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Stick and Leaf-Insects of the World

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https://www.napeditions.com/en/71-stick-and-leaf-insects-of-the-world.html Over 260 stick and leaf-insects (phasmids) are described and illustrated by biogeographical zone. Descriptions allow non-specialists to help identify the presented species, and learn about habitat, food plants and distribution. Phasmid systematics are approached from perspectives of taxonomy and phylogeny, along with bibliographic notes on major phasmid explorers and main publications. An extensive checklist of currently described species and their distribution is included. A comprehensive guide to the biology of phasmids. This book covers phasmid biology (anatomy, main biological functions, ecology) as well as basic advice on collecting and rearing phasmids. Trivia like myths and legends on phasmids and pop culture references are also presented.
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... Their life history makes phasmids interesting study subjects for ageing research, as this lineage represents some of the largest insects known and species that have a prolonged life expectancy of up to three years after imaginal molt [41]. After this last molt, phasmids do not molt anymore and, hence, their cuticle is subject to continuous ageing. ...
... We investigated the change in attachment ability and tarsal morphology in the species Sungaya aeta Hennemann, 2023 (Heteropterygidae). Members of Heteropterygidae can reach impressive life expectancies [41,44], with anecdotal reports extending over five years. The change in attachment performance was quantified through attachment force measurements. ...
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Many insect species have found their way into ageing research as small and easy-to-keep model organisms. A major sign of ageing is the loss of locomotory functions due to neuronal disorders or tissue wear. Soft and pliable attachment pads on the tarsi of insects adapt to the substrate texture to maximize their real contact area and, thereby, generate attachment during locomotion. In the majority of stick insects, adhesive microstructures covering those pads support attachment. Stick insects do not molt again after reaching the imaginal stage; hence, the cuticle of their pads is subject to continuous ageing. This study aims to quantify how attachment ability changes with age in the stick insect Sungaya aeta Hennemann, 2023 and elucidate the age effects on the material and microstructure of the attachment apparatus. Attachment performance (adhesion and friction forces) on substrates with different roughnesses was compared between two different age groups, and the change of attachment performance was monitored extending over a larger time frame. Ageing effects on the morphology of the attachment pads and the autofluorescence of the cuticle were documented using light, scanning electron, and confocal laser scanning microscopy. The results show that both adhesion and friction forces decline with age. Deflation of the pads, scarring of the cuticle, and alteration of the autofluorescence, likely indicating stiffening of the cuticle, were observed to accumulate over time. This would reduce the attachment ability of the insect, as pads lose their pliant properties and cannot properly maintain sufficient contact area with the substrate.
... Whilst the great variety of phasmid egg phenotypes has received increasingly more interest over the last years , the presence and mechanism of structural coloration-such as iridescence in phasmid eggs-has not been studied before. This does not mean that iridescence or any other structural coloration is not present, as Phasmatodea is a diverse order of over 3500 species, with highly species-specific egg phenotypes (Brock et al., 2023;Brock & Büscher, 2022). Iridescent coloration is also found in some species of neotropical stick insects (Heteronemiinae) (Crispino et al., 2022) and potentially in some species of Diapheromerinae. ...
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Colours in nature can be pigmentary, structural or a combination of both. The prevalence, function and nanostructural origin of structural coloration in eggs is largely unknown. Stick and leaf insect eggs display a wide variety of colours, most of which are produced by pigments. The eggs of Myronides glaucus (Phasmida: Lonchodidae; Hennemann, 2021), however, show a clear purple to green iridescence. Here, we use micro‐spectrophotometry, Fourier‐transform infrared reflectance, transmission‐ and scanning electron microscopy, atomic force microscopy, finite‐difference time‐domain optical simulations and experimental approaches to elucidate the mechanism for iridescence in M. glaucus eggshells, which together reveal that iridescence is caused by thin‐film interference by a 200‐ to 450‐nm‐thick outermost layer. These results highlight the diversity of phasmid eggs and the need to study the different mechanisms and functions of structural coloration.
... Brock et al. (2018) discusses parthenogenesis and those interested can see references relating to the Indian Stickinsect Carausius morosus and the mainly Mediterranean genus Bacillus, also discussions on Clitarchus hookeri. For further detail of the subject see Brock & Büscher 2022. In species where males occur only rarely, they may not be able to fertilise females, as is the case with cultured Carausius morosus. ...
