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Die Phylogenie der Stab- und Gespenstschrecken (Insecta: Phasmatodea)
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... Although the number of taxa is much less than that of other groups of insects, such as the Diptera or Coleoptera, many details of their evolution, such as changes in morphology, behaviour or lifestyle remain unresolved. One reason lies in the notable differences in the subgroups of Polyneoptera and their strong ecological differentiation that has impeded a reliable reconstruction of the internal relationships for many years (Bradler, 2009;Beutel et al., 2013;Wipfler et al., 2019). Attachment pad morphology is both an indicator and a result of the complex mesh of adaptations. ...
... Stick and leaf insects are an impressive model group for the exploration of many evolutionary questions, especially convergence. Limited spatial dispersion, and extensive adaptive radiation, have led to a high degree of convergent traits in Phasmatodea, such as those related to visual camouflage (Bradler, 2003(Bradler, , 2009(Bradler, , 2015Bradler & Buckley, 2018), oviposition techniques (Carlberg, 1983(Carlberg, , 1987Sellick, 1997a, b;Goldberg et al., 2015;Robertson et al., 2018;, different degrees of wing loss Whiting et al., 2003;Zeng et al., 2019), and ecomorphs (morphological forms with similar ecological occupancy) with specific vertical stratification within the vegetation (Büscher et al., 2018a;Buckley et al., 2010;Bank et al., 2021). Phasmids are predominantly nocturnal insects that are distributed nearly worldwide and exclusively feed on plants (Bradler, 2003(Bradler, , 2009(Bradler, , 2015Bradler & Buckley, 2018;Robertson et al., 2018;Brock et al., 2020). ...
... Limited spatial dispersion, and extensive adaptive radiation, have led to a high degree of convergent traits in Phasmatodea, such as those related to visual camouflage (Bradler, 2003(Bradler, , 2009(Bradler, , 2015Bradler & Buckley, 2018), oviposition techniques (Carlberg, 1983(Carlberg, , 1987Sellick, 1997a, b;Goldberg et al., 2015;Robertson et al., 2018;, different degrees of wing loss Whiting et al., 2003;Zeng et al., 2019), and ecomorphs (morphological forms with similar ecological occupancy) with specific vertical stratification within the vegetation (Büscher et al., 2018a;Buckley et al., 2010;Bank et al., 2021). Phasmids are predominantly nocturnal insects that are distributed nearly worldwide and exclusively feed on plants (Bradler, 2003(Bradler, , 2009(Bradler, , 2015Bradler & Buckley, 2018;Robertson et al., 2018;Brock et al., 2020). Their greatest diversity occurs in the tropics. ...
Functional systems that evolve as a response to specific environmental challenges often exhibit convergent traits. Organs adapted for attachment to a surface are tuned to a general requirement independent of the phylogenetic position of the organism. The different strategies employed for solving similar problems often represent the same physical principles, and that is why the morphology of attachment structures (and also many other functional systems) is channelled by physical rules. Different animals, therefore, employ similar mechanisms to attach to the broad variety of substrates with different surface conditions. There are two main types of attachment devices that occur on animal legs: hairy and smooth. They differ greatly in their morphology and ultrastructure, but both solve the same problem of proper mechanical adaptation to the variety of natural roughnesses by maximising real contact area with them. Adaptation to specific surface conditions within these groups resulted in several different solutions to the specific ecological surroundings the lineages radiated into. As the conditions are similar in the discrete environments, the adaptations of the attachment systems of different animal groups reveal similar mechanisms. For this reason, on the one hand, similar attachment organs evolved in different lineages of animals, and, on the other hand, different attachment organ modifications occur within the same lineages. In this chapter we summarise the data published in the literature on the structural and functional principles of hairy and smooth attachment pads with a special focus on insects. We describe ultrastructure, surface patterns, the origin of different pads and their evolution, discuss the results of mechanical testing of material properties (elasticity, viscoelasticity, adhesion, friction) and basic physical forces contributing to adhesion, show the influence of different factors, such as substrate roughness and pad stiffness, on contact forces, and review the chemical composition of pad fluids, which are an important component of adhesive function. The structure of this chapter is a kind of fractal. It starts with the omnipresence of the pads in animals. Then we zoom into the phylogeny focusing on insects as the largest animal group on earth, showing convergent evolution of attachment pads therein. In the subsequent step we further zoom in on the phylogeny of one insect group, Phasmatodea, and explore convergent evolution of attachment pads at an even finer scale. Such a hierarchical structure of the chapter helps us to show that convergent evolution occurs at different levels within the animal tree of life. Since convergent events might be potentially interesting for engineers in revealing a kind of optimal solution by nature. Finally, the biomimetic implications of the discussed results are briefly presented.
