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1

Jintsu, Yoshie, Toshiki Uchifune, and Ryuichiro Machida. "Structural features of eggs of the basal phasmatodean Timema monikensis Vickery & Sandoval, 1998 (Insecta: Phasmatodea: Timematidae)." Arthropod Systematics & Phylogeny 68, no. 1 (2010): 71–78. http://dx.doi.org/10.3897/asp.68.e31716.

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Structural features of the eggs of a basal phasmatodean, Timema monikensis Vickery & Sandoval, 1998 (Timematidae) were examined. The eggs of this species are soft and deposited coated with soil and/or other extraneous particles. The chorion, which is transparent and weakly sclerotized, is composed of an endochorion and an exochorion. The non-inclined operculum is located at the anterior pole of the egg. The chorion in the marginal region of the operculum is thinned to form an opercular collar together with the chorion of the egg body. An inverted triangular micropylar plate is on the ventral side of the egg attached to the opercular collar. The micropylar plate is without external differentiations but is specialized inside the chorion. A single micropyle, with a simple funnel-shaped chorionic opening, occurs on either side of the micropylar plate. The posterior mound, located at the posterior pole, is a thickened chorion rich in fine vertical striations, and the serosal cuticle beneath is thickened and highly specialized. The eggs of Timematidae were characterized and compared with those of Euphasmatodea and Embioptera. A phylogenetic discussion is presented, strongly supporting the assemblage of Timematodea, Euphasmatodea and Embioptera as monophyletic.
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2

Jintsu, Yoshie, Toshiki Uchifune, and Ryuichiro Machida. "Structural features of eggs of the basal phasmatodean Timema monikensis Vickery & Sandoval, 1998 (Insecta: Phasmatodea: Timematidae)." Arthropod Systematics & Phylogeny 68 (February 28, 2010): 71–78. https://doi.org/10.3897/asp.68.e31716.

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Structural features of the eggs of a basal phasmatodean, Timema monikensis Vickery & Sandoval, 1998 (Timematidae) were examined. The eggs of this species are soft and deposited coated with soil and/or other extraneous particles. The chorion, which is transparent and weakly sclerotized, is composed of an endochorion and an exochorion. The non-inclined operculum is located at the anterior pole of the egg. The chorion in the marginal region of the operculum is thinned to form an opercular collar together with the chorion of the egg body. An inverted triangular micropylar plate is on the ventral side of the egg attached to the opercular collar. The micropylar plate is without external differentiations but is specialized inside the chorion. A single micropyle, with a simple funnel-shaped chorionic opening, occurs on either side of the micropylar plate. The posterior mound, located at the posterior pole, is a thickened chorion rich in fine vertical striations, and the serosal cuticle beneath is thickened and highly specialized. The eggs of Timematidae were characterized and compared with those of Euphasmatodea and Embioptera. A phylogenetic discussion is presented, strongly supporting the assemblage of Timematodea, Euphasmatodea and Embioptera as monophyletic.
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3

Forni, Giobbe, Federico Plazzi, Alex Cussigh, et al. "Phylomitogenomics provides new perspectives on the Euphasmatodea radiation (Insecta: Phasmatodea)." Molecular Phylogenetics and Evolution 155 (February 2021): 106983. http://dx.doi.org/10.1016/j.ympev.2020.106983.

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4

Luo, Ting, Qianwen Zhang, Siyu Pang, Yanting Qin, Bin Zhang, and Xun Bian. "Comparative Mitochondrial Genomic and Phylogenetic Study of Eight Species of the Family Lonchodidae (Phasmatodea: Euphasmatodea)." Genes 16, no. 5 (2025): 565. https://doi.org/10.3390/genes16050565.

