External macro- and micromorphology of the male of the stick insect Hermarchus leytensis (Insecta: Phasmatodea) with phylogenetic considerations

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.     

Giant stick insects reveal unique ontogenetic changes in biological attachment devices

A strong modification of tarsal and pretarsal attachment pads during the postembryonic development is described for the first time. In the exceptionally large thorny devil stick insect Eurycantha calcarata a functional arolium is only present in the immature instars, enabling them to climb on smooth surfaces, especially leaves. Nymphs are also characterized by greyish and hairy euplantulae on tarsomeres 1–4. The gradual modifications of the arolium and the euplantula of tarsomere 5 in the nymphal development are probably mainly related to increased weight. The distinct switch in the life style between the leaf-dwelling nymphal stages and the ground-dwelling adults results in the final abrupt change of the adhesive devices, resulting in a far-reaching reduction of the arolium, the presence of a fully-developed, elongated euplantula on tarsomere 5, and white and smooth euplantulae on tarsomeres 1–4. The developmental remodelling of attachment pads also reflects a phylogenetic pattern. The attachment devices of the earlier instars are similar to those found in the basalmost lineage of extant stick insects, Timema, which is characterized by a very large pan-shaped arolium and a hairy surface of the tarsal and pretarsal attachment pads.     

Regulation of cuticular hydrocarbon profile maturation by Drosophila tanning hormone,bursicon, and its interaction with desaturase activity

Shortly after emergence the exoskeleton (cuticle) of adult insects is rapidly expanded, hardened (sclerotized), and pigmented (melanized). In parallel with this process, the oenocytes, which are large polyploid cells located below the abdominal epidermis, secrete onto the cuticle a cocktail of cuticular hydrocarbons (CHs) and waxes. These improve the waterproofing of the cuticle, and also provide important chemosensory and pheromonal cues linked with gender, age, and species differentiation. The hardening and pigmentation of the new cuticle are controlled by the neurohormone, bursicon, and its receptor, encoded by the DLGR2 receptor, rickets (rk); by contrast, little is known about the timecourse of changes in CH profile and about the role of bursicon in this process. Here we show in Drosophila that rk function is also required for the normal maturation of the fly's CH profile, with flies mutant for rk function showing dramatically elevated levels of CHs. Interestingly, this effect is mostly abrogated by mutations in the Δ9 desaturase encoded by the desaturase1 gene, which introduces a first double bond into elongated fatty-acid chains, suggesting that desaturase1 acts downstream of rk. In addition, flies mutant for rk showed changes in the absolute and relative levels of specific 7-monoenes (in males) and 7,11-dienes (in females). The fact that these differences in CH amounts were obtained using extractions of very different durations suggests that the particular CH profile of flies mutant for rk is not simply due to their unsclerotized cuticle but that bursicon may be involved in the process of CH biosynthesis itself.     

Revision of the rhinoceros beetle genus Oryctophileurus Kolbe with description of a new species,the male of O. varicosus Prell,and notes on biogeography (Scarabaeoidea,Dynastinae, Phileurini)

The genus Oryctophileurus is reviewed and its validity is supported by a combination of the following apomorphic characters: a single cephalic horn with lateral carina, pronotal cavity with ocellate punctures and two teeth or tubercles close behind the anterior pronotal margin. The male of Oryctophileurus varicosus Prell, 1934, is described for the first time. A new species, Oryctophileurus guerrai Perger & Grossi sp. n., from subhumid Tucuman-Bolivian forest in the Southern Bolivian Andes is described. The new species is distinguished from its closest relative, O. armicollis Prell, 1911, by a narrower distance between the inner teeth of the dorsal pronotal protuberances and a reduced area of weakly developed ocellate punctures above the posterolateral pronotal margin. The occurrence of Oryctophileurus species in areas of endemism along the eastern slope of the tropical Andes suggests that these populations represent biogeographic “relicts”, and the discovery of Oryctophileurus guerrai sp. n. in the southern Bolivian Andes suggests that this area is underrated with respect to insect diversity and endemism.     

