No full-text available
To read the full-text of this research,
you can request a copy directly from the authors.
Taxonomic revision of Paraphasma Redtenbacher, 1906 (Phasmatodea, Pseudophasmatidae) based on phallic and external morphology
Abstract
Paraphasma Redtenbacher, 1906 is a genus of fully-winged stick insects occurring in central and northern South America. We carried out a morphology-based taxonomic revision of this genus with emphasis on the phallic organ, a structure that has been poorly explored for taxonomic purposes in Phasmatodea. Additionally, pairwise genetic distances between mitochondrial COI gene sequences were calculated for ten Paraphasma specimens representing six species. We recognize nine valid species in the genus plus one nomen dubium, Paraphasma fasciatum Gray, 1835. We redescribe Paraphasma and the species previously assigned to it, describe Paraphasma indistinctum Chiquetto-Machado sp. nov., Paraphasma sooretama Chiquetto-Machado sp. nov. and Paraphasma spinicauda Chiquetto-Machado sp. nov., and provide a key to the species in the genus. The male of Paraphasma minus Redtenbacher, 1906 is described for the first time, as well as the eggs of six species. We transfer Paraphasma amabile Redtenbacher, 1906 to Pseudophasma Kirby, 1896 (comb. nov.) and synonymize Pseudophasma xanthotaenidium Günther, 1930 under this species (syn. nov.). In addition, Phasma perspicillaris Stoll, 1813 is removed from the synonymy of Paraphasma laterale (Fabricius, 1775) and synonymized under Parastratocles xanthomela (Olivier, 1792) (syn. nov.). The examination of the phallic organ was essential for species delimitation, as most species of Paraphasma are very similar in the external morphology of both sexes. The analysis of the COI sequences supported the species delimitation, resulting in remarkably lower pairwise distances between conspecific individuals (p-distance ≤ 2.0%) than between different species (p-distance 6.9–17.5%). We hope that this paper will stimulate further studies exploring the taxonomic and phylogenetic potential of the internal male genitalia of stick insects.
... Neotropical stick insects are generally poorly studied: many species are known only from their original descriptions and relatively few have been covered in recent taxonomic works (see, e.g., Gutiérrez-Valencia et al. 2017;Conle et al. 2020;Crispino et al. 2020;Madeira-Ott et al. 2020;Chiquetto-Machado & Cancello 2021;Ghirotto 2021;Chiquetto-Machado et al. 2022). Compared to the fauna of other continents, Neotropical phasmids are underrepresented in phylogenetic analyses, reflecting the lack of basic scientific work concerning them (Robertson et al. 2018;Simon et al. 2019). ...
... More specific studies must be conducted to attend to these questions. Nevertheless, this is one more of recent recurring examples of the benefits of an integrative taxonomy in Phasmatodea (Gutiérrez-Valencia et al. 2017;Crispino et al. 2020;Chiquetto-Machado & Cancello 2021;Ghirotto 2021;Chiquetto-Machado et al. 2022). ...
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.).
... The Phasmatodea is one of the least-studied insect orders in Brazil; recently, the state of the art of this group was reviewed by Madeira-Ott et al. [8]. New initiatives have been increasing the knowledge about this group, especially regarding taxonomy, morphology, biology, and ecology [9][10][11][12][13][14][15][16][17][18][19]. However, several factors bring difficulties to the study of this order in Brazil, such as intraspecific morphological variation, the low number of specimens collected, and a lack of specialised literature [20]. ...
Cladomorphus petropolisensis sp. nov., a new species of stick insect from Petrópolis, Rio de Janeiro, Brazil, is herein described and compared to the other sympatric species, C. phyllinus Gray, 1835 (Phasmatidae, Cladomorphinae). The description of the new species is supported by morphological and molecular evidence. Kimura-2-parameter (K2P) intraspecific COI divergences among the holotype of C. petropolisensis sp. nov. and C. phyllinus individuals ranged from 2.9% to 4.4%, which are suggestive of distinct species, especially when considering that all Cladomorphus individuals studied were collected in the Petrópolis municipality. The new species can be distinguished from C. phyllinus Gray, 1835 by several characteristics: smaller size, the presence of two spines on the hind femora, the relative longer length of the ovipositor, and spiny tegument, especially in the mesonotum, sculpturing of the operculum of the egg.
Based on the observation of specimens kept in captivity for 69 days, here we describe in detail for the first time the predatory behavior of an assassin bug Harpactor angulosus (Lepeletier & Serville, 1825) (Hemiptera: Heteroptera: Reduviidae: Harpactorinae), on a stick insect Cladomorphus phyllinus Gray, 1835 (Phasmatodea: Phasmatidae: Cladomorphinae). The behavior of the generalist predator H. angulosus as well as the evasive reactions of C. phyllinus were detailedly described and illustrated. Because both species occur in the Atlantic Forest biome and were recorded in the same geographic coordinates, in an urbanized green area in the city of Petrópolis, Rio de Janeiro State, Brazil, the ecological relationship here described is also plausible of occurring in nature and opens new venues to be explored. The unprecedented phytophagy of H. angulosus was also recorded.
