibiae 7.4, mesotibiae 7.1, metatibiae 8.5.
Measurements of paratype male Coll RC 17-222 [mm]. Length of body (including cerci and head, excluding antennae) 52.1, length/width of head 2.8/2.9, antennae damaged and missing most segments, pronotum 2.8, mesonotum 3.8, length/width of tegmina 18.6/5.7, length of alae 36.4, greatest width of abdomen 10.8, profemora 10.4, mesofemora 8.8, metafemora 11.3, protibiae 7.4, mesotibiae 7.1, metatibiae 8.6.
Noun, Alangan in origin, meaning “shy leaf”. We wish to honor the original inhabitants of the area that this species is native to by using a language local to northern Mindoro. We choose the threatened language, Alangan, which is spoken by as few as 2,150 speakers in north-central Mindoro (according to Ethnologue: Languages of the World; ethnologue.com). The name is formed from the Alangan words iyá (meaning shy) + daon (meaning leaf;
Currently only known from the type locality Puerto Galera, Mindoro Island, Philippines.
Mindoro, as a smaller island, was originally thought to only have one Phyllium species present, Phyllium mindorense Hennemann, Conle, Gottardo & Bresseel, 2009, and based upon examination of multiple specimens from this island, Phyllium mindorense appears to be the most commonly collected species. However, a morphologically distinct set of males which did not match with the known male Phyllium mindorense morphology were reviewed. These specimens were included within the analyses of
One possible identification which was explored is that these could represent the unknown male sex of Phyllium bilobatum Gray, 1843, which unfortunately is only known from a single holotype specimen and the inexact locality of “Philippine Islands” (
Holotype
♀: “Philippines, Mindanao, Lanao Del Sur, Wao, February 2019; Collection SLT; DNA no. 35”. Deposited in the Montreal Insectarium, Quebec, Canada (IMQC; Fig.
Females are morphologically most similar to Phyllium bilobatum due to their small size, strongly lobed abdomen, broad profemoral exterior lobe, large triangular teeth of the profemoral interior lobe, and similar thorax spination. The antennae morphology can differentiate these species as Phyllium ortizi sp. nov. has the terminal antennal segment notably longer (approximately the same length as the preceding two and a half or three segments; Fig.
For male Phyllium ortizi sp. nov. they are morphologically most similar to Phyllium mabantai and Phyllium iyadaon sp. nov. based upon their size, wing venison, and general lobes of the legs. The abdomen of Phyllium ortizi sp. nov. appears to be relatively stable in shape based upon reared and wild collected specimens and within Phyllium mabantai male abdominal shape is also rather stable despite female Phyllium mabantai having drastically different forms. Between these two species the abdominal general shape and the size of the eye sports appears to be a reliable feature for differentiation as the Phyllium mabantai abdomen has smooth abdominal edges (either giving the abdomen a spade shaped or slightly ovoid appearance) and abdominal segment V has small eye spots vs Phyllium ortizi sp. nov. which has somewhat of a rectangular abdomen (with segments V and VI approximately even in width) and the margins of VI, VII, and VIII gently rounded giving the abdomen a slightly scalloped edge and the eye spots of abdominal segment V are always large taking up at least ½ of the segment length (Fig.
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.
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.
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).
The eggs of Phyllium ortizi sp. nov. are rather unique as they have two general types of pinnae on their surfaces (of which all surfaces have the same types) some that are the more typical feather-like pinnae and a second type which is thinner and more filament-like, lacking projections like is seen in the feather-like pinnae. These thinner filament-like pinnae are not seen in other phylliids eggs, and the closest types are those seen in Phyllium tobeloense bhaskarai
Freshly hatched nymphs are only known for a handful of Phyllium species at present, but from what is known Phyllium ortizi sp. nov. can be differentiated from congenerics reliably. Phyllium ortizi sp. nov. (Figs
Female. Coloration. Coloration description is based upon living individuals in culture (Fig.
Morphology. Head. Head capsule slightly longer than wide, vertex moderately wrinkled with evenly spaced, small nodes across most of the surface (most distinct on the posterior ½), with the posteromedial tubercle the most prominent feature on the head capsule, many times larger and broader than any of the nodes (Fig.
Measurements of holotype female [mm]. Length of body (including cerci and head, excluding antennae) 66.8, length/width of head 6.0/5.4, antennae (broken in the holotype), pronotum 5.9, mesonotum 6.2, length of tegmina 43.5, greatest width of abdomen 31.6, profemora 12.3, mesofemora 11.0, metafemora 13.8, protibiae 7.3, mesotibiae 8.5, metatibiae 12.1.