... Pantel (1915) discussed the structure and variability of the vomer in different subgroups of Phasmatodea and detected a vomer in three genera of "Ascepasmini" (= Aschiphasmatini) including Presbistus. Bradler (1999Bradler ( , 2009 argued that the vomer was already present in the phasmatodean ground plan and that it represents a plesiomorphic character for all Euphasmatodea. Most authors have only described the external part of the vomer, and a standardized nomenclature for the different parts of this organ is still needed. ...
... The characters of the male vomer are frequently used when describing or differentiating stick insect taxa (Bradler 2009, Bresseel and Constant 2018a, Cumming et al. 2021. The nomenclature proposed here for the dissected vomer can be used in nearly all species, and the structure of the vomer provides useful differential characters when studied from different angles. ...
... The body of the vomer shows considerable variation in shape but most often narrows towards the posterior. This organ can be functionally replaced or can be strongly reduced in genera with a modified tergum X or specialized cerci for clasping the female (Pantel 1915, Bradler 2003, 2009, Hennemann and Conle 2008. ...
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.
... comm., 2022). (Caudell, 1904) Arumatia dubia (Caudell, 1904) -Brock & Hasenpusch 2007: 7, 70;2009: 151. -Araujo & Garraffoni 2012 235. ...
... Several lineages of Phasmatodea present males with specialisations for grasping the female terminalia, often involving modifications of the cerci or tergum X (Bradler 2009). However, we are not aware of any specialisation similar to that of Arumatia motenata gen. ...
... Interestingly, some representatives of Neotropical Diapheromerini, including Phantasca and Arumatia gen. nov., share a few female features with some representatives of the Diapheromerini clade Eusermyleformia (see Bradler 2009;Hennemann & Conle 2012), more specifically with species of the North American genera Manomera Rehn & Hebard, 1907 and Diapheromera Gray, 1835, which could be better compared through available literature. These common features consist of the short subgenital plate, the prominent epiproct visible in dorsal view, the parallel, posteriorly pointing and often elongate cerci, the very long gonoplac and absent or reduced gonangulum, the reduction or lack of a praeopercular organ, the incurved and widened posterior region of tergum VII and the often elongate basitarsi. ...
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.).
... Long tegmina are in fact considered the plesiomorphic condition of Polyneoptera, which are suggested to have been independently reduced in various lineages such as in stoneflies or webspinners as well as stick and leaf insects [79], albeit it remains unclear when and how often forewing shortening occurred in Phasmatodea. Also the fossils' tegmina venation is different and its modification in modern phasmatodeans might be due to a convergent evolutionary process of wing shortening and not because of common ancestry, especially when considering the few extant, unrelated and strongly subordinated species that exhibit long tegmina with venation differing from other recent forms (i.e., Heteropteryx, Prisopus, Phylliidae) [78,80,81]. While the large tegmina of the flightless leaf insect females (Phylliidae) contribute to their leaf mimicry and therefore might have been secondarily elongated [82,83] (the volant males possess shorter tegmina), the wing morphology of male Heteropteryx is most similar to that of stem phasmatodeans [36,78,81]. ...
... Also the fossils' tegmina venation is different and its modification in modern phasmatodeans might be due to a convergent evolutionary process of wing shortening and not because of common ancestry, especially when considering the few extant, unrelated and strongly subordinated species that exhibit long tegmina with venation differing from other recent forms (i.e., Heteropteryx, Prisopus, Phylliidae) [78,80,81]. While the large tegmina of the flightless leaf insect females (Phylliidae) contribute to their leaf mimicry and therefore might have been secondarily elongated [82,83] (the volant males possess shorter tegmina), the wing morphology of male Heteropteryx is most similar to that of stem phasmatodeans [36,78,81]. As subordinated lineage within the ground-dwelling and predominantly wingless Heteropterygidae, it was suggested that Heteropteryx secondarily became arboreal and the male regained the capability of active flight. ...
... Our results substantiate this hypothesis, particularly, when considering the results of the traitdependent diversification analysis (HiSSE) from which we concluded that flight or flightlessness might not have been the main driver of euphasmatodean diversification. Alternatively, the evolution of hard-shelled eggs [55,81,87] and the acquisition of endogenous pectinase genes [88] further shaping the co-evolution with their food plants are most likely to have contributed to the success of the early euphasmatodean lineages. Also the recurrent opportunities for colonising new land masses (i.a., the Indo-Pacific region) appears to have promoted speciation [80,89] and might explain the increased diversification rate recovered for Lonchodidae, Lanceocercata and relatives within Oriophasmata (square symbol; Additional file 1: Fig. S7). ...
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.