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Background: Lonchodidae is the largest family within the order Phasmatodea, and although many studies have been conducted on this family, the monophyly of the family has not been established. Methods: Eight mitogenomes from Lonchodidae, including the first complete mitogenomes of four genera, were sequenced and annotated to explore their features and phylogenetic relationships. Results: The total length ranged from 15,942–18,021 bp, and the mitogenome consisted of 13 protein-coding genes (PCGs), 22 tRNA genes, 2 rRNA genes, and a control region (CR). atp8 had the highest A + T content in Lonchodidae, except for Neohirasea stephanus and Asceles clavatus, in which the highest A + T contents were detected in nad6. The phylogenetic trees were reconstructed via Bayesian inference (BI) and maximum likelihood (ML) based on the PCG123 and PCG12 datasets. As the phylogenetic trees show, Necrosciinae is recognized as monophyletic, but the monophyly of Lonchodinae has not been supported. Gene deletion and rearrangement have occurred mainly in Lonchodidae and Aschiphasmatidae. The most common reason for gene rearrangements was tandem duplication random loss (TDRL), but trnI of Stheneboea repudiosa inverted into the CR. In addition, genes within the same family or genus share related sequences and conserved gene blocks. Conclusions: we expanded the mitochondrial genomic data for this family, thereby establishing a foundational dataset for future studies.
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Chen, Sha, Xiangchu Yin, Xiaodan Lin, et al. "Stick insect in Burmese amber reveals an early evolution of lateral lamellae in the Mesozoic." Proceedings of the Royal Society B: Biological Sciences 285, no. 1877 (2018): 20180425. http://dx.doi.org/10.1098/rspb.2018.0425.

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Extant stick and leaf insects commonly imitate twigs or leaves, with lateral lamellae used to enhance crypsis or achieve mimicry for protection. However, the origin and early evolution of such lateral expansions among Phasmatodea are unknown, because all known Mesozoic phasmatodeans hitherto lack preserved evidence of such structures. We report here the first Mesozoic stick insect, Elasmophasma stictum gen. et sp. nov., with well-preserved, thin, lateral lamellae on the thoracic pleura, the terga of abdominal segments I–X and the ventrolateral margins of all femora. This new species, from the mid-Cretaceous amber of northern Myanmar, has a clear, stick-like body and is assigned to Euphasmatodea. The abdominal structures of E. stictum exhibit traces of multiple expansions of the terga, suggesting that such structure might have been an early development of body expansions used to improve crypsis for stick or leaf insects when they sprawled on twigs or leaves.
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6

Archibald, S. Bruce, and Sven Bradler. "Stem-group stick insects (Phasmatodea) in the early Eocene at McAbee, British Columbia, Canada, and Republic, Washington, United States of America." Canadian Entomologist 147, no. 6 (2015): 744–53. http://dx.doi.org/10.4039/tce.2015.2.

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AbstractStem-group Phasmatodea, known as the Susumanioidea, are previously established from the Jurassic through the Paleocene. Here, we extend this record to the early Eocene with five new fossils: two forewings from the Klondike Mountain Formation exposures at Republic, Washington, United States of America, and three partially complete specimens from the McAbee locality in southern British Columbia, Canada. We assign both of the Republic specimens to the new genus and species Eoprephasma hichensinew genus, new species. Two of the McAbee fossils appear to represent two further new species, which we refer to as Susumanioidea species A and B for lack of clearly preserved diagnostic species-level character states. The third might belong to one of these two species, but this is unclear. In all three, the mesothorax and metathorax are not notably extended, the forewings are not shortened, the foreleg femur is straight, and species A possesses an extended, external ovipositor with an operculum (unknown in the other specimens). These conditions are rare and never found in combination in Euphasmatodea. All other stem-group Phasmatodea younger than the Early Cretaceous of China are only known from isolated wings.
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7

Ghirotto, Victor Morais, Edgar Blois Crispino, Phillip Watzke Engelking, Pedro Alvaro Barbosa Aguiar Neves, Júlia De Góis, and Pedro Ivo Chiquetto-Machado. "Arumatia, a new genus of Diapheromerinae stick insects (Insecta, Phasmatodea) from Brazil, with description of five new species and reassessment of species misplaced in Australian genera." European Journal of Taxonomy 827 (July 4, 2022): 1–85. http://dx.doi.org/10.5852/ejt.2022.827.1849.