New Afrotropical species belonging to genera never recorded before for the Afrotropical fauna (Diptera: Tachinidae)

Five remarkable new Afrotropical species belonging to four Old World genera never recorded before for the Afrotropical region (Neophryxe, Calliethilla, Metadrinomyia, Pseudalsomyia) are described and compared with congeners. Existing keys to tropical African tachinid genera are implemented. A brief discussion on the apomorphic support to each genus is provided.     

A new species of <Emphasis Type="Italic">Eurycletodes</Emphasis> Sars, 1909 (Copepoda: Harpacticoida: Argestidae) from the southern hemisphere including remarks on the phylogeny of the genus and its subgenera

This paper deals with taxonomy and phylogenetics of the genus Eurycletodes Sars, 1909 (Copepoda: Harpacticoida: Argestidae). Samples, collected from the southeast Atlantic on board RV “Meteor” during the cruises DIVA-1 (M48/1) and DIVA-2 (M63/2), contain specimens of Eurycletodes. Eurycletodes is characterized as a monophylum by A1 segments III + IV fused, basal seta of md palp lost, exp of md palp reduced to 1 seta or completely lost. Similarly, the subgenera Eurycletodes (Eurycletodes) and Eurycletodes (Oligocletodes) are characterized as monophyletic by the loss of the inner seta on P1 exp2 (apomorphic to E. (E.)) and the absence of the inner seta on P5 endopodal lobe (apomorphic to E. (O.)). Eurycletodes profundus is renamed as E. (O.) profundus. Eurycletodes ephippiger is the only species of the genus without subgeneric designation. Eurycletodes (O.) diva sp. nov. is described. The new species differs from described species of the genus by a larger body size, P5 endopodal lobe only slightly protruding, last segment of A1 with 2 outer setae, furcal rami elongated between setae VII and IV. The occurrence of 2 specimens of Eurycletodes (O.) diva sp. nov. at 2 sites separated by the Walvis Ridge supports the hypothesis that geographic obstacles do not prevent harpacticoid copepods from spreading in the deep sea.     

The spermatodesm of Cloeon dipterum (L.): fine structure and sperm movement

Sperm bundles of the ephemeropteran Cloeon dipterum (L.) were found in the seminal vesicle. They are apically protected by cyst cells and sperm anterior regions are embedded in a secretion produced by these cells. Cyst cells further degenerate making sperm cells free. The sperm bundles are able to perform a forward progression only when sperm are stick together with their tail posterior tips. On the contrary, when sperm have their tail ends free, they beat but are unable to progress. The functional significance of the sperm bundles is discussed.     

Structure and evolution of stridulatory mechanisms in New Zealand wetas (Orthoptera : Stenopelmatidae)

Four types of stridulatory mechanisms were found in 35 species (7 genera) of New Zealand wetas (Orthoptera : Stenopelmatidae). These include: (1) tergo-femoral; (2) tergo-tergal; (3) mandbbuao-mandibular; and (4) pleuro-coxal. These fell into 12 groups, based upon shape, density and pattern of tergal and femoral pegs, number of terga bearing pegs and presence of mandibular tusks bearing stridulatory tubercles. The plesiomorphic condition, consisting of spinose peg patches on 3 or more abdominal tergites rubbed by bands of pegs on the femur, is found in Hemiandrus spp. Apomorphically-derived tergal files have arisen separately in 3 other genera. In Hemideina, (2 species groups) well-formed files are very similar among species. In Deinacrida, (4 species groups), the trend is toward file reduction into one or 2 massive tergal ridges, and embellished femoral pegs. In the tusked wetas (2 gen., 2 spp.), one species has a crude file of many broken ridges. Stridulatory structures on mandibular tusks of both New Zealand species are unique, although the tusks appear to have their origin in African stenopelmatids. The plesiomorphic condition appears to have been deployed originally for defense stridulation (inter-specific communication). Additional intea-specific stridulatory communication developed in Hemideina and Deinacrida, using the apomorphic file and peg mechanisms. Here, the stridulation is associated with defense, calling, mating and disturbance behaviors.     

