Morphogenèse de l'appareil génital ectodermique femelle d'Eurygaster austriaca (Schrank, 1776) (Heteroptera : Scutelleridae)

Female ectodermal genital apparatus of Eurygaster austriaca (Heteroptera : Scuttelleridae) originates from 3 primary buds, which are situated on 3 different regions and from one secondary bud. The external genitalia arise from 8–9th and 9–10th buds. The common oviduct with its fingershaped tubules, results from a median invagination of the 7–8th intersegmental area. The spermatheca is a dorso-apical invagination of the 8–9th bud. The vagina and its annexes originate from fused 8–9th and 9–10th buds and then, they have a bisegmental origin. The genital orifice, or secondary gonopore, also has the same origin.     

THE HIGHER CLASSIFICATION OF THE ORDER EMBIOPTERA: A CLADISTIC ANALYSIS

Abstract— A matrix of 41 Embiid taxa (representing the 8 formally recognized families of the Order) and 36 characters were cladistically analysed as a first attempt for understanding the higher classification of the Order Embioptera. The resulting trees were rooted with Clothodidae as the sister group of the other Embioptera. The results suggest that the current classification contains several artificial groups. With the rooting used, only Anisembiidae and Australembiidae are monophyletic. Embiidae is polyphyletic, as Australembiidae+ Notoligotomidae, Enveja (incertae sedis) and Oligotomidae+Teratembiidae appear within Embiidae, and the "embiid" Microembia appears within Notoligotomidae. Oligotomidae is paraphyletic in terms of Teratembiidae. Four of the genera included in the analysis are paraphyletic: Mesembia, Chelicerca (in terms of Dactylocerca and Pelorembia ), Aposthonia (in terms of Oligotoma ), and Metoligotoma (in terms of Australembia ). Pelorembia and Dactylocerca are synonymized with Chelicerca .     

The exoskeleton of the female genitalic region in Petrobiellus takunagae (Insecta: Archaeognatha): Insect-wide terminology,homologies, and functional interpretations

The exoskeleton of the female genitalic region (abdominal venters 7–9) in Petrobiellus takunagae (Machilidae-Petrobiellinae) is studied using light microscopy and SEM. Sclerites are distinguished from membrane by the degree of cuticular flexibility. However, the microsculpture of the cuticle is shown to be useful in characterising the heterogeneity of the cuticle and in detecting weak sclerotisations. The morphology of Petrobiellus is compared with that in Trigoniophthalmus alternatus (Machilidae-Machilinae) described previously. While venter 7 is similar, venters 8 and 9 show many differences in the presence/absence or fusion/separation of particular sclerites. This suggests female genitalic morphology to be a valuable character system for phylogenetic and taxonomic work in Archaeognatha. Comparison with other insect orders is aimed at detecting homologous structures and conditions. Important points are: (1) Petrobiellus has a sclerotised genital lobe posteriorly on venter 7, similar to Zygentoma and Dictyoptera; it bears the gonopore. (2) Petrobiellus has a posterior sclerite on venter 9 that is very similar to a sclerite of Odonata. (3) The morphology of the coxal lobes of venter 9 (gonoplacs) suggests their function as a sheath of the ovipositor. From female genitalic morphology we deduce the process of oviposition, describing an external egg transportation tract.     

The phylogeny and classification of Embioptera (Insecta)

