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Phylogenetic scheme of the Phasmatodea according to Whiting et al. (2003); Terry and Whiting (2005); Bradler (2009). Sperm characters have been mapped onto the cladogram in order to reconstruct the possible pattern of evolution of the sperm structure in the two major extant lineages of the order, Timema and Euphasmatodea. Squares represent apomorphic (black) and plesiomorphic (white) character states
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The ultrastructure of spermatogenesis and spermatozoa was studied in Timema poppensis Vickery & Sandoval, 1999, a putative basal taxon of Phasmatodea. The apical portion of testis follicles consists of spermatogonial cells with polymorphic nuclei. Primary spermatocytes display very short primary cilia originating from the peripheral centrosomes. Ea...
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... spermatozoa, we identified flat membranous cisterns close to the mitochondrial derivatives and axoneme, and between them were positioned the accessory bodies. These membranous structures seem to correspond to remnants of the cisterns (membranous sacs) observed between the mitochondrial derivatives and the flagellar axoneme during spermiogenesis, which have been described for the mature spermatozoa of other insect orders, such as Mantophasmatodea (Dallai et al., 2003), Embioptera (Dallai et al., 2007), and Phasmatodea (Gottardo et al., 2012). ...
Spermatozoa may occur as physical aggregates of two or more cells, which is an array that frequently occurs in Adephaga. We investigated the spermatozoa of Gyretes sp. using light, scanning, and transmission electron microscopy techniques. The spermatozoa of this whirligig beetle were approximately 850 μm in length and mainly exhibit a three-layered acrosome, flagellum with tiny amorphous accessory bodies, a paraflagellar body, membranous cisterns close the mitochondrial derivatives and axoneme. The spermatozoa occurred in two conformations depending on the organ along the male reproductive tract where they are found. Sperm bundles featuring up to 512 cells bond for the sperm heads formed at the end of the spermatogenesis and leave the testes in this conformation; however, before the vasa deferentia, the spermatozoa were in disarray but became
arranged upon attachment to the spermatostyle in the vasa deferentia. The spermatostyle has only been identified in two Adephaga groups, the aquatic Gyrinidae and the terrestrial Carabidae, whereas the other aquatic “Hydradephaga” species have other mechanisms of sperm aggregation. The formation of a secondary sperm aggregate may be important for increasing the number of spermatozoa that are grouped and transferred to the female, increasing the gene pool of the ejaculate, or improving the selection of viable spermatozoa (ejaculate quality). The spermatozoa of Gyretes sp. share characters with other Gyrinidae species, and present a set of sperm features that distinguish them from those in other Adephaga families, such as apical three-layered acrosome, trapezoid nucleus in cross-section, the presence of a paraflagellar body, tiny and amorphous
accessory bodies near membranous cisterns, symmetrical mitochondrial derivatives, among others.
... Sperm cells are inherently connected to copulation and fecundation and are constantly under evolutionary pressure. Therefore, spermatozoa are commonly understood as holders of a species-specific set of features (Jamieson, 1987, Dallai et al., 2011Gottardo et al., 2012). ...
In this study, the sperm ultrastructure of three species of Idarnes genus was investigated using light and transmission electron microscopy. Spermatozoon morphology of the three species was similar to that of most Chalcidoidea, with helicoidally twisted nucleus and flagellum. The head region consists of an acrosome and a nucleus; the nucleus-flagellum transition region characterized by the presence of mitochondrial derivatives and the centriolar adjunct; a flagellum region, which includes the axoneme with microtubular arrangement 9 + 9 + 2 and two mitochondrial derivatives. However, the sperm of these three species exhibit features that discriminate one species from each other: (1) only one species, Idarnes sp. 2 (carme group) exhibited an extracellular sheath surrounding the anterior portion of the nucleus, which extends to the anterior region of the flagellum, but it did not present filaments; (2) the acrosome in the three species was quite different, Idarnes sp. 1 and Idarnes sp. 2 (carme group) has two compartments (acrosomal and subacrosomal vesicles) while Idarnes sp. 3 (flavicollis group) has a third compartment (perforatorium); (3) the centriolar adjunct elongated and its location among the mitochondrial derivatives is similar for the three species analyzed; (4) mitochondrial derivatives differ between the species, with triangular (Idarnes sp. 1 and sp. 3) and elongated or flat shaped (Idarnes sp. 2) appearance. These data shows that sperm structure may differ within the same genus and confirms the potential of these cells in phylogenetic and taxonomic analyses in the Chalcidoidea superfamily, as well as in Hymenoptera in general.
... The number, size and cross-sectional shape of mitochondrial derivatives are different in different insect groups (Phillips 1970, Dallai et al. 1995, Lino-Neto and Dolder 2001, Gracielle et al. 2009, Mancini et al. 2009, Gottardo et al. 2012, Dias et al. 2013, Zhang et al. 2016. For most Sternorrhyncha species, individualised sperm is stored in the seminal vesicle. ...
