Abstract
The genus Sadyattes Stål, 1875 is a poorly understood clade of stick insects of the subfamily Platycraninae and tribe Stephanacridini, that currently comprises three species from Borneo as well as type-species S. borrii Stål, 1875 whose locality is as yet a mystery. In its present recognition the genus is here shown to be paraphyletic and research has revealed that a number of species currently attributed to other genera, one of which is not even closely related, actually belong in Sadyattes. To elucidate these shortcomings a taxonomic review of Sadyattes at the species level is presented along with a generic redescription. Sadyattes is distributed throughout the land masses of the Sunda Shelf (Borneo, Peninsular Malaysia, Java and Sumatra with some of its surrounding islands) and the Philippine Islands. Differential diagnoses as well as notes on intraspecific variability and the distributions are provided for all known species, including new descriptions and combinations, with most species also illustrated. Keys are presented to distinguish females and males. The genus Eucarcharus Brunner v. Wattenwyl, 1907 is found to be a synonym of Sadyattes and thus synonymised (syn. nov.). The two Philippine species previously contained in Eucarcharus, E. fallax Brunner v. Wattenwyl, 1907 and E. feruloides (Westwood, 1859), are automatically transferred to Sadyattes, which results in two new generic combinations (comb. nov.). Four species are transferred to Sadyattes from other genera, which results in another four new generic combinations (comb. nov.): the Bornean S. decoris (Seow-Choen, 2016) and S. incertus (Brunner v. Wattenwyl, 1907) from Peninsular Malaysia, Bangka Island and the Andaman Islands from the genus Phobaeticus Brunner v. Wattenwyl, 1907 (Clitumninae: Pharnaciini), the Bornean S. nigricornis (Redtenbacher, 1908) from the genus Pharnacia Stål, 1877 (Clitumninae: Pharnaciini), and the Philippine S. leytensis (Zompro, 1997) from the genus Hermarchus Stål, 1875 (Platycraninae: Stephanacridini). Six new species from the Philippines are described: S. banwaon sp. nov. from the island of Mindanao from the female, S. maganda sp. nov. from the island of Mindoro from the female and male, S. matipuno sp. nov. from the island of Luzon from the female, male and egg, S. mindanaense sp. nov. from the island of Mindanao from the female, male and egg, S. panayense sp. nov. from the island of Panay from the female only, and S. tubaense sp. nov. from the island of Luzon from the male. A redescription of the male of the type-species S. borrii Stål, 1875 is presented, whose distribution is here shown to comprise Java and Sumatra, along with a description of the previously unrecorded female, which has proven to be represented by Nesiophasma zanus Hennemann, 1998 (syn. nov.). This latter synonymy renders the questionable locality “New Guinea” of Nesiophasma zanus as erroneous. The previously unknown males of S. feruloides comb. nov., S. fallax comb. nov., and S. incertus comb. nov. are described and those of the latter two species are illustrated for the first time. Also, the eggs of S. fallax comb. nov. and S. feruloides comb. nov. are described and illustrated for the first time. Nearchus grubaueri Redtenbacher, 1908 is re-synonymised with S. incertus (Brunner v. Wattenwyl, 1907; rev. syn.). Holotypes of the newly described species are deposited in the collections of Institute Royal des Sciences Naturelles de Belgique, Brussels, Belgium (RBINS) and the Natural History Museum, London, U.K. (NHMUK).
References
- Bank, S. & Bradler, S. (2022) A second view on the evolution of flight in stick and leaf insects (Phasmatodea). BMC Ecology and Evolution, 22, 62, 1–17. https://doi.org/10.1186/s12862-022-02018-5
- Bank, S., Buckley, T.R., Büscher, T.H., Bresseel, J., Constant, J., de Haan, M., Dittmar, D., Dräger, H., Kahar, R.S., Kang, A., Kneubühler, B., Langton-Myers, S.S. & Bradler, S. (2021) Reconstructing the nonadaptive radiation of an ancient lineage of ground-dwelling stick insects (Phasmatodea: Heteropterygidae). Systematic Entomology, 46, 487–507. https://doi.org/10.1111/syen.12472
- Beier, M. (1968) 10. Phasmida (Stab- oder Gespenstheuschrecken). In: Helmcke, J.G., Starck, D. & Wermuth, H. (Eds.), Handbuch der Zoologie. Eine Naturgeschichte der Stämme des Tierreiches, 4 (2), 2/19, pp. 1–56.