A phylogenetic analysis of the order Embioptera is presented with a revised classification based on results of the analysis. Eighty‐two species of Embioptera are included from all families except Paedembiidae Ross and Embonychidae Navás. Monophyly of each of the eight remaining currently recognized families is tested except Andesembiidae Ross, for which only a single species was included. Nine outgroup taxa are included from Blattaria, Grylloblattaria, Mantodea, Mantophasmatodea, Orthoptera, Phasmida and Plecoptera. Ninety‐six morphological characters were analysed along with DNA sequence data from the five genes 16S rRNA, 18S rRNA, 28S rRNA, cytochrome c oxidase I and histone III. Data were analysed in combined analyses of all data using parsimony and Bayesian optimality criteria, and combined molecular data were analysed using maximum likelihood. Several major conclusions about Embioptera relationships and classification are based on interpretation of these analyses. Of eight families for which monophyly was tested, four were found to be monophyletic under each optimality criterion: Clothodidae Davis, Anisembiidae Davis, Oligotomidae Enderlein and Teratembiidae Krauss. Australembiidae Ross was not recovered as monophyletic in the likelihood analysis in which one Australembia Ross species was recovered in a position distant from other australembiids. This analysis included only molecular data and the topology was not strongly supported. Given this, and because parsimony and the Bayesian analyses recovered a strongly supported clade including all Australembiidae, we regard this family also as monophyletic. Three other families – Notoligotomidae Davis, Archembiidae Ross and Embiidae Burmeister, as historically delimited – were not found to be monophyletic under any optimality criterion. Notoligotomidae is restricted here to include only the genus Notoligotoma Davis with a new family, Ptilocerembiidae Miller and Edgerly, new family, erected to include the genus Ptilocerembia Friederichs. Archembiidae is restricted here to include only the genera Archembia Ross and Calamoclostes Enderlein. The family group name Scelembiidae Ross is resurrected from synonymy with Archembiidae (new status) to include all other genera recently placed in Archembiidae. Embiidae is not demonstrably monophyletic with species currently placed in the family resolved in three separate clades under each optimality criterion. Because taxon sampling is not extensive within this family in this analysis, no changes are made to Embiidae classification. Relationships between families delimited herein are not strongly supported under any optimality criterion with a few exceptions. Either Clothodidae Davis (parsimony) or Australembiidae Ross (Bayesian) is the sister to the remaining Embioptera taxa. The Bayesian analysis includes Australembiidae as the sister to all other Embioptera except Clothididae, suggesting that each of these taxa is a relatively plesiomorphic representatative of the order. Oligotomidae and Teratembiidae are sister groups, and Archembiidae (sensu novum), Ptilocerembiidae, Andesembiidae and Anisembiidae form a monophyletic group under each optimality criterion. Each family is discussed in reference to this analysis, diagnostic combinations and taxon compositions are provided, and a key to families of Embioptera is included.     

三种蚤生殖系统的细微结构:雌性外生殖器的发育

本文研究了缓慢细蚤 Leptopsylla segnis(Schōnherr),不等单蚤 Monopsyllus anisus(Rothschild)和猫栉首蚤指名亚种Ctenocephalides felis felis(Bouché)雌性外生殖器的结构,观察了从幼虫、前蛹、蛹至成虫各发育时期的雌性外生殖器的内部结构变化.对一直悬而未决的雌蚤中输卵管等的起源问题,进行了详细的观察和探讨.认为从晚期3龄幼虫开始出现的雌性外生殖器芽,在前蛹期成为四个部分:1.第7腹板后缘腹壁内陷形成的一对外胚层囊;2.紧接外胚层囊后并延伸至第8腹板的外胚层增厚;3.在第8腹板后部,外胚层增厚两侧的产卵器芽;4.第8腹板后缘腹壁内陷形成的受精囊芽.并认为,这三种蚤的中输卵管由一对外胚层囊和其后的外胚层增厚前端的一小部分内陷形成,阴道由外胚层增厚的大部分和第8、9腹板腹壁内陷形成,受精囊由受精囊芽内陷形成.     

Comparative study of the morphology of the female seminal receptacles of Ilia nucleus and Persephona mediterranea (Decapoda,Brachyura, Leucosiidae)

Because of the poor knowledge of the morphology of the female reproductive organs of most brachyuran crabs, this study investigated two Atlantic representatives of the family Leucosiidae, Ilia nucleus (Linnaeus, 1758) and Persephona mediterranea (Herbst, 1794), using histological methods and magnetic resonance imaging (MRI). While the vagina conforms to the concave type, the arrangement of the two chambers of the seminal receptacle differs strongly from that of other eubrachyuran sperm storage organs. Both chambers are oriented laterally within the crab's body. This is in contrast to the dorso-ventral orientation described in most other known brachyuran crabs. The lateral chamber is covered by cuticle, whereas the medial chamber is covered by a holocrine glandular epithelium. The oviduct connection is located ventrally, posterior to the vagina. The oviduct orifice is characterized by a transition from the epithelium lining the oviduct to the seminal receptacle's holocrine glandular epithelium. Moreover, muscle fibres are attached to the oviduct orifice and to the sternal cuticle. This musculature can be interpreted as an important feature in the fertilization and egg-laying process by supporting and controlling the inflow of eggs into the seminal receptacle lumen. The results of this study are compared to the morphology of the seminal receptacle of another leucosiid crab, Ebalia tumefacta (Montagu, 1808), and to those of other known eubrachyuran crabs.     

