The male postabdomen and reproductive system of Bibio marci Linnaeus, 1758 (Hexapoda: Diptera: Bibionidae)

The male postabdomen and the internal parts of the male genital system of Bibio marci (Bibionomorpha) were examined and reconstructed 3‐dimensionally. Several features differ from the presumptive dipteran groundplan. The bases of the gonopods are fused with each other and with tergite IX. The penis is not tube‐shaped and only sclerotized on the ventral side. The vasa deferentia are S‐shaped, and two pairs of accessory glands are present. In contrast to these characteristics, the arrangement of the internal parts is probably close to the ancestral condition. With its specific shape, the penis is well suited for the transfer of a spermatophore. The dorsal sclerite of the copulatory organ probably represents the medially fused parameres. A cladistic analysis of 27 characters of the postabdomen yielded two most parsimonious trees, with the strict consensus as follows: Nannochoristidae (outgroup) + (Culicidae [Culicomorpha] + ((Nymphomyiidae + (Tipulidae + Trichoceridae)) + (Tabanidae [Brachycera] + (Bibionidae, Anisopodidae, Axymyiidae [Bibionomorpha])))). Potential synapomorphies of Bibionomorpha (including Axymyiidae) and Brachycera are the fusion of sternum IX with the gonocoxites, the fusion of the parameres forming the dorsal sclerite and the presence of an entire series of postabdominal muscles (M4, M20, M23, M26, M27, M31, M35 and M37). The results of the analysis are preliminary as it is based on a single‐character system with a limited taxon sampling. However, the main result – a clade Bibionomorpha + Brachycera – is fully compatible with current hypotheses on dipteran phylogeny.     

The thoracic morphology of Archostemata and the relationships of the extant suborders of Coleoptera (Hexapoda)

Thoracic structures of Tetraphalerus bruchi are described in detail. The results were compared with features found in other representatives of Archostemata and other coleopteran suborders. Differences between thoracic structures of Tetraphalerus and members of other archostematan subgroups are discussed. External and internal characters of larval and adult representatives of 37 genera of the coleopteran suborders are outlined, coded and analysed cladistically, with four groups of Neuropterida as outgroup taxa. The results strongly suggest the branching pattern Archostemata + [Adephaga + (Myxophaga + Polyphaga)]. Coleoptera excluding Archostemata are supported with a high Bremer support. Important evolutionary changes linked with this branching event are simplifications of the thoracic skeleton resulting in reduced degrees of freedom (i.e. a restricted movability, especially at the leg bases), and a distinct simplification of the muscle system. This development culminates in Polyphaga, which are also strongly supported as a clade. Internalization of the partly reduced propleura, further muscle losses, and the fusion of the mesoventrites and metaventrites—with reversal in Scirtoidea and Derodontidae—are autapomorphies of Polyphaga. Archostemata is a small relict group in contrast to highly successful xylobiontic groups of Polyphaga. The less efficient thoracic locomotor apparatus, the lack of cryptonephric Malpighian tubules, and the rise of angiosperms with beetle groups primarily adjusted to them may have contributed to the decline of Archostemata.
© The Willi Hennig Society 2008.     

The male postabdomen and genital apparatus of †Mengea tertiaria,a strepsipteran amber fossil (Insecta)