- Bollens, T., Krijns, R. & Bresseel, J. (2010a) Collectietrip op de Filipijnen: deel 2: Mindanao. Phasma Newsletter, 20 (78), 7–14.
- Bollens, T., Krijns, R. & Bresseel, J. (2010b) Collectietrip op de Filipijnen: vervolg. Phasma Newsletter, 20 (79), 22–26.
- Bradler, S. (2001) The Australian stick insects, a monophyletic group within the Phasmatodea? Zoology, 104 (Supplement IV, DZG 94.1), 69.
- Bradler, S. (2009) Phylogenie der Stab- und Gespenstschrecken (Phasmatodea). Species, Phylogenie and Evolution, 2, 3–139. https://doi.org/10.17875/gup2009-710
- Bradler, S. & Buckley, T.R. (2018) “Biodiversity of Phasmatodea”. In: Foottit, R.G. & Adler, P.H. (Eds.), Insect Biodiversity: Science and Society. Vol. II. Wiley-Blackwell, Hoboken, New Jersey, pp. 281–313. https://doi.org/10.1002/9781118945582.ch11
- Bradler, S., Cliquennois, N. & Buckley, T.R. (2015) Single origin of Mascarene stick insects: ancient radiation on sunken islands? BMC, Evolutionary Biology, 15, 196. https://doi.org/10.1186/s12862-015-0478-y
- Bradley, J.C. & Galil, B.S. (1977) The taxonomic arrangement of the Phasmatodea with keys to the subfamilies and tribes. Proceedings of the Entomological Society of Washington, 79, 176–208.
- Bragg, P.E. (2001) Phasmids of Borneo. Natural History Publications (Borneo), Kota Kinabalu, 772 pp.
- Brock, P.D. (1995) Catalogue of Stick and Leaf-Insects (Insecta: Phasmida) associated with Peninsular Malaysia and Singapore. Malayan Nature Journal, 49, 83–102.
- Brock, P.D. (1996) Changes of taxonomy in giant stick-insects. Phasmid Studies, 5 (1), 25–31.
- Brock, P.D. (1998) Catalogue of type specimens of Stick and Leaf-Insects in the Naturhistorisches Museum Wien (Insecta: Phasmida). Kataloge der wissenschaftlichen Sammlungen des Naturhistorischen Museums in Wien, 13 (5), 5–72.
- Brock, P.D. & Büscher, T.H. (2022) Stick and Leaf Insects of the world. Phasmids. N.A.P. Editions, Paris, 609 pp.
- Brock, P.D., Marshall, J.A., Beccaloni, G.W. & Harman, A.J.E. (2016) The types of Phasmida in the Natural History Museum, London, UK. Zootaxa, 4179 (2), 151–208. https://doi.org/10.11646/zootaxa.4179.2.1
- Bruner, L. (1915) A preliminary catalogue of the orthopteroid insects of the Philippine Islands. University of Nebraska Studies, 15 (2), 195–281.
- Brunner von Wattenwyl, C. (1907) n.k. In: Die Insektenfamilie der Phasmiden, II. Phasmidae Anareolatae (Clitumnini, Lonchodini, Bacunculini). Verlag W. Engelmann, Leipzig, pp. 181–340, pls. 7–15.
- Brunner von Wattenwyl, C. & Redtenbacher, J. (1906–1908) Die Insektenfamilie der Phasmiden I–III. Verlag Wilhelm Engelmann, Leipzig, 589 pp., 27 pls.
- Buckley, T.R., Attanayake, D. & Bradler, S. (2009) Extreme convergence in stick insect evolution: phylogenetic placement of the Lord Howe Island tree lobster. Proceedings of the Royal Society of London, B, 276, 1055–1062. https://doi.org/10.1098/rspb.2008.1552
- Buckley, T.R., Attanayake, D., Nylander, J.A.A. & Bradler, S. (2010) The phylogenetic placament and biogeographical origins of the New Zealand stick insects (Phasmatodea). Systematic Entomology, 35, 207–225. https://doi.org/10.1111/j.1365–3113.2009.00505.x
- Caudell, A.N. (1927) On a collection of Orthopteroid Insects from Java made by Owen Bryant and William Palmer in 1909. Proceedings of the United States National Museum, 71 (3), 1–42. https://doi.org/10.5479/si.00963801.71-2675.1
- Clark-Sellick, J.T. (1997) Descriptive terminology of the phasmid egg capsule, with an extended key to the genera based on egg structure. Systematic Entomology, 22, 97–122. https://doi.org/10.1046/j.1365-3113.1997.d01–30.x
- Clark-Sellick, J.T. (1998) The micropylar plate of the eggs of Phasmida, with a survey of the range of plate form within the order. Systematic Entomology, 23, 203–228. https://doi.org/10.1046/j.1365-3113.1998.00056.x
- Cumming, R.T., Le Tirant, S., Linde, J.B., Solan, M.E., Foley, E.M., Eulin, N.E.C., Lavado, R., Whiting, M.F., 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
- Dickerson, R.E. (1928) Distribution of life in the Philippines. Bureau of Science Monograph, 21, 1–322, pls. 1–42.