Morphology of the female genital ducts of the blue land crab Cardisoma guanhumi (Crustacea: Brachyura: Gecarcinidae)

This is a histological and histochemical analysis of the terminal portion of the female reproductive system and genital ducts of the blue land crab (Cardisoma guanhumi). Animals were collected in the Jaguaribe estuary (Ceará, Brazil) and dissected. Genital duct fragments were fixed and submitted to different staining techniques. The female reproductive system consists of a pair of ovaries and a pair of genital ducts. In the mid‐posterior portion of each lobe, the ovaries communicate with the genital ducts, which are subdivided into oviduct, spermatheca, vagina, and gonopore. Histologically, the spermatheca of C. guanhumi is composed of columnar secretory epithelium and is divided into a dorsal zone and a ventral zone, the latter covered internally by a cuticle layer. Both zones are enveloped by a thin layer of loose connective tissue. Histological cross sections revealed the vagina to be concave, a pattern considered phylogenetically more advanced than the simple, tubular form. Our findings suggest fertilization is internal, favoring sperm from the most recent copulation.     

Internal reproductive organs of the female humpbacked fly,Megaselia scalaris Loew (Diptera : Phoridae)

The female reproductive system of the humpbacked fly Megaselia scalaris Loew (Diptera : Phoridae) was examined in whole mount preparations and serial sections. The system includes 2 ovaries, paired lateral oviducts, a common oviduct, and a genital chamber, opening externally through a gonopore, anteriad and ventrad to the anus. The ducts of the 2 accessory glands open independently into the dorsal region of the genital chamber. The terminal duct of a 2-armed spermatheca joins the right posterior and ventral wall of the genital chamber, immediately inside the gonopore. Passing dorally, the spermathecal duct lies immediately ventral to the duct of the right accessory gland. A short distance posteriad, it divides into two branches, each supplying an arm of the spermatheca. The genital chamber extends both anteriorly and posteriorly from its junction with the common oviduct, creating anterior and posterior compartments. In the right lateral wall of the genital chamber, a distinctive loop-shaped thickening (plate) resembles a darkened thread when it is observed through the integument. Features likely to have taxonomic utility include the posterior and ventral location of the terminal portion of the spermathecal duct; and the asymmetrically arranged, loop-shaped plate.     

The female postabdomen and internal genitalia of the basal moth genus Agathiphaga (Insecta: Lepidoptera: Agathiphagidae): morphology and phylogenetic implications

The female postabdomen of Agathiphaga vitiensis terminates in a telescope‐type extensible oviscapt with an apial ‘oviscapt probe’ composed of fused segments behind VIII. Exceptionally within the Lepidoptera two pairs of long ‘anterior apophyses’ arise from segment VIII, from the dorsum and venter. Agathiphaga has the most elaborate postabdominal musculature recorded from female Lepidoptera, comprising 24 muscle sets of which nine may be family autapomorphies. Apophysis musculature does not permit unambiguous homologizing of the single anterior apophysis in Lepidoptera–Glossata with either the dorsal or ventral pair in Agathiphaga, but is compatible with an interpretation of the glossatan anterior apophyses as a composite formation. Nine muscle sets shared with rhyacophilid caddisflies are ascribed to the amphiesmenopteran ground plan. The spermathecal duct represents an intermediate stage between the simple type present in Micropterigidae and the ‘two‐compartment type’ characteristic of almost all other Lepidoptera. The spermatheca has no lagena. The bursa copulatrix is small and simple. Accessory glands are very large, simple sacs. There are 40 + ovarioles per ovary. A terminal cloaca is extremely short. The numerous ovarioles potentially support a sister‐group relationship between Agathiphagidae and all other Lepidoptera, whereas the spermathecal duct histology supports the alternative conservative placement of the family as sister group of all nonmicropterigid Lepidoptera. © 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 159 , 905–920.     