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Two species of stick insect with a distinctive morphology, Candovia evoneobertii (Zompro & Adis, 2001) and Echetlus fulgens Zompro, 2004, were considered to be native to Australia and introduced into Brazil. However, Heteronemia dubia (Caudell, 1904) and Heteronemia fragilis (Brunner von Wattenwyl, 1907), both described more than a hundred years ago from South America, exhibit striking similarities with the two purportedly introduced species and are found to be conspecific with C. evoneobertii. Careful analysis of the literature and specimens revealed that these species belong to the Neotropical tribe Diapheromerini (Diapheromeridae) and represent a new genus, Arumatia Ghirotto gen. nov. We therefore propose Arumatia fulgens (Zompro, 2004) gen. et comb. nov. and Arumatia dubia (Caudell, 1904) gen. et comb. nov. We further redescribe A. dubia (Caudell, 1904) gen. et comb. nov. based on several specimens and synonymize Heteronemia fragilis syn. nov. and Candovia evoneobertii syn. nov. under it. Additionally, five new Brazilian species are described: Arumatia diamante Ghirotto gen. et sp. nov. from Abaíra, Bahia; Arumatia aramatia Ghirotto gen. et sp. nov. from Porto Nacional, Tocantins; Arumatia motenata Ghirotto gen. et sp. nov. from Serra do Cipó, Minas Gerais; Arumatia crassicercata Ghirotto, Crispino & Engelking gen. et sp. nov. from Alto Paraíso de Goiás, Goiás; and Arumatia anyami Ghirotto, Crispino & Neves gen. et sp. nov. from Costa Marques, Rondônia. Species of Arumatia gen. nov. occur mostly in the Cerrado domain, and represent the first Diapheromeridae recorded in this area. Most species are known exclusively from females with only A. aramatia gen. et sp. nov. and A. motenata gen. et sp. nov. known from both sexes. Adult and egg morphology are described and illustrated in detail for all species, as well as the nymph stages for A. dubia. Biological observations are presented, including parthenogeny in A. dubia and one of the few detailed accounts of sexual behaviour in Euphasmatodea (for A. motenata gen. et sp. nov.). Finally, a species of Diapheromerini described in error from Brazil, Diapheromera armata Piza, 1973, is synonymized under the North American Megaphasma denticrus (Stål, 1875) (syn. nov.).
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8

de, Luna Manuel, Roberto García-Barrios, Gerardo Cuéllar-Rodríguez, and Ulises López-Mora. "A new record of a winged stick insect (Phasmatodea) from Mexico, with a checklist and key to the species of the family Pseudophasmatidae from continental North America." Journal of Orthoptera Research 32, no. (2) (2023): 171–76. https://doi.org/10.3897/jor.32.98203.

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The winged stick insect Metriophasma iphicles (Redtenbacher, 1906) (Phasmatodea: Pseudophasmatidae) is recorded for the first time from Mexico (state of Veracruz), making this the northernmost record of both the species and genus. A checklist of species in the family Pseudophasmatidae from Mexico and the USA is presented, and a key to the species listed is proposed. With the current record, the number of continental North American species of Phasmatodea increases to 108, and the number of genera in the region increases to 23.
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9

Gao, Hao-Ran, and Chong-Xin Xie. "Pylaemenes gulinqingensis sp. nov., a new species of subfamily Dataminae (Phasmida, Heteropterygidae) from Yunnan Province, China." Journal of Orthoptera Research 31, no. (2) (2022): 125–29. https://doi.org/10.3897/jor.31.79783.

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Pylaemenes gulinqingensis sp. nov., from Gulinqing Nature Reserve, Maguan County, Yunnan, China, is described for the first time from two females and six eggs. The types of the new species are deposited in Southwest Forestry University (SWFU), China, as well as in the first author's collection.
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10

Xie, ChongXin, Jun Cai, and YuHan Qian. "New species of Andropromachus (Phasmatodea: Lonchodidae: Necrosciinae: Necrosciini) from Yunnan Province, China." Biodiversity Data Journal 10 (January 6, 2022): e78080. https://doi.org/10.3897/BDJ.10.e78080.