Stick and leaf insects (Phasmatodea) are large terrestrial herbivorous arthropods known for masquerading as plant parts such as bark, twigs and leaves. Their evolutionary history is largely shaped by convergent evolution associated with adaptive radiations on geographically isolated landmasses that have repeatedly generated ground-dwelling ecomorphs. The members of one lineage, however, the Oriental Heteropterygidae, are morphologically rather uniform, and have a predominantly ground-dwelling lifestyle. The phylogeny of Heteropterygidae that comprises approximately 130 described species is controversial and remains uncertain. In particular, the systematic position of the giant Jungle Nymph Heteropteryx dilatata, whose males are capable of flight and exhibit the most plesiomorphic wing morphology among extant phasmatodeans, is of major interest to the scientific community. Here, we analysed a set of seven nuclear and mitochondrial genes to infer the phylogeny of Heteropterygidae covering the group's overall diversity. The divergence time estimation and reconstruction of the historical biogeography resulted in an ancestral distribution across Sunda-land with long distance dispersal events to Wallacea, the Philippines and the South Pacific. We were able to resolve the relationships among the three principal subgroups of Heteropterygidae and revealed the Dataminae, which contain entirely wingless small forms, as the sister group of Heteropteryginae + Obriminae. Within Heteropteryginae, Haaniella is recovered as paraphyletic in regard to Heteropteryx. Consequently, Heteropteryx must be considered a subordinate taxon deeply embedded within a flightless clade of stick insects. Within Obriminae, the Bornean Hoploclonia is strongly supported as the earliest diverging lineage. Based on this finding, we recognize only two tribes of equal rank among Obriminae, the Hoplocloniini trib. nov. and Obrimini sensu nov. Within the latter, we demonstrate that previous tribal assignments do not reflect phylogenetic relationships and that a basal splitting event occurred between the wing-bearing clade Miroceramia + Pterobrimus and the remaining wingless Obrimini. The Philippine genus Tisamenus is paraphyletic with regard to Ilocano hebardi, thus, we transfer the latter species to Tisamenus as Tisamenus hebardi comb. nov. and synonymize Ilocano with Tisamenus. We discuss character transformations in the light of the new phylogenetic results and conclude that the current taxonomic diversity appears to be mainly driven by allopatry and not to be the result of niche differentiation. This radiation is thus best described as a nonadaptive radiation.
While the leaf insects (Phylliidae) are a well-supported group within Phasmatodea, the genus Phyllium Illiger, 1798 has repeatedly been recovered as paraphyletic. Here, the Phyllium (Phyllium) celebicum species group is reviewed and its distinctiveness from the remaining Phylliini genera and subgenera in a phyloge- netic context based on morphological review and a phylogenetic analysis of three genes (nuclear gene 28S and mitochondrial genes COI and 16S) from most known and multiple undescribed species is shown. A new genus, Cryptophyllium gen. nov., is erected to partially accommodate the former members of the celebicum species group. Two species, Phyllium ericoriai Hennemann et al., 2009 and Phyllium bonifacioi Lit & Eusebio, 2014 morphologically and molecularly do not fall within this clade and are therefore left within Phyllium (Phyllium). The transfer of the remaining celebicum group members from Phyllium Illiger, 1798 to this new genus creates the following new combinations; Cryptophyllium athanysus (Westwood, 1859), comb. nov.; Cryptophyllium celebicum (de Haan, 1842), comb. nov.; Cryptophyllium chrisangi (Seow-Choen, 2017), comb. nov.; Cryptophyllium drunganum (Yang, 1995), comb. nov.; Cryptophyl- lium oyae (Cumming & Le Tirant, 2020), comb. nov.; Cryptophyllium parum (Liu, 1993), comb. nov.; Cryptophyllium rarum (Liu, 1993), comb. nov.; Cryptophyllium tibetense (Liu, 1993), comb. nov.; Crypto- phyllium westwoodii (Wood-Mason, 1875), comb. nov.; Cryptophyllium yapicum (Cumming & Teemsma, 2018), comb. nov.; and Cryptophyllium yunnanense (Liu, 1993), comb. nov.
The review of specimens belonging to this clade also revealed 13 undescribed species, which are described within as: Cryptophyllium animatum gen. et sp. nov. from Vietnam: Quang Nam Province; Cryptophyllium bankoi gen. et sp. nov. from Vietnam: Quang Ngai, Thua Thien Hue, Da Nang, Gia Lai, Quang Nam, and Dak Nong Provinces; Cryptophyllium bollensi gen. et sp. nov. from Vietnam: Ninh Thuan Province; Cryptophyllium daparo gen. et sp. nov. from China: Yunnan Province; Cryptophyllium echidna gen. et sp. nov. from Indonesia: Wangi-wangi Island; Cryptophyllium faulkneri gen. et sp. nov. from Vietnam: Quang Ngai and Lam Dong Provinces; Cryptophyllium icarus gen. et sp. nov. from Vi- etnam: Lam Dong and Dak Lak Provinces; Cryptophyllium khmer gen. et sp. nov. from Cambodia: Koh Kong and Siem Reap Provinces; Cryptophyllium limogesi gen. et sp. nov. from Vietnam: Lam Dong, Dak Lak, and Dak Nong Provinces; Cryptophyllium liyananae gen. et sp. nov. from China: Guangxi Province; Cryptophyllium nuichuaense gen. et sp. nov. from Vietnam: Ninh Thuan Province; Cryptophyllium phami gen. et sp. nov. from Vietnam: Dong Nai and Ninh Thuan Provinces; and Cryptophyllium wennae gen. et sp. nov. from China: Yunnan Province. All newly described species are morphologically described, il- lustrated, and molecularly compared to congenerics.
With the molecular results revealing cryptic taxa, it was found necessary for Cryptophyllium westwoo- dii (Wood-Mason, 1875), comb. nov. to have a neotype specimen designated to allow accurate differen- tiation from congenerics. To conclude, male and female dichotomous keys to species for the Cryptophyl- lium gen. nov. are presented.