... It is interesting that A. vetulum has a slightly slender body shape and unique male genitalia that differs from susumaniids but with a similar wing venation. It may be that Pterophasmatidae intermingle plesiomorphic traits from Susumaniidae with modern S8-S11 online) are apterous nymphs (without wing buds), have slender bodies, emarginate labra, short prothoraces, elongate meso-and metathoraces, metasterna fused with abdominal sternum I, basally curved profemora, pentamerous tarsi, and undivided cerci, all of which supports their phylogenetic placement within Euphasmatodea (family Incertae sedis) [6]. It is important that these species had already evolved twig mimicry during the mid-Cretaceous, exhibiting the same morphological modifications of the thorax as other extant Euphasmatodea. ...
... The fusion of the metasternum with abdominal sternum I is considered to be a synapomorphy for Timematodea and Euphasmatodea [6], and is easily discerned in A. grossa of Susumaniidae (Fig. S1e online), three groups of Pterophasmatidae (Pterophasma, Leptophasma, Meniscophasma) [5], and the mid-Cretaceous species of Euphasmatodea reported herein (Fig. S7d online), indicating that this may be a character shared more broadly by all Phasmatodea s.l. In addition, a clear boundary can be observed between the metanotum and abdominal tergum I in Susumaniidae (Fig. 1e and Fig. S1d online), Pterophasmatidae ( Fig. 1f and Fig. S2g online), and Timematodea, while the metanotum is slightly fused with abdominal tergum I in those mid-Cretaceous species of Euphasmatodea (Fig. 1g and Figs. ...
... Las descripciones del cuerpo siguen la terminología estándar de Grimaldi y Engel (2005), enfocándose en los términos utilizados para insectos palo de Bragg (1997). El poculum es definido por Bragg (1997) como "La tapa de la abertura genital: el esternito noveno" en algunos casos dividido transversalmente (Bradler 2009). Se reconoce que el poculum está dividido en dos esternitos, se nombra esclerito basal del poculum al esclerito ubicado en contacto con el esternito VIII; Conle et al. (2007) lo reconocen como esternito IX. ...
Se describe Nooxapty gen. nov. y dos especies: Nooxapty isabelae sp. nov. y Nooxapty mixe sp. nov. para el norte de Oaxaca, México. Este género nuevo es colocado en la tribu Diapheromerini Kirby, 1904 y está relacionado con varias especies de Pseudosermyle Caudell, 1903. Nooxapty clavigerum comb. n. es cambiado de Pseudosermyle y colocado en Nooxapty gen. n., debido a que comparte sus caracteres principales.
... Las descripciones del cuerpo siguen la terminología estándar de Grimaldi y Engel (2005), enfocándose en los términos utilizados para insectos palo de Bragg (1997). El poculum es definido por Bragg (1997) como "La tapa de la abertura genital: el esternito noveno" en algunos casos dividido transversalmente (Bradler 2009). Se reconoce que el poculum está dividido en dos esternitos, se nombra esclerito basal del poculum al esclerito ubicado en contacto con el esternito VIII; Conle et al. (2007) lo reconocen como esternito IX. ...
Nooxapty gen. nov. and Nooxapty isabelae sp. nov. and Nooxapty mixe sp. nov. were described from Oaxaca, Mexico.
... The presence of two free cells in the anal area is also unlikely for an attribution to the Paoliida and Dictyoptera (Prokop et al. 2014), Phasmatodea (Bradler 2009), Embioptera (Ross 2000), Zoraptera (Kukalová-Peck and Peck 1993), and the majority of the winged insects currently attributed to the 'Grylloblattida' (sensu Storozhenko 1998) or the 'Eoblattida' (Aristov 2015), even if the Paoliida, Dictyoptera, and 'Grylloblattida' have a m-cua vein and a convex CuA emerging from a stem Cu. Few 'Grylloblattida' (e.g. the Geinitziidae Prosepididontus calopteryx Handlirsch, 1920 or Sinosepididontus chifengensis Huang and Nel, 2008) also have two cells in the anal area, but they also have much more crossveins in the radial and median areas than in the new fossil, a different organisation of the cells of the anal area, and a different pattern of the vein CuA clearly subdivided in two main branches close to crossvein m-cua (Ansorge and Rasnitsyn 2000;Huang and Nel, 2008). ...