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The genus <i>Andropromachus</i> (Lonchodidae: Necrosciinae: Necrosciini) is a small genus of stick insects including four species with a distribution in northern Vietnam and south-western China.A new species of stick insects <i>Andropromachus gulinqingensis</i> <b>sp. n.</b> is described from Yunnan Province of China. Diagnostic characters of the new species are illustrated and an identification key to male and female <i>Andropromachus</i> is provided along with a checklist for all described species.
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11

Cumming, Royce T., Tirant Stéphane Le, Jackson B. Linde, et al. "On seven undescribed leaf insect species revealed within the recent "Tree of Leaves" (Phasmatodea, Phylliidae)." ZooKeys 1173 (August 3, 2023): 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.
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12

Xu, Fangling, Yingjie Jiang, and Maofa Yang. "Descriptions of two new stick insect species of Cnipsomorpha Hennemann, Conle, Zhang & Liu (Phasmatodea) from China based on integrative taxonomy." ZooKeys 1176 (August 22, 2023): 37–53. https://doi.org/10.3897/zookeys.1176.75490.

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Accurate taxonomical identification is an extremely important basis for stick insect research, including evolutionary biology but also applied biology such as pest control. In addition, genetic methods are a valuable identification auxiliary technology at present. Therefore, this paper used morphological and molecular data to investigate five stick insect specimens from the genus Cnipsomorpha in Yunnan, successfully identifying two new species: Cnipsomorpha yunnanensis Xu, Jiang &amp; Yang, sp. nov. and C. yuxiensis Xu, Jiang &amp; Yang, sp. nov. A phylogenetic tree was constructed through their 28S and COI genes in order to infer the phylogenetic position of the two new species. Photographs of the new species and a key to all known Cnipsomorpha species are provided.
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13

Conle, Oskar V., Pablo Valero, and Frank H. Hennemann. "A new Trychopeplus species (Phasmatodea, Diapheromerinae, Cladomorformia) discovered from Ecuador's enigmatic Chocó ecoregion." ZooKeys 1217 (November 12, 2024): 309–26. https://doi.org/10.3897/zookeys.1217.130397.

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This study presents a description of a new stick insect species belonging to the genus <i>Trychopeplus</i> Shelford, 1909, discovered by the authors in the Chocó ecoregion of northwestern Ecuador. <i>Trychopeplus mashpiensis</i> sp. nov. is described and illustrated based on males, females, and eggs. The distinctive features of this new species, such as its unique body ornamentation and the morphology of its egg structure without fringes, clearly differentiate it from other known species within the genus. Photographs of the new species are provided, along with an updated key and distribution map for all <i>Trychopeplus</i> species. These findings enhance our understanding of the genus's diversity.
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Bresseel, Joachim, and Jérôme Constant. "Extension of the phasmid genus Presbistus to Cambodia with a new species and notes on genitalia and captive breeding (Phasmida, Aschiphasmatidae, Aschiphasmatinae)." Journal of Orthoptera Research 31, no. (2) (2022): 105–17. https://doi.org/10.3897/jor.31.78520.

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A new species of Presbistus Kirby, 1896, Presbistus vitivorus sp. nov., is described from Cambodia based on both sexes, nymphs, and eggs. Male genitalia and vomer are described and figured. Illustrations of adults, nymphs, specimens in situ, host plants, a distribution map and records on biology and breeding in captivity are provided. The host plants of the species belong to the family Vitaceae. The genus Presbistus and the family Aschiphasmatidae are recorded from Cambodia for the first time. The species diversity and the distribution of the genus are discussed, and it is shown that the genus is restricted to Sundaland. A nomenclature for the morphology of the dissected vomer is proposed and tries to homologize the previously used terms.
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Qian, YuHan, ChongXin Xie, and Cui Li. "First record of the genus Medaura Stål (Phasmatodea, Phasmatidae, Clitumninae) from China, with description of a new species." Biodiversity Data Journal 10 (December 9, 2022): e96341. https://doi.org/10.3897/BDJ.10.e96341.