The male genital apparatus of a fossil insect including internal soft parts is described in detail for the first time. The conditions found in an approximately 42‐My‐old specimen of ?Mengea tertiaria embedded in Baltic amber are compared to what is found in other extinct and extant strepsipterans, notably members of the basal ‘Mengenillidae’ (probably paraphyletic). The postabdomen of ?Mengea is very similar to what is present in other fossil and extant members of Strepsiptera. Only few structural features vary within the group, but it differs strongly from the apparatus in other holometabolan lineages. The slender, exposed tergite IX, the complete absence of parameres, a sperm pump formed by a strongly developed muscularis (layers of mainly circular muscle fibres) around the proximal part of the ejaculatory duct and the presence of four specific muscles are potential autapomorphies of the Strepsiptera. A presumptive strepsipteran ground plan feature found in ?Mengea is the nearly straight, simple penis, which is also present in ?Protoxenos, ?Cretostylops, Bahiaxenos, Mengenilla, Eoxenos and Congoxenos. This strongly suggests that males of ?Mengea (and other stem group strepsipterans) copulated in a very similar way as males of extant members of the group with free‐living females (e.g. Mengenilla). In contrast, the penis of stylopidian males, which copulate with females parasitizing in pterygote hosts, is hook shaped. A sister group relationship between ?Mengea and Strepsiptera s.s. (extant groups) is supported by a distinctly weaker sclerotization of the abdominal tergites, compared to the corresponding sternites. The study of other stem group strepsipterans using μ‐computer tomography should have high priority. This technique has a great potential to facilitate morphological reconstruction and phylogenetic placement of amber fossils.     

A new species of Laranda Walker 1869 (Orthoptera: Grylloidea, Phalangopsidae) from remnant patches of the Brazilian Atlantic Forest

The genus Laranda has six described species and is confined to South and Southeast of Brazil. We describe a new species and discuss the biology and distribution of the genus. The new species can be distinguished from its known congeners by the following characteristics: absence of yellow spots on pronotum and base of posterior tibiae; female copulatory papilla: sclerotization in dorsal view forming opposing acute angles, apical lobes narrow and small; male genitalia: pseudepiphallic median process short and wide; pseudepiphallic paramere with apex incurved and ectophallic fold surpassing apex of the parameres. The genus is distributed within the Atlantic Forest biome; the new species is found on tree trunks, as well as on forest leaf litter.     

Genital feelers: the putative role of parameres and aedeagal sensilla in Coleoptera Phytophaga (Insecta)

The male copulatory organ (aedeagus) of the Curculionoidea and the Chrysomeloidea is originally composed of a median lobe and a tegmen with basal struts and distal parameres. Within the Phytophaga (=Pseudotetramera), the parameres have been reduced several times. Comparison of different types of parameres, median lobes, aedeagi lacking parameres, and investigation of dissected pairs in copula revealed that (1) parameres do not provide mechanical coupling, (2) mechanical footing is provided by the endophallus, (3) median lobes of Phytophaga bear different kinds of sensilla. Mechanical and behavioural interaction between male and female copulatory organs were studied morphologically and by observation of live, copulating pairs. For the first time, copulation of a Sagrinae-species (Chrysomelidae: Sagrinae: Mecynodera coxalgica) was investigated in detail.     

The morphology and evolution of the female postabdomen of Holometabola (Insecta)

In the present article homology issues, character evolution and phylogenetic implications related to the female postabdomen of the holometabolan insects are discussed, based on an earlier analysis of a comprehensive morphological data set. Hymenoptera, the sistergroup of the remaining Holometabola, are the only group where the females have retained a fully developed primary ovipositor of the lepismatid type. There are no characters of the female abdomen supporting a clade Coleopterida + Neuropterida. The invagination of the terminal segments is an autapomorphy of Coleoptera. The ovipositor is substantially modified in Raphidioptera and distinctly reduced in Megaloptera and Neuroptera. The entire female abdomen is extremely simplified in Strepsiptera. The postabdomen is tapering posteriorly in Mecopterida and retractile in a telescopic manner (oviscapt). The paired ventral sclerites of segments VIII and IX are preserved, but valvifers and valvulae are not distinguishable. In Amphiesmenoptera sclerotizations derived from the ventral appendages VIII are fused ventromedially, forming a solid plate, and the appendages IX are reduced. The terminal segments are fused and form a terminal unit which bears the genital opening subapically. The presence of two pairs of apophyses and the related protraction of the terminal unit by muscle force are additional autapomorphies, as is the fusion of the rectum with the posterior part of the genital chamber (cloaca). Antliophora are supported by the presence of a transverse muscle between the ventral sclerites of segment VIII. Secondary egg laying tubes have evolved independently within Boreidae (absent in Caurinus) and in Tipulomorpha. The loss of two muscle associated with the genital chamber are likely autapomorphies of Diptera. The secondary loss of the telescopic retractability of the postabdomen is one of many autapomorphies of Siphonaptera.     