Objective synonyms of phasmids involving Olivier, 1792, Stoll, 1788 and Lichtenstein, 1796 are listed and discussed.
Stick insects (Phasmatodea) are among the least studied insect groups in Brazil. Although in recent years there has been an increase in the number of published studies on taxonomy and morphology of Brazilian stick insects, they remain very little-known concerning biological and bionomic aspects. From five females of Cladoxerus cryphaleus (Phasmatidae: Cladomorphinae) collected in the Iguaçu National Park, Paraná, Brazil, a captive colony of parthenogenetic females was started, based on which data on longevity, fertility, and viability of eggs of this species were recorded for the first time. After imaginal molting, the 21 females studied had an average longevity of 103.7 days. Each female produced on average 125.3 eggs, which had a hatching rate of 34.2%. In addition, we recorded a C. cryphaleus male nymph being parasitized by a biting midge of the subgenus Forcipomyia (Microhelea) (Diptera: Ceratopogonidae). The male, in pre-imaginal instar and already parasitized, was collected in the municipality of Petrópolis, Rio de Janeiro.
Stick and leaf insects (Phasmatodea) are a distinctive insect order whose members are characterized by mimicking various plant tissues such as twigs, foliage and bark. Unfortunately, the phylogenetic relationships among phasmatodean subfamilies and the timescale of their evolution remain uncertain. Recent molecular clock analyses have suggested a Cretaceous–Palaeogene origin of crown Phasmatodea and a subsequent Cenozoic radiation, contrasting with fossil evidence. Here, we analysed transcriptomic data from a broad diversity of phasmatodeans and, combined with the assembly of a new suite of fossil calibrations, we elucidate the evolutionary history of stick and leaf insects. Our results differ from recent studies in the position of the leaf insects (Phylliinae), which are recovered as sister to a clade comprising Clitumninae, Lancerocercata, Lonchodinae, Necrosciinae and Xenophasmina. We recover a Permian to Triassic origin of crown Phasmatodea coinciding with the radiation of early insectivorous parareptiles, amphibians and synapsids. Aschiphasmatinae and Neophasmatodea diverged in the Jurassic–Early Cretaceous. A second spur in origination occurred in the Late Cretaceous, coinciding with the Cretaceous Terrestrial Revolution, and was probably driven by visual predators such as stem birds (Enantiornithes) and the radiation of angiosperms.
Phasmatodea (Arthropoda, Insecta) comprises phytophagous insects, popularly known as stick bugs because most of them resemble dry branches. Currently, more than 3000 species are known, the majority of which can be found in tropical regions of the world. In Brazil, it is estimated that there are approximately 200 species, but these numbers may not truly reflect the group’s richness. A high degree of intraspecific polymorphism, a low number of specimens sampled, a shortage of specialized literature, and having only a small number of experts have amplified the difficulties in studying this order. Therefore, this study aims to present a historical survey of the Phasmatodea studies that have been conducted in Brazil thus far, indicating gaps in knowledge and discussing perspectives to expand understanding within this group. Furthermore, to the best our knowledge, the molecular characterization of the cytochrome oxidase subunit I gene of mitochondrial DNA of two species of phasmids found in Brazil, Cladomorphus phyllinus Gray, 1835, and Pseudophasma missionum Piza, 1981, is presented for the first time. In addition, this study records for the first time the occurrence of P. missionum in Brazilian territory.
Morphology of Phasmatodea eggs is remarkably diverse and highly valuable in taxonomic research. Two alterative hypotheses have been proposed to describe the phylogenetic relationship of the species from the genus Agathemera Stål. Additionally, descriptions of the egg morphology within Agathemera have been done based on the eggs of two species. This small sample size does not represent the diverse egg morphology along the genus, thus we attempt to describe the eggs from all the known Agathemera species. The main goal of the present study is to determine whether the evolution of the eggs occurred through either divergent or convergent evolution. We based our descriptions on morphometrics, morphology and the ultrastructure. For data analysis, principal component analysis (PCA) was performed on morphometric variables and the characters emerged from the morphological and ultrastructure were mapped over the molecular phylogeny. The results show that it is possible to discriminate among species using the morphology of the different egg structures, and furthermore, a divergent event at the base of the tree, differentiate the overall egg shape and the internal micropylar plate shape. Finally, we conclude that both divergent and convergent evolution are shaping the different structures of the Agathemera eggs.
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.
Achrioptera is a taxon of extremely large and exceptionally colorful stick insects endemic to Madagascar and the Comoros Archipelago. We studied the phylogenetic position of the Achriopterini, comprising the genera Achrioptera and Glawiana, based on a multigene phylogeny and concluded that it is a sister group to other Madagascan phasmids (Anisacanthidae) rather than to Neotropical or Australo-Pacific groups as was suggested in a previous study based on morphology. Our results also point to unresolved relationships (potential paraphyly of Achrioptera), taxonomic issues (elevation of A. punctipes cliquennoisi to species level), and detection of cryptic diversity (in A. impennis), demonstrating the need of additional research. A DNA barcoding approach based on COI sequences of Achrioptera species revealed a clear discrimination between closely related and morphologically similar species. Applying integrative taxonomy using multiple lines of evidence, we demonstrated that the well-known species with blue males from Montagne des Français and Foret d'Orangea in the far north of Madagascar, previously attributed to Achrioptera fallax, represents a new species, which we describe as Achrioptera manga sp. nov. based on morphological, chromatic, and genetic (mitochondrial and nuclear) differences. We also describe a second new giant species from this massif: Achrioptera maroloko sp. nov. is among the largest insects (females reaching up to 24 cm total length) and differs from its sister species A. spinosissima from western Madagascar in morphology, coloration, and substantial DNA barcode divergence. These magnificent new species confirm the significance of the Montagne des Français area as a hotspot of biodiversity and microendemism. The biogeographic pattern of the species pair A. fallax/A. manga is paralleled by species pairs of reptiles and amphibians suggesting a similar evolutionary history. Finally, we discuss the sexual dichromatism of Achrioptera species with conspicuous males and mostly cryptic females. As possible reasons, we consider female mate choice and divergent habits of males and females, but aposematism combined with toxic substances produced in defense glands or accumulated in the insect's body from nutritional plants are more plausible explanations for this phenomenon.