The sperm ultrastructure of two ricaniid species, Pochazia shantungensis (Chou & Lu) and Ricania speculum (Walker), was investigated using light and transmission electron microscopy. Both species have monoflagellate sperm, the shape and ultrastructure of the mature spermatozoon of these two species are similar in morphology, and 128 spermatozoa are organized into sperm bundles with their heads embedded in a homogenous matrix forming the spermatodesmata. The individual sperm is filiform and includes the head, neck and flagellum. The head is needle-like, with a bilayer acrosome and an inferior elongated nucleus which is formed of homogeneously compact and electron-dense chromatin. The neck region is indistinct and is comprised of the centriole and centriole adjunct with a homogeneous dense substance. The long flagellum has the typical 9 + 9 + 2 axoneme microtubule pattern and two symmetrical mitochondrial derivatives with an orderly array of cristae flanking both sides, and a pair of well-developed fishhook-shaped accessory bodies. Current evidence shows that ricaniid species have D-shaped mitochondrial derivatives in cross-section and a serrated electron-dense region. The phylogenetic relationship of Fulgoroidea with other superfamilies in Auchenorrhyncha is briefly discussed.
... We separately analyzed two datasets: (1) The DNA barcoding dataset consisting of 49 sequences of Achrioptera and Glawiana with Heteropteryx dilatata as outgroup, and (2) the multigene dataset comprising sequences of 71 taxa of Phasmatodea. The multigene dataset was rooted with the North American stick insect taxon Timema, which has been repeatedly demonstrated to form the sister group of all remaining stick and leaf insects (= Euphasmatodea) (Whiting et al., 2003;Klug and Bradler, 2006;Bradler, 2009;Tomita et al., 2011;Gottardo et al., 2012). Phylogenetic trees of both datasets were reconstructed with the GTR substitution model selected for all partitions using Maximum Likelihood (ML) analyses in MEGA 5.2.2 (Tamura et al., 2011) and Bayesian Inference (BI) analyses in MrBayes 3.1.247 ...
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.
... 55 Membranous cistern adhering to the outer side of the accessory body: (0) absent; (1) present. Present in Embioptera and Phasmatodea Gottardo et al. 2012). 56 Puff-like expansion associated to the accessory bodies: (0) absent; (1) present. ...
... 69 *Shape of the flagellum: (0) cylindrical; (1) flattened. A flattened flagellum was observed in Timematodea and Euphasmatodea(Baccetti 1987;Gottardo et al. 2012). 70 Flagellum with prominent sidefins: (0) absent; (1) present.Prominent sidefins are present in Scutigerella(Dallai and Afzelius 2000). ...
Spermatozoa provide an unusual character system, with a limited number of components organized in a single cell. Similar spermatozoa occur in groups widely separated in the phylogenetic tree of Metazoa. Nevertheless, the character system contains phylogenetic information. Hexapoda have acquired spermatophores along with the switch from aquatic to terrestrial habitats, and related to this, a multitude of different sperm types. The aim of this study is a formal evaluation of the phylogenetic information content of spermatozoa. For the first time, sperm characters are coded for formal phylogenetic analyses. Different approaches are used and compared. Mainly due to a high level of homoplasy, the evaluation of sperm characters alone is insufficient for a reconstruction of the phylogeny of the group. Yet, a reliable reconstruction of the evolution of insect sperm is possible when character transformations are assessed using a phylogeny based on extensive molecular data. Important changes took place in the early evolution of Hexapoda. Sperm characters support some major clades (e.g. Hexapoda, Dicondylia, Polyneoptera, Psocodea), but important steps in the evolution are not reflected by transformations of spermatozoa, notably the rise of Pterygota or Holometabola. Important innovations are the formation of mitochondrial derivatives and the acquisition of accessory microtubules. Some features are conservative, whereas others evolved rapidly (e.g. presence or absence of the acrosome vesicle). Some groups are conservative in their sperm features (e.g. Odonata, Heteroptera), whereas the evolution of spermatozoa was distinctly accelerated in others (e.g. Ephemeroptera). The rate of evolution can change drastically in closely related groups. Profound changes in the morphologically uniform Zoraptera underline that sperm evolution can follow a pattern very different from the general somatic morphology. The mode of character reconstruction preferred here will be useful for the evaluation of specialized morphological character systems and strengthen the concept of evolutionary morphology.
... The order Phasmatodea (leaf and stick insects) consists of approximately 3000 known species often characterized by an elongated body cavity and procryptic camouflage (Azevedo et al. 2013). The relatively short and stout Timematidae are considered the sister group to the Euphasmatodea (Whiting et al. 2003, Terry & Whiting 2005, Friedemann et al. 2012, Gottardo et al. 2012, Tilgner et al. 1999). Recent works suggest Phasmatodea are a monophyletic group based on morphological (Tilgner et al. 1999, Friedemann et al. 2012, Beutel et al. 2013), mtDNA (Plazzi et al. 2011, Bradler et al. 2014), nuclear genomic (Buckley et al. 2009, Bradler et al. 2014), and transcriptomic (Misof et al. 2014) datasets. ...