Comparative morphology of the male genitalia of Aphididae (Insecta,Hemiptera): part 1

The present study provides new data concerning the morphology of the male genitalia of Aphididae and unifies their nomenclature. The structure of the male genitalia of 31 species from 26 genera of Aphididae was studied with light and scanning electron microscopy. In the studied species, the genitalia of males consist of a phallus composed of the sclerotized basal part with its articulation and a membranous apical part—an aedeagus. Laterally of the phallus, there is a pair of setose parameres. The shape of the aedeagus, the shape and length of the sclerotized basal part and its articulation as well as the variability of parameres in their form and the number of setae are recognized as important systematic signs of the genitalia. These characters are considered in conjunction with the phylogenetic relationships among the studied taxa.     

Head structures of Priacma serrata Leconte (Coleptera, Archostemata) inferred from X-ray tomography

Internal and external features of the head of Priacma serrata were studied with X-ray microtomography and with histological methods. The comparison of both techniques shows that X-ray tomography is a promising new technique for the investigation of insect anatomy. The still somewhat coarse resolution of the X-ray data is compensated for by advantages like the nondestructive and artifact-free data acquisition. The head of P. serrata and other adults of Archostemata is characterized by many derived features. Muscular features of Priacma, especially muscles of the labium and pharynx, differ strongly from what is found in other groups of Coleoptera. Several character states are considered as autapomorphies of Archostemata: scale-like surface structures, constricted neck, strongly reduced tentorium, and the plate-like, enlarged prementum. The scales provide a protecting surface pattern and may have evolved with a more exposed lifestyle. The enlarged prementum forms a lid, which closes the mouth and covers the ligula when it is pulled back by contraction of the unusually strong submento-premental muscle. The presence of four cone-shaped protuberances on the dorsal side of the head is considered an autapomorphy of Cupedidae. The galea with a narrow stalk and a round and pubescent distal galeomere is another autapomorphy of this family. It has probably evolved as an adaptation to pollen-feeding. The shape of the mandible of Cupedidae is plesiomorphic compared to what is found in adults of Ommatidae. The vertical arrangement of apical teeth is an autapomorphy of the latter family. The lateral insertion of the antenna in Priacma is a groundplan feature of Cupedidae. The dorsal shift is a synapomorphy of all other cupedid genera. A cladistic analysis of characters of the head and additional data resulted in the following branching pattern: ((Crowsoniella + (Omma + Tetraphalerus)) + (Micromalthus + (Priacma + (Paracupes + (Cupes + Tenomerga + Prolixocupes + Rhipsideigma + Distocupes + (Adinolepis + Ascioplaga)))))).     

Comparative morphology of the tibial flexor and extensor tendons in insects

Abstract. The relative size, orientation, and degrees of sclerotization of the tibial flexor and extensor tendons are compared in nineteen orders of insects. The sclerotized, independently movable tibial flexor sclerite, known previously only from Alticinae and Carabidae (Coleoptera), is found in some other Coleoptera, Megaloptera, Neuroptera, Hymenoptera and Heteroptera. The Heteroptera also have another small sclerite at the base of the tibial extensor tendon. The tibial flexor sclerite is presumed to provide additional strength and leverage to the flexion of the tibia in certain insect groups; it may also provide protection for the ventral side of the femoro-tibial joint of the leg.     