Stick and leaf insects (Phasmatodea) are large, tropical, predominantly nocturnal herbivores, which exhibit extreme masquerade crypsis, whereby they morphologically and behaviorally resemble twigs, bark, lichen, moss, and leaves. Females employ a wide range of egg-laying techniques, largely corresponding to their ecological niche, including dropping or flicking eggs to the forest floor, gluing eggs to plant substrate, skewering eggs through leaves, ovipositing directly into the soil, or even producing a complex ootheca. Phasmids are the only insects with highly species-specific egg morphology across the entire order, with specific egg forms that correspond to oviposition technique. We investigate the temporal, biogeographic, and phylogenetic pattern of evolution of egg-laying strategies in Phasmatodea. Our results unequivocally demonstrate that the ancestral oviposition strategy for female stick and leaf insects is to remain in the foliage and drop or flick eggs to the ground, a strategy that maintains their masquerade. Other major key innovations in the evolution of Phasmatodea include the (1) hardening of the egg capsule in Euphasmatodea; (2) the repeated evolution of capitulate eggs (which induce ant-mediated dispersal, or myrmecochory); (3) adapting to a ground or bark dwelling microhabitat with a corresponding shift in adult and egg phenotype and egg deposition directly into the soil; and (4) adhesion of eggs in a clade of Necrosciinae that led to subsequent diversification in oviposition modes and egg types. We infer at minimum 16 independent origins of a burying/inserting eggs into soil/crevices oviposition strategy, 7 origins of gluing eggs to substrate, and a single origin each of skewering eggs through leaves and producing an ootheca. We additionally discuss the systematic implications of our phylogenetic results. Aschiphasmatinae is strongly supported as the earliest diverging extant lineage of Euphasmatodea. Phylliinae and Diapheromerinae are both relatively early diverging euphasmatodean taxa. We formally transfer Otocrania from Cladomorphinae to Diapheromerinae and recognize only two tribes within Diapheromerinae: Diapheromerini sensu nov. and Oreophoetini sensu nov. We formally recognize the clade comprising Necrosciinae and Lonchodinae as Lonchodidae stat. rev. sensu nov.
Stick and leaf insects (order Phasmatodea) are a mesodiverse lineage of large terrestrial herbivores with predominantly tropical distribution and few species inhabiting more temperate regions. The phylogenetic position of the Phasmatodea among the lower neopteran insects has been debated for many years, with basically every orthopteroid insect order proposed as the potential sister taxon. The stick and leaf insects exhibit a remarkably poor fossil record. The fascinating and variable biology of stick insects has made them excellent model systems for investigating a number of evolutionary phenomena, including speciation and reproductive isolation, evolution of parthenogenesis and alternative reproductive strategies, and more recently the evolution of cold tolerance. Evidence for monophyletic Phasmatodea is undisputed and has grown stronger in recent years, with evidence coming from various sources. The chapter lists and discusses the currently recognized, non‐encaptic monophyletic groups. The contributions of amateur taxonomists play a crucial role in describing the phasmatodean diversity.
Insects have developed specialized structures on their feet for adhering to surfaces, with stick and leaf insects or Phasmatodea exhibiting an unexpectedly high diversity of these structures. In Phasmatodea, attachment on different substrates is achieved by two types of pads on the legs: the euplantulae on the tarsomeres and the arolium on the pretarsus. The euplantulae are adhesive structures capable of adaptability to the substrate profile and generation of the required attachment strength. The diversity of euplantular microstructures of 56 species that represent all major lineages recognized within Phasmatodea and the whole biogeographical distribution of the group are examined using scanning electron microscopy (SEM). Nine different types of attachment structures can be distinguished whereby one, the nubby type, can be further divided into three different distinct types based on the specific ratio of each conical outgrowth. We mapped the morphological data from the SEM onto a phylogenetic tree we reconstructed based on molecular data. Previously, the evolution of different adhesive microstructures (AMs) on these pads has been suggested to reflect phylogenetic groups. However, different types of AMs are found within monophyletic groups, and our ancestral character state reconstruction suggests smooth euplantulae in the ground pattern of Euphasmatodea and multiple independent origins of other forms. The type of AM appears to be strongly associated with ecomorphs, e.g., smooth euplantular surfaces are more frequently found in tree-dwellers than in ground-dwellers, whilst the attachment pads of ground-dwelling species primarily bear conical cuticular outgrowths (nubby euplantulae).