Mating behavior of the parasitic honeybee mite Tropilaelaps clareae

The mating behavior of adult Tropilaelaps clareae males and females was observed in glass test tubes. The male jumped on the dorsum of the female, stretched legs I forward, and hooked their distal ends on the frontal margin of the female dorsum. He then slipped sideways to the female venter. In the venter-to-venter position, the male clasped the female between her legs I and II, with his legs I. The male then moved backwards until his gnathosoma reached about 1/2 way along the epigynial plate of the female. The male vibrated and probably pushed the spermatophore out of his body. Next he moved laterally and placed the gnathosoma between coxae III and IV where the gonopore is located. Probably the sperm was introduced into the gonopore by means of the spermatodactyl. Subsequently the male moved to one side and stroked the anterior part of the epigynial plate and the gonopore area with his leg II. He repeated these contacts on average 282 times (variation 100 up to 522). The male then moved on the other side of the female venter and repeated the whole procedure. Finally, the male withdrew himself backwards from the female. The total mating process lasted between 3 and 42 min, with an average of 23 min. Multiple matings of both, males and females were observed.     

A newly discovered sperm transport system in the female of Lygaeidae bugs

In the female reproductive system of the relatively large hemipteran, the western conifer seed bug Leptoglossus occidentalis (Heidemann), a cuticle‐lined tube extends medially along the surface of the vagina from the proximal end of the spermathecal complex anteriorly to the base of the common oviduct. This medial tube houses the proximal end of the spermathecal duct, thereby enabling the transport of material from the spermatheca at the distal end of the spermathecal complex, past the vagina (or bursa copulatrix) and directly to the common oviduct. The proximal portion of the spermathecal complex also contains an insemination duct that is separate from the spermathecal duct. The insemination duct allows the male intromittent organ to extend from the vagina to the spermatheca without navigating through the spermathecal duct. The reproductive systems of two previously studied Hemiptera, the milkweed bug Oncopeltus fasciatus (Dallas) and the box elder bug Leptocoris trivittatus (Say), possess a similar cuticle‐lined medial tube housing the spermathecal duct. This new information provides a clearer understanding of sperm transport in the female reproductive system of Lygaeidae bugs, and helps to clarify the path of the male organ during copulation, as well as the movement of sperm during egg laying.     

Morphological analysis of the female reproductive system of Stenorhynchus seticornis (Brachyura: Inachoididae) and comparisons with other Majoidea

We examined the female reproductive system of the yellowline arrow crab Stenorhynchus seticornis by means of histological and histochemical techniques. Mature specimens obtained in the field were kept in the laboratory for mating experiments. After 24 h, newly mated females were dissected, and their reproductive trait routinely processed for embedding in historesin. The specimens examined each possessed a pair of kidney‐shaped seminal receptacles (SR), and these we classified as ventral type, based on the location of the oviduct opening. The mesodermal dorsal region (DR) of SR consisted of a stratified epithelium with scaly cells, while the ectodermal ventral region (VR) was composed of a simple epithelium covered by a cuticle. The oviduct opened at the transition region (TR) between DR and VR and had no velum. The simple epithelium of TR had more folds on the face of the oviduct opening. The vagina exhibited the same features as the TR epithelium and was contiguous to VR, anchored by muscles. In the lumen, from one to three strata of sperm packets were observed, the dorsal one containing free sperm, and the most ventral stratum, spermatophores. An acidophilic glycoprotein layer enclosed the strata. Spermatophores in the ventral stratum were enclosed in a voluminous secretion, composed by acid polysaccharides most likely from the last male mated. The ventral‐type receptacle, stratified sperm packets, and polyandry, usually observed in females of Majoidea, suggest the occurrence of sperm competition in S. seticornis, favoring the sperm of the last male mated, as its sperm mass is located near the opening of the female oviduct.     