- Dohrn, H. (1910) Beitrag zur Kenntnis der Phasmiden. Stettiner Entomologische Zeitschrift, 71, 397–414.
- Forni, G., Martelossi, J., Valero, P., Hennemann, F.H., Conle, O., Luchetti, A. & Mantovani, B. (2022) Macroevolutionary analyses provide new evidence of phasmid wing evolution as a reversal process. Systematic Biology, 71 (6), 1471–1486. https://doi.org/10.1093/sysbio/syac038
- Gottardo, M (2008) New record of Hermarchus leytensis Zompro, with notes on its life history (Phasmatodea: Phasmatidae). Phasmid Studies, 17, 11–15.
- Gottardo, M. & Vallotto, D. (2014) External macro and micromorphology of the male of the stick insect Hermarchus leytensis (Insecta: Phasmatodea) with phylogenetic considerations. Comptes Rendus Biologies, 337, 258–268. https://doi.org/10.1016/j.crvi.2014.02.005
- Günther, K. (1935a) Die von Gerd Heinrich 1930-1932 auf Celebes gesammelten Phasmoiden. Mitteilungen des Zoologischen Museums Berlin, 21 (1), 1–29, pls. 1 & 2.
- Günther, K. (1935b) Über einige Phasmoiden aus der Sammlung des Herrn Dr. C. Willemse, Eijgelshoven. Natuurhistisch Maandblad, 24 (12), 123–126 + 138–140.
- Günther, K. (1935c) Phasmoiden aus Centralborneo. Gesammelt von Dr. Mjöberg 1925-26. Arkiv for Zoologi, 28A (9), 1–29.
- Günther, K. (1953) Über die taxonomische Gliederung und die geographische Verbreitung der Insektenordnung der Phasmatodea. Beiträge zur Entomologie, Berlin, 3, 541–563.
- Harman, A.J.E. (2013) The development of the Phasmid Species List Part Six: PSG No. 251 – PSG No. 300. Phasmid Study Group Newsletter, 131, 12–14.
- Harman, A.J.E. (2015) The development of the Phasmid Species List Part Seven: PSG No. 301 – PSG No. 350. Phasmid Study Group Newsletter, 135, 26–27.
- Hennemann, F.H. (1999) On two species of Phasmatodea in the collection of the Institut royl des Sciences naturelles de Belgique, Bruxelles, including the description of a new species. Bulletin S.R.B.E./K.B.V.E., 135, 213–215.
- Hennemann, F.H. (2020) Megacraniinae – The Palm Stick Insects: A new subfamily of Old World Phasmatodea and a redefinition of Platycaraninae Brunner v. Wattenwyl, 1893 (Phasmatodea: “Areolatae”). Zootaxa, 4896 (2), 151–179. https://doi.org/10.11646/zootaxa.4896.2.1
- Hennemann, F.H. (2021) Stick insects of Sulawesi, Peleng and the Sula Islands, Indonesia – a review including checklists of species and descriptions of new taxa (Insecta: Phasmatodea). Zootaxa, 5073 (1), 1–189.
- https://doi.org/10.11646/zootaxa.5073.1.1
- Hennemann, F.H. & Conle, O.V. (2004) Revision of the tribe Achriopterini Bradley & Galil, 1977, with the description of new genus, three new species and a new subspecies from Madagascar (Phasmatodea: Phasmatidae: Phasmatinae). Mitteilungen der Münchner Entomologischen Gesellschaft, 94, 5–54.