The South American genus Phantasca Redtenbacher, 1906 (Phasmatodea: Diapheromeridae: Diapheromaerinae) is re-diagnosed and revised at the species level. The precedingly unknown eggs are described for the first time. The genus Pterolibethra Günther, 1940 (type species: P. heteronemia Günther, 1940) is re-synonymised, with Phantasca (syn. nov.) and consequently the two species originally contained, P. heteronemia Günther, 1940 and P. poeciloptera Günther, 1940, are transferred to Phantasca (comb. rev.). P. laeta Conle, Hennemann & Gutierréz, 2011 is not congeneric and is transferred to the genus Jeremiodes Hennemann & Conle, 2007 (Cladomorphinae: Cladomorphini; comb. nov.). Two species are removed from Bacteria Berthold, 1827 and transferred to Phantasca; these are B. quadrilobata Chopard, 1911 and B. montana Redtenbacher, 1906 (comb. nov.). Six new species are described: P. adiposa sp. nov., P. amabile sp. nov., P. femorata sp. nov., P. guianensis sp. nov., P. nigrolineata sp. nov. and P. ruboligata sp. nov. The male and egg of P. quadrilobata (Chopard, 1911) are described and illustrated for the first time. The genus now contains 13 species that are distributed throughout the northern half of South America. A key as well as detailed descriptions and illustrations are presented for all known species.
The molecular evolutionary genetics analysis (Mega) software implements many analytical methods and tools for phylogenomics and phylomedicine. Here, we report a transformation of Mega to enable cross-platform use on Microsoft Windows and Linux operating systems. Mega X does not require virtualization or emulation software and provides a uniform user experience across platforms. Mega X has additionally been upgraded to use multiple computing cores for many molecular evolutionary analyses. Mega X is available in two interfaces (graphical and command line) and can be downloaded from www.megasoftware.net free of charge.
Examination of unidentified Phylliidae specimens revealed a number of undescribed species from the island of Luzon, Philippines. Morphological and molecular study of specimens from the obscure phasmid genus Microphyllium Zompro, 2001, revealed a new species, which we describe as Microphyllium haskelli Cumming sp. nov.. It is here described and differentiated from the two other species in the genus, both currently only known from adults of a single sex. Pseudomi-crophyllium Cumming gen. nov. is described as a new genus within Phylliidae with the type species Pseudomicrophyllium faulkneri Cumming gen. et sp. nov. as the sole known species in the genus. As is unfortunately often the case in the leaf-mimicking family Phylliidae, this new genus and species is only known from a single specimen. In addition to the new genus, two new Phyllium (Phyllium) species from the siccifolium species-group are named and described as Ph. (Ph.) an-tonkozlovi Cumming sp. nov. and Ph. (Ph.) bourquei Cumming & Le Tirant sp. nov.. In addition to the newly described species, Phyllium (Phyllium) geryon Gray, 1843 is redescribed from a nearly perfect specimen, completing some of the morphological knowledge gaps currently missing because of the severely damaged holotype specimen. A key to all known species of Phylliidae from Luzon is included. Holotype specimens for all four new species will be deposited in the National Museum of the Philippines type collection and paratype specimens will be deposited into the San Diego Natural History Museum collection or retained within the first author's collection.
The stick insect Paraphasma paulense was described based on one male from the vicinity of São Paulo city, state of São Paulo, Brazil. Here we describe and illustrate the female, egg and first instar nymph of the species, including scanning electron micrographs of the egg. In addition, we give brief behavioural reports and present a few comments on the biology and morphology of P. paulense.
Type specimens of 67 taxa of Phasmida (including probable type specimens of 24 taxa) have been located in the Zoological Institute, Russian Academy of Sciences, St. Petersburg. The species are listed alphabetically, with the number of specimens, sex and locality data.
Type specimens of 437 Phasmida taxa have been located in the Natural History Museum, London (NHMUK, formerly BMNH), including 480 primary types of 372 taxa. Taxa with types are listed alphabetically by their specific or subspecific name, and the number of specimens, sex and locality data are given.
Background: The study of islands as model systems plays a key role in understanding many evolutionary processes. Knowledge of the historical events leading to present-day island communities is pivotal for exploring fundamental mechanisms of speciation and adaptation. The remote Mascarene archipelago (Mauritius, Réunion, Rodrigues), considered to be the product of an age-progressive trend of north-to-south volcanic activity in the Indian Ocean, hosts a remarkably diverse, endemic and threatened concentration of flora and fauna that has traditionally been considered to be biogeographically related to Madagascar and Africa. To explore the evolutionary diversity of the Mascarene stick insects (Phasmatodea), we constructed a global phylogeny from approximately 2.4 kb of mitochondrial and nuclear sequence data of more than 120 species representing all major phasmatodean lineages.
Results: Based on the obtained time-calibrated molecular tree we demonstrate that the current phasmid community of the Mascarene archipelago, which consists of members of four presumably unrelated traditional subfamilies, is the result of a single ancient dispersal event from Australasia and started radiating between 16–29 million years ago, significantly predating the age of Mauritius (8–10 million years).
Conclusions: We propose that the Mascarene stick insects diversified on landmasses now eroded away, presumably to the north of Mauritius. In consequence, ancient islands have probably persisted in the Indian Ocean until the emergence of Mauritius and not only served as stepping stones for colonisation events during sea-level lowstands, but as long-lasting cradles of evolution. These ancient landmasses most likely allowed for adaptive speciation and served as significant sources of diversity that contributed to the biomes of the Mascarene archipelago and the megadiverse Madagascar.