Morphology of the abdominal skeleton and muscles of the mosquito,Culiseta inornata (Williston) (Diptera: Culicidae)

The morphology of the abdominal skeleton and muscles of the adult mosquito is incompletely known. The objectives of this study were to investigate these features in a common species, Culiseta inornata (Williston). Preserved specimens were stained lightly with methylene blue and studied with a dissecting microscope at 70 × and lower. The sclerites of the pregenital segements are best developed in segment II. The base of segment VIII in the male is narrow and semicircular in shape. This modification aids in rotation of the terminalia. Two new names are introduced for parts of the terminalia. Apodeme of sternum 9 is proposed for atrial plate of the female. Gonocoxital apodeme is a new term for a structure in the male. Both of these structures serve for attachement of muscles. Terms preferred for parts of the male terminalia are: (1) gonocoxite and gonostylus for the clasping organ; (2) paramere for the sclerotized plates on each side of and joined to the aedeagus; (3) sternum 10 for paraprocts. Sternum 10 is used because the occurrence of true paraprocts in the Nematocera is questionable. Thirty-four muscles are illustrated, and the origin and insertion of each is described. Eighteen of the muscles are newly described for the mosquito. The rotational muscles of the male terminalia were identified. The results are presented in 21 text figures.     

The unique locomotor apparatus of whirligig beetles of the tribe Orectochilini (Gyrinidae,Coleoptera)

Whirligig beetles, which are known for their rapid gliding on the water surface, have evolved a unique locomotor apparatus. External and internal thoracic structures of Orectochilus villosus (Orectochilini) are described in detail and documented with microcomputed tomography, computer‐based 3D reconstructions, and scanning electronic microscopy (SEM). The results are compared with conditions found in other genera of Gyrinidae and other groups of Coleoptera. The focus is on structures linked with locomotion, especially on the unusual flight apparatus, which differs strongly from that of other beetles. As in the other Orectochilini, the prothorax of Orectochilus displays characters typical for Gyrinidae, with triangular procoxae and forelegs transformed into elongated, sexually dimorphic grasping devices. The musculature of this segment is similar to the pattern found in other Coleoptera. Similar to all other extant Gyrinidae, the mesothorax is characterized by an extensive and flat mesoventrite, suitable for gliding on the water surface. As in Heterogyrinae and the other Gyrininae, the pterothoracic legs are transformed into paddle‐like structures, enabling the beetles to move with high speed on the surface film. The musculature of the mesothorax is reduced compared to other Coleoptera, but similar to what is found in the other Gyrininae. The metathoracic skeleton and musculature are simplified in Orectochilini compared to other Gyrininae and other groups of Coleoptera. In O. villosus, only 10 metathoracic muscles are preserved. 36 are present in an archostematan beetle, a condition probably close to the coleopteran ground plan. The metathoracic dorsal longitudinal bundles are absent in Gyrininae, muscles that play a role as indirect flight muscles in most other neopteran insects. The rest of the posteromotoric flight apparatus is distinctly modified, with a limited number of skeletomuscular elements taking over more functions. The large muscle M84 (IIIdvm7) M. noto–trochanteralis, for instance, functions as dominant wing levator, but is also responsible for the powerful and rapid backstroke of the hind legs. The presence of this muscle is a synapomorphy of Heterogyrinae and Gyrininae. The narrow metafurca in the latter group is likely linked to its large size. The elytra likely contribute to the control of the flight of the beetle, whereas they shield and inhibit the flight apparatus during swimming.     

Phylogenetic Relationships of the Suborders of Coleoptera (Insecta)