Measurements of paratype females [mm] (ex culture). Length of body (including cerci and head, excluding antennae) 74.2–76.4, length/width of head 6.2–6.4/5.9– 6.1, antennae 4.1–4.4, pronotum 3.9–4.2, mesonotum 6.4–6.6, length of tegmina 47.8–48.2, length of alae 2.4–2.6, greatest width of abdomen 35.5–36.8, profemora 14.0–14.4, mesofemora 12.8–13.2, metafemora 15.7–16.1, protibiae 7.5–8.2, mesotibiae 8.4–8.9, metatibiae 13.7–14.1.
Male. Coloration. Coloration description is based upon living specimens in captivity (Fig.
Morphology. Head. Head capsule approximately as long as wide, with a vertex that is slightly lumpy and marked by a few irregularly spaced nodes (Fig.
Measurements of paratype males [mm] (ex culture). Length of body (including cerci and head, excluding antennae) 53.5–55.8, length/width of head 2.9–3.0/2.8– 2.9, antennae 2.6.1–27.9, pronotum 2.3–2.4, mesonotum 2.8–2.9, length of tegmina 17.6–19.0, length of alae 39.7–41.2, greatest width of abdomen 12.3–12.9, profemora 8.7–9.4, mesofemora 8.9–9.3, metafemora 10.4–10.8, protibiae 5.8–6.2, mesotibiae 5.3–6.0, metatibiae 7.8–7.9.
Eggs. (Fig.
Measurements including the extended pinnae [mm]. Length (including operculum): 4.5–4.6; maximum width of capsule when viewed from lateral aspect 2.9–3.0; length of micropylar plate 2.2–2.3.
(Fig.
Patronym. Named after Maxime Ortiz (France) who is a well-known phasmid breeder who established this new species in culture and who shared type material with the authors for this study. Thanks to his efforts we were able to understand the morphological diversity within the males, females, freshly hatched nymphs, and eggs.
At present this species is known from four provinces in Mindanao; Lanao Del Sur Province (with the exact type locality of Wao Municipality), South Cotabato Province, Davao del Sur Province, and from the inexact locality of Bukidnon Province from captive reared specimens.
This species was first thought to represent a small, strongly lobed Phyllium mabantai or possibly the first record of Phyllium bilobatum Gray, 1843 in more than a century, a species with which Phyllium ortizi sp. nov. shares many general similarities. However, thanks to the rearing of additional specimens by Maxime Ortiz (France), the intraspecific variation was revealed, and the fine details instead reliably differentiate these species. Phyllium ortizi sp. nov. was sampled within
The eggs of Phyllium ortizi sp. nov. are primarily covered with pinnae “type 5” (feather-like) as defined by
Pulchriphyllium pulchrifolium (Audinet-Serville, 1838).
Historically, many Pulchriphyllium populations have been the subject of repeated taxonomic adjustments, oftentimes without being properly assessed or clearly defined. Originally, many of the populations were treated as distinct species, but eventually, as phylliids became better known and more common in museums, authors started to assume phylliids were species-poor with wide geographic ranges (
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
Within the Pulchriphyllium there appear to be several recurring, leaf-shape mimicry patterns; broad leaf (like in Pulchriphyllium giganteum), boxy leaf (like in Pulchriphyllium pulchrifolium), and tapered leaf (like in Pulchriphyllium bioculatum Gray, 1832). Of particular difficulty to differentiate are the numerous species where females have a tapered abdominal shape and males have rounded/ovoid abdomen, herein referred to as “bioculatum-like species”. These “bioculatum-like species” are a subset of the “bioculatum species group”, the only species group within the Pulchriphyllium proposed by
The “bioculatum-like species” are difficult to morphologically differentiate, have wide-ranging geographic distributions (Fig.
(♀): “Silhet” (OUMNH; Fig.
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.
Presently, Pulchriphyllium scythe stat. rev., comb. nov. is one of the species we lack molecular data for, but thankfully there are consistent morphological differences which allow differentiation from congenerics. At the moment, the morphology of the freshly hatched nymph and the egg are not known, which would likely yield additional features for differentiation. The female Pulchriphyllium scythe stat. rev., comb. nov. are the easiest to consistently differentiate, and when coupled with their geographic isolation, are the reason why we are reinstating this population as a valid species.