The female postabdomen and genitalia of the basal moth family Heterobathmiidae (Insecta: Lepidoptera): Structure and phylogenetic significance

Female Heterobathmia have the segments behind VIII forming a compact ‘terminal unit’ with a large saddle-shaped dorsal plate and a membranous ventroposterior surface bearing the separate gonopore and anus. While females of most of the nine known species are overall similar, Heterobathmia valvifer is unique amongst lepidopterans in possessing paired ventral appendages (‘ovipositor valves’) arising from the intersegmental groove following segment VIII; evidence from musculature contradicts an interpretation of these appendages structures as ‘true’ ovipositor valves. The ventroposterior wall of the terminal unit in H. valvifer bears paired sclerites, possible homologues of the ‘ventral rods’ in basal Lepidoptera-Glossata. In Heterobathmia megadecella sclerites on paired longitudinal elevations in comparable positions probably are/include homologues of these sclerites. Their similarity with paired sclerotizations in the corresponding region of hydrobiosid caddisflies is noted. A prominent frame-like sclerotization in the genital chamber, located in front of the spermathecal duct origin, is present only in H. megadecella.Putative heterobathmiid autapomorphies include an enlarged ‘subgenital plate’ on venter VIII, absence of apophyses on segment VIII, shortened apophyses on the terminal unit, multilobed accessory glands (but their ‘type 1’ secretory epithelium is plesiomorphic at this level), a conspicuous papilla in the chamber cuticle bearing the opening of the ductus bursae on its apex, and inwards-pointing spines in the ductus bursae. A variably developed thickening of the anterior genital chamber intima is another putative family autapomorphy, while an extreme thickening of the posterior intima seen in Heterobathmia pseuderiocrania is not of general occurrence in heterobathmiids. A sistergroup relationship between Heterobathmiidae and Glossata is supported by their fully developed ‘2-compartment section’ of the spermathecal duct and losses of some likely lepidopteran groundplan muscles.     

Myo-inositol in the reproductive tract of the female rat

• 1.1. myo-Inositol concentrations in oviduct, ovary and uterus were many-fold those of blood serum during all four stages of the estrous cycle of the female rat.
• 2.2. Inositol concentration was higher in oviduct than in ovary or uterus and was lower in uterine fluid than in uterus.
• 3.3. Estrus uteri had higher inositol concentrations than uteri in other phases of the cycle.
• 4.4. In order to measure dynamic aspects of the distribution of inositol. the distribution of radioactivity among organs of the reproductive tract of mature female rats was measured 45 min after i.p, injection of [2-³H]myo-inositol.
• 5.5. These organs concentrated inositol from the blood, and the tissue radioactivity (expressed as dpm/mg wet wt of tissue) increased in the sequence: vagina < cervix < uterus < ovary < oviduct.
• 6.6. The uterus and ovary concentrated myo-inositol more strongly during proestrus than during metestrus. diestrus or estrus.
• 7.7. The contents of proestrus follicles were more highly radioactive than was the ovary itself, whereas proestrus uterine fluid was less radioactive than the uterine tissue.

     

The second introduction of a non-native Embioptera to Korea via ornamental plants: Evaluating DNA barcoding to prepare for a global inflow of unfamiliar taxa

This study reports a webspinner, Aposthonia ceylonica, found in imported ornamental Tillandsia plants in greenhouse gardens in Paju and Seoul. This is the second record of a non-native Embioptera in Korea. The insects were found in the gaps between the basal leaves of the plants, and lived as a group in galleries of woven silk produced from the forelegs. The morphological characters and silk structures were investigated using scanning electron microscopy. The mitochondrial COI gene was sequenced for molecular identification, and to evaluate the validity of DNA barcoding for Embioptera. These insects can only survive and breed in greenhouse gardens because they are subtropical to tropical. However, they might ultimately settle after introduction in the Korean Peninsula if tropicalization continues.     

The sperm structure of Embioptera (Insecta) and phylogenetic considerations

The sperm structure of two species of Embioptera, Embia savignyi Westwood 1837 and Aposthonia japonica (Okajima 1926), was studied. Spermatozoa of both species exhibit a monolayered acrosome and a layer of material surrounding the sperm cells for most of their length. The presence of a 9+9+2 axoneme provided with accessory microtubules with 16 protofilaments, two accessory bodies and two crystallized mitochondrial derivatives are characters shared with other polyneopteran taxa. The supposed close relationship between Embioptera and Phasmatodea is not supported by characters of the sperm ultrastructure.     