L'association ASPER a été missionnée par le Parc Amazonien de Guyane pour réaliser l'inventaire des Phasmatodea de Saül, village isolé au centre de la Guyane. Le matériel et la méthode utilisés sont détaillés et les espèces observées sont listées. Au total ce sont 32 espèces de 21 genres différents qui ont été inventoriées. Abstract. – Contribution to the inventory of Phasmatodea of Saül, French Guiana. The organization ASPER has been engaged by the Parc Amazonien de Guyane to make the inventory of Phasmatodea of Saül, isolated village in the center of French Guiana. Material and method are detailed. A total of 32 species were observed, belonging to 21 different genera.
The eggs of stick and leaf insects (Phasmatodea) bear strong resemblance to plant seeds and are commonly dispersed by females dropping them to the litter. Here we report a novel egg-deposition mode for Phasmatodea performed by an undescribed Vietnamese species of the enigmatic subfamily Korinninae that produces a complex egg case (ootheca), containing numerous eggs in a highly ordered arrangement. This novel egg-deposition mode is most reminiscent of egg cases produced by members of unrelated insect orders, e.g. by praying mantises (Mantodea) and tortoise beetles (Coleoptera: Cassidinae). Ootheca production constitutes a striking convergence and major transition in reproductive strategy among stick insects, viz. a shift from dispersal of individual eggs to elaborate egg concentration. Adaptive advantages of ootheca formation on arboreal substrate are likely related to protection against parasitoids and desiccation and to allocation of specific host plants. Our phylogenetic analysis of nuclear (28S, H3) and mitochondrial (COI, COII) genes recovered Korinninae as a subordinate taxon among the species-rich Necrosciinae with Asceles as sister taxon, thus suggesting that placement of single eggs on leaves by host plant specialists might be the evolutionary precursor of ootheca formation within stick insects.
We present a phylogenetic analysis that includes all known species of Agathemera, using as outgroup four species of the genera Heteronemia, Spinonemia, Monticomorpha, and Anisomorpha. Phylogenetic inference was based on three genes, 16S, COI (mitochondrial markers) and H3 (nuclear marker), based on the maximum-parsimony, maximum-likelihood and Bayesian inference methods. Our results show that the genus Agathemera is monophyletic; six of its eight species showed phylogenetic support, while the group A. claraziana + A. millepunctata was not resolved. The resulting topology shows two major clades, the first with A. maculafulgens, A. luteola, A. crassa, A. millepunctata, and A. claraziana, and the second including A. grylloidea, A. mesoauriculae, and A. elegans. Species of the first clade have large mesonotal processes and are found mainly east of the Andes, except for A. crassa. Members of the second clade have small or absent mesonotal processes and are distributed mainly to the west of the Andes.
© 2012 The Linnean Society of London, Zoological Journal of the Linnean Society, 2012, 165, 63–72.
The genus Malacomorpha Rehn, 1906 is revised at the species-level, based upon examination of all necessary type-material and extensive material housed in ANSP, CMNH and USNM mainly collected on nine international expeditions and collections at 280 sites distributed throughout the geography of the Dominican Republic from 2001 through 2004. A re-description of the genus and detailed descriptions of all 13 known species are provided. Seven new species are described and illustrated: Malacomorpha bastardoae n. sp., Malacomorpha macaya n. sp., Malacomorpha hispaniola n. sp., Malacomorpha minima n. sp., Malacomorpha multipunctata n. sp. & Malacomorpha obscura n. sp. from Hispaniola and Malacomorpha sanchezi n. sp. from Puerto Rico. The eggs of M. bastardoae n. sp., M. cyllarus (Westwood, 1859), M. jamaicana (Redtenbacher, 1906), M. multipunctata n. sp., M. obscura n. sp., M. sanchezi n. sp., and M. spinicolle (Burmeister, 1838) are described and illustrated, those of the four latter species for the first time. According to the original description and distribution Phasma graveolens King, 1867 is obviously a synonym of Malacomorpha cyllarus (Westwood, 1859), and not a synonym of Anismorpha buprestoides (Stoll, 1813) as stated by previous authors (n. syn.). A lectotype is designated for Phasma spinicolle Burmeister, 1838. The newly described species, but M. longipennis (Redtenbacher, 1906) and M. hispaniola n. sp. in particular, prove the genera Pseudolcyphides Karny, 1923 (Type-species: Phasma spinicolle Burmeister, 1838) and Alloeophasma Redtenbacher, 1906 (Type-species: Anophelepis poeyi Saussure, 1868) to be synonyms of Malacomorpha Rehn, 1906 (n. syn.). Consequently, the type species of both genera are here transferred to Malacomorpha Rehn, 1906 (n. comb.). The genus now contains apterous, brachypterous and pterous species restricted to the Greater Antilles and Bahamas.
American Pseudophasmatinae stick insects are often colorful, winged or apterous, with or without scale-shaped tegmina, with an area apicalis on all tibiae, and either with a ventromedian carina on the meso and metafemora, or with broad profemora, as long as head, pro- and mesonotum combined (Zompro, 2004). In the egg, micropylar plate is externally located in the middle of the dorsal surface, and internally appears of the open type. Among Pseudophasmatinae, the genus Tithonophasma Zompro, 2004 has a single species, Tithonophasma tithonus (Gray, 1835). Although this species is frequent in museum collections (Zompro, 2004), accurate morphological and biological information potentially useful in phylogenetic approaches are still missing.