From all other bioculatum-like species, Pulchriphyllium scythe stat. rev., comb. nov. females can be differentiated by their notably larger size (10.8 to 11.0 cm (vs the other bioculatum-like species which range from 6.7 to 9.5 cm long). Females of Pulchriphyllium scythe stat. rev., comb. nov. are so large they actually fall within the size range of Pulchriphyllium giganteum (Hausleithner, 1984) (but can easily be differentiated by the mesopleural spination as Pulchriphyllium scythe stat. rev., comb. nov. does not have the notable medial projection like in Pulchriphyllium giganteum). Male Pulchriphyllium scythe stat. rev., comb. nov. are poorly known (few males have been located in museum collections and few living observations have been located, coupled with the original male syntype specimen being lost), therefore, differences have been difficult to identify as consistent. At the moment the most notable feature we have been able to identify is that the profemoral exterior lobe appears to be slightly narrower than in most bioculatum-like species, giving the exterior lobe a more obtuse angle appearance.
Thanks to iNaturalist (Figs
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.
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.
The only remaining syntype female (Fig.
Pulchriphyllium scythe stat. rev., comb. nov. is an uncommonly recorded taxon, which has mostly been recognized as a valid species by some authors, and as a synonym or subspecies/form by others (
(2♀♀, 2♂♂; OUMNH).
Lectotype designation
: 1♀: “crurifolium ♀ Seychelles; Seychelles; Wings O; e coll. Serville” (Fig.
Paralectotype designations
: 1♂: “Seychelles; phyllium crurifolium Serville ♂; Phyllium crurifolium, teste Serville. E coll. Marchal, Mae. Hist. Nat. Orth. p. 292” (Fig.
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.
“Phyllium dardanus” Holotype (1♂; Fig.
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 (
Phyllium gelonus
Gray, 1843 (
Female Pulchriphyllium crurifolium stat. rev., comb. nov. are the easiest to differentiate from the other “bioculatum”-like species by comparing the profemoral exterior lobe shape. In Pulchriphyllium crurifolium stat. rev., comb. nov. the anterior margin of the exterior profemoral lobe is nearly straight, giving the lobe a boxy appearance (Fig.
Adult female profemoral lobes from various Pulchriphyllium species for comparison A Pulchriphyllium crurifolium stat. rev., comb. nov. (
Males are difficult to differentiate, and even after reviewing numerous specimens in series there appears to be enough intraspecific variation to make reliable differentiation impossible. On average males tend to have slightly narrower profemoral exterior lobes with a strongly obtuse angle (Fig.
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).
Eggs of Pulchriphyllium crurifolium stat. rev., comb. nov. are slightly smaller than the other “bioculatum”-like species and have a great deal more raised nodes throughout the surfaces (Fig.
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.
To date, the authors have yet to see an observation of a freshly hatched nymph of this taxon. With how unique and species-specific congenerics are (Fig.
(Fig.
This taxon was originally described as a unique species from the Seychelles (
In the decades following the description of Pulchriphyllium crurifolium stat. rev., comb. nov., there were several other taxa described from the Seychelles, but since their description they have been synonymized with Pulchriphyllium bioculatum (
The eggs of Pulchriphyllium crurifolium stat. rev., comb. nov. are very similar to most Pulchriphyllium species therefore this species adds credibility to the phylogeny of
Holotype
♂: “Vietnam: Da Nang Province, Ba Na Mt. 1,450 m. elv. May 2015; Coll RC 16-007” (Fig.
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.
Female, freshly hatched nymph, and egg unknown. Males are morphologically most similar to Pulchriphyllium maethoraniae (Delfossee, 2015) and Pulchriphyllium sinense (Liu, 1990). These three species are difficult to morphologically differentiate, especially because Pulchriphyllium maethoraniae and Pulchriphyllium sinense males are only known from a few specimens, therefore the intraspecific variation is not well understood. On average it appears the mainland species (Pulchriphyllium maethoraniae and Pulchriphyllium heracles sp. nov.) are slightly larger than Pulchriphyllium sinense from Hainan Island, but that could simply be due to our limited knowledge of Pulchriphyllium sinense variation. One morphological feature which does appear to be consistent for differentiating Pulchriphyllium heracles sp. nov. from the other two is the interior lobe of the mesofemora. In Pulchriphyllium heracles sp. nov. the lobe is slightly thinner, with a maximum width of ca 1 ¼ the width of the mesofemoral shaft, whereas in the other two species this lobe is broader, ca 1½× as wide or wider.
Male. Coloration. Coloration description based on the dried type specimens, but despite slight discoloration from the preservation process, the natural patterns appear rather consistent. The coloration throughout is pale green with brown highlights/markings (Fig.
Morphology. Head. Head capsule ca ¼ longer than wide, with a vertex that is smooth, lacking distinct granulation; the posteromedial tubercle is not prominent, only slightly raised above the head capsule (Fig.