- Hennemann, F.H. & Conle, O.V. (2006) Studies on New Guinean giant stick-insects of the tribe Stephanacridini Günther, 1953, with the description of a new genus and three new species of Stephanacris Redtenbacher, 1908 (Orthoptera: Phasmatodea: „Anareolatae“). Zootaxa, 1283 (1), 1–24. https://doi.org/10.11646/zootaxa.1238.1.1
- Hennemann, F.H. & Conle, O.V. (2008) Revision of Oriental Phasmatodea: The tribe Pharnaciini Günther, 1953, including the description of the world’s longest insect, and a survey of the family Phasmatidae Gray, 1835 with keys to the subfamilies and tribes (Phasmatodea: „Anareolatae“: Phasmatidae). Zootaxa, 1906 (1), 1–316. https://doi.org/10.11646/zootaxa.1906.1.1
- Hennemann, F.H. & Conle, O.V. (2009) Studies on the genus Phasmotaenia Navás, 1907, with the descriptions of five new species from the Solomon Islands, a revised key to the species and notes on its geographic distribution (Phasmatodea: „Anareolatae“: Phasmatidae s.l.: Stephanacridini). Zootaxa, 2011 (1), 1–46. https://doi.org/10.11646/zootaxa.2011.1.1
- Hennemann, F.H., Conle, O.V., Gottardo, M. & Bresseel, J. (2009) On certain species of the genus Phyllium Illiger, 1898, with proposals for an intra-generic systematization and the descriptions of five new species from the Philippines and Palawan (Phasmatodea: Phylliidae; Phylliinae: Phylliini). Zootaxa, 2322 (1), 1–83. https://doi.org/10.11646/zootaxa.2322.1.1
- Hennemann, F. H., Conle, O. V. & Suzuki, S. (2015) A study of the members of the tribe Phasmatini Gray, 1835, that occur within the boundaries of Wallacea (Phasmatodea: Phasmatidae: Phasmatinae: „Lanceocercata“). Zootaxa, 4008 (1), 1–74. https://doi.org/10.11646/zootaxa.4008.1.1
- Hennemann, F.H., Damastra, G.B, Damaledo, D.M., Cumming, R.T. & Le Tirant, S. (2023) Nesiophasma sobesonbaii n. sp.-a new giant stick insect from the island of Timor, Indonesia (Insecta: Phasmatodea). Faunitaxys, 11 (20), 1–13. https://doi.org/10.57800/faunitaxys-aa(20)
- Huxley, T.H. (1868) December 10, 1868. Proceedings of the Zoological Society of London, 36 (1), 622–637. https://doi.org/10.1111/j.1469-7998.1868.tb00455.x
- Kevan, D.K.M. (1977) The higher classification of orthopteroid insects: a general view. Lyman Entomological Museum and Research Laboratory Memoirs, 1977, 4–31.
- Kevan, D.K.M. (1982) Phasmatodea (Stick-Insects). In: Naumann, I.D., Came, P.B., Lawrence, J.F., Nielsen, E.S., Spradbery, J.P., Taylor, R.W., Whitten, M.J. & Littlejohn, M.J. (Eds.), The Insects of Australia. Vol. 1. 2nd Edition. CSIRO Publishing, Melbourne University Press, Melbourne, pp. 394–404.
- Kirby, W.F. (1904) A Synonymic Catalogue of Orthoptera, 1. Orthoptera Euplexoptera, Cursoria et Gressoria. (Forficulidae, Hemimeridae, Blattidae, Mantidae, Phasmidae). British Museum, London, 501 pp. [http://biodiversitylibrary.org/page/6196392]
- Otte, D. & Brock, P. (2005) Phasmid Species File. Catalog of Stick and Leaf Insects of the World. CafePress.com, The Insect Diversity Association and the Academy of Natural Sciences, Philadelphia, Pennsylvania, 414 pp.
- Redtenbacher, J. (1908) n.k. In: Die Insektenfamilie der Phasmiden. Vol. III. Phasmidae Anareolatae (Phibalosomini, Acrophyllini, Necrosciini). Verlag W. Engelmann, Leipzig, pp. 341–589, pls. 16–27.
- Robertson J.A., Bradler S. & Whiting M.F. (2018) Evolution of oviposition techniques in stick and leaf insects (Phasmatodea). Frontiers in Ecology and Evolution, 6 (216), 1–16. https://doi.org/10.3389/fevo.2018.00216
- Seow-Choen, F. (2000) An illustrated guide to the Stick and Leaf Insects of Peninsular Malaysia and Singapore. Natural History Publications (Borneo), Kota Kinabalu, 173 pp.