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The first male of Acanthoxyla prasina is described and illustrated. Comments are made on the genus Acanthoxyla with a comparison of the males of A. prasina and A. inermis, each so far only known from a single UK specimen.
... The genus Calvisia (Necrosciinae: Necrosciini) in China is known as "jade stick insects (翠䗛)", showing beautiful and aposematic coloration among species in this subfamily. At present, this genus contains 6 subgenera and 44 species distributed in temperate and tropical Asia (Brock & Büscher 2022, Brock et al. 2024, Seow-Choen 2016. ...
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Calvisia is a colorful winged stick insect genus consisting of 6 subgenera and 44 species widely distributed in temperate and tropical Asia. C. medogensis syn. nov. was discovered in Mêdog, Xizang (Tibet), China and is so far the only species recorded from China. We here propose that C. medogensis syn. nov. is a synonym of C. fuscoalata after checking type specimens of both species. New materials studied are deposited in Yunnan Agricultural University, China (YNAU).
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The male of the Rossel Island Stick-insect, Eurycantha portentosa Kirby, 1904 is described and figured.
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Several stick insects occurring in Brazil belong to the Diapheromerinae. Before this work, the Brazilian genus Exocnophila contained three species known only from females, Exocnophila exintegra Zompro, 2001, Exocnophila tuberculata (Brunner von Wattenwyl, 1907) and Exocnophila cornuta (Brunner von Wattenwyl, 1907). Based on recently collected material, we found that the males of Exocnophila are assigned to another genus in the Diapheromerinae, the heterogeneous Bacteria. Here we show that the type species Exocnophila exintegra Zompro, 2001 syn. nov. is a junior synonym of Bacteria brevitarsata Brunner von Wattenwyl, 1907, which is transferred and redescribed as Exocnophila brevitarsata comb. nov.. We also transfer and redescribe Bacteria hastata as Exocnophila hastata comb. nov. and propose a new species, Exocnophila ovifuscum sp. nov., from Rio de Janeiro state, Brazil, with descriptions of females, males and eggs. We provide the first biological observations for the genus and a short discussion about its relationships.
Article
We here present an updated and annotated catalogue of the stick and leaf insect (Phasmatodea) types currently housed in the Zoological Museum Hamburg (ZMH), part of the Leibniz Institute for the Analysis of Biodiversity Change (LIB). We found a total of 305 type specimens, belonging to 130 species. This includes 110 primary types (45 holotypes, 13 lectotypes, 52 syntypes) and 195 paratypes and paralectotypes. Most of the species were described by Redtenbacher, others come from Zompro, Brunner von Wattenwyl, Weidner and others. We provide updated information on the taxonomy of all species. This catalogue will represent an important resource for future taxonomic work on the group.
Article
Stick and leaf insects (Phasmatodea) are a moderately diverse order that comprises almost 3,500 extant species of large to very large often impressively camouflaged nocturnal herbivores. The order also stands out as one of the few insect orders that have until lately lacked a robust higher-level phylogeny and still the relationships between many New World taxa in particular remain unknown. The Diapheromerinae is one of the main lineages of the Occidophasmata and comprises a bulk of the diversity of New World stick insects. The clade is endemic to the Americas and includes the longest insects of that region. During the past 25 years Diapheromerinae has experienced various often inconsistent attempts of a classification, which have resulted in a complicated history that has meanwhile constituted a heterogenous mixture of historical and contemporary views. To counter these challenges workers have made use of rank-free taxa to provisionally group supposedly monophyletic clades. Currently the Diapheromerinae are sub-divided into the two tribes Diapheromerini and Oreophoetini and recent molecular analyses have shown taxa of the Cladomorphinae Günther, 1953 to belong to Diapheromerinae. Certainly, the clade still deserves much work to delimit meaningful sub-groups within Diapheromerinae reflective of their evolutionary history and the results of the latest comprehensive molecular-based phylogenetic studies already imply sub-divisions within the Diapheromerini. The rank-free taxon Cladomorformia tax. n. is established to comprise former Cladomorphini, Cranidiini, Otocrania Redtenbacher, 1908 as well as sections of Diapheromerini sensu Robertson et al., 2018, which are the genera that have previously been placed in the “Phanocles group” sensu Zompro, 2001 as well as two genera of the “Bacteria group” and one genus of the “Clonistria group”: Alienobostra, Bostriana, Calynda, Globocalynda, Laciphorus, Phanocles, Phanocloidea and Trychopeplus. The genus Cranidium and all genera of Cladomorphini sensu Hennemann et al., 2016 are here formally transferred to Diapheromerinae, this is Aplopocranidium, Cladomorphus, Jeremia, Jeremiodes, Otocraniella and Xylodus. A holistic review of Cladomorformia at the genus level is conducted herein and new diagnoses, differentiations and keys to all 25 genera that are now contained in that clade are presented. Lists of species are provided for all genera, which include detailed type data, synonymies and distributional records. Moreover, identification keys to all 163 known valid species are provided to render identification of species within the 25 known genera possible. In total, 232 species are covered if the 69 synonymous taxa are included. The subfamily Haplopodinae is introduced to comprise all genera of the tribes Haplopodini, Hesperophasmatini and Pterinoxylini as classified by Hennemann et al. (2016) as well as the recently described Teruelphasmini. Renaming former Cladomorphinae sensu Robertson et al. (2018) into Haplopodinae is necessary, because the type-genus Cladomorphus can no longer be considered a member of that lineage and belongs in Diapheromerinae. Haplopodinae is the only New World clade that belongs into the Old World Oriophasmata. Bostranova Villet, 2023 has been introduced to replace the preoccupied Bostra Stål, 1875. The type-species of Bostranova, Bacteria turgida Westwood, 1859, however is here shown to belong in Phanocloidea Zompro, 2001. Thus, Bostranova is synonymised under Phanocloidea (n. syn.) and all species are transferred to Phanocloidea and other genera. Paraphanocles Zompro, 2001 (Type-species: Mantis keratosqueleton Olivier, 1792) is synonymised with Phanocles Stål, 1875 (syn. n.). Seven new genera are described: Globocrania gen. n. (Type-species: Bacteria emesa Westwood, 1859), Hirtuleiodes gen. n. (Type-species: Phibalosoma gibbosa Chopard, 1911), Lanceobostra gen. n. (Type-species: Bacteria aetolus Westwood, 1859), Ocreatophasma gen. n. (Type-species: Ocreatophasma elegans sp. n.), Parotocrania gen. n. (Type-species: Parotocrania panamae sp. n.), Phanoclocrania gen. n. (Type-species: Bostra dorsuaria Stål, 1875) and Spinocloidea gen. n. (Type-species: Spinocloidea panamaense sp. n.). All seven genera are described from both sexes and the eggs, with the exception of Ocreatophasma gen. n., which is known from the females only. Forty-one new species are described: Globocalynda cornuta sp. n. from Ecuador, Globocalynda marcapatae sp. n. from Peru, Globocalynda ruficollis sp. n. from Bolivia, Hirtuleiodes peruanus sp. n. from Peru, Jeremiodes costaricensis sp. n. from Costa Rica, Jeremiodes ecuadoricus sp. n. from Ecuador, Jeremiodes peruanus sp. n. from Peru, Lanceobostra chapalaense sp. n. from Mexico, Lanceobostra glabra sp. n. from Mexico, Lanceobostra oaxacaee sp. n. from Mexico, Lanceobostra ornata sp. n. from Mexico, Lanceobostra torquata sp. n. from Mexico, Lanceobostra tuckerae sp. n. from Mexico, Ocreatophasma elegans gen. n., sp. n. from Peru, Ocreatophasma fragile gen. n., sp. n. from Peru, Ocreatophasma modestum gen. n., sp. n. from Peru, Parotocrania acutilobata gen. n., sp. n. from Ecuador, Parotocrania curvata gen. n., sp. n. from Peru, Parotocrania panamae gen. n., sp. n. from Panama, Phanocles acutecornutus sp. n. from Ecuador, Phanocles barbadosense sp. n. from Barbados, Phanocles berezini sp. n. from Mexico, Phanocles brevipes sp. n. from Peru, Phanocles chiapasense sp. n. from Mexico, Phanocles cuzcoense sp. n. from Peru, Phanocles ecuadoricus sp. n. from Ecuador, Phanocles falcatus sp. n. from Ecuador, Phanocles maximus sp. n. from Panama, Phanocles mexicanus sp. n. from Mexico, Phanocles pleurospinosus sp. n. from Costa Rica, Phanocles rehni sp. n. from Honduras, Phanocles solidus sp. n. from Ecuador, Phanocles spectabilis sp. n. from Costa Rica, Phanocles superbus sp. n. from Ecuador, Phanocloidea sanguinea sp. n. from Ecuador, Phanocloidea semiptera sp. n. from Venezuela, Phanocloidea venezuelica sp. n. from Venezuela, Spinocloidea panamaense gen. n., sp. n. from Panama, Spinocloidea splendida gen. n., sp. n. from Colombia and Spinocloidea tumescens gen. n., sp. n. from Costa Rica. With a maximum recorded body length of 285.0 mm the new species Phanocles maximus sp. n. from Panama is the longest extant insect of the Americas and the Occidophasmata clade. A total of 132 taxonomic changes are conducted: 64 species are transferred to other genera and 43 new synonyms are established. Lectotypes are designated for 25 taxa to ensure stability of the concerned names or new synonymies here established. Moreover, the previously unknown females of five and males of twelve species are described and illustrated for the first time. Colour illustrations are presented of the eggs of 39 species, of which those of 26 species are formally described and illustrated for the first time.