Measurements of holotype male Coll RC 16-007 [mm]. Length of body (including cerci and head, excluding antennae) 66.0, length/width of head 4.1/3.7, antennae 27.3, pronotum 2.9, mesonotum 3.6, length/width of tegmina 13.8/5.1, length of alae 52.3, greatest width of abdomen 25.0, profemora 14.0, mesofemora 9.7, metafemora 10.2, protibiae 8.1, mesotibiae 7.0, metatibiae 8.6.
Measurements of paratype males, given as a range for smallest to largest [mm]. Length of body (including cerci and head, excluding antennae) 59.4–71.4, length/width of head 4.0–4.5/3.3–4.0, antennae 25.4–27.6, pronotum 3.0–3.7, mesonotum 3.3–5.1, length/width of tegmina 11.9–15.7/4.8–6.0, length of alae 45.0–57.2, greatest width of abdomen 24.2–26.7, profemora 11.4–14.6, mesofemora 9.0–9.3, metafemora 9.0–9.5, protibiae 6.9–8.0, mesotibiae 6.7–7.8, metatibiae 6.6–8.6.
Proper noun, Greek in origin. Following a mythological trend started by
Records for this species are primarily from southern and central Vietnam (from the provinces of Kon Tum, Da Nang, Gia Lai, Lam Dong, Quang Nam, and Thua Thien Hue) and additionally one record from the northern Vietnam province of Nghe An is known (Fig.
Specimens of this Vietnamese population were originally thought to represent range expansions for Pulchriphyllium maethoraniae. Therefore, this population was tentatively identified within
Interestingly, despite near identical morphology with Pulchriphyllium maethoraniae and Pulchriphyllium sinense, Pulchriphyllium heracles sp. nov. has been recovered in the molecular phylogeny as entirely unrelated to Pulchriphyllium maethoraniae and Pulchriphyllium sinense (which were recovered as sister to each other; Fig.
Holotype
♀: “Indonesia, West Java, Sukabumi, Kabandungan Village, Mt. Halimun, found 19th August 2020, first started laying eggs 16th September 2020, died 8th December 2020; laid a total of 78 eggs” (Fig.
Adult males and females have proven difficult to differentiate morphologically from Pulchriphyllium giganteum. Within both sexes of both species there is significant overlap in overall size and due to intraspecific variation, few to no features have allowed confident differentiation. Instead, the only reliable and easily observed morphological differences are the freshly hatched nymphs and egg morphology.
Females are very similar to Pulchriphyllium giganteum but can be differentiated by the mesopleurae. In Pulchriphyllium giganteum the posterior two or three spines are grouped together as a broad set projecting away from the mesopleuron margin, while in Pulchriphyllium bhaskarai sp. nov. these posterior-most spines are less prominent, only slightly projecting away from the mesopleuron margin (Fig.
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).
Males (Fig.
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).
Freshly hatched nymphs (Figs
The eggs of Pulchriphyllium bhaskarai sp. nov. are rather unique and can easily differentiate this species from Pulchriphyllium giganteum by the presence of five laterally running semi-hollow tubes which are open on the anterior end (two tubes are located on each of the lateral surfaces and one tube is located on the ventral surface: Fig.
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).
Female. Coloration. The coloration description is based upon the holotype living individual when it was alive (Fig.
Morphology. Head. Head capsule is slightly longer than wide, with a vertex that is marked throughout by moderately formed granulation and a singularly pointed posteromedial tubercle which is notably larger than the other granulation on the head capsule (Fig.
Measurements of holotype female [mm]. Length of body (including cerci and head, excluding antennae) 97.0, length/width of head 8.3/7.1, antennae 5.2, pronotum 6.2, mesonotum 4.9, length of tegmina 55.5, greatest width of abdomen 50.7, profemora 21.3, mesofemora 15.3, metafemora 18.9, protibiae 10.6, mesotibiae 11.1, metatibiae 15.6.
Measurements of paratype females [mm]. Length of body (including cerci and head, excluding antennae) 95.6–114.5, length/width of head 8.2–10.8/7.0–8.3, antennae 5.1–6.2, pronotum 6.1–6.8, mesonotum 4.8–5.5, length of tegmina 55.3–67.0, greatest width of abdomen 50.5–56.1, profemora 21.0–27.6, mesofemora 15.2–17.5, metafemora 18.7–21.3, protibiae 10.5–12.6, mesotibiae 11.0–13.0, metatibiae 15.4–17.8.
Male. Coloration. Coloration description based upon images of live specimens. Overall coloration is pale green throughout with somewhat variable patches of tan, brown, or black (Fig.