The type material of Phasmatodea deposited in Brazilian museums and institutions is listed for the first time. New synonyms are proposed: Phibalosoma paulense Toledo Piza, 1938, Phibalosoma rochai Toledo Piza, 1938, Bacteria tuberculata Toledo Piza, 1938 and Bacteria tuberculata var. argentina Toledo Piza, 1938 are junior synonyms of Cladomorphus phyllinus (Gray, 1835). Nineteen new combinations are established.O material-tipo de Phasmatodea depositado em museus e instituições brasileiras é listado pela primeira vez. Novas sinonímias são propostas: Phibalosoma paulense Toledo Piza, 1938, Phibalosoma rochai Toledo Piza, 1938, Bacteria tuberculata Toledo Piza, 1938 e Bacteria tuberculata var. argentina Toledo Piza, 1938 são sinônimos júnior de Cladomorphus phyllinus (Gray, 1835). Dezenove novas combinações são estabelecidas.
A catalogue of all Phasmatodea type material lodged in the Staatliche Museum für Naturkunde in Stuttgart (SMNS) is provided (28 species: 10 described by BRUNNER V. WAT- TENWYL (1907), 2 by HENNEMANN, CONLE & BRÜCKNER (1996a, 1996b), 1 by HENNEMANN, GEHLER & CONLE (1995a), 1 by KEVAN (1955) and 14 by REDTENBACHER (1906, 1908)). The taxa are listed alphabetically by species for each author along with the number of specimens, sex, locality and additional information if necessary. A few of the specimens, marked as types, are very doubtful to represent type material; their status is clarified. Some background infor- mation on the collection of the SMNS, important collectors and the type material is provided. The type material of four taxa described by BRUNNER V. WATTENWYL (1907) and four taxa described by REDTENBACHER (1908) from the SMNS could not be traced; it is listed separate- ly. Lectotypes are designated for Diardia battak Redtenbacher, 1908 and Ocnophila ornatissi- ma Brunner v. Wattenwyl, 1907. Bacteria imitans Redtenbacher, 1908 is found to be a junior synonym of Phanocloidea muricata (Burmeister, 1838). Keywords: Phasmatodea, type material, Staatliches Museum für Naturkunde in Stuttgart, SMNS, lectotypes, new synonym.
Asymmetry is a common phenom among invertebrates. A consequence of this phenom is the rise of chiral forms (or enantiomorphs), which are structures that are mirror images of each other. The asymmetry can be divided into two: directional asymmetry, when only one of the enantiomorphs exists in nature, and antisymmetry, when both chiral forms exist in nature in equal frequencies, a much rarer phenom than the prior. In this paper, I describe the unknown male and redescribe the female of Isagoras paxillus (Westwood) and a new species of Isagoras Stål from the Brazilian Atlantic Forest, Isagoras sobrali sp. nov. (based on males from Rio Grande do Norte: Natal). Furthermore, I also describe the first case of antisymmetry in Phasmatodea in these two species and discuss the relevance of this finding within a morphological, behavioral, and systematic context.
The internal male genitalia have been poorly investigated in Phasmatodea, remaining virtually unexplored in phylogenetic studies. Here we describe and illustrate the main phallic elements in several Neotropical stick insects, with emphasis on Paraphasma (Pseudophasmatidae), and present a phylogenetic analysis of this genus. The analysis included ten terminals in the ingroup and 18 in the outgroup, and was based on 32 characters of the phallic organ and 48 of external morphology. In order to compare these datasets in terms of phylogenetic signal and level of homoplasy, the consistency and retention indices of the cladogram were calculated separately for each of them, and partial analyses were also conducted using each dataset alone. The phylogenetic reconstruction revealed Paraphasma as polyphyletic and led us to propose a new, monotypic genus, Ecuadoriphasma gen. nov., three new combinations (Ecuadoriphasma cognatum, Paraphasma trianguliferum and Tithonophasma cancellatum) and place Oestrophora as a synonym of Paraphasma. Additionally, Olcyphides hopii and Paraphasma dentatum are synonymized with Paraphasma laterale. Both external and phallic characters were determinant for the topology obtained, and the latter were less homoplastic in the phylogenetic tree. Our results highlight the usefulness of phallic morphology for inferring phylogenetic relationships in Phasmatodea, especially among closely related genera and species.
Stick insects have received little attention in Brazil, with many taxa lacking taxonomic, biological and morphological information on the literature. This represents a gap of knowledge for inclusive groups in Phasmatodea and for the Neotropical diversity as a whole, including members of poorly known Pygirhynchini (Heteronemiidae). Canuleius similis Redtenbacher belongs to that lineage and is redescribed based on 123 individuals recently collected or raised in captivity. Bacteria ornata Brunner von Wattenwyl is found to be a junior synonym of C. similis. Lectotypes and paralectotypes are designated for Canuleius inermis Redtenbacher, of which part of the syntype series is assigned to C. similis. The male, female, nymphs and eggs were analyzed, illustrated and described. External macromorphology of mouthparts, cerci, tarsi, antennae and internal morphology of genitalia of both sexes, in special the male genitalia were accounted for. The findings on male and female genitalia are discussed considering available information for Phasmatodea, including so far infrequently referenced works on Chilean species, hoping to shed more light in the understanding of genital structures in these insects. The male genitalia have common characters between Heteronemiidae and Pseudophasmatidae, indicating that the former may be a member of Occidophasmata. Female and male internal genitalia vary interespecifically and appears to be conservative in the same population. Traditional taxonomic characters, mostly related to camouflage, are shown to vary whereas detailed morphology is emphasized to be more conservative and encouraged to be included in future analyses. Additional information on habitat, behavior and development are given.
http://phasmida.speciesfile.org
The Phasmida Species File (PSF) is a taxonomic database of the world's Phasmida (stick and leaf insects, known as walking sticks and walking leaves in the U.S.). There is full synonymic and taxonomic information for 3,350 valid species and 5,300 taxonomic names, 37,500 citations to 3,178 references, over 7,600 specimen records and 16,800 images of 75% of valid species, with more being added to on a regular basis. Another future aim of this database is to provide high quality images of living phasmids in the wild. The PSF is annually in fed into the catalogue of life (Species 2000: Naturalis, Leiden, the Netherlands).