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The genus Oxyartes currently comprises 17 taxa, of which 12 are found in China, representing the high diversity in China. This list includes the two species from China as described in this paper. The first is a new remarkable species O. bouxraeuz sp. nov. collected from Gulinqing township, Yunnan. The second is a newly recorded species, O. cresphontes. This species is reported from Mêdog, Xizang, China. A key to this genus from China is presented. Type specimens are deposited in the Yunnan Agricultural University (YNAU).
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The diversity of insect eggs is astounding but still largely unexplained. Here, we apply phylogenetic analyses to over 210 species of stick and leaf insects (order Phasmatodea), coupled with physiological measurements of metabolic rate and water loss, to evaluate several major classes of factors that may drive egg morphological diversification: life history constraints, material costs and mechanical constraints, and ecological circumstances. We show support for all three classes, but egg size is primarily influenced by female body size and strongly trades off with egg number. Consequently, females that lay relatively fewer but larger eggs, which develop more slowly because of disproportionately low metabolic rates, tend to bury or glue them in specific locations, instead of simply dropping them from the foliage (ancestral state). This form of parental care then directly favors relatively elongated eggs, which may facilitate their specific placement and allow easier passage through the oviducts in slender species. In addition, flightless females display a higher reproductive output and consequently lay relatively more and larger eggs compared to flight-capable females. Surprisingly, local climatic conditions had only weak effects on egg traits. Overall, our results suggest that morphological diversification of stick insect eggs is driven by a complex web of causal relationships among traits, with dominant effects of resource allocation strategies and mechanical constraints.
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En base a un estudio, principalmente, de las genitalia, se redescribe la hembra y por primera vez el macho de Bacunculus comutus (Philippi) y se establece la especie Bacunculus blanchardi n. sp.
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A synonymic catalog of species, essential for researchers. 414 pages, spiral bound [replaces a CD issued in 2003 (First Edition)] OUT OF PRINT
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Mimicry and secondary defense are staples among predator-prey interactions. Among insects, the stick and leaf insects are masters of camouflage. Nonetheless, a meager understanding of their origin and early mimetic evolution persists. Here, we report the earliest mimetic and defensive strategies of a stick insect from the Middle Jurassic of China, Aclistophasma echinulatum gen. et sp. nov., exquisitely preserving abdominal extensions and femoral spines. The distribution of these characteristics mapped onto the phylogeny of Phasmatodea reveals that abdominal extensions and femoral spines developed multiple times during the evolution of stick insects, and indicates that the origin of abdominal extensions predates other modifications, while tergal extensions predate other expansions of the body, such as those of the sterna and pleura, as well as defensive femoral spines. The new fossil provides clues into early antipredator defensive strategies, allows inferences as to the potential environment and predators, and reveals the mimetic and defensive mechanisms of stick insects from 165 million years ago.
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The evolutionary transition between winglessness and a full-winged morphology requires selective advantage for intermediate forms. Conversely, repeated secondary wing reductions among the pterygotes indicates relaxation of such selection. However, evolutionary trajectories of such transitions are not well-characterized. The stick insects (Phasmatodea) exhibit diverse wing sizes at both interspecific and intersexual levels, and thus provide a system for examining how selection on flight capability, along with other selective forces, drives the evolution of flight-related morphology. Here, we examine variation in relevant morphology for stick insects using data from 1,100+ individuals representing 765 species. Although wing size varies along a continuous spectrum, taxa with either long or miniaturized wings are the most common, whereas those with intermediate-sized wings are relatively rare. In a morphological space defined by wing and body size, the aerodynamically relevant parameter termed wing loading (the average pressure exerted on the air by the wings) varies according to sex-specific scaling laws; volant but also flightless forms are the most common outcomes in both sexes. Using phylogenetically-informed analyses, we show that relative wing size and body size are inversely correlated in long-winged insects regardless of sexual differences in morphology and ecology. These results demonstrate the diversity of flight-related morphology in stick insects, and also provide a general framework for addressing evolutionary coupling between wing and body dimensions. We also find indirect evidence for a “fitness valley” associated with intermediate-sized wings, suggesting relatively rapid evolutionary transitions between wingless and volant forms.