Morphology. Head. Head capsule approximately as wide as long, with a vertex that is slightly wrinkled/rough textured. The posteromedial tubercle is small but distinctly raised above the head capsule. Frontal convexity stout and ending with a blunt point and marked with only a few short setae. Compound eyes bulbous, occupying slightly > ⅖ of the head capsule lateral margins (Fig.
Measurements of paratype males [mm]. Length of body (including cerci and head, excluding antennae) 73.4–74.0, length/width of head 4.5–4.6/2.1–2.2, antennae 25.7–25.9, pronotum 3.2–3.3, mesonotum 2.9–3.0, length of tegmina 14.3–14.5, length of alae 56.8–57.0, greatest width of abdomen 29.0–29.2, profemora 14.8–14.9, mesofemora 10.5–10.6, metafemora 11.2–11.3, protibiae 7.4–7.6, mesotibiae 6.9–7.0, metatibiae 9.0–9.2.
Eggs. (Fig.
Measurements [mm]. Length (including operculum): 7.9–8.0; maximum width of capsule when viewed from lateral aspect 4.2–4.3; length of micropylar plate 5.0–5.1.
(Fig.
Patronym. Named to honor Edy Bhaskara who supplied the specimens which solved the mystery surrounding this species. Previously only males were known, which are morphologically indistinguishable from Pulchriphyllium giganteum, until Edy Bhaskara supplied the authors with fresh specimens (which allowed DNA analyses) as well as adult females, freshly hatched nymphs, and eggs, all of which allowed a full understanding to be generated for this species and allowed differentiation from congenerics.
At present only known from Java, Indonesia, with records from throughout the island (Suppl. material
Originally, it was thought that the “giganteum”-like population on Java, Indonesia was simply a range expansion for Pulchriphyllium giganteum (
Within the DNA-based analyses of
Thanks to DNA analyses, the geographic distribution of Pulchriphyllium giganteum has been clarified to encompass West Malaysia, West Borneo, and North Borneo as a single species. Unfortunately, Sumatra and Nias island leave an area of significant uncertainty as no DNA samples are yet available, and to date only male specimens have been observed/collected (which look identical to both Pulchriphyllium giganteum and Pulchriphyllium bhaskarai sp. nov.). Interestingly, despite Pulchriphyllium bhaskarai sp. nov. being presently understood as endemic to Java, Pulchriphyllium giganteum has a wide range from South Thailand, West Malaysia, and throughout Borneo (
The eggs of Pulchriphyllium bhaskarai sp. nov. are quite unique, as they lack the typical Pulchriphyllium dense fins, and instead have rigid, open tubes running along the egg surfaces (Fig.
Holotype
♂: “WS 415; India: Karnataka, Agumbe Ghats, Canopy Light Trap, N 13 29.386’ E 75 04.537’ 10-X-2004” (Fig.
Pulchriphyllium anangu sp. nov. are morphologically most similar to Pulchriphyllium bioculatum and Pulchriphyllium agathyrsus, of which the adult morphology is difficult to differentiate, but the freshly hatched nymphs (Fig.
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.
Female Pulchriphyllium anangu sp. nov. (Fig.
Male Pulchriphyllium anangu sp. nov. has proven impossible to differentiate morphologically from Pulchriphyllium agathyrsus due to notable coloration and abdominal shape variation and therefore no consistent differences have been identified (Fig.
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.
Freshly hatched nymphs of Pulchriphyllium anangu sp. nov. are very similar to Pulchriphyllium bioculatum and only subtle differences have been identified in their coloration. Typically, the overall coloration of Pulchriphyllium bioculatum tends to be a bit darker (Fig.
Female. Coloration. Coloration description is based upon numerous images of live individuals from iNaturalist (Fig.