Anatomy and ultrastructure of the female reproductive system of Pleioplana atomata (Platyhelminthes: Polycladida)

The ultrastructure of the female reproductive system of the polyclad flatworm Pleioplana atomata is described. Numerous ovaries are scattered throughout the entire body but are mainly concentrated on the dorsal side. Within an ovary, a germinative zone with oogonia and prefolicular cells is located in the dorsal part of the ovary. The remaining part of the gonad is filled with previtellogenic and early vitellogenic oocytes enwrapped by follicular cells. During previtellogenesis, oocytes produce numerous eggshell globules, which are distributed into the cortical area of the cell in later stages. Eventually, these globules release their contents into the space between the eggshell cover and oolemma. Similar types of globules are also found in others flatworms, and may represent useful phylogenetic characters. Entolecital, vitellogenic oocytes pass to paired uteri, where vitellogenesis is completed. The remainder of the female reproductive system consists of paired thin uterine ducts that join a vagina. The distal part of the long, curved vagina forms a large Lang's vesicle, while the proximal part is connected to a female atrium leading to a female gonopore. We hypothesize that Lang's vesicle functions in the digestion of excess sperm received. Two kinds of different shell (cement) glands that release their secretion into the vagina are identified. Both are unicellular glands and each gland cell connects to the lumen of the vagina via an individual canal. Similar glands in other acotylean polyclads have been implicated in the formation of eggshell covers. J. Morphol. 2009. © 2008 Wiley‐Liss, Inc.     

The effect of oestradiolum benzoicum and progesterone on AChE activity in the nerves of the female reproductive system of immature pigs

The present paper describes the effect of exogenous oestradiol and progesterone on the AChE activity in nerves of the ovary, oviduct, uterus, and vagina, and on morphological features of the AChE-positive nerves of these organs in sexually immature pigs. Studies were carried out on 30 sexually immature pigs, 37 to 43 kg in weight, at the age of 115 to 125 d, of the BWP (Big White Polish) race. The animals were divided into 4 groups. Group I (n = 6) constituted the control. Animals in group II (n = 8) received oestradiolum benzoicum. Those in group III (n = 8) were given progesterone, and in group IV (n = 8) oestradiol and progesterone. Oestradiol injections resulted in a 1.2-fold increase of AChE activity in the ovary, 1.3-fold in the oviduct, 1.7-fold in the uterine horn, and a 1.2-fold decrease of this activity in the cervix, while no differences were noted in the vagina. Progesterone injections did not affect AChE activity in the ovary, while a 1.1-fold increase was observed in the oviduct, 1.2-fold in the uterine horn and 1.1-fold in the cervix compared to the control. A 1.1-fold decrease of this activity was observed in the vagina. Joint injections of oestradiol and progesterone resulted in about a 1.4-fold increase of AChE activity in the ovary, 1.3-fold in the oviduct, 1.4-fold in the uterine horn, 1.6-fold in the vagina, while no changes were observed in the cervix. The results suggest the effect of the administered hormones upon AChE activity in the organs under study, mainly in the oviduct, uterine horn, and vagina. They also point to the possibility of a synergic effect of oestradiol and progesterone as regards to an increase of the activity.     

Ultrastructure and functional morphology of the female reproductive organs inProtodrilus (Polychaeta,Annelida)

The morphology and function of the female reproductive organs in 6Protodrilus species are investigated by light- and transmission electron microscopy. Possible ways in which spermatozoa may enter the female coelom after leaving the spermatophore are discussed for species with and without special female reception organs. Only femaleP. rubropharyngeus andP. flavocapitatus have “dorsal organs” for spermatophore reception. The structure and function of these organs are described, as well as those of the oviduct found in 3 of the species investigated. The possible phylogenetic origin of gonoducts and different modes of oviposition within the genus are discussed. Finally, the high taxonomic significance of female traits such as dorsal organs, oviducts, cocoon glands and lateral ciliary rows in this genus is stressed.