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The stick insect tribe, Medaurini in subfamily Clitumniae, contains 11 genera and 75 known species, with species diversity of this tribe being rich in southeast Asia and China, as is reflected in the Phasmida Species File Online [PSF]. The genus <i>Medaura</i> includes four named species and they are distributed over Bangladesh, Indonesia and India. The discovery of this new species in Xishuangbanna Dai Autonomous Prefecture marks the first identification of the genus <i>Medaura</i> in China.The genus <i>Medaura</i> Stål is reported for the first time from China, based on a new species <i>M. aculeiformis</i> Xie &amp; Qian sp. n. and the identification characteristics of species are described and illustrated in this paper.
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Xie, Chong-Xin, Jun Wen, and Yu-Han Qian. "Nanhuaphasma Chen, He & Li, 2002 is a junior synonym of Dajaca Brunner von Wattenwyl, 1893 (Phasmatodea, Aschiphasmatidae, Dajacini)." ZooKeys 1082 (January 18, 2022): 51–62. https://doi.org/10.3897/zookeys.1082.73272.

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The genus Nanhuaphasma Chen, He &amp; Li, 2002 was established as a member of the family Pseudophasmatidae Rehn, 1904 (now belonging to Aschiphasmatidae Brunner von Wattenwyl, 1893) based on the male of N. hamicercum Chen &amp; He, 2002. We review the status of Nanhuaphasma and N. hamicercum by examining the holotype and male and female non-types which were collected in same location as the holotype. We find that Nanhuaphasma is a junior synonym of Dajaca Brunner von Wattenwyl, 1893 and N. hamicercum is a junior synonym of D. napolovi Brock, 2000. Complementing egg morphology of D. napolovi and keys to eight species of Dajaca are provided.
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17

Villet, Martin H. ""Bactricia nematodes Kby., 1894" (Phasmida, Diapheromeridae, Diapheromerinae) is a nomen nudum." African Invertebrates 65, no. (1) (2024): 1–7. https://doi.org/10.3897/AfrInvertebr.65.115507.

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A review of published evidence indicates that <i>Bactricia nematodes</i> Kirby, 1894 is a <i>nomen nudum</i> because it is an unavailable name. The specimen collected during the Lund University Swedish South African Expedition and reported by this name is a male of <i>Bactricia bituberculata</i> (Schaum, 1857).
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Villet, Martin H. "A replacement name for Bostra Stål (Insecta, Phasmida, Diapheromeridae), a junior homonym of Bostra Walker (Insecta, Lepidoptera, Pyralidae)." Journal of Orthoptera Research 32, no. (2) (2023): 149–52. https://doi.org/10.3897/jor.32.97047.

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Bostra Stål (Phasmida, Diapheromeridae) is a junior homonym of Bostra Walker (Lepidoptera, Pyralidae). The replacement name Bostranova Villet nomen nov. is proposed for Bostra Stål, and new combinations are proposed for the species-group names currently included in that phasmid genus.
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19

Cumming, Royce T., and Tirant Stéphane Le. "Three new genera and one new species of leaf insect from Melanesia (Phasmatodea, Phylliidae)." ZooKeys 1110 (July 5, 2022): 151–200. https://doi.org/10.3897/zookeys.1110.80808.

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With the first large-scale Phylliidae molecular phylogeny recently published adding a great deal of clarity to phylliid diversity, several of the rarer species which could not be included were methodically and morphologically reviewed. This review resulted in identification of numerous substantial morphological features that suggest there are Melanesian clades that create polyphyletic groups within the phylliids which should instead be taxonomically recognized as unique. These rarer Melanesia species have historically been considered to be southern representatives of the Pulchriphyllium Griffini, 1898 sensu lato. However, there are notable morphological differences between the Pulchriphyllium sensu stricto and the "schultzei" group. Therefore, two new genera are erected, Vaabonbonphyllium gen. nov. from the Solomon Islands and Papua New Guinea and Rakaphyllium gen. nov. from New Guinea and the Aru Islands. Erection of these two new genera warrants the following new combinations: Rakaphyllium schultzei (Giglio-Tos, 1912), comb. nov., Rakaphyllium exsectum (Zompro, 2001b), comb. nov., and Vaabonbonphyllium groesseri (Zompro, 1998), comb. nov. Additionally, while reviewing material an undescribed Vaabonbonphyllium gen. nov. specimen was located and is herein described as Vaabonbonphyllium rafidahae gen. et sp. nov. from Mt. Hagen, Papua New Guinea. Additionally, a morphologically unique clade of several species recovered as sister to the Nanophyllium sensu stricto was recognized and their numerous unique morphological features and monophyly leads the authors to erect the new genus Acentetaphyllium gen. nov. which warrants the following new combinations: Acentetaphyllium brevipenne (Größer, 1992), comb. nov., Acentetaphyllium larssoni (Cumming, 2017), comb. nov., Acentetaphyllium miyashitai (Cumming et al. 2020), comb. nov., and Acentetaphyllium stellae (Cumming, 2016), comb. nov. With the addition of several new genera, a key to phylliid genera is included for adult males and females.
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Chen, Zhi-Teng, and Xiao-Han Ye. "Taenionema sinensis sp. n., the first endemic species of Taenionema Banks, 1905 (Plecoptera, Taeniopterygidae) from China." Biodiversity Data Journal 11 (June 8, 2023): e104618. https://doi.org/10.3897/BDJ.11.e104618.