- Seow-Choen, F. (2016) A Taxonomic Guide to the Stick Insects of Borneo. Natural History Publications, Kota Kinabalu, 454 pp.
- Seow-Choen, F. (2017) A Taxonomic Guide to the Stick Insects of Borneo. Vol. II. Natural History Publications, Kota Kinabalu, 261 pp.
- Seow-Choen, F. (2018) A Taxonomic Guide to the Stick Insects of Sumatra. Vol. I. Natural History Publications, Kota Kinabalu, 703 pp.
- Seow-Choen, F. (2019) A Taxonomic Guide to the Stick Insects of Borneo. Vol. III. Natural History Publications, Kota Kinabalu, 278 pp.
- Seow-Choen, F. (2021) A Taxonomic Guide to the Stick Insects of Peninsular Malaysia. Vol. I. Natural History Publications, Kota Kinabalu, 944 pp.
- Seow-Choen, F. (2023) A Taxonomic Guide to the Stick Insects of Java. Vol. I. Natural History Publications, Kota Kinabalu, 652 pp.
- Simon, S., Letsch, H., Bank, S., Buckley, T.R., Donath, A., Liu, S., Machida, R., Meusemann K., Misof, B., Podsiadlowski, L., Zhou, X., Wipfler, B. & Bradler, S. (2019) Old World and New World Phasmatodea: Phylogenomics Resolve the Evolutionary History of Stick and leaf Insects. Frontiers in Ecology and Evolution, 7 (345), 1–13. https://doi.org/10.3389/fevo.2019.00345
- Stål, C. (1875) Recensio Orthopterorum.Revue critique des Orthoptères déscrits par Linné, de Geer et Thunberg. Vol. 3. P. A. Norstedt & Söner, Stockholm. [unknown pagination]
- Stål, C. (1877) Espèces nouvelles de Phasmides. Annales de la Société Entomologique de Belgique, Comptes Rendues, 20, 62–68.
- Tilgner, E.H. (2002) Systematics of Phasmida. Dissertation, University of Georgia, Athens, Georgia. Available from: https://esploro.libs.uga.edu/esploro/outputs/doctoral/Systematics-of-phasmida/9949333911102959 (accessed 3 February 2025)
- Vallejo, B. (2011) The Philippines in Wallacea. In: Telnov, D.M. (Ed.), Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, Vol. 1. Latvian Society of Entomology, Riga, Latvia, pp. 27–42.
- Vanschuytbroeck, C. & Cool, J. (1981) Catalogue et liste du materiel typique des Phasmatodea conservé dans les collections entomologiques de l’Institut Royal des Sciences Naturelles de Belgique. Bulletin de l’Institut Royal des Sciences Naturelles de Belgique, 53 (23), 1–26.
- Wallace, A.R. (1876) The Geographical Distribution of Animals. 2 Vols. Harper, New York, New York, 503 pp. & 607 pp.
- Westwood, J.O. (1859) Catalogue of Orthopterous insects in the collection of the British Museum. Part 1. Phasmidae. British Museum, London, 196 pp., 40 pls. https://doi.org/10.5962/bhl.title.65981
- Whiting. M.F., Bradler, S. & Maxwell, T. (2003) Loss and recovery of wings in stick insects. Nature, 421 (6920), 264–267. https://doi.org/10.1038/nature01313
- Zompro, O. (1997) Hermarchus leytensis n. sp., eine neue Phasmide von den Philippinen. Entomologische Zeitschrift, 107 (1), 38–40.
- Zompro, O. (2004) Revision of the genera of the Areolatae, including the status of Timema and Agathemera (Insecta, Phasmatodea). Abhandlungen des Naturwissenschaftlichen Vereins in Hamburg, N.F., 37, 1–327.
- Zompro, O. (2005) Catalogue of type material of the insect order Phasmatodea, housed in the Museum für Naturkunde der Humboldt-Universität zu Berlin, Germany and in the Institut für Zoologie der Martin-Luther-Universität in Halle (Saale), Germany. Mitteilungen aus dem Museum für Naturkunde Berlin, Deutsche entomologische Zeitschrift, 52 (2), 251–290. https://doi.org/10.1002/mmnd.200410018