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Phasmatodea comprises over 3,000 extant species and stands out as one of the last remaining insect orders for which a robust, higher-level phylogenetic hypothesis is lacking. New research suggests that the extant diversity is the result of a surprisingly recent and rapid radiation that has been difficult to resolve with standard Sanger sequence data. In order to resolve the early branching events of stick and leaf insects, we analyzed transcriptomes from 61 species, including 38 Phasmatodea species comprising all major clades and 23 outgroup taxa, including all other Polyneoptera orders. Using a custom-made ortholog set based on reference genomes from four species, we identified on average 2,274 orthologous genes in the sequenced transcriptomes. We generated various sub-alignments and performed maximum-likelihood analyses on several representative datasets to evaluate the effect of missing data and matrix composition on our phylogenetic estimates. Based on our new data, we are able to reliably resolve the deeper nodes between the principal lineages of extant Phasmatodea. Among Euphasmatodea, we provide strong evidence for a basal dichotomy of Aschiphasmatodea and all remaining euphasmatodeans, the Neophasmatodea. Within the latter clade, we recovered a previously unrecognized major New World and Old World lineage, for which we introduce the new names Oriophasmata tax. nov. (“Eastern phasmids”) and Occidophasmata tax. nov. (“Western phasmids”). Occidophasmata comprise Diapheromerinae, Pseudophasmatinae, and Agathemera, whereas all remaining lineages form the Oriophasmata, including Heteropterygidae, Phylliinae, Bacillus, Lonchodidae (Necrosciinae + Lonchodinae), Clitumninae, Cladomorphinae, and Lanceocercata. We furthermore performed a divergence time analysis and reconstructed the historical biogeography for stick and leaf insects. Phasmatodea either originated in Southeast Asia or in the New World. Our results suggest that the extant distribution of Phasmatodea is largely the result of dispersal events in a recently and rapidly diversified insect lineage rather than the result of vicariant processes.
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This edited volume is provides an authoritative synthesis of knowledge about the history of life. All the major groups of organisms are treated, by the leading workers in their fields. With sections on: The Importance of Knowing the Tree of Life; The Origin and Radiation of Life on Earth; The Relationships of Green Plants; The Relationships of Fungi; and The Relationships of Animals. This book should prove indispensable for evolutionary biologists, taxonomists, ecologists interested in biodiversity, and as a baseline sourcebook for organismic biologists, botanists, and microbiologists. An essential reference in this fundamental area.
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To better understand insect evolution, fossils – mainly known by their wings – must be used as terminals in phylogenetic analyses. Such analyses are, however, rarely performed because of a lack of consensus on the homology of venation in insects. Researchers do not agree with the current concept on the exact number and identity of the main veins. Here, we confirm the presence, which has been in question since the early 20th century, of an independent main postcubital vein (PCu vein) between the cubital and anal veins (29 fossil and extant examined orders; > 85% of observed insects). The PCu vein corresponds to the so‐called vein 1A or first anal vein. It is easily identified by the unique shape of its bulla. It may have several branches and be partially fused with the cubital and anal veins. Once the PCu vein was identified, we reconsidered as an example the particular case of the Phasmatodea, showing that extant stick insects have a unique venation among insects, with a reduced median vein and a simple cubital vein adjacent or fused to the PCu vein. This study is a new approach towards resolving wing vein homology issues, crucial for future large‐scale phylogenetic analyses in insects combining extant and extinct taxa. Insect wing venation is pivotal in systematic entomology but is still controversial, different venation patterns being used according to taxonomic groups. Using microcomputed tomography, we provide a unified wing venation pattern and demonstrate the presence of a postcubital vein (PCu) in all Neoptera between the cubital and anal veins. We clarify wing venation homologies across insect orders and use phasmids as an example to illustrate the implications of this unified wing venation pattern.