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The taeniopterygid genus <i>Taenionema</i> Banks, 1905 currently contains 14 species distributed in the Nearctic and the eastern Palearctic Regions. <i>Taenionema japonicum</i> (Okamoto, 1922) is the only species known from the Eastern Hemisphere, specifically in Japan, Korea, Mongolia, Russia and north-eastern China. The authors recently described the larvae of an undetermined <i>Taenionema</i> species, which was supposed to represent a second Palaearctic species.This paper reports the first endemic species of <i>Taenionema</i> Banks, 1905, <i>Taenionema sinensis</i> sp. n. from China, which also represents the second species of <i>Taenionema</i> from the Eastern Hemisphere. Description and illustrations based on male and female adults are provided. The new species is easily distinguished from all congeners by the bilobed abdominal sternum 9 of the male adult. The female adult is characterised by the posteriorly truncate postgenital plate. The male larva is distinguished by the emarginate subgenital plate and hook-shaped paraprocts.
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21

Yang, Hongru, Michael S. Engel, Chungkun Shih, et al. "Independent wing reductions and losses among stick and leaf insects (Phasmatodea), supported by new Cretaceous fossils in amber." BMC Biology 21, no. 1 (2023). http://dx.doi.org/10.1186/s12915-023-01720-0.

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Abstract Background Phasmatodea (stick and leaf insects) play a central role on the debate regarding wing reduction and loss, and its wings are putative reacquisition from secondarily wingless ancestors based solely on extant species. A pivotal taxon in this respect is the species-poor Timematodea, consisting of approximately 21 wingless extant species, which form the sister group of all remaining winged or wingless stick and leaf insects, the Euphasmatodea. Results Herein, the new fossils of Timematodea from mid-Cretaceous Kachin amber are reported, with winged and wingless species co-occurring. The palaeogeographic distributions of all fossils of Holophasmatodea are summarized, showing their wide paleo-distributions. The phylogenetic analysis based on morphological characters confirms the earliest-diverging lineage of winged Breviala cretacea gen. et sp. nov. in Timematodea, and the possible relationships among all families of Holophasmatodea. These are critical for the reconstruction of patterns of wing evolution in early Phasmatodea. Conclusions The new fossils suggest that Timematodea once had wings, at least during the mid-Cretaceous. The palaeogeographic occurrences imply that Timematodea probably have been widely distributed since at least the Jurassic. The phylogenetic analysis with the ancestral-state reconstruction of wings indicates that the common ancestors of Holophasmatodea were winged, the reductions and losses of wings among Timematodea and Euphasmatodea have occurred independently since at least the Cretaceous, and the reduction or loss of the forewing earlier than the hind wings.
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Li, Yanfei, Shujing Wang, Jiayue Zhou, et al. "The phylogenic position of aschiphasmatidae in euphasmatodea based on mitochondrial genomic evidence." Gene, September 2021, 145974. http://dx.doi.org/10.1016/j.gene.2021.145974.

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23

Ghirotto, Victor M., Edgar B. Crispino, Pedro I. Chiquetto‐Machado, Pedro A. B. A. Neves, Phillip W. Engelking, and Guilherme C. Ribeiro. "The oldest Euphasmatodea (Insecta, Phasmatodea): modern morphology in an Early Cretaceous stick insect fossil from the Crato Formation of Brazil." Papers in Palaeontology 8, no. 3 (2022). http://dx.doi.org/10.1002/spp2.1437.