A new Late Jurassic species of the rare synechodontiform shark, <Emphasis Type="Italic">Welcommia</Emphasis> (Chondrichthyes,Neoselachii)

A new Jurassic species of the very rare and incompletely known synechodontiform shark, Welcommia, is described. The new species, Welcommia cappettai, is represented only by a single tooth, precluding reconstruction of its dentition in detail. Nevertheless, this specimen provides sufficient information and characteristics to establish its taxonomic status. Welcommia cappettai n. sp. occurs in the middle Oxfordian (Upper Jurassic) of south-western Germany. This is the first unambiguous record and named species of Welcommia from the Late Jurassic, substantially reducing the rather large gap in the fossil record of this synechodontiform taxon. So far, two Welcommia species from the Lower Jurassic of Belgium and the Lower Cretaceous of southern France have been described. An additional, still unnamed species seemingly occurs in the Oxfordian of southern France. The new species has plesiomorphic and apomorphic characteristics and, probably, an intermediate dental pattern that tentatively enables reconstruction of evolutionary trends in the dentition of this shark from small and compact teeth with broad, almost triangular cusps, to mesio-distally lengthened teeth with elongated mesial heels resulting in an extremely extended mesial cutting edge in addition to more delicate cusp and cusplets in advanced forms. These differences might be related to improved feeding mechanisms. It is hypothesized that Welcommia was predominantly a component of the Mediterranean faunal province. The disappearance of Welcommia in the Early Cretaceous remains ambiguous and might be related to competition by other sharks, for example hexanchiforms, or might represent a collecting bias and/or taxonomic misidentification of isolated teeth.     

A new spider species of the genus Argenna (Aranei,Dictynidae) from Western Siberia

A new dictynid species, Argenna sibirica sp. n., is described based on a male from the environs of Tobolsk. The new species has a combination of somatic criteria typical of the genera Argenna Thorell, 1870 and Altella Simon, 1874 but differs from them in the presence of numerous warts on the chelicerae, an elongated labium, and an elongated crescent-shaped palp conductor.     

FISH identification of Helicoverpa armigera and Mamestra brassicae chromosomes by BAC and fosmid probes

Since the Bombyx mori genome sequence was published, conserved synteny between B. mori and some other lepidopteran species has been revealed by either FISH (fluorescence in situ hybridization) with BAC (bacterial artificial chromosome) probes or linkage analysis. However, no species belonging to the Noctuidae, the largest lepidopteran family which includes serious polyphagous pests, has been analyzed so far with respect to genome-wide conserved synteny and gene order. For that purpose, we selected the noctuid species Helicoverpa armigera and Mamestra brassicae, both with n = 31 chromosomes. Gene-defined fosmid clones from M. brassicae and BAC clones from a closely related species of H. armigera, Heliothis virescens, were used for a FISH analysis on pachytene chromosomes. We recognized all H. armigera chromosomes from specific cross-hybridization signals of 146 BAC probes. With 100 fosmid clones we identified and characterized all 31 bivalents of M. brassicae. Synteny and gene order were well conserved between the two noctuid species. The comparison with the model species B. mori (n = 28) showed the same phenomenon for 25 of the 28 chromosomes. Three chromosomes (#11, #23 and #24) had two counterparts each in H. armigera and M. brassicae. Since n = 31 is the modal chromosome number in Lepidoptera, the noctuid chromosomes probably represent an ancestral genome organization of Lepidoptera. This is the first identification of a full karyotype in Lepidoptera by means of BAC cross-hybridization between species. The technique shows the potential to expand the range of analyzed species efficiently.     

The male reproductive system of Zorotypus caudelli Karny (Zoraptera): Sperm structure and spermiogenesis