Morphology. Head. Head capsule is slightly longer than wide, with a vertex that is marked throughout by moderately formed granulation (that is typically a paler green than the head capsule base color) and a posteromedial tubercle which is notably larger than the other granulation (and is often brown in color). Frontal convexity finely pointed, with several short setae on the apex. Compound eyes slightly protruding from the head capsule, not overly large, taking up ca ¼ of the head capsule lateral margins. Ocelli absent. Antennal fields slightly wider than the width of the first antennomere. Antennae. Antennae consist of nine segments (including scapus and pedicellus), with the terminal segment approximately the same length as the preceding 2⅓ segments’ lengths combined. Antennomeres I–VIII are marked with slight granulation and sparse short setae. Thorax. Pronotum with slightly convex anterior margin and gently converging lateral margins, which converge to the posterior margin that is ⅔ the width of the anterior margin. The pronotum surface is slightly lumpy and marked with a distinct sagittal furrow and slight central perpendicular furrow. The pronotum has distinct, smooth anterior and lateral rims. Prosternum, mesosternum, and metanotum are covered sparsely with nodes that are irregularly spaced throughout the surface, but all relatively small. Prescutum approximately as long as wide, with lateral rims marked by 10–12 small tubercles relatively evenly sized and spaced, giving the lateral rims a rough appearance. The prescutum anterior rim is distinct but not strongly protruding and the rim surface is relatively smooth. Prescutum surface slightly raised along the sagittal plane and the surface is only slightly marked with granulation. The mesopleura start to diverge slightly posterior to the prescutum anterior rim and diverge uniformly with straight margins. The mesopleuron margin is marked throughout with five or six larger tubercles and six or seven smaller tubercles interspersed throughout. Mesopleuron surface slightly granular. Wings. Tegmina long, typically reaching significantly onto abdominal segments VII or VIII. Tegmen venation; the subcosta (Sc) is the first vein in the forewing and runs parallel with the margin for the anterior ½, and then bends slightly and runs towards the margin where it terminates ca ¼ of the way through the tegmen length. The radius (R) spans the central portion of the forewing with two subparallel branched veins; the first radius (R1) branches ca ¼ of the way through the tegmen length and terminates slightly proximal to the midline, and the radial sector (Rs) branches ca ⅓ of the way through the tegmen length and terminates ca ⅗ of the way through the length. The media (M) is bifurcate with media anterior (MA) splitting near the middle of the tegmen length and the media posterior (MP) splits ca ⅗ of the way through the length. The media anterior and media posterior terminate near to the posterior ¼ of the tegmen after arcing gently and running parallel with each other. The cubitus (Cu) is bifurcate, branching near the posterior ¼ of the wing into the cubitus anterior (CuA) and cubitus posterior (CuP) which both terminate near the tegmen apex. The first anal vein (1A) is simple and fuses with the cubitus ca ¼ of the way through the tegmen length. Ala rudimentary, no more than a nub. Abdomen. Abdominal segments II through the anterior ⅔ of segment IV diverge strongly, the remainder of IV diverges only slightly to segment V which is the widest segment, followed by segments VI and VII which converge slightly with straight margins to VIII through X which converge more significantly to the broadly rounded apex. Genitalia. Subgenital plate starts on the anterior margin of tergum VIII, is moderately broad, and extends to the anterior margin of tergum X, ending in a blunt apex. Gonapophyses VIII are moderate in length, extending halfway onto tergum X, and are moderately broad; gonapophyses IX are smaller and hidden under gonapophyses VIII. The cerci are flat, not strongly cupped, with a granular surface and numerous small setae. Legs. Profemoral exterior lobe broad, and somewhat recurved with a slightly acute angle and a greatest width ca 5 ½ to 6× the width of the profemoral shaft. Profemoral exterior lobe proximal margin is marked with four or five distinct serrate teeth, and the distal margin is smooth, lacking serration. Profemoral interior lobe is only situated on the distal ⅔ of the profemora, ca 3× as wide as the greatest width of the profemoral shaft and is roundly arcing. The proximal margin of the profemoral interior lobe is relatively smooth and the distal margin is marked with three or four dulled, weakly formed teeth with looping gaps between them. The mesofemoral exterior lobe is roundly triangular and arcs from end to end. The mesofemoral exterior lobe proximal margin is straight and smooth, while the distal margin is slightly rounded and marked with four or five serrate teeth. The mesofemoral interior lobe is slightly wider than the mesofemoral shaft, and the exterior lobe is ca 1½× wider than the mesofemoral shaft. The mesofemoral interior lobe arcs unevenly end to end with seven or eight dulled teeth throughout the length, and the distal end slightly wider than the proximal end. The metafemoral exterior lobe is thin and smooth, hugging the metafemoral shaft and lacks dentition. The metafemoral exterior lobe is thin on the proximal ⅓ and then a gently arcing lobe on the distal ⅔ approximately as wide as the metafemoral shaft width. The metafemoral exterior lobe has moderate serration throughout with those on the distal ½ more prominent. Protibial exterior lobe unevenly arcing with a smooth margin from end to end with a steady increase in width from the proximal to the distal ends. Protibial interior lobe also spans the entire length in a smooth, rounded triangle, with the widest point on the distal ⅓. Mesotibiae and metatibiae lacking interior lobes, with exterior lobes having a similar rounded triangular shape weighted towards the distal end and ca 2× as wide as the shafts on which they are found.