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24

Niekampf, Marco, Paul Meyer, Felix S. C. Quade, Alexander R. Schmidt, Tim Salditt, and Sven Bradler. "High disparity in repellent gland anatomy across major lineages of stick and leaf insects (Insecta: Phasmatodea)." BMC Zoology 9, no. 1 (2024). http://dx.doi.org/10.1186/s40850-023-00189-2.

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Abstract Background Phasmatodea are well known for their ability to disguise themselves by mimicking twigs, leaves, or bark, and are therefore commonly referred to as stick and leaf insects. In addition to this and other defensive strategies, many phasmatodean species use paired prothoracic repellent glands to release defensive chemicals when disturbed by predators or parasites. These glands are considered as an autapomorphic trait of the Phasmatodea. However, detailed knowledge of the gland anatomy and chemical compounds is scarce and only a few species were studied until now. We investigated the repellent glands for a global sampling of stick and leaf insects that represents all major phasmatodean lineages morphologically via µCT scans and analyzed the anatomical traits in a phylogenetic context. Results All twelve investigated species possess prothoracic repellent glands that we classify into four distinct gland types. 1: lobe-like glands, 2: sac-like glands without ejaculatory duct, 3: sac-like glands with ejaculatory duct and 4: tube-like glands. Lobe-like glands are exclusively present in Timema, sac-like glands without ejaculatory duct are only found in Orthomeria, whereas the other two types are distributed across all other taxa (= Neophasmatodea). The relative size differences of these glands vary significantly between species, with some glands not exceeding in length the anterior quarter of the prothorax, and other glands extending to the end of the metathorax. Conclusions We could not detect any strong correlation between aposematic or cryptic coloration of the examined phasmatodeans and gland type or size. We hypothesize that a comparatively small gland was present in the last common ancestor of Phasmatodea and Euphasmatodea, and that the gland volume increased independently in subordinate lineages of the Occidophasmata and Oriophasmata. Alternatively, the stem species of Neophasmatodea already developed large glands that were reduced in size several times independently. In any case, our results indicate a convergent evolution of the gland types, which was probably closely linked to properties of the chemical components and different predator selection pressures. Our study is the first showing the great anatomical variability of repellent glands in stick and leaf insects.
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Bank, Sarah, and Sven Bradler. "A second view on the evolution of flight in stick and leaf insects (Phasmatodea)." BMC Ecology and Evolution 22, no. 1 (2022). http://dx.doi.org/10.1186/s12862-022-02018-5.

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Abstract Background The re-evolution of complex characters is generally considered impossible, yet, studies of recent years have provided several examples of phenotypic reversals shown to violate Dollo’s law. Along these lines, the regain of wings in stick and leaf insects (Phasmatodea) was hypothesised to have occurred several times independently after an ancestral loss, a scenario controversially discussed among evolutionary biologists due to overestimation of the potential for trait reacquisition as well as to the lack of taxonomic data. Results We revisited the recovery of wings by reconstructing a phylogeny based on a comprehensive taxon sample of over 500 representative phasmatodean species to infer the evolutionary history of wings. We additionally explored the presence of ocelli, the photoreceptive organs used for flight stabilisation in winged insects, which might provide further information for interpreting flight evolution. Our findings support an ancestral loss of wings and that the ancestors of most major lineages were wingless. While the evolution of ocelli was estimated to be dependent on the presence of (fully-developed) wings, ocelli are nevertheless absent in the majority of all examined winged species and only appear in the members of few subordinate clades, albeit winged and volant taxa are found in every euphasmatodean lineage. Conclusion In this study, we explored the evolutionary history of wings in Phasmatodea and demonstrate that the disjunct distribution of ocelli substantiates the hypothesis on their regain and thus on trait reacquisition in general. Evidence from the fossil record as well as future studies focussing on the underlying genetic mechanisms are needed to validate our findings and to further assess the evolutionary process of phenotypic reversals.
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