One hundred seven external and internal characters of larval and adult representatives of 28 genera of the coleopteran suborders were analyzed cladistically. Four groups of Neuropterida were introduced as outgroup. The analysis yielded 18 trees with a minimum of 194 steps (CI 0.691). All trees support the monophyly of all four suborders and a branching pattern (Archostemata + (Adephaga + (Myxophaga + Polyphaga))). The presence of elytra with meso- and metathoracic locking devices, the specific hind-wing folding, the close connection of exposed sclerites, the absence of the mera, the absence of eight thoracic muscles, the reduced abdominal sternite I, and the invagination of terminal segments are autapomorphies of Coleoptera. The monophyly of Coleoptera excl. Archostemata is supported by further transformations of the thoracic sclerites such as absence of the mesothoracic discriminal line and katepisternal joint, by an internalized or absent metathoracic trochantin, by the presence of a bending zone in the hind-wing, and by eight further muscle losses. Fusion of tibia and tarsus and presence of a single claw are larval synapomorphies of Myxophaga and Polyphaga. Adults are characterized by fusion of protrochantin and propleura and by the rigid connection of the meso- and metathoracic ventrites. The eucinetoid lineage of Polyphaga is characterized by the secondary absence of the bending zone of the alae. This results in a distinctly simplified wing folding mechanism. The monophyly of Cucujiformia (+ Bostrichoidea) is supported by the presence of cryptonephric Malpighian tubules. Transformations of fore-and hind-wings, reinforcement and simplification of the thoracic exoskeleton, and an efficient use of a distinctly reduced set of thoracic muscles play an important role in the early evolution of Coleoptera. Many different larval character transformations take place in the earlier Mesozoic within the suborders.     

A new species of the genus <Emphasis Type="Italic">Trixagus</Emphasis> Kugelann, 1794 (Coleoptera,Throscidae) from Iran

Trixagus baluchicus sp. n. from southeastern Iran is described. The new species is similar to T. duvalii (Bonvouloir, 1859), but differs from the latter in the narrower body more strongly tapered posteriorly, pale rustybrown coloration, less transverse pronotum, sparser punctation, distinct punctures on the elytral interstriae, somewhat narrower antennal club, wider parameres evenly convex at sides, and more elongate phallobase enlarged only in the basal third.     

Chlorinated tyrosine derivatives in insect cuticle

A method for quantitative measurement of 3-monochlorotyrosine and 3,5-dichlorotyrosine in insect cuticles is described, and it is used for determination of their distribution in various cuticular regions in nymphs and adults of the desert locust, Schistocerca gregaria. The two chlorinated tyrosine derivatives were present in all analyzed regions in mature adult locusts, the highest concentrations were found in the sclerotized cuticle of femur and tibia, but significant amounts were also present in the unsclerotized arthrodial membranes. Small amounts of the two amino acids were obtained from pharate, not-yet sclerotized cuticle of adult femur and tibia, the amounts increased rapidly during the first 24 h after ecdysis and more slowly during the next two weeks. Control analyses using stable isotope dilution mass spectrometry have confirmed that the chlorinated tyrosines are not artifacts formed during sample hydrolysis. Mono- and dichlorotyrosine are also present in cuticular samples from other insect species, such as the beetle, Tenebrio molitor, the moth Hyalophora cecropia, the cockroach Blaberus craniifer, and the bug Rhodnius prolixus, but not in the sclerotized puparial cuticle of the blowfly, Calliphora vicina, or in sclerotized ootheca from the cockroach, Periplaneta americana. Cuticular sclerotization and formation of chlorotyrosines occur simultaneously in locust legs; sclerotized cuticles tend to have a higher content of chlorotyrosines than unsclerotized cuticles, but it is concluded that the chlorotyrosines are not just a by-product from the sclerotization process.     

The sperm pumps of Strepsiptera and Antliophora (Hexapoda)

Male genital structures of representatives of Strepsiptera, Siphonaptera and Diptera are described in detail, with special emphasis on sperm pumps. The parts involved in the apparatus are evaluated with respect to their homology. Functional interpretations are presented based on the morphological observations. The phylogenetic significance of characters related to the male genital apparatus is discussed. The sperm pumps differ strongly in Strepsiptera and Antliophora (s.s.) and are not homologous. The strepsipteran type, which lacks any sclerotized parts, has evolved independently. Autapomophies of the male genital apparatus are the compact testes, the large balloon‐shaped vesicula seminalis, the strongly developed musculature of the proximal ductus ejaculatorius, the strongly simplified copulatory organ, the unusual muscles of segments VIII and IX, and the complete absence of accessory glands. The median fusion and almost globular shape of the vesicula are potential autapomorphies of Corioxenidae. The absence of the furrow separating the testes from the vesicula seminalis is a derived condition found in Xenos and Myrmecolax. A sperm pump is absent in Boreus (Mecoptera) and Culicomorpha and the functionally relevant parts and their arrangement differ strongly in Siphonaptera, Pistillifera and Diptera (excl. Culicomorpha). The presence of a functional and homologous pumping apparatus does not belong to the groundplan of Antliophora, which implies that this alleged autapomorphy of the clade is invalid. A sperm pump belongs to the groundplan of Diptera and was secondarily reduced in Culicomorpha, many representatives of Bibionomorpha, and in Diopsidae. It was very likely primarily absent in Mecoptera. However, the precise reconstruction of the groundplan depends on the position of Nannochoristidae within Mecoptera and on the possible affinities of Siphonaptera and Boreidae. Sperm pumps should be considered as a functional term and not be used as a character for phylogenetic reconstruction, unless specific similarities are included in the character definition.     