Endopterygote insects are very scarcely found in the Carboniferous strata while their occurrence during the Permian becomes more common. Here, we present the earliest evidence of the stem group Amphiesmenoptera (clade Mecopterida) from the Pennsylvanian (Moscovian) of the Piesberg quarry near Osnabrück (Lower Saxony, Germany), which shares a number of venation traits with representatives of the Permian family Microptysmatidae. Microcarbonella paradoxa gen. nov. et sp. nov. is based on the wing venation diagnosed by the characteristic branching pattern of radial veins and particularly looping anal area in form of two anal cells. This tiny insect species with wing length reaching over 6 mm demonstrates that early caddisfly-like endopterygotes co-existed in coal swamp ecosystems along with far more abundant hemimetabolous insects.
urn:lsid:zoobank.org:pub:47768C95-782C-4FBA-829D-D62E5F4A4154
The genus Pterinoxylus Serville, 1838 is redescribed and revised at the species level. It is distributed throughout most of Central America, the northern half of South America and also has one species on the Lesser Antilles. Two new species are described from Costa Rica: P. cocoense n. sp. from both sexes and the eggs and P. speciosus n. sp. from both sexes. The female of P. perarmatus (Redtenbacher, 1908) is described and illustrated for the first time, as are the eggs of the type-species P. eucnemis Serville, 1838 and P. perarmatus (Redtenbacher, 1908).
The process of biological evolution has resulted in a wide variety of forms,
functions and strategies and this has led to distinct optimization of certain traits in
organisms. Technical adoption of some “inventions of nature” might be highly
beneficial for innovative developments in materials science and engineering through
biomimetic studies. One prominent field of biomimetic research is related to
prehension and manipulation mechanisms in robotics since these tasks are just as
ubiquitous in technical environments as they are in nature. Biological end effectors
with purposes ranging from simple locomotion, mating and prey catching up to delicate
object manipulation have been realized there, with innumerable, sometimes subtle
variants of structure and properties between them. Even though there is some coarse
biological classification of biological gripping devices, the latter represent certain core
principles, which evolved convergently due to the underlying basic physical
phenomena. This chapter aims to categorize the most common physical principles, their
advantages and shortcomings, and present a range of biological examples. Furthermore,
possible transfers of functional principles from biological systems into technical
environments are discussed.
A synonymic catalog of species, essential for researchers. 414 pages, spiral bound [replaces a CD issued in 2003 (First Edition)] OUT OF PRINT
Timema coffmani sp. nov. is described from southwestern Arizona. It is a specialist feeder on juniper (Juniperus). Morphologically it is similar to T. ritensis Hebard from southeastern Arizona. The previously unknown female sex of T. ritensis also is described and both sexes are compared with T. coffmani. Measurements and distinguishing features are given for both species.
Amino acid sequence data are available for ribulose biphosphate carboxylase, plastocyanin, cytochrome c, and ferredoxin for a number of angiosperm families. Cladistic analysis of the data, including evaluation of all equally or almost equally parsimonious cladograms, shows that much homoplasy (parallelisms and reversals) is present and that few or no well supported monophyletic groups of families can be demonstrated. In one analysis of nine angiosperm families and 40 variable amino acid positions from three proteins, the most parsimonious cladograms were 151 steps long and contained 63 parallelisms and reversals (consistency index = 0.583). In another analysis of six families and 53 variable amino acid positions from four proteins, the most parsimonious cladogram was 161 steps long and contained 50 parallelisms and reversals (consistency index = 0.689). Single changes in both data matrices could yield most parsimonious cladograms with quite different topologies and without common monophyletic groups. Presently, amino acid sequence data are not comprehensive enough for phylogenetic reconstruction among angiosperms. More informative positions are needed, either from sequencing longer parts of the proteins or from sequencing more proteins from the same taxa.
Wing-dimorphic insects are excellent subjects for a study of the evolution of dispersal since the nondispersing brachypterous morph is easily recognized. The purpose of this paper is to develop a framework within which the evolution of wing dimorphism can be understood. A review of the literature indicates that the presence or absence of wings may be controlled by a single locus, two-allele genetic system or a polygenic system. Both types of inheritance can be subsumed within a general threshold model. An increase in the frequency of a brachypterous morph in a population may result from an increased relative fitness of this morph or the emigration of the macropterous type. The abundance of wing-polymorphic species argues for an increased fitness of the brachypterous form. An analysis of the life-history characteristics of 22 species of insects indicates that the brachypterous morph is both more fecund and reproduces earlier that the macropterous morph. Unfortunately, data on males are generally lacking. It is suggested that suppression of wing production results when some hormone, perhaps juvenile hormone, exceeds a threshold value during a critical stage of development. Further, it is known that in the monomorphically winged species Oncopeltus fasciatus both flight and oviposition are regulated by the titer of juvenile hormone. These observations are used to construct a possible pathway for the evolution of wing dimorphism. This suggests that evolution to a dimorphic species requires both an increase in the rate of production of the wing suppressing hormone and a change in the threshold level at which wing and wing-muscle production are suppressed. The stage in this evolutionary sequence that an organism will reach depends on the stability of the habitat.