The sparse descriptions of the stick insect (Phasmatodea) digestive system as reported/provided in the literature are highly contradictory. This paper describes the digestive systems of several families of Phasmatodea (Timematidae, Heteropterygidae, Diapheromeridae, Pseudophasmatidae, and Phasmatidae) plus the gut microbiome of these and one other (Phylliidae) to both verify past findings and provide a general description of the Phasmatodea alimentary canal. The constrictions imposed by this anatomy on phasmid gut microbiology, its connections to recently released Phasmatodea transcriptomes, and how it differs from the anatomy of related orders in the Polyneoptera are discussed. All Phasmatodea have ridged proventriculi lined or covered with small spines. Anterior projections of the midgut, sometimes described as gastric caeca, are only found in Euphasmatodea and often obscure the proventriculus. We define the cardia as the complex of foregut and midgut tissue where the type II peritrophic matrix is produced. Appendices of the midgut are an autapomorphy for Phasmatodea, but Timema have fewer and larger appendices relative to body size. We suggest caeca-like projections and the loss of large, proventricular teeth are apomorphies of Euphasmatodea. We identify a possible facultative symbiosis in Eucalyptus-feeding species that requires further study.
... In the examined chrysomelids, accessory bodies have a pyriform shape (Baccetti and Daccordi, 1988), while in curculionoids they have a hook-like appearance surrounding the axoneme (Burrini et al., 1988; Dallai et al., 1998; Name et al., 2007). Similar diversification of accessory bodies were found in Phasmatodea (Baccetti, 1987; Gottardo et al., 2012) and within Zoraptera (see Dallai et al., 2011 Dallai et al., , 2012 Dallai et al., , 2014a,b,c). S. cordigera also exhibits an interesting feature at the centriolar level. ...
... A derived feature, dense laminae radiating in a cartwheel array between neighboring centriolar triplets, is shared with Phasmatodea and Embioptera. An apomorphy shared with Phasmatodea is the presence of 17 protofilaments in the tubular wall of the outer accessory microtubules ( Dallai et al. 2011Dallai et al. , 2012bGottardo et al. 2012). ...
Zoraptera are a cryptic and enigmatic group of insects. The species diversity is lower than in almost all other groups of Hexapoda, but may be distinctly higher than presently known. Several new species were described from different regions recently. The systematic placement was discussed controversially since the group was discovered 100 years ago. Affinities with Isoptera and Psocoptera were discussed in earlier studies. A sistergroup relationship with Acercaria (Psocodea, Thysanoptera, Hemiptera) was proposed by W. Hennig, for the first time based on a strictly phylogenetic argumentation. More recent studies consistently suggest a placement among the “lower neopteran orders” (Polyneoptera). Close affinities to Dictyoptera were proposed and alternatively a sistergroup relationship with Embioptera or with Embioptera + Phasmatodea (Eukinolabia), respectively. The precise placement is still controversial and the intraordinal relationships are largely unclear. Recent transcriptome analyses tentatively suggest a clade Zoraptera + Dermaptera as sistergroup of all other polyneopteran orders.
The oldest fossils are from Cretaceous amber. An extinct genus from this era may be the sistergroup of all the remaining zorapterans. The knowledge of the morphology, development and features related to the reproductive system greatly increased in recent years. The general body morphology is very uniform, whereas the genitalia differ strongly between species. This is likely due to different kinds of selection, i.e. sexual selection in the case of the genital organs. The mating pattern also differs profoundly within the order. A unique external sperm transfer occurs in Zorotypus impolitus. A species-level phylogeny and more investigations of the reproductive system should have high priority.
... Over this evolutionary time Timema and Euphasmatodea have acquired significant morphological differences [1]. Timema, with 21 Neartic species [9] , has been morphologically characterized in various character systems101112131415. In comparison the morphology of the diverse Euphasmatodea is less well documented, as accurate studies of key systematic characters are available for only a few taxa among the $3000 species known worldwide. ...
External morphological features of adult males are described in the stick insect Hermarchus leytensis from the Philippines, a species belonging to the little-known euphasmatodean lineage Stephanacridini. Mouthparts are characterized by some likely specialized features: (1) a dentate dorsal cutting edge on the mandibles; (2) distinct differences in size and shape between the galeae; (3) absence of an apical field of trichomes on the galeae; and (4) lacinial setae not protruding from the mesal margin, which features three bearing-like protuberances. The latter character state represents a very unusual condition, not known in other phasmatodeans or even in polyneopteran insects. A distinctive characteristic of attachment devices is that each euplantula is divided into two separated pads with a smooth microstructure, as it also occurs in some members of the clade Schizodecema. Male terminalia exhibit character states previously unknown in Stephanacridini, including (1) a symmetrical type of vomer and (2) claspers equipped with ∼70 very minute ventral teeth on each thorn pad. Potential implications for the systematic placement of H. leytensis are discussed. The results also underline the importance of microanatomical investigations as source of substantial characters for future analyses on phasmatodean systematics.
... As pointed out above, accessory microtubules with 16 protofilaments are part of the insect groundplan (Jamieson et al., 1999). The presence of 17 protofilaments represents an unusual apomorphic condition that has evolved in few other neopteran lineages (Phasmatodea: e.g. Gottardo et al., 2012; Zoraptera: e.g. Dallai et al., 2011 Dallai et al., , 2012; and Trichoptera: e.g. ...