Two new species of Cryptocephalum n. gen. (Monogenoidea: Dactylogyridae) from the cephalic lateral line of Percichthys trucha (Perciformes: Percichthyidae) in Patagonia, Argentina

Two new species of Monogenoidea were found parasitizing the cephalic lateral line canals of Percichthys trucha (Valenciennes) (Perciformes: Percichthyidae). These species are described as members of a newly proposed genus of Dactylogyridae. Cryptocephalum n. gen. is characterized by the site of infection and the combination of the several features: ventral and dorsal anchor/bar complexes, anchors with strongly elongated shaft and recurved point, shaft and point of dorsal anchors protruding laterally from haptor, hooks with 2 subunits and with pair 5 smaller than the others; gonads overlapping; coiled male copulatory organ with counterclockwise rings, accessory piece formed by 2 distinct parts, and a tubular, sclerotized ventral vagina. C ryptocephalum petreum n. sp. is characterized by having both anchor pairs protruding laterally from haptor, male copulatory organ with a coil of 2-1/2 rings, accessory piece tweezers-shaped, and sclerotized vaginal vestibule. Cryptocephalum spiralis n. sp. has ventral anchors protruding ventrally and dorsal ones protruding laterally, male copulatory organ with a coil of 1-1/2 rings, the antero-dorsal part of the accessory piece saddle-shaped, vaginal vestibule not present, and coiled vagina. This is the first record of Dactylogyridae species parasitizing the cephalic lateral line of fishes.     

中国蒙新区土蝽科一新种及四新纪录：半翅目：土蝽科

本文记录了我国蒙新区土蝽科１新种：新疆光土蝽Ｓｅｈｉｒｕｓ ｘｉｎｊｉａｎｇｅｎｓｉｓ ｓｐ．ｎｏｖ。及４个中国新纪录种：黄圆土蝽Ｂｙｓｉｎｕｓ ｐｅｎｉｃｉｌｌａｔｕｓ Ｅ．Ｗａｇｎｅｒ,小圆土蝽Ｂ．ｍｉｎｏｒ Ｅ．Ｗａｇｎｅｒ,光头伊土蝽Ａｅｔｈｕｓ ｌａｅｖｉｃｅｐｓ Ｋｅｒｚｈｅｒ及斜光土蝽Ｓ．ｐａｒｅｎｓ Ｍｕｌｓａｎｔ ｅｔ Ｒｅｙ。     

A new species of Kritskyia (Dactylogyridae,Ancyrocephalinae) parasite of urinary bladder of Prochilodus lineatus (Prochilodontidae,Characiformes) from the Floodplain of the High Paraná river,Brazil

A new species of Kritskyia inhabits the urinary bladder of the "curimba", Prochilodus lineatus in the floodplain of the high Paraná river. The new species resembles others members of Kritskyia in the following features: haptor lacking anchors and with 14 hooks marginal, posterior male copulatory organ non articulated with the accessory piece and vagina a sclerotized tube. However, it differs from the known species mainly by the shape of the copulatory complex. This is the third endoparasitic monogenean species reported from freshwater Neotropical fish.