Considering the overall uniformity of the morphology of Zoraptera, the structural diversity of the male genital system is remarkable. Structures related to the male reproductive system of Zorotypus caudelli differ profoundly from those of Zorotypus hubbardi. The testes are elongated rather than spherical, the seminal vesicle is apparently absent, and the deferent ducts are very long. A feature shared by these two species and other zorapterans examined is that the two accessory glands are closely adherent to each other and form a single large structure, from which the ejaculatory duct originates. This is a potential zorapteran autapomorphy. Another feature possibly present in the groundplan of the order is the strong elongation of the sperm cells. This may be connected with a reproductive strategy of males trying to avoid re-mating of females with other males after the first copulation. The extremely long and coiled spermathecal duct of Z. caudelli and other zorapteran species is possibly correlated with the sperm elongation, and both features combined may result in a sexual isolating mechanism. The short duration of mating of Zorotypus barberi and Zorotypus gurneyi suggests that the male introduces sperm into the female tract up to the opening of the spermathecal duct using their long coiled aedeagus. A thick glycocalyx around the sperm in the distal part of the deferent ducts probably protects the sperm cells during their forward progression towards the long spermathecal duct, and is removed when they reach the apical receptacle. The spermatogenesis of Z. caudelli follows a pattern commonly found in insects, but differs distinctly from that of Z. hubbardi in the number of spermatids in each sperm cyst. An unusual and possibly autapomorphic feature of Z. caudelli is a disconnection of sub-tubules A and B at the level of microtubule doublets 1 and 6 of the mature sperm cells. It is conceivable that this results in a shorter period of sperm motility. The character combination found in different zorapteran species supports the view that the sperm, a very compact functional unit, does not evolve as a unit, but like in other more complex body regions, sperm components can also be modified independently from each other. This results in different mosaic patterns of plesiomorphic and derived features in a very compact entity in different species of the very small and otherwise uniform order Zoraptera. In Z. caudelli, for instance, the bi-layered acrosome and small accessory bodies are plesiomorphic states among several others, whereas the mitochondrial derivatives and the elongate nucleus are apparently derived conditions. Other combinations likely occur in other zorapteran species. Only few but noteworthy sperm characters indicate possible phylogenetic affinities of Zoraptera. A possible synapomorphic feature, the presence of dense laminae radiating in a cartwheel array between neighbouring centriolar triplets, is shared with Phasmatodea and Embioptera. Another potential synapomorphy shared with Phasmatodea is the presence of 17 protofilaments in the tubular wall of the outer accessory microtubules.     

Spermatogenesis and Spermatozoa in <Emphasis Type="Italic">Stygocapitella subterranea</Emphasis> (Annelida,Parergodrilidae), an Enigmatic Supralittoral Polychaete

The intertidal polychaete species Stygocapitella subterranea (Parergodrilidae) is characterised by extraordinary biology and morphology, resembling those of clitellates and Hrabeiella periglandulata, a terrestrial species of Annelida. An ultrastructural study of the spermatogenesis and spermatozoa was undertaken to elucidate whether these similarities might exhibit adaptive characters typical of annelids with highly derived reproductive modes. A second goal was to find out whether there are some common apomorphic features between S. subterranea and its sister taxon, Parergodrilus heideri, instead of the differences observed on the light-microscopic level, as well as to look for potential synapomorphies to support a suggested relationship to Orbiniidae and Questidae. Spermatogenesis conforms to the general pattern typical of Annelida. Spermatids develop on large cytophores comprising at least 128 cells. The spermatozoa are extremely thread-like and, with a length of about 320 μm, are among the longest spermatozoa known for annelids. The acrosome is elongated and consists only of an acrosomal vesicle with a large subacrosomal space. A conspicuous feature is the incomplete chromatin condensation, resembling late spermatids. In the long midpiece, there is a single ring-shaped mitochondrial derivative, which develops by fusion out of a multiple array of eight mitochondria surrounding the axoneme. There is a distinct annulus between midpiece and tail. The proximal part of the tail is immobile; the axoneme is surrounded by a thick layer of cytoplasm and bears a velum-like extension. In addition to characters apomorphic for S. subterranea, these latter three features exhibit certain similarities to P. heideri that are likely to be synapomorphic. Unfortunately, a relationship of Parergodrilidae to an orbiniid/questid clade does not receive additional support from spermatozoal characters. Similarities with either Clitellata or H. periglandulata are likely to be primarily related to corresponding features of their reproductive biology rather than to phylogenetic relationship.     