Measurements of paratype females [mm]. Length of body (including cerci and head, excluding antennae) 76.6–78.7, length/width of head 8.9–9.4/6.9–7.1, antennae 4.0–4.1, pronotum 5.2–5.5, mesonotum 4.5–4.7, length of tegmina 44.7–47.6, greatest width of abdomen 34.9–35.4, profemora 19.5–19.9, mesofemora 14.0–14.4, metafemora 15.8–16.1, protibiae 9.8–10.0, mesotibiae 9.0–9.4, metatibiae 11.9–12.2.
Male. Coloration. Coloration description based upon numerous images of live individuals recorded on iNaturalist (Fig.
Morphology. Head. Head capsule approximately as long as wide, with a vertex that is weakly granular; posteromedial tubercle small but distinctly raised from the capsule surface (Fig.
Measurements of holotype male [mm]. Length of body (including cerci and head, excluding antennae) 63.1, length/width of head 3.7/3.9, antennae (missing from holotype specimen), pronotum 3.1, mesonotum 2.7, length of tegmina 13.2, length of alae 49.1, greatest width of abdomen 22.5, profemora 14.2, mesofemora 11.6, metafemora 9.2, protibiae 6.9, mesotibiae 6.8, metatibiae 9.1.
(Fig.
Proper noun, named after Anangu, one of the Yakshi, or tree nymphs of Tamil mythology from southern India (
At present known from multiple states in southwestern India (Fig.
Despite being an undescribed species, leaf insect observations from India are commonly recorded on the citizen science platform iNaturalist. These submitted observations allowed a significant look into adult male (Fig.
Within the southwestern Indian state of Kerala, the Malayali people call this species “Pera Rani” (meaning “Queen of Guava plant” in Malayalam) a fitting common name as phylliids are frequently observed by farmers of guava (pers. comm. Gavas Ragesh, Kerala Agricultural University). Local farmers additionally report this species as being observed on cashew and mango trees, but never in significant enough populations to be considered a pest. Of these three common agricultural plants, only mango is native to Asia (with the other two native to the tropics of the western hemisphere;
Holotype
♀: “Indonesia: South Kalimantan, Mt. Meratus, VIII.2020; DNA sample SLT050” (Fig.
Paratype
♀: “Indonesia, Kalimantan, Mt. Besar VII.2018; DNA sample SLT006” (Fig.
Males, freshly hatched nymphs, and eggs are currently unknown. At present only known from two female specimens, therefore little is known of the intraspecific variation of this species. Additionally, throughout the “bioculatum”-like species ranges other species show a decent amount of variation in their abdominal shapes/femoral lobe spination, so a reliable morphological feature has yet to be identified for differentiation. Also, with two congenerics from Borneo with the “bioculatum”-like morphology only known from males, and this species only known from a female, nothing can be said about morphological differentiation presently for these Bornean species. Only DNA analyses have allowed differentiation at this time (Fig.
Female. Coloration. The coloration description is based upon the dried type specimens which appears to have been well-preserved with minimal discoloration (Fig.
Morphology. Head. Head capsule is as long as wide, with a vertex that is marked throughout by weakly formed granulation (Fig.
Measurements of holotype female [mm]. Length of body (including cerci and head, excluding antennae) 84.5, length/width of head 7.6/6.0, antennae 3.8, pronotum 5.4, mesonotum 4.7, length of tegmina 50.6, greatest width of abdomen 48.9, profemora 18.5, mesofemora 8.4, metafemora 14.7, protibiae 8.0, mesotibiae 7.5, metatibiae 13.3.
Measurements of paratype female [mm]. Length of body (including cerci and head, excluding antennae) 78.9, length/width of head 7.7/6.2, antennae 4.5, pronotum 5.4, mesonotum 4.8, length of tegmina 44.1, greatest width of abdomen 40.0, profemora 15.6, mesofemora (missing from paratype), metafemora 15.2, protibiae 9.5, mesotibiae (missing from paratype), metatibiae 12.6.
Patronym, named after Gilles Delisle (Canada). Friend of the authors and enthusiastic entomologist. Gilles Delisle is a birdwing butterfly specialist and research associate of the Montreal Insectarium. His passion for birdwing butterflies allowed him to travel many times throughout Indonesia and Papua New Guinea. He donated his butterfly collection after many years of studies to the Montreal Insectarium, Canada.
At present only known from the localities of Mt. Meratus and Mt. Besar, Kalimantan, Indonesia.
When the specimen was preliminarily reviewed, three possible identifications were assumed. First, it was considered that this might represent the unknown female Pulchriphyllium agnesagamaae or Pulchriphyllium fredkugani (both species currently only known from males from northern Borneo). Or secondly, that this specimen might simply be a range expansion for Pulchriphyllium bioculatum, a species which has been confirmed via molecular analyses to occur on southwestern Borneo as well as peninsular Malaysia (Fig.
Due to the limited sampling of leaf insect specimens in collections and the challenges of species delimitation, often poorly known/rarely encountered species are misidentified as members of better-known clades. Additionally, the pronounced sexual dimorphism has sometimes led to males and females having been occasionally mistakenly identified as separate species. Here, we have described seven new phylliid species: Phyllium iyadaon sp. nov. from Mindoro Island, Philippines; Phyllium samarense sp. nov. from Samar Island, Philippines; Phyllium ortizi sp. nov. from Mindanao Island, Philippines; Pulchriphyllium heracles sp. nov. from Vietnam; Pulchriphyllium delislei sp. nov. from South Kalimantan, Indonesia; Pulchriphyllium bhaskarai sp. nov. from Java, Indonesia; and Pulchriphyllium anangu sp. nov. from southwestern India. These species were uncovered within the phylogenetic tree inferred by
Molecular work has been especially useful in the differentiation within Pulchriphyllium. Historically, many species with “bioculatum”-like morphology (females that have tapered abdomens and males with rounded/ovoid abdomens) have been called forms or subspecies of Pulchriphyllium bioculatum. This previous classification of Pulchriphyllium bioculatum had the species occurring within a markedly wide geographic range, from the Seychelles to north India and Bangladesh, through mainland Asia to Thailand, peninsular Malaysia, and south into Sumatra. However, DNA sequencing data has revealed that what has previously been considered “bioculatum” does not form a monophyletic group, and instead the true Pulchriphyllium bioculatum (from West Malaysia and adjacent areas: Fig.
These newly described species not only help to expand our knowledge of phylliid diversity, but also help to highlight recent and yet to be undertaken areas of research. For example, investigation into the morphological and functional diversity of phylliid eggs is revealing interesting stories of abiotic interactions and biotic mimicry (
In addition to describing new species based upon freshly collected material, the extensive review of type specimens from historic, little-known taxa has allowed the designation of lectotype specimens and clarified the status of Pulchriphyllium scythe and Pulchriphyllium crurifolium as distinct species to reflect their unique morphology and geographic isolation. Continued sampling and molecular sequencing will be crucial in resolving remaining uncertainties and identifying new species in the future.
Thank you to many individuals who shared photos, specimen data, and specimens with us for this work: Maxime Ortiz (France), Edy Bhaskara (Indonesia), Frank Hennemann (Germany), Robert Douglas (Oxford), Judith Marshall (
The authors have declared that no competing interests exist.
No ethical statement was reported.
This study was supported by the German Research Foundation (DFG grants BR 2930/4-2 and BR 2930/7-1) and the National Science Foundation (DEB- 1557114 to Michael F. Whiting and Sven Bradler).
Conceptualization: RTC, SBr, SBa. Investigation: RTC, SLT, SBr, SBa. Resources: SLT, SBr, SBa, NECE, MFW. Visualization: RTC, SLT, JBL, NECE. Writing - original draft: RTC, MES, EMF, SBa. Writing - review and editing: RTC, SLT, JBL, MES, EMF, MFW, SBr, SBa.
Royce T. Cumming https://orcid.org/0000-0001-7930-1292
Stéphane Le Tirant https://orcid.org/0000-0001-7463-5621
Jackson B. Linde https://orcid.org/0000-0003-1480-0212
Megan E. Solan https://orcid.org/0000-0002-2298-5336
Evelyn Marie Foley https://orcid.org/0009-0007-9754-212X
Ramon Lavado https://orcid.org/0000-0002-2774-5104
Michael F. Whiting https://orcid.org/0000-0003-1157-888X
Sven Bradler https://orcid.org/0000-0001-9307-1032
Sarah Bank https://orcid.org/0000-0001-6952-1590
All of the data that support the findings of this study are available in the main text or Supplementary Information.
Examined specimens and paratype data with depositories
Data type: Specimen/observation data
Explanation note: Taxon information for each specimens examined including sex, type designation, sample ID, collection data, and links to observational data available online.
All sampled specimens are listed with their collection data, deposition location, and GenBank accession numbers for the generated sequences.
Data type: Specimen data
Explanation note: Taxon information for each specimen used in the phylogenetic analysis including sex, type designation, sample ID, sampling site, and GenBank accession numbers. Newly generated molecular data in bold.
Multiple sequence alignment of concatenated dataset
Data type: phylogenetic data
Full-tree figure with outgroups
Data type: Resulting tree based on re-analysis of the phylogenetic relationships of the 169 taxa from
Explanation note: Fully recovered tree with outgroups.