The present paper describes 16 new species and one new genus from French Guiana and numerous taxonomic changes are proposed to the Phasmatodea of French Guiana. Fifty-six new combinations, twenty-six new synonymies, two lectotypes and four neotypes are designated.
The taxonomy of Dinelytron Gray (Prisopodidae: Prisopodinae) is controversially discussed in the literature, as well as the delimitation between Prisopodini genera. The phylogenetic relationships between Prisopodini genera were never studied. Attempting to resolve these problems, Dinelytron was reviewed and a phylogeny was conducted to test the monophyly of the genus and better delimitate the limits of the tribe genera. Resulting, Prisopoides gen. nov., with four species was described. In Dinelytron, one species is redescribed and five new Brazilian species are described. Dinelytron unilineatus (Redtenbacher) comb. nov. was transferred from Damasippus Stål. New diagnosis for Damasippus and Prisopus are proposed, based on external characters and of the male genitalia. The phylogeny recovered two most parsimonious trees, both supporting the taxonomic changes proposed in the present study. Prisopodini and each of its genera are shown to be monophyletic. Novelties on the male genitalia are described and discussed, as well as their impact over Phasmatodea systematics.
The male genitalia have been neglected in Phasmatodea studies over the years. There are few works describing and illustrating this organ and no studies demonstrating its usage. Concerned by this matter, this paper aimed to investigate whether the male genitalia of Phasmatodea is useful for systematics. First, by describing a new species of Creoxylus Audinet-Serville (Creoxylus duckei sp. nov.) and including the description of its male genitalia, along with the differentiation from the male genitalia of the know species Creoxylus spinosus (Fabricius). By comparing these two genitalia, the first case of chirality in Phasmatodea was reported. Secondly, by describing the genitalia of two Prexaspes Stål species (Prexaspes viridipes Stål and another identified to genus level) and making the differentiation between them. Finally, by comparing all four described genitalia. Results showed that the male genitalia are useful to differentiation and identification of species/genera. It is discussed what knowledge can be drawn from these genitalia descriptions in terms of phylogeny of Xerosomatinae and Phasmatodea as a whole, as well as homologies with previous literature.
The stick insect Pseudophasma cambridgei Kirby, 1904 was described from a single male collected in the region of Santarém, state of Pará, Brazil. Based on specimens recently collected near the type locality, the species is redescribed here, including the first description of the female and egg. In addition, Pseudophasma septemtrionalis Toledo Piza, 1977 is synonymised with P. cambridgei, and morphological features of the genus Pseudophasma are discussed.
The phasmatodeans or stick and leaf insects are considered to be a mesodiverse insect order with more than 3000 species reported mainly from the tropics. The stick insect subfamily Necrosciinae comprises approximately 700 described species in more than 60 genera from the Oriental and Australian region, forming the most species-rich subfamily traditionally recognized within Phasmatodea. However, the monophyly of this taxon has never been thoroughly tested and the evolutionary relationships among its members are unknown. We analyse three nuclear (18S and 28S rDNA, histone 3) and three mitochondrial (CO II, 12S and 16S rDNA) genes to infer the phylogeny of 60 species of stick insects that represent all recognized families and major subfamilies sensu Günther and the remarkable diversity within Necrosciinae. Maximum parsimony, maximum likelihood and Bayesian techniques largely recover the same substantial clades, albeit with highly discordant relationships between them. Most members of the subfamily Necrosciinae form a clade. However, the genus Neohirasea – currently classified within Lonchodinae – is strongly supported as subordinate to Necrosciinae, whereas Baculofractum, currently classified within Necrosciinae, is strongly supported within Lonchodinae. Accordingly, we formally transfer Neohirasea and allied taxa (namely Neohiraseini) to Necrosciinae sensu nova (s.n.) and Baculofractum to Lonchodinae s.n. We also provide further evidence that Leprocaulinus, until recently recognized as Necrosciinae, belongs to Lonchodinae, and forms the sister taxon of Baculofractum. Furthermore, Lonchodinae is paraphyletic under exclusion of Eurycantha and Neopromachus. We reinstate the traditional view that Neopromachus and related taxa (Neopromachini sensu Günther) are a subgroup of Lonchodinae and transfer those taxa + the New Guinean Eurycanthinae accordingly. Morphological evidence largely corroborates our molecular-based findings and also reveals that Menexenus fruhstorferi is a member of the genus Neohirasea and is thus transferred from Menexenus (Lonchodinae) to Neohirasea, as Neohirasea fruhstorferi comb.n. (Necrosciinae s.n.). Other phylogenetic results include Areolatae and Anareolatae each supported as polyphyletic, Heteropteryginae and Lanceocercata (Bayesian analysis) are monophyletic, albeit with low support, and Necrosciinae s.n. and Lonchodinae s.n. are recovered as sister taxa (Bayesian analysis).
An attempt is made to standardize further the descriptive terminology of the phasmid egg capsule by introducing stricter definitions and standard abbreviations. In addition, the various forms of the internal micropylar plate are categorized. Eophasma Sellick is replaced by Eophasmodes nov.n. A key to 131 generic forms of these eggs is provided. Where more than one egg form is associated with a genus, a diagnosis of the subgroups is provided.