Morphology of the terminalia of the stick insect Dajaca napolovi from Vietnam (Insecta: Phasmatodea)

Aschiphasmatinae is a small group of stick insects from the Oriental region whose genital morphology has been rarely described in detail. The subfamily is of particular interest, as phylogenetic studies have shown Aschiphasmatinae to be the sister group to the remaining Euphasmatodea. In this paper, the male and female terminalia are described for the first time in Dajaca napolovi Brock, a little known aschiphasmatine species from Vietnam. In the male, the transversally undivided abdominal sternum IX and gently incurved cerci with a conspicuous apical tooth represent apomorphies of Aschiphasmatinae. Thorn pads on the hind margin of abdominal tergum X consist of only a single row of 6–7 ventrally oriented teeth. The simple thorn pad structure of Aschiphasmatinae can represent an ancestral condition for Euphasmatodea. The vomer on venter X is well‐developed and features two unusually large basal apodemes and a strongly developed apical spine showing a specialized streaked surface micropattern. Female terminalia are characterized by an unkeeled abdominal sternum VIII covering the reduced primary ovipositor. Gonapophysis VIII does not extend beyond the tip of gonapophysis IX. An asymmetry concerning the size of the paired gonapophyses is reported for the first time in Phasmatodea.     

One proboscis,two tasks: Adaptations to blood-feeding and nectar-extracting in long-proboscid horse flies (Tabanidae,Philoliche)

Female Pangoniinae in the tabanid fly genus Philoliche can display remarkably elongated proboscis lengths, which are adapted for both blood- and nectar-feeding. Apart from their role as blood-sucking pests, they represent important pollinators of the South African flora. This study examines the morphology of the feeding apparatus of two species of long-proboscid Tabanidae: Philoliche rostrata and Philoliche gulosa – both species display adaptations for feeding from a diverse guild of long-tubed flowers, and on vertebrate blood. The heavily sclerotised proboscis can be divided into two functional units. The short, proximal piercing part is composed of the labrum-epipharynx unit, the hypopharynx and paired mandible and maxilla. The foldable distal part is composed of the prementum of the labium which solely forms the food canal and is responsible for nectar uptake via the apical labella. The proboscis works as a drinking straw, relying on a pressure gradient provided by a two-part suction pump in the head. Both proboscis and body lengths and suction pump dimensions show a significantly correlated allometric relationship with each other. This study provides detailed insights into the adaptations for a dual diet using an elongated sucking proboscis, and considers these adaptations in the context of the evolution of nectar feeding in Brachycera.     

The saurian malarias of Venezuela: Haemosporidian parasites of gekkonid lizards

Telford S. P., Jr. 1978. The saurian malarias of Venezuela: haemosporidian parasites of gekkonid lizards. International Journal for Parasitology8: 341–353. Five haemosporidian species were found among 185 gekkonid lizards from Estados Portuguesa, Cojedes and Aragua, Venezuela, four of which were new to science. A pigmented Plasmodium species is described from Gonatodes taniae of Estado Aragua. It produces 8–20 merozoites in variably shaped schizonts, and elongate, irregularly margined prematuration gametocytes which contract to form round to broadly elongate mature gametocytes. Phyllodactylus ventralis of Estado Portuguesa is parasitized by two new unpigmented malarial species. One produces 11–35 merozoites in schizonts which are often rounded or elongated, occasionally fan-shaped. Gametocytes are always elongated and usually lie diagonally across one end of the host cell or laterally to the nucleus. The second species forms rounded mature schizonts nearly filled with 14–32 merozoites. The sexual stages are usually round or oval, rarely elongate. Plasmodium aurulentum Telford, 1971 was found in Thecadactylus raplcaudus of Estados Portuguesa and Cojedes. A single Thecadactylus from Cojedes was infected by a haemosporidian species of uncertain generic identity which resembles a parasite found earlier in a Panamanian gecko.     

Bark and ambrosia beetles (Curculionidae: Scolytinae), their phoretic mites (Acari) and associated Geosmithia species (Ascomycota: Hypocreales) from Virgilia trees in South Africa

Bark and ambrosia beetles are ecologically and economically important phloeophagous insects that often have complex symbiotic relationships with fungi and mites. These systems are greatly understudied in Africa. In the present study we identified bark and ambrosia beetles, their phoretic mites and their main fungal associates from native Virgilia trees in the