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Revision of the genera of the Areolatae, including the status of Timema and Agathemera (Insecta, Phasmatodea)

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... Most of the information on stick insects from the country is limited to original descriptions of species, mainly in the works by Gray (1835), Burmeister (1838), Westwood (1859), Bates (1865) and Kirby (1904), in the monograph by Brunner von Wattenwyl (1907) and Redtenbacher (1906Redtenbacher ( , 1908, and in various papers by Toledo Piza (1936, 1937, 1938, 1944, among others). Recently, only a few studies have addressed Brazilian stick insects, such as: three broad taxonomic revisions that included some Brazilian taxa (Zompro 2001, 2004, Hennemann et al. 2016; the redescription of Tithonophasma tithonus (Gray, 1835) (Lima et al. 2013); and the descriptions of one new species in the genus Cladomorphus Gray, 1835 (Kumagai and Fonseca 2009), and another in Agrostia Redtenbacher, 1906(Heleodoro et al. 2017. Furthermore, few Brazilian species have the egg described in spite of the importance of egg morphology and egg-laying mechanism in Phasmatodea taxonomy and phylogenetics (Sellick 1997, Clark Sellick 1998, Zompro 2004, Goldberg et al. 2015. ...
... Recently, only a few studies have addressed Brazilian stick insects, such as: three broad taxonomic revisions that included some Brazilian taxa (Zompro 2001, 2004, Hennemann et al. 2016; the redescription of Tithonophasma tithonus (Gray, 1835) (Lima et al. 2013); and the descriptions of one new species in the genus Cladomorphus Gray, 1835 (Kumagai and Fonseca 2009), and another in Agrostia Redtenbacher, 1906(Heleodoro et al. 2017. Furthermore, few Brazilian species have the egg described in spite of the importance of egg morphology and egg-laying mechanism in Phasmatodea taxonomy and phylogenetics (Sellick 1997, Clark Sellick 1998, Zompro 2004, Goldberg et al. 2015. ...
... Paraphasma currently contains ten species of which eight are recorded from Brazil (Zompro 2004, Brock et al. 2017). The de-scription of the genus was based on characters of the femora, tibiae, tegmina and mesosternum (Redtenbacher 1906). ...
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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.
... Indeed, numerous phylogenetic studies based on molecular data (e.g. Whiting et al., 2003;Buckley et al., 2009;Kômoto et al., 2011;Bradler et al., 2014Bradler et al., , 2015Goldberg et al., 2015;Büscher et al., 2018;Robertson et al., 2018;Simon et al., 2019;Song et al., 2020;Zeng et al., 2020), in addition to the morphological analysis of Bradler (2009), have been contesting many of the groups established by Günther (1953) and other taxa proposed by Bradley & Galil (1977) and Zompro (2004). As a consequence, Phasmatodea is currently in a process of general taxonomic rearrangement, aiming towards a phylogeny-based classification. ...
... Conle et al., 2008;Lima et al., 2013;Chiquetto-Machado & Albertoni, 2017;Büscher et al., 2019;Heleodoro & Rafael, 2019). It is composed of small-to medium-sized stick insects, most of which have well-developed wings, but also includes many brachypterous and apterous species (Zompro, 2004;. Representatives of the group occupy various microhabitats, from the forest canopy to the ground, and use different oviposition strategies, i.e. dropping the eggs, gluing them to the substrate or burying them in the soil (Robertson et al., 2018). ...
... Representatives of the group occupy various microhabitats, from the forest canopy to the ground, and use different oviposition strategies, i.e. dropping the eggs, gluing them to the substrate or burying them in the soil (Robertson et al., 2018). In recent phylogenetic studies, Pseudophasmatidae was repeatedly supported as a monophyletic group that was usually referred to as 'Pseudophasmatinae' (Bradler, 2009;Bradler et al., 2015;Goldberg et al., 2015;Büscher et al., 2018;Robertson et al., 2018;Simon et al., 2019;Zeng et al., 2020), either corresponding to Pseudophasmatidae sensu Zompro (2004) (i.e. comprising Pseudophasmatinae, Stratocleinae and Xerosomatinae) or including also Prisopodinae, which had already been regarded by Günther (1953) as a member of Pseudophasmatidae. ...
Article
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.
... Over recent decades, the debate dealing with phasmatodean systematics was mostly devoid of an explicit phylogenetic basis. Instead, unmethodical classifications and numerous artificial groupings at various taxonomic ranks (e.g., Bradley and Galil, 1977;Zompro, 2004;Brock and Marshall, 2011) resulted in a highly chaotic taxonomy with incorrectly appointed taxa still recognized in the current Phasmida Species File database (Brock et al., 2017). This situation has now been improved by numerous phylogenetic analyses of molecular datasets being published Buckley et al., 2009Buckley et al., , 2010Bradler et al., 2014Bradler et al., , 2015Goldberg et al., 2015;Robertson et al., 2018). ...
... The phylogenetic position of Agathemera has been an evolutionary enigma. Based on morphological evidence, Agathemera was repeatedly placed as sister group to all remaining Euphasmatodea (Bradler, 2000(Bradler, , 2009; = Neophasmatidae sensu Bradler, 2003; = Verophasmatodea sensu Zompro, 2004;Klug and Bradler, 2006;Klug, 2008;Friedemann et al., 2012). However, this assumption and consequently the monophyly of Verophasmatodea have never been supported by molecular studies, which place Agathemera as a subordinate taxon, albeit with unstable position among stick insects Buckley et al., 2009;Bradler et al., 2014Bradler et al., , 2015Goldberg et al., 2015;Büscher et al., 2018). ...
... In contrast, Bradler et al. (2015) obtained a topology Dataminae + (Heteropteryginae + Obriminae), which was also favored based on phylogenetic analysis of morphological characters by Bradler (2009) and also in our study supported by the trees inferred from the datasets NT decisive and AA SOS (Supplementary Figures S4, S5). The third possible topology Heteropteryginae + (Obriminae + Dataminae), as proposed by Zompro (2004), was favored by one previous molecular phylogeny (Goldberg et al., 2015) but was not recovered by any of our phylogenetic inferences. Additional FcLM analyses in combination with permutation tests revealed that the inferred relationship by the dataset AA SOS (Heteropteryx and Obriminae being closest relatives) is strongly biased by confounding signal. ...
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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.
... Prisopodidae is a group of stick insects (Phasmatodea) with 52 described species which are assigned to two subfamilies, the Asiatic Korinninae and the Neotropical Prisopodinae (Brock et al. 2019). Historically, the Prisopodidae was considered to belong in the Pseudophasmatidae (Günther 1953; and was erected as a different family by Zompro (2004), who proposed the current taxonomic classification. However, this arrangement is recently demonstrated to be artificial, as many authors presented evidence (mouth-parts morphology, tarsal attachment morphology, egg-shell morphology, egg-laying strategy, molecular data) of Korininae belonging under Necrosciinae (Lonchodidae) and the Prisopodinae under Pseudophasmatidae, as previous authors proposed (Bradler 2009;Goldberg et al. 2015;Büscher & Gorb 2017;Robertson et al. 2018). ...
... Prisopodinae is subdivided in two tribes, the Paraprisopodini and the Prisopodini (Zompro 2004;Brock et al. 2019). These two tribes are differentiated by the short tegmen of the paraprisopodines, which is less than half the length of the posterior wing, in comparison with the relatively long tegmen of the prisopodines, which exceeds half the length of the posterior wing (Zompro 2004). ...
... Prisopodinae is subdivided in two tribes, the Paraprisopodini and the Prisopodini (Zompro 2004;Brock et al. 2019). These two tribes are differentiated by the short tegmen of the paraprisopodines, which is less than half the length of the posterior wing, in comparison with the relatively long tegmen of the prisopodines, which exceeds half the length of the posterior wing (Zompro 2004). Robertson et al. (2018) provided the first evidence of these two tribes forming a monophyletic group, as two Melophasma Redtenbacher, 1906 (Paraprisopodini) species were recovered along two species of Prisopus Fargeau & Audinet-Serville, 1828 (Prisopodini). ...
Article
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.
... Another strongly supported Neotropical clade (PP = 0.96) comprises the species poor South American Agathemera and Heteronemiini + the species rich Pseudophasmatidae (Figure 4), often referred to as Pseudophasmatinae (Bradler et al., 2014Goldberg et al., 2015). These taxa were considered as Pseudophasmatinae by Günther (1953), but its subgroups Agathemera, Heteronemiini (= Bacunculini sensu Günther, 1953) and Prisopodini were subsequently removed from Pseudophasmatinae (or Pseudophasmatidae, Zompro, 2004). Based on morphological evidence, Agathemera was repeatedly placed as sister group to all remaining Euphasmatodea (Klug and Bradler, 2006;Bradler, 2009;Friedemann et al., 2012), but this hypothesis is in sharp contrast to all molecular studies which support Agathemera as a subordinate euphamatodean taxon (Whiting et al., 2003;Buckley et al., 2009;Bradler et al., 2014Bradler et al., , 2015Goldberg et al., 2015;Büscher et al., 2018), albeit with no consensus regarding its placement among the remaining stick insects. ...
... Based on morphological evidence, Agathemera was repeatedly placed as sister group to all remaining Euphasmatodea (Klug and Bradler, 2006;Bradler, 2009;Friedemann et al., 2012), but this hypothesis is in sharp contrast to all molecular studies which support Agathemera as a subordinate euphamatodean taxon (Whiting et al., 2003;Buckley et al., 2009;Bradler et al., 2014Bradler et al., , 2015Goldberg et al., 2015;Büscher et al., 2018), albeit with no consensus regarding its placement among the remaining stick insects. Zompro (2004) erected the subfamily Prisopodinae for Prisopus and related species (e.g., Melophasma) and the family Prisopodidae for Prisopodinae + Korinninae. Prisopus and allies are enigmatic taxa. ...
... We recovered Prisopodini nested within Pseudophasmatinae (see also Goldberg et al., 2015), but in some analyses it was supported as an early divergent neophasmatodean lineage far removed from Pseudophasmatinae. The internal phylogeny of Pseudophasmatidae does not reflect the current classification following the Phasmida Species File (Brock et al., 2017, mainly based on Zompro, 2004). The subgroup Xerosomatinae is supported (PP = 1) but not its subdivision into tribes, e.g., Xerosomatini (represented here by Acanthoclonia and Creoxylus) and Prexaspini (represented by Isagoras and Metriophasma) (see also Bradler et al., 2015). ...
Article
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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.
... Heteronemiidae (sensu Zompro 2004) is a family of "Areolatae" stick insects, with many species historically classified in "Anareolatae" groups (see Zompro 2001a,b;2004). The family currently includes about 80 species (Brock et al. 2019) and is poorly known in both biological and phylogenetic aspects. ...
... Heteronemiidae (sensu Zompro 2004) is a family of "Areolatae" stick insects, with many species historically classified in "Anareolatae" groups (see Zompro 2001a,b;2004). The family currently includes about 80 species (Brock et al. 2019) and is poorly known in both biological and phylogenetic aspects. ...
... The genus is the most diverse of the tribe and many of its species are poorly defined, the original descriptions are superficial, and often only one sex is known. According to Zompro (2004), some traits of Canuleius include: granulated or tuberculated body; third antennomere triangular in cross section; strongly elongated mesonotum; femora often bearing lobes; micropylar plate of egg with a broad and raised margin projecting beyond polar area. The only information available on the eggs of Canuleius consists of a brief comment and a schematic illustration (Zompro 2004: Figure 10.1) of an egg not identified to species level. ...
Article
We present new information on the poorly known genus of stick insects, Canuleius Stål, 1875, with redescriptions of two Brazilian species, Canuleius grandis Toledo Piza, 1936 and Canuleius sanguinolentus (Brunner von Wattenwyl, 1907) comb. nov., the latter herein transferred from Bacteria Berthold, 1827. Type material of both species was examined, as well as additional specimens recently collected and deposited at the Museu de Zoologia da Universidade de São Paulo (MZUSP). The types of two other species in the genus, Canuleius affinis Toledo Piza, 1936 and Canuleius brevipes Toledo Piza, 1936, were also examined and both species are herein synonymised under C. grandis (syn. nov.). The study includes the first descriptions of the male of C. grandis, the female of C. sanguinolentus and the eggs of both species, which are the first descriptions of eggs of Canuleius. Some notes are presented on the biology of both species, including the first observations of mating and oviposition in Heteronemiidae. Taxonomic aspects of Canuleius are discussed.
... It is important to note that although studies such as those by Zompro (2001Zompro ( , 2004Zompro ( , 2004b and Hennemann et al. (2016Hennemann et al. ( , 2018 are broad taxonomic revisions, not restricted to Brazilian taxa, they do include relevant contributions to the taxonomy of some Brazilian phasmids genera and species. For example, the latest major changes to the phasmids classification, in particular to the re-arrangement of the Areolatae, have been conducted by Zompro (2001Zompro ( , 2004Zompro ( , 2004b. ...
... It is important to note that although studies such as those by Zompro (2001Zompro ( , 2004Zompro ( , 2004b and Hennemann et al. (2016Hennemann et al. ( , 2018 are broad taxonomic revisions, not restricted to Brazilian taxa, they do include relevant contributions to the taxonomy of some Brazilian phasmids genera and species. For example, the latest major changes to the phasmids classification, in particular to the re-arrangement of the Areolatae, have been conducted by Zompro (2001Zompro ( , 2004Zompro ( , 2004b. For Brazil, Zompro (2001Zompro ( , 2004Zompro ( , 2004b proposed two new genera, Exocnophila Zompro, 2001, andOtocraniella Zompro, 2004, and described seven new species. ...
... For example, the latest major changes to the phasmids classification, in particular to the re-arrangement of the Areolatae, have been conducted by Zompro (2001Zompro ( , 2004Zompro ( , 2004b. For Brazil, Zompro (2001Zompro ( , 2004Zompro ( , 2004b proposed two new genera, Exocnophila Zompro, 2001, andOtocraniella Zompro, 2004, and described seven new species. The revision works by Hennemann et al (2016Hennemann et al ( , 2018) did much to clarify the current status of the nine species of Cladoxerus Peletier de Saint Fargeau & Serville, 1828, and the four species of Phantasca Redtenbacher, 1906, in our territory. ...
Article
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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.
... Nevertheless, the scheme suggested by Günther (1953) gained broad acceptance by subsequent authors (Bradley andGalil 1977, Kevan 1982), who proposed several substantial alterations and rank escalations, generally considered to be unwarranted (Key 1991, Klug and Bradler 2006, Bradler 2009). It did not come as a surprise that the only available phylogenetic analyses of morphological data (Tilgner 2002, Bradler 2009) did not corroborate the traditional classification proposed by Günther (1953) or successive authors (Bradley and Galil 1977, Kevan 1982, Key 1991, Zompro 2004a. ...
... Günther (1953), however, treated the Phylliinae as one subfamily among many. The idea of leaf insects representing a basal, high-ranking lineage among the Phasmatodea was revitalized by Zompro (2003Zompro ( , 2004a, but was criticized by Bradler et al. (2003). In stark contrast, the majority of published studies recovered the Phylliinae as a subordinate taxon within the Euphasmatodea (Tilgner 2002;Whiting et al. 2003;Terry and Whiting 2005;Bradler 2009;Buckley et al. 2009a;Kômoto et al. 2012;Bradler et al. 2014Bradler et al. , 2015Goldberg et al. 2015), albeit with constantly changing positions among stick insects. ...
... Most taxa are wingless ground-dwellers that are well camouflaged in leaf litter. The Heteropteryginae were traditionally divided into four subgroups (Günther 1953, Klante 1976, of which the Malagasy Anisacanthini were excluded from the group by Zompro (2004a). Molecular data have supported this view , Goldberg et al. 2015, but monophyly of the remaining Heteropteryginae, consisting of the Datamini, Heteropterygini, and Obrimini, is not well established either. ...
Chapter
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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.
... Previously, the genus was monotypic and is only known from the Ecuadorean Andes. It is characterized by its robust shape, trapezoidal mesonotum, and the presence of a transverse sensorial area on the prosternum (Zompro, 2004). Also, we synonymize Grylloclonia n. syn. ...
... Subgenital plate with posterior margin rounded or triangular in females; males present variable poculum that covers the vomer completely. Type species: Pachyphloea aberrans Redtenbacher, 1906 by original monotypy Comments: In his taxonomic key, Zompro (2004) differentiated Pachyphloea and the Grylloclonia-Dicranoclonia Zompro, 2004complex (synonymized under Acanthoclonia Stål, 1875by Conle et al., 2011 based on the arrangement of lateral margins of the mesonotum and the presence/absence of a sensorial area on the prosternum. The arrangement of the lateral margins of the mesonotum however is a variable character even within species, normally with females presenting relatively broader margins of mesonotum than males. ...
... Subgenital plate with posterior margin rounded or triangular in females; males present variable poculum that covers the vomer completely. Type species: Pachyphloea aberrans Redtenbacher, 1906 by original monotypy Comments: In his taxonomic key, Zompro (2004) differentiated Pachyphloea and the Grylloclonia-Dicranoclonia Zompro, 2004complex (synonymized under Acanthoclonia Stål, 1875by Conle et al., 2011 based on the arrangement of lateral margins of the mesonotum and the presence/absence of a sensorial area on the prosternum. The arrangement of the lateral margins of the mesonotum however is a variable character even within species, normally with females presenting relatively broader margins of mesonotum than males. ...
Article
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We describe a new species of Pachyphloea from the Natural National Park (NNP) Chicaque, Cundinamarca in the Colombian Andes, providing data on the structure and variability of its eggs. In addition, we synonymize Grylloclonia n. syn. under Pachyphloea, and provide a key for distinguishing males and females of the known species. Finally, we discuss the taxonomical delimitation of genera within the tribe Xerosomatini, and highlight the necessity of carrying out additional studies in this particular taxon of stick insects.
... Since the clade was originally introduced as Heteropteryginae by Kirby (1896), various taxa have been added and transferred in a rather disorderly way to and within the four traditional subgroups Anisacanthini, Datamini, Heteropterygini and Obrimini (e.g. Redtenbacher, 1906;Rehn & Rehn, 1938;Günther, 1953;Beier, 1968;Klante, 1976;Bradley & Galil, 1977;Zompro, 2004) and Heteropteryginae was eventually elevated to the rank of a family by Zompro (1996) with its aforementioned tribes considered as subfamilies afterwards (see Hennemann et al., 2016a, for a detailed summary). The Anisacanthinae comprising a group of Malagasy stick insects was later considered to be unrelated to the remaining Heteropterygidae and excluded by Zompro (2004). ...
... Redtenbacher, 1906;Rehn & Rehn, 1938;Günther, 1953;Beier, 1968;Klante, 1976;Bradley & Galil, 1977;Zompro, 2004) and Heteropteryginae was eventually elevated to the rank of a family by Zompro (1996) with its aforementioned tribes considered as subfamilies afterwards (see Hennemann et al., 2016a, for a detailed summary). The Anisacanthinae comprising a group of Malagasy stick insects was later considered to be unrelated to the remaining Heteropterygidae and excluded by Zompro (2004). This view was recently supported by molecular data, which simultaneously provided evidence for a monophyletic group combining all stick insects from Madagascar (Glaw et al., 2019;Simon et al., 2019). ...
... According to the phylogenetic studies of Heteropterygidae based on morphological data (Klante, 1976;Bradler, 2009), Dataminae are the sister group to Heteropteryginae + Obriminae. This combination is favoured by only one molecular analysis , whereas other studies hypothesize either Heteropteryginae + (Dataminae + Obriminae) (Zompro, 2004;Goldberg et al., 2015;Glaw et al., 2019) or Obriminae + (Dataminae + Heteropteryginae) (Robertson et al., 2018;Büscher et al., 2018a;Simon et al., 2019;Forni et al., 2020). The lack of a robust phylogeny for the Heteropterygidae impedes the reconstruction of its biogeographic history as well as the evolution of certain key traits such as wings or secondary ovipositors within this group. ...
Article
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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.
... Nevertheless, some morphological characteristics with high taxonomic potential are poorly explored in Phasmatodea research in general, not only for Brazilian species. Their unique and heterogeneous eggs lack descriptions for most species, including many Brazilian ones (Bedford 1978;Clark Sellick 1997;1998;Zompro 2004). The internal genitalia are even less studied. ...
... The Americas are home to several endemic lineages of stick insects (Phasmatodea), mostly occurring in the Neotropics (Zompro 2001a;Zompro 2004). Recently, phylogenomic data pointed that several New World lineages form a monophylum, the Occidophasmata (Simon et al., 2019). ...
... Among poorly investigated phasmid groups the Heteronemiidae is distributed all over America and stands out with historical taxonomic misplacing and vagueness. Several species are known only by brief descriptions of one or few specimens or by just one sex, even after extensive reviews in this century (Zompro 2001a,b;2004;Crispino et al., 2020). This phasmid group is consistently poorly sampled in phylogenetic studies, so their relationships with other lineages remain uncertain (Buckley et al., 2009;Bradler et al., 2015;Goldberg et al., 2015;Büscher et al., 2018a;Robertson et al., 2018;Glaw et al., 2019;Simon et al., 2019), but some studies indicate a close relation with Pseudophasmatidae (Robertson et al., 2018). ...
Article
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.
... The present paper is the eighth part of an on-going study of the neotropical Phasmatodea and provides a revision of the rarely known Colombian genus Decidia Stål, 1875 (Pseudophasmatidae: Pseudophasmatinae: Anisomorphini). The genus was erroneously placed in the tribe Psdeudophasmatini by Zompro (2004) but is obviously very close to the Central American Autolyca Stål, 1875, a member of the tribe Anisomorphini. Hence, Decidia is here re-transferred to the tribe Anisomorphini (rev. ...
... Decidia was previously known only from the female of its type-species, Decidia soranus (Westwood, 1859). Brachyelena hirsuta Hebard, 1933, the type-species of Brachyelena Hebard, 1933 and also described merely upon a female from central Colombia, was synonymized with D. soranus (Westwood) by Zompro (2004). Examination of material in the collections of the Academy of Natural Sciences, Philadelphia (ANSP) and the Museo Entomológico Francisco Luís Gallego of the Universidad Nacional de Colombia in Medellín, ♀ ♀ small and much shorter than anal segment; round in cross-section and constricted towards the apex; finely bristled. ...
... Zompro (2004: 140) in contrast claimed relation to Neophasma Redtenbacher, 1906, stated that "Decidia species appear like very broad Neophasma species […]" and erroneously transferred Decidia to the tribe Pseudophasmatini. The inaccuracies and resulting problems of the arrangement of the subfamily Pseudophasmatinae and contained tribes Pseudophasmatini and Anisomorphini proposed by Zompro (2004) have already been pointed out and partly corrected by Murányi (2007: 57) and Conle & Hennemann (2008: 8). Zompro (2004: 131) distinguished between Pseudophasmatini and Anisomorphini merely on the basis that the profemora were at least as long or distinctly longer (Pseudophasmatini), or equal in length to shorter than the combined length of the head, pro-and mesonotum (Anisomorphini). ...
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The genus Decidia Stål, 1875 is revised with an updated description of the genus, descriptions of all four known species, illustrations and keys are provided. The male of the type-species Decidia soranus (Westwood, 1859) is described and illustrated for the first time and a re-description of the females is given. Examination of the Colombian Peruphasma doylei (Caudell, 1906) has shown this to belong in Decidia (n. comb.); the male is re-described and the previously unknown female described and illustrated for the first time. Also Autolyca blapoides Redtenbacher, 1906 from South Ecuador has proven to belong in Decidia and is here transferred (n. comb.) with the female re-described and unknown male described and illustrated for the first time. These two species show the genus Decidia to contain not only winged but apterous species as well. A very colourful new species, Decidia magnifica n. sp. from the central Cordillera of Colombia, is described and illustrated from both sexes. The eggs of Decidia remain as yet unknown. Decidia appears to be restricted to the Andean regions of Central and Southern Colombia and Ecuador above an altitude of 2000 metres, referred to as the biogeographical province Cauca. The genus shows close relation to the Central American Autolyca Stål, 1875 and is hence re-transferred from Pseudophasmatini to the tribe Anisomorphini (rev. stat.).
... Paraphasma umbretta (Lichtenstein, 1796) is thus the valid name of this species, notwithstanding that its junior synonym Paraphasma maculatum (Gray) has been more commonly used in the literature (see e.g. Handlirsch 1930;Zompro 2004;Conle et al. 2011Conle et al. , 2020Brock et al. 2016;Chiquetto-Machado & Cancello 2021), possibly as a consequence of Redtenbacher (1906) having used it as a valid name. ...
... Kirby, 1904a: 410;Rehn, 1906: 278;Rehn, 1907: 165. Paraphasma fasciatum, Redtenbacher, 1906Giglio-Tos, 1910: 10;Passerin d'Entrèves, 1981: 55;Zompro, 2004 Remarks. The type material of Paraphasma fasciatum is likely to be lost (Brock et al. 2022). ...
... Pseudophasma xanthotaenidium Günther, 1930 was erroneously regarded as a synonym of Paraphasma marginale by Klante (1960) and was later listed as a valid species of Paraphasma by Zompro (2004), who also pointed out that it was probably a synonym of Paraphasma amabile. Comparisons between the types of P. xanthotaenidium and P. amabile confirmed this hypothesis, so this species is here synonymized under Pseudophasma amabile comb. ...
Article
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.
... Most studies based on morphology, transcriptome, and mitochondrial genome data have shown that the monophyly of Phasmatodea can be confirmed, and Timema is recognized as the sister group to all remaining phasmids [28,[30][31][32][33]. Nevertheless, data on mitochondrial genomes considered that Timema did not belong to the Euphasmatodea, but grouped with other orders (e.g., Orthoptera and Embioptera) [6,34]. This result coincided with the study about the morphology of Timema species in egg that indicated that Timema was a separate lineage [35]. Based on 16S, 12S RNA genes, and protein codon genes, Song et al. [22] constructed nine phylogenetic trees, and eight of these failed to recover the monophyly of Phasmatodea because the clade of Embioptera and Zoraptera that existed in long-branch attraction (LBA) was a sister group to Euphasmatodea, whereas only one phylogenetic tree supported the monophyly of Phasmatodea, as found in Song et al. [36]. ...
... In this study, O. guangxiensis was the sister clade to O. mouhotii belonging to Dataminae, and Heteropteryginae was the sister clade to (Dataminae + Obriminae). Meanwhile, the same conclusion was also obtained by some research that utilized morphological data [35,114] and molecular data [115]. By contrast, Bank et al., based on three nuclear data sets (18S, 28S and H3) and four mitochondrial data genes At the family level, our data supported the monophyly of Heteropterygidae but Diapheromeridae, Phasmatidae, and Lonchodidae were not recovered. ...
... In this study, O. guangxiensis was the sister clade to O. mouhotii belonging to Dataminae, and Heteropteryginae was the sister clade to (Dataminae + Obriminae). Meanwhile, the same conclusion was also obtained by some research that utilized morphological data [35,114] and molecular data [115]. By contrast, Bank et al., based on three nuclear data sets (18S, 28S and H3) and four mitochondrial data genes (COX1, COX2, 12S, and 16S), were in favor of the clade of Dataminae + (Heteropteryginae + Obriminae) [116], consistent with previous results [117][118][119]. ...
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Insects of the order Phasmatodea are mainly distributed in the tropics and subtropics and are best known for their remarkable camouflage as plants. In this study, we sequenced three complete mitochondrial genomes from three different families: Orestes guangxiensis, Peruphasma schultei, and Phryganistria guangxiensis. The lengths of the three mitochondrial genomes were 15,896 bp, 16,869 bp, and 17,005 bp, respectively, and the gene composition and structure of the three stick insects were identical to those of the most recent common ancestor of insects. The phylogenetic relationships among stick insects have been chaotic for a long time. In order to discuss the intra- and inter-ordinal relationship of Phasmatodea, we used the 13 protein-coding genes (PCGs) of 85 species for maximum likelihood (ML) and Bayesian inference (BI) analyses. Results showed that the internal topological structure of Phasmatodea had a few differences in both ML and BI trees and long-branch attraction (LBA) appeared between Embioptera and Zoraptera, which led to a non-monophyletic Phasmatodea. Consequently, after removal of the Embioptera and Zoraptera species, we re-performed ML and BI analyses with the remaining 81 species, which showed identical topology except for the position of Tectarchus ovobessus (Phasmatodea). We recovered the monophyly of Phasmatodea and the sister-group relationship between Phasmatodea and Mantophasmatodea. Our analyses also recovered the monophyly of Heteropterygidae and the paraphyly of Diapheromeridae, Phasmatidae, Lonchodidae, Lonchodinae, and Clitumninae. In this study, Peruphasma schultei (Pseudophasmatidae), Phraortes sp. YW-2014 (Lonchodidae), and species of Diapheromeridae clustered into the clade of Phasmatidae. Within Heteropterygidae, O. guangxiensis was the sister clade to O. mouhotii belonging to Dataminae, and the relationship of (Heteropteryginae + (Dataminae + Obriminae)) was recovered.
... Isagoras Stål, 1875 and Planudes Stål, 1875 belong to Prexaspini (Phasmatodea: Pseudophasmatidae: Xerosomatinae) and are closely related (Conle et al. 2011). Redtenbacher (1906) and Zompro (2004) attempted to distinguish these genera but had no success due to their morphological resemblance, especially between males. Redtenbacher (1906) suggested that the difference would be the developed posterior wing of Isagoras in contrast to the brachypterous posterior wing of Planudes. ...
... Redtenbacher (1906) suggested that the difference would be the developed posterior wing of Isagoras in contrast to the brachypterous posterior wing of Planudes. Zompro (2004) tried to improve this differentiation by delimitating this difference on wings to females. However, there are species of Isagoras in which females are apterous (i,e I. nitidus Redtenbacher, 1906) and a species of Planudes with fully developed wings (P. ...
... crenulipes Rehn, 1904). Nonetheless, Conle et al. (2011) pointed out that Zompro (2004) listed only the apterous and omitted the brachypterous species and the one with developed wings. Furthermore, Conle et al. (2011) stated that the male terminalia of both genera are similar, making the differentiation between them even harder. ...
Article
Isagoras aurocaudata sp. nov. is being described from two female specimens from the States of Minas Gerais, Brazil. Diagnostic characters for the new species are the yellowish compound eye with brown spots, the yellowish spot at basal third of tegmina and the yellowish abdomen segments 8–11. Furthermore, Isagoras aurocaudata sp. nov. is compared to species of Isagoras Stål and Planudes Stål.
... Timema), its phylogenetic position has long remained unclear. Zompro 23 proposed Phylliidae as a sister group to all other Euphasmatodea (excl. Agathemera = Verophasmatodea therein), while more recent molecular analyses recovered them as sister to the remaining Neophasmatodea (=Euphasmatodea excl. ...
... As stated above, all genera are recovered as distinct clades in both ML and BI inferences. In contrast to morphological studies 11,23,48 , where Nanophyllium as the only member of the tribe Nanophylliini 48 was hypothesised as the sister group to the remaining phylliids (Phylliini), our results reveal the genus to be a subordinate clade within Phylliidae. The proposition of Nanophyllium as a high-ranking phylliid clade was based solely on males, which bear morphologically unique characteristics 39,48 . ...
Article
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The insect order Phasmatodea is known for large slender insects masquerading as twigs or bark. In contrast to these so-called stick insects, the subordinated clade of leaf insects (Phylliidae) are dorso-ventrally flattened and therefore resemble leaves in a unique way. Here we show that the origin of extant leaf insects lies in the Australasian/Pacific region with subsequent dispersal westwards to mainland Asia and colonisation of most Southeast Asian landmasses. We further hypothesise that the clade originated in the Early Eocene after the emergence of angiosperm-dominated rainforests. The genus Phyllium to which most of thẽ 100 described species pertain is recovered as paraphyletic and its three non-nominate subgenera are recovered as distinct, monophyletic groups and are consequently elevated to genus rank. This first phylogeny covering all major phylliid groups provides the basis for future studies on their taxonomy and a framework to unveil more of their cryptic and underestimated diversity.
... Multiple expansions of the terga might have been an early development used by phasmatodeans in their cryptic strategiesThe expansions of the abdomen lateral lamellae of living stick insects (e.g. Extatosoma Gray 1833) are normally produced by the terga[33,34], but in extant leaf insects, both the abdominal terga and sterna are extended simultaneously, converging and forming the lamellate structures involved in their mimicry, a condition unique among Phasmatodea[33,34]. Wedmann ...
... Multiple expansions of the terga might have been an early development used by phasmatodeans in their cryptic strategiesThe expansions of the abdomen lateral lamellae of living stick insects (e.g. Extatosoma Gray 1833) are normally produced by the terga[33,34], but in extant leaf insects, both the abdominal terga and sterna are extended simultaneously, converging and forming the lamellate structures involved in their mimicry, a condition unique among Phasmatodea[33,34]. Wedmann ...
Article
Wingless and shorter winged stick insects are very common today, but most known extinct stick insects had fully developed wings, leading to contentious affinities among the extinct winged and extant groups. We report herein three male winged stick insects, assigned to Pterophasmatidae fam. nov., from mid-Cretaceous Myanmar (Burmese) amber. Pterophasmatidae fam. nov. are regarded as transitional taxa from extinct winged to modern wingless and shorter winged stick insects based on their similar tegmina venation with extinct Susumanioidea and some body features the same as extant Phasmatodea. However, their symmetric phallic organs comprising two consistent phallomeres are different from those of all living groups. Phylogenetic analyses suggest that the extinct winged taxa, including the new family, are the stem groups of modern stick and leaf insects, and all of them constitute the clade of Phasmatodea. New findings indicate winged and wingless stick insects' morphologies diversified significantly during or before the mid-Cretaceous.
... Morphological terms follow Bragg (2001), Zompro (2004) and Bradler (2009). The eggs of P. (Paragongylopus) cheni sp. ...
... n. were extracted from the abdomen of the holotypic female. Ootaxonomic terminology refers to Clark (1976aClark ( , b, 1979Clark ( , 1988Clark ( , 1998, Clark-Sellick (1997) and Zompro (2004). The descriptions of coloration are based on dried specimens. ...
Article
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This study provides a taxonomic review of Paragongylopus Chen & He, 1997 with descriptions of one new subgenus, four new species and one new subspecies. They are namely Paragongylopus (Paragongylopus) cheni sp. n., P. (Paragongylopus) sinensis pingbianensis subsp. n., P. (Planoparagongylopus) lii subgen. n. and sp. n., P. (Planoparagongylopus) abramovi sp. n., and P. (Planoparagongylopus) nabanheensis sp. n. The occurrence of P. (Paragongylopus) plaumanni Zompro, 2000 in China is reconfirmed. Paragongylopus is firstly recognized in Vietnam. Keys to the species of both subgenera and checklists of known species are also provided. (https://jor.pensoft.net/article/15291)
... The type specimens of new taxa were deposited in CAU, RBINS, and the Hong Kong Entomological Society collection, Hong Kong, China (HKES). Morphological terms follow Bragg (2001), Zompro (2004) and Bradler (2009). The descriptions of coloration are based on dried specimens. ...
... n. were naturally laid by an adult female. Ootaxonomic terminology refers to Clark (1976aClark ( , b, 1979Clark ( , 1988Clark ( , 1998, Clark-Sellick (1997) and Zompro (2004). Measurements are given in millimeters (mm), with min-max values if appropriate. ...
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This paper provides a taxonomic study for the phasmid genus Parasinophasma Chen & He, 2008; reports for the first time Parasinophasma from Hong Kong, China and Vietnam; describes six new species and two new subspecies: P. bresseeli sp. n., P. constanti sp. n., P. laifanae sp. n., P. liui sp. n., P. luchunense luchunense sp. n. and subsp. n., P. luchunense xingyuei subsp. n. and P. sparsigranulatum sp. n.; proposes one new combination: Parasinophasma bouvieri (Redtenbacher, 1908) comb. n. transferred from Orthonecroscia Kirby, 1904; and gives a checklist of known species and a revised key to species. (https://jor.pensoft.net/articles.php?id=15289)
... With both taxa being endemic to Madagascar and the adjacent Comoro Islands, this finding makes sense from a biogeographic point of view. Nonetheless, this result is particularly noteworthy, since Anisacanthidae and Achriopterini are considered to belong to the two different suborders of Phasmatodea, Areolatae, and Anareolatae (Günther, 1953;Zompro, 2004;Cliquennois, 2007Cliquennois, , 2008. This traditional subdivision is based on the presence (areolate) or absence (anareolate) of the area apicalis, a demarcated triangular area located ventrally on the apex of the tibiae (Günther, 1953). ...
... The presence of the area apicalis is considered to represent the plesiomorphic condition among stick insects (Bradler, 2009). The hypothesis of a close relationship between Anisacanthidae and Heteropteryginae, with Anisacanthini forming a subgroup of Heteropteryginae, is old (Günther, 1953;Klante, 1976), but was revised by Zompro (2004) who excluded Anisacanthini from Heteropteryginae. This assumption gained acceptance in subsequent studies (Cliquennois, 2007(Cliquennois, , 2008 and is also supported here. ...
Article
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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.
... Bragg (2001) revised Dajaca and provided an identification key. Zompro (2004) revised Phaeophasma as a junior synonym of Dajaca. Seow-Choen (1998, 2017, 2020 systematically worked through Dajaca based on specimens from Borneo and Sumatra. ...
... Fore femora more or less straight. Tibiae unarmed (Bragg 2001;Zompro 2004). After comparing the diagnostic features, Nanhuaphasma shows similar characters to the Dajaca, and we could not find significant morphological differences between the two and therefore consider Nanhuaphasma to be a junior synonym of Dajaca. ...
Article
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The genus Nanhuaphasma Chen, He & Li, 2002 was established as a member of the family Pseudophas-matidae Rehn, 1904 (now belonging to Aschiphasmatidae Brunner von Wattenwyl, 1893) based on the male of N. hamicercum Chen & He, 2002. We review the status of Nanhuaphasma and N. hamicercum by examining the holotype and male and female non-types which were collected in same location as the holotype. We find that Nanhuaphasma is a junior synonym of Dajaca Brunner von Wattenwyl, 1893 and N. hamicercum is a junior synonym of D. napolovi Brock, 2000. Complementing egg morphology of D. napolovi and keys to eight species of Dajaca are provided.
... The relevance of the stick insect eggs in the clarification of its taxonomy has been previously discussed (Camousseight and Bustamante 1991;Sellick 1997;Zompro 2004). Additionally, these traits are commonly used by researchers in descriptions of genera or species (Mantovani and Scali 1987;Salmon 1991;Zompro 2004), even being used in the identification of extinct species (Sellick 1994). ...
... The relevance of the stick insect eggs in the clarification of its taxonomy has been previously discussed (Camousseight and Bustamante 1991;Sellick 1997;Zompro 2004). Additionally, these traits are commonly used by researchers in descriptions of genera or species (Mantovani and Scali 1987;Salmon 1991;Zompro 2004), even being used in the identification of extinct species (Sellick 1994). In this study we described for first time the eggs from the eight species of the South American endemic genus Agathemera. ...
Article
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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 Phasmatodea, commonly known as leaf and stick insects, are terrestrial herbivorous insects with approximately 3.000 species described worldwide (Brock et al., 2018). Throughout the taxonomic history of the order, different authors proposed different taxonomic arrangements (e.g., Redtenbacher 1906;Bradley & Galil 1977;Zompro 2004). Despite the differences between these taxonomic arrangements, all these authors based their works on the external body morphology and on the egg morphology in the case of Zompro (2004). ...
... Throughout the taxonomic history of the order, different authors proposed different taxonomic arrangements (e.g., Redtenbacher 1906;Bradley & Galil 1977;Zompro 2004). Despite the differences between these taxonomic arrangements, all these authors based their works on the external body morphology and on the egg morphology in the case of Zompro (2004). Another common feature shared by these classifications is the lack of investigation of the male phallic complex (herein treated as "genitalia"; the group of external sclerites of the apex of the abdomen are going to be referred as "terminalia"). ...
Article
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.
... However, these new findings clearly differ from the latter based on these characters: stridulum and auditory tympanum absent, pronotum without lateral lobes, base of profemur square, metatibia without denticulations or serrations along dorsal margins and tarsal claws lack teeth (Gurney & Rentz, 1978). Zompro (2004Zompro ( , 2005 supposed that Timematodea should be a sister group of webspinners rather than remaining stick insets based on the structure of egg, reduced ocelli and the male right cercus of two parts in Timematodea (Zompro, 2004(Zompro, , 2005, which is not recognized yet (Bradler, 2009). The new taxa share some characters with Embioptera, for example, ocelli absent and enlarged metafemur, but these features are also present in some groups of Euphasmatodea. ...
... However, these new findings clearly differ from the latter based on these characters: stridulum and auditory tympanum absent, pronotum without lateral lobes, base of profemur square, metatibia without denticulations or serrations along dorsal margins and tarsal claws lack teeth (Gurney & Rentz, 1978). Zompro (2004Zompro ( , 2005 supposed that Timematodea should be a sister group of webspinners rather than remaining stick insets based on the structure of egg, reduced ocelli and the male right cercus of two parts in Timematodea (Zompro, 2004(Zompro, , 2005, which is not recognized yet (Bradler, 2009). The new taxa share some characters with Embioptera, for example, ocelli absent and enlarged metafemur, but these features are also present in some groups of Euphasmatodea. ...
Article
Many extant insects have developed pad structure, euplantulae or arolia on their tarsi to increase friction or enhance adhesion for better mobility. Many polyneopteran insects with euplantulae, e.g., Grylloblattodea, Mantophasmatodea and Orthoptera, have been described from the Mesozoic. However, the origin and evolution of stick insect's euplantulae are poorly understood due to rare fossil records. Here, we report the earliest fossil records of Timematodea hitherto, Tumefactipes prolongates gen. et sp. nov. and Granosicorpes lirates gen. et sp. nov., based on three specimens from the mid‐Cretaceous Burmese amber. Specimens of Tumefactipes prolongates gen. et sp. nov. have extremely specialized and expanded euplantulae on their tarsomere II. These new findings are the first known and the earliest fossil records about euplantula structure within Phasmatodea, demonstrating the diversity of euplantulae in Polyneoptera during the Mesozoic. Such tarsal pads might have increased friction and helped these mid‐Cretaceous stick insects to climb more firmly on various surfaces, such as broad leaves, wetted tree branches or ground. These specimens provide more morphological data for us to understand the relationships of Timematodea, Euphasmatodea, Orthoptera and Embioptera, suggesting that Timematodea might be monophyletic with Euphasmatodea rather than Embioptera and Phasmatodea should have a closer relationship with Orthoptera rather than Embioptera. This article is protected by copyright. All rights reserved
... The function of these "sensory" areas currently remains unknown. ZOMPRO (2004) raised Datamini to subfamily level as Dataminae, based on the position of these sensory areas, as well as the absence of a spine on the area apicalis and the absence of a beak-like ovipositor in females. ...
... & REHN (1939) erected the tribe Datamini and differentiated it from Obrimini, the only other tribe they included in Obriminae, by having in addition to the intercoxal specialized area of the prosternum, a similarly "specialized area" on the anterior portion of the mesosternum. The authors referred to what would later be known as the "sensory areas" on the prosternum and profurcasternum (ZOMPRO, 2004;BRADLER, 2009;HENNEMANN et al., 2016a). Sensory ...
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The genus Microrestes gen. nov. is erected to accommodate the new species Microrestes robustus sp. nov. from North Vietnam. Pylaemenes trapezius Ho, 2016 from southern China is transferred to Microrestes and the new combination M. trapezius (Ho, 2016) comb. nov. is proposed. A third species of Microrestes is recorded from northern Thailand based on photographs. The genus is compared with the other genera of Dataminae. Microrestes robustus sp. nov., the type species, is described and figured from both sexes and the egg. The genus is recorded from Vietnam, China and Thailand. A key and distribution map are provided for its species. A standardized nomenclature of the cephalic armature for Microrestes gen. nov. is proposed.
... The earliest work on the taxonomy of Hong Kong stick insects was provided by Brock and Seow-Choen (2000), who recorded seven genera and nine species. Later, four species including Entoria hei Ho, 2013, Necroscia shukayi (Bi, Zhang & Lau, 2001), Planispectrum hongkongense Zompro, 2004, andSinophasma mirabile Günther, 1940 were added to the list of Hong Kong stick insects (Bi et al., 2001;Zompro, 2004: Ho, 2008, 2013a. The most recent comprehensive work on the Hong Kong stick insects was provided by Ho (2013b), in which 15 genera and 18 species were recorded. ...
... The earliest work on the taxonomy of Hong Kong stick insects was provided by Brock and Seow-Choen (2000), who recorded seven genera and nine species. Later, four species including Entoria hei Ho, 2013, Necroscia shukayi (Bi, Zhang & Lau, 2001), Planispectrum hongkongense Zompro, 2004, andSinophasma mirabile Günther, 1940 were added to the list of Hong Kong stick insects (Bi et al., 2001;Zompro, 2004: Ho, 2008, 2013a. The most recent comprehensive work on the Hong Kong stick insects was provided by Ho (2013b), in which 15 genera and 18 species were recorded. ...
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Ho, G.W.C. 2017. First record of Neososibia brevispina Chen & He, 2000 from Hong Kong (Phasmida: Diapheromeridae: Necrosciinae). Hong Kong Entomological Bulletin, 9(2): 9–11. [http://hkentsoc.org/bulletin/HKEB9(2)_Oct_2017_Ho_Neososibia_brevispina.pdf]
... The genus was redescribed by ZOMPRO (2004) upon both sexes and the egg. ZOMPRO's description is based on material of O. mouhotii (Bates, 1865) from Nakhon Ratchasima, Thailand. ...
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Six new species of the genus Orestes Redtenbacher, 1906, O. bachmaensis sp. nov., O. botot sp. nov., O. diabolicus sp. nov., O. dittmari sp. nov., O. draegeri sp. nov. and O. krijnsi sp. nov. are described and figured from Vietnam. Descriptions are based on both sexes, except for O. diabolicus sp. nov., of which only the male is known. The type species O. mouhotii (Bates, 1865) is redescribed and its distribution range is extended. Dares (Dares) subcylindricus Redtenbacher, 1906 is removed from the synonymy with Orestes mouhotii (Bates,1865), transferred to Orestes and redescribed from both sexes and eggs. The new combination Orestes subcylindricus Redtenbacher, 1906 stat. rev., comb. nov. is proposed. The generic description of Orestes is slightly improved and the delimitation from Pylaemenes Stål, 1875 is precised. A plate of the type species of Pylaemenes, P. coronatus (Haan, 1842) is presented for comparison. Pylaemenes guangxiensis Bi & Li, 1994, P. japonicus Ho, 2016 and P. shirakii Ho & Brock, 2013 are transferred to Orestes and the new combinations O. guangxiensis (Bi & Li, 1994) comb. nov., O. japonicus (Ho, 2016) comb. nov. and O. shirakii (Ho & Brock, 2013) comb. nov. are proposed. The cephalic armature is discussed and figured. A key to the species of Orestes from Vietnam and Cambodia is presented with accompanying plates, as well as a distribution map and live in nature pictures.
... Cependant, rien ne permet de décider pour l'heure si les caractères partagés avec les Leiophasmatinae constituent des synapomorphies ou des symplésiomorphies. J'avais tenté [Cliquennois, 2008] d'établir des états de caractères apomorphes afin de fonder la monophylie de la famille des Anisacanthidae, ainsi que celle des trois sous-familles qui la composent, à savoir les Anisacanthinae, les Leiophasmatinae et les Xerantherinae, mais cet essai se fondait sur l'appartenance de cette famille à la superfamille des Bacilloidea sensu Zompro [2004], superfamille que les analyses moléculaires révèlent polyphylétique [Bradler et al., 2014 ;Bradler et al., 2015]. Selon Bradler et al. [2015], les Phasmes de Madagascar semblent former un groupe monophylétique (si l'on excepte Antongilia muricata Redtenbacher, 1906, au placement incertain), au sein duquel les Anisacanthidae auraient les Achriopterini pour groupe-frère. ...
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Résumé. – L’auteur décrit une nouvelle espèce de Phasme récoltée sur l’île d’Anjouan (Comores) dont elle est endémique. Les caractères qu’elle présente nécessitent l’érection d’un nouveau genre. Sa position taxinomique apparaît incertaine : sa morphologie la place cependant au sein de la famille des Anisacanthidae, le plus sûrement parmi les Leiophasmatinae. L’histoire géologique des Comores s’avère être mal connue, ce qui ne permet pas de comprendre l’histoire évolutive de cette nouvelle espèce de Phasme qui constitue un exemple sans précédent de taxon original au sein de la faune comorienne. - - - - - - - - Summary. – Adelophasma anjouanense gen. n., sp. n., an enigmatic stick insect of Anjouan island, Comoro archipelago (Phasmatodea). A new species collected on Anjouan island (Comoros), of which it is endemic, is described. The characters it presents require the erection of a new genus. Its taxonomic position appears uncertain: its morphology places it within the family of Anisacanthidae most surely in Leiophasmatinae. The geological history of the Comoros proves to be poorly known, which does not allow to understand the evolutionary history of this new species of stick insect which constitutes an unprecedented example of an original taxon within the Comorian fauna Keywords. – Phasmatodea, Anisacanthidae, Leiophasmatinae, Comores, Anjouan, Madagascar.
... The eggs were presumed to be mature according to general egg structure of Lonchodinae ( . Ootaxonomic description refers to Clark (1976aClark ( , 1976bClark ( , 1979Clark ( , 1988Clark ( , 1998, Clark-Sellick (1997) and Zompro (2004 Vertex with a pair of apically pointed horns; mesonotum distinctly granulated; antero-ventral and postero-ventral carinae of femora with two to three spines …… ..........M. asperatum comb. nov. ...
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Ho, G.W.C. 2014. Taxonomic note on the genus Megalophasma Bi, 1995 (Phasmida: Phasmatidae: Lonchodinae). Hong Kong Entomological Bulletin, 6(1): 15–18. [http://hkentsoc.org/bulletin/HKEB6(1)_Ho_Megalophasma.pdf]
... Ootaxonomic description is based on the eggs obtained from two adult females which laid their eggs on the surface of a rearing cage. Ootaxonomic terminology refers to Clark (1976aClark ( , 1976bClark ( , 1979Clark ( , 1988Clark ( , 1998, Clark-Sellick (1997) and Zompro (2004 Head: Oval, longer than wide. Sparsely covered with minute granules. ...
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Ho, G.W.C. 2014. On the discovery of male Paragongylopus sinensis Chen & He, 1997 and the first report of Paragongylopus plaumanni Zompro, 2000 from China (Phasmida: Diapheromeridae: Pachymorphinae: Gratidiini). Hong Kong Entomological Bulletin, 6(2): 12–15. [http://hkentsoc.org/bulletin/HKEB6(2)_Ho_Paragongylopus.pdf]
... Leaf similarity is further enhanced by the wing venation of the large female forewings resembling the nerves of angiosperm leaves (Klante 1976). Historically, Phylliidae have been considered to be an early lineage among Phasmatodea branching off from a basal node of the phasmatodean tree of life (Zompro 2004). However, more recent phylogenomic analyses place them as a subordinate lineage among the Euphasmatodea as members of the Old World clade Oriophasmata (Simon et al. 2019). ...
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Within the last two years, the leaf insects of the genus Phyllium of both the islands of Java and Sumatra have been reviewed extensively based on morphological observations. However, cryptic species which cannot be differentiated morphologically may be present among the various populations. Since it has frequently been demonstrated that analyses based on molecular data can bring clarity in such cases, we conducted a phylogenetic analysis based on three genes (nuclear gene 28S and mitochondrial genes COI and 16S) from the Phyllium species of these islands. The results show distinct molecular divergence for several populations and suggest the presence of two new cryptic species, morphologically inseparable from Phyllium hausleithneri Brock, 1999. From Sumatra, the population originally thought to be a range expansion for Phyllium hausleithneri, is now here described as Phyllium nisus sp. nov., with the only consistent morphological difference being the color of the eggs between the two populations (dark brown in P. hausleithneri and tan in P. nisus sp. nov.). Further, an additional population with purple coxae from Java was morphologically examined and found to have no consistent features to separate it morphologically from the other purple coxae species. This cryptic species from Java was however shown to be molecularly distinct from the other purple coxae populations from Sumatra and Pen-insular Malaysia and is here described as Phyllium gardabagusi sp. nov. In addition, Phyllium giganteum is here officially reported from Java for the first time based on both historic and modern records of male specimens.
... Besides this peculiar case, no phasmid species is regarded as introduced to the Neotropics (Baker 2015;Brock et al. 2022). More common, however, are mislabelling of specimens or the historical inclusion of species in genera from other continents due to overall resemblance Zompro 2001Zompro , 2004aZompro , 2004b. These historical associations among taxa from different regions of the world are further influenced by the high phenotypic convergences across Phasmatodea Jacobson & Bianchi, 1902(Buckley et al. 2009). ...
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Two species of stick insect with a distinctive morphology, Candovia evoneobertii (Zompro & Adis, 2001) and Echetlus fulgens Zompro, 2004, were considered to be native to Australia and introduced into Brazil. However, Heteronemia dubia (Caudell, 1904) and Heteronemia fragilis (Brunner von Wattenwyl, 1907), both described more than a hundred years ago from South America, exhibit striking similarities with the two purportedly introduced species and are found to be conspecific with C. evoneobertii. Careful analysis of the literature and specimens revealed that these species belong to the Neotropical tribe Diapheromerini (Diapheromeridae) and represent a new genus, Arumatia Ghirotto gen. nov. We therefore propose Arumatia fulgens (Zompro, 2004) gen. et comb. nov. and Arumatia dubia (Caudell, 1904) gen. et comb. nov. We further redescribe A. dubia (Caudell, 1904) gen. et comb. nov. based on several specimens and synonymize Heteronemia fragilis syn. nov. and Candovia evoneobertii syn. nov. under it. Additionally, five new Brazilian species are described: Arumatia diamante Ghirotto gen. et sp. nov. from Abaíra, Bahia; Arumatia aramatia Ghirotto gen. et sp. nov. from Porto Nacional, Tocantins; Arumatia motenata Ghirotto gen. et sp. nov. from Serra do Cipó, Minas Gerais; Arumatia crassicercata Ghirotto, Crispino & Engelking gen. et sp. nov. from Alto Paraíso de Goiás, Goiás; and Arumatia anyami Ghirotto, Crispino & Neves gen. et sp. nov. from Costa Marques, Rondônia. Species of Arumatia gen. nov. occur mostly in the Cerrado domain, and represent the first Diapheromeridae recorded in this area. Most species are known exclusively from females with only A. aramatia gen. et sp. nov. and A. motenata gen. et sp. nov. known from both sexes. Adult and egg morphology are described and illustrated in detail for all species, as well as the nymph stages for A. dubia. Biological observations are presented, including parthenogeny in A. dubia and one of the few detailed accounts of sexual behaviour in Euphasmatodea (for A. motenata gen. et sp. nov.). Finally, a species of Diapheromerini described in error from Brazil, Diapheromera armata Piza, 1973, is synonymized under the North American Megaphasma denticrus (Stål, 1875) (syn. nov.).
... -Since the original description, where Phyllium drunganum was not placed within a particular subgenus (Yang, 1995& Chen, 1999, there has been significant confusion about the taxonomic placement of this species. Some authors have placed Phyllium drunganum within Phyllium (Pulchriphyllium) based on all tibiae with small exterior lobes (Zompro, 2004;Größer, 2008) and others in the Phyllium (Phyllium) subgenus (Otte & Brock, 2005;Hennemann, et al., 2009;Chen & He, 2008). Most recently with the arrangement of species groups by Hennemann et al. (2009), Phyllium drunganum was returned to the Phyllium (Phyllium) subgenus, but unfortunately it was placed within the siccifolium species group without morphological justification. ...
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Phyllium oyae, new species from Laos.
... -Since the original description, where Phyllium drunganum was not placed within a particular subgenus (Yang, 1995& Chen, 1999, there has been significant confusion about the taxonomic placement of this species. Some authors have placed Phyllium drunganum within Phyllium (Pulchriphyllium) based on all tibiae with small exterior lobes (Zompro, 2004;Größer, 2008) and others in the Phyllium (Phyllium) subgenus (Otte & Brock, 2005;Hennemann, et al., 2009;Chen & He, 2008). Most recently with the arrangement of species groups by Hennemann et al. (2009), Phyllium drunganum was returned to the Phyllium (Phyllium) subgenus, but unfortunately it was placed within the siccifolium species group without morphological justification. ...
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Review of leaf-insects collected in Northeastern Laos has revealed a previously unnamed species, which we here describe as Phyllium (Phyllium) oyae species nova. This new species falls within the celebicum species group as described in Hennemann et al. (2009) due to the presence of well developed alae in females and males with a wide exterior profemoral lobe. Upon a review of congenerics, we find that Phyllium (Phyllium) drunganum Yang, 1995 is misplaced and transfer it from the siccifolium species group to the celebicum species group based on the presence of developed alae in females. With so many of the species in this group only known from a single sex, we differentiate Phyllium oyae n. sp. from all species. The species name Phyllium rayongii Thanasinchayakul, 2006 is determined to be a nomen nudum and therefore unavailable according to ICZN Article 16.4.1. To conclude, we present dichotomous keys to males and females known within the celebicum species group.
... Pygirhynchini is formed by four poorly delimited genera: Canuleius Stål, 1875, Ceroys Audinett-Serville, 1838, Paraceroys Gutiérrez, 2011 andPygirhynchus Audinet-Serville, 1838, of which the first three occur in Brazil (Zompro 2004, Conle et al. 2011. The taxonomy of Pygirhynchini is still needing revision (see Crispino et al. 2020;Ghirotto 2021). ...
Article
Several lineages of stick insects (Phasmatodea) are poorly studied, especially in the Neotropics. Recently, Brazilian stick insects have been subject of significant research and, among them, the Heteronemiidae, although several genera remain to be explored. Pygirhynchini is an Heteronemiidae lineage currently containing four genera, of which three occur in Brazil: Ceroys, Canuleius and Pygirhynchus. The former is subdivided in Ceroys (Ceroys) and Ceroys (Miroceroys) based on ornamentation variations absence or presence of a spine on the scapus. We describe two new species, Ceroys (Miroceroys) cancelloae sp. nov. and Ceroys (Miroceroys) indicattii sp. nov. based on both sexes and eggs. The new species inhabit submontane, ombrophilous Atlantic Forest in Southeast Brazil and can be differentiated from other Ceroys (Miroceroys) by conspicuous features of external morphology of adults and eggs. We also briefly describe and illustrate nymphal stages of C. cancelloae sp. nov. and present notes on the biology of both species, and a distribution map for the genus, increasing the scarce knowledge of the natural history of phasmids in Brazil.
... The monophyly of Phasmatodea is well supported by morphological traits [5,6] and molecular evidence [7][8][9]. Nevertheless, the assumption of monophyletic Phasmatodea has also been challenged by some authors [10,11]. ...
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Phasmatodea represents an order of hemimetabolous insects. This group includes species with extreme forms of masquerade crypsis, whereby they imitate twigs, bark, lichen, moss, and leaves. In this study, we sequenced and annotated three mitochondrial genomes (mitogenomes) from Phasmatodea. The lengths of the novel mitogenomes range from 14,162 bp to 15,879 bp. The gene content and organization correspond to those inferred for the ancestral insect. We conducted phylogenetic analyses together with the existing mitogenomes of polyneopterans and mayflies. In most cases, the Phasmatodea was non-monophyletic, with Embioptera and Zoraptera nested inside. The mitogenome sequences from Embioptera and Zoraptera suffered from high substitution rates and displayed very long branches in phylogenetic trees. The monophyletic Phasmatodea was recovered only when the analysis employed the site-heterogeneous CAT-GTR model in PhyloBayes and used the nucleotide dataset PCG_nt. The Euphasmatodea was well established by various data types and inference methods. In addition, the clade Heteropterygidae and the subfamilies Lonchodinae and Necrosciinae were strongly supported. The Australasian clade Lanceocercata was recovered across analyses. However, the Clitumninae was non-monophyletic.
Article
The mitochondrial genome (mitogenome) has been regarded as significant source of data to better understand the phylogenetic relationships within the Euphasmatodea, but no mitogenome in Aschiphasmatoidea has been sequenced to date. In this study, two mitogenomes of Orthomeria smaragdinum and Nanhuaphasma hamicercum of Aschiphasmatidae were sequenced and annotated for the first time. The same mitochondrial gene rearrangement structure was present in the two mitogenomes sequenced, showing as the translocation of tRNA-Arg and tRNA-Asn, which conformed to the tandem duplication-random loss and could be used as a possible synapomorphy for Aschiphasmatidae. The phylogenetic results based on the maximum likelihood (ML) and bayesian inference (BI) methods both showed that Aschiphasmatidae and Neophasmatodea in Euphasmatodea are sister taxa. Although the monophyly of Oriophasmata, Occidophasmata, Diapheromeridae, Phasmatidae, Lonchodidae and Bacilloidea has not been solved, the monophyly of Neophasmatodea and Phyllioidea was well supported.
Article
Stick insects (Phasmatodea) are herbivorous, mostly nocturnal insects known for their camouflage specialization , constituting a moderately diverse group with around 3400 extant described species. The overall poor and often confusing fossil record of Phasmatodea, especially regarding older taxa, has muddled the knowledge of the early evolution of the lineage and hindered reliable calibration points for ordinal-level phylogenies. The phylogenetic relationships within the order remain unresolved, but recent research sheds more light on the subject. Here we report on the oldest known fossil of Euphasmatodea, by revising the taxo-nomic identity of one fossil species (Eoproscopia reliquum Mendes, Vasconcelos & Oliveira) previously described as a stick grasshopper (family Proscopiidae). Based on the finding of a more complete specimen the taxon is redescribed and a new genus, Araripephasma, is erected to accommodate this species in Euphasmatodea. Araripephasma reliquum comb. nov. is a remarkably modern-looking stick insect species that provides a new minimum age for Euphasmatodea and sets a good and reliable calibration point for phylo-genetic and evolutionary studies. The implications of this rare finding for the evolutionary history of Phasmatodea are discussed.
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The new species Pylaemenes scrupeus sp. nov. from Ratchaburi province, Thailand is described and illustrated based on both sexes and the egg. The new species is diagnosed and differentiated from closely related species. The constitution of Pylaemenes Stål, 1875 and the problems arising with generic attribution of species, considering recent phylogenetic studies, are discussed. The genus is for the first time recorded from Thailand and pictures of living specimens and the habitat as well as a distribution map of continental Pylaemenes species are provided. Dares ziegleri Zompro & Fritzsche, 1999 is transferred to Orestes Redtenbacher, 1906 leaving the genus Dares Stål, 1875 restricted to Borneo and Palawan.
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The Oriental stick insect genus Trachythorax Redtenbacher, 1908 is diagnosed, compared to closely related taxa, and reviewed based on examination of type material, collection material and photographic records, including citizen science sourced data. Each species is discussed and two new species are described from Vietnam: T. albomaculatus sp. nov. from Kon Chu Rang National Park in Central Vietnam and T. auranticollis sp. nov. from Dong Nai Biosphere Reserve in Southern Vietnam. Trachythorax yunnanensis Gao & Liang, 2021 stat. nov. is elevated to valid species, from status of subspecies of T. maculicollis (Westwood, 1848). Trachythorax illaesa (Redtenbacher, 1908) stat. rev. comb. nov. is reinstated as a valid species from previous status of junior synonym of T. maculicollis. As a result the genus Trachythorax now contains 15 species. New distribution records are provided for several species including new country records: Thailand and Myanmar for T. gohi Brock, 1999, Sri Lanka for T. illaesa Redtenbacher, 1908, Cambodia for T. maculicollis, India for T. sparaxes (Westwood, 1859) as well as Laos, Thailand, and Vietnam for T. yunnanensis. Distribution maps are provided for T. albomaculatus sp. nov., T. auranticollis sp. nov., T. gohi, T. maculicollis and T. yunnanensis. Egg morphology and egg deposition are described, discussed and illustrated for T. maculicollis, T. albomaculatus, T. yunnanensis and T. illaesa, the three latter species for the first time. Morphological adaptations of eggs are compared to those observed in other closely related genera such as Asceles Redtenbacher, 1908, Calvisia Stål, 1875, Korinnis Günther, 1932, Loxopsis Westwood, 1859, Marmessoidea Brunner von Wattenwyl, 1893, Sipyloidea Brunner von Wattenwyl, 1893 and Tagesoidea Redtenbacher, 1908, and egg parasitism is hypothesized as a potential evolutionary driver. Egg parasitism by Hymenoptera: Chalcidoidea is documented for the first time in nature for T. maculicollis and T. illaesa; it is hypothesised that several morphological characters are counteradaptations to the egg parasitism.
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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. Here, we revisit 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 lineage. The disjunct distribution of ocelli substantiates the hypothesis on trait reacquisition and that wings were regained in Phasmatodea.
Article
Extant stick and leaf insects commonly imitate twigs or leaves, with lateral lamellae used to enhance crypsis or achieve mimicry for protection. However, the origin and early evolution of such lateral expansions among Phasmatodea are unknown, because all known Mesozoic phasmatodeans hitherto lack preserved evidence of such structures. We report here the first Mesozoic stick insect, Elasmophasma stictum gen. et sp. nov., with well-preserved, thin, lateral lamellae on the thoracic pleura, the terga of abdominal segments I-X and the ventrolateral margins of all femora. This new species, from the mid-Cretaceous amber of northern Myanmar, has a clear, stick-like body and is assigned to Euphasmatodea. The abdominal structures of E. stictum exhibit traces of multiple expansions of the terga, suggesting that such structure might have been an early development of body expansions used to improve crypsis for stick or leaf insects when they sprawled on twigs or leaves.
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A new subgenus Orthomeria (Parorthomeria) is established with Orthomeria (Parorthomeria) alexis (Westwood, 1859) as the type species. Orthomeria (Parorthomeria) turneri n.sp. is described from a male and female collected at Danum Valley, Sabah, one specimen from the Baram river region of Sarawak, and from two specimens previously recorded by the author as variations of O. alexis. The new species is closely related to alexis.
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All five known species of stick insects (Insecta: Phasmida) from the Virgin Islands were systematically studied for the first time
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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.
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The enigmatic, monotypic genus Laciphorus Redtenbacher, 1908 (type species: Laciphorus lobulatus Redtenbacher, 1908) is redescribed and its taxonomic position is explained. It belongs to a subgroup of ‘anareolate’ Phasmatodea currently referred to as Diapheromerinae: Diapheromerini: ‘Bacteria-group’ and now members of a clade termed Occidophasmata (Simon et al., 2019). Globocalynda Zompro, 2001 is shown to be the morphologically most similar and presumably most closely related genus. Two new synonymies are revealed. The previously believed lost female holotype of Laciphorus lobulatus Redtenbacher, 1908 is traced in the collection of the Naturhistorisches Museum, Vienna (NHMW) and is shown to be the same specimen as the holotype of Bostra scabrinota Redtenbacher, 1908, which consequently is a junior objective synonym (n. syn.). Both species are here shown to be synonyms of Ocnophila capitata Brunner v. Wattenwyl, 1907, because this represents the male of Redtenbacher’s two aforementioned species (n. syn.). Thus, the valid name for this species now is Laciphorus capitatus (Brunner v. Wattenwyl, 1907) n. comb. A lectotype is designated for Ocnophila capitata Brunner v. Wattenwyl, 1907. Laciphorus appears to be geographically restricted to the ‘Coastal Lomas’ of western Peru, which biogeographically lie within the Coastal Peruvian Desert, and seems to be peculiar to its host-plant Ophryosporus pubescens Small, 1814 (Asteraceae).
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Phasmatodea, commonly known as “stick or leaf insects”, are a rather small insect order with more than 3000 known species in the world and most of them live in tropics and subtropics. Phasmatodea are divided into two suborders of Timematodea and Euphasmatodea. Stick insects usually have an elongated, stick‐like body and the longest insect in the world is female Phryganistria chinensis, with a body length (excluding antennae and legs) of 370mm. Leaf insects (Phyllidae), instead, bear a flat, leaf‐like body. Phasmatodea have a hemimetabolous life cycle with three stages: eggs, nymphs and adults. Phasmatodea are renowned for their camouflage to imitate the stems, twigs, and leaves. Hagiphasmatidae are typified by large body size, not leaf‐like, long and filiform antenna, and well‐developed forewing. Susumaniidae, along with Hagiphasmatidae, formed Susumanioidea, which have been considered as stem‐Phasmatodea.
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A new species of Presbistus Kirby, 1896, Presbistus vitivorus sp. nov., is described from Cambodia based on both sexes, nymphs, and eggs. Male genitalia and vomer are described and figured. Illustrations of adults, nymphs, specimens in situ, host plants, a distribution map and records on biology and breeding in captivity are provided. The host plants of the species belong to the family Vitaceae. The genus Presbistus and the family Aschiphasmatidae are recorded from Cambodia for the first time. The species diversity and the distribution of the genus are discussed, and it is shown that the genus is restricted to Sundaland. A nomenclature for the morphology of the dissected vomer is proposed and tries to homologize the previously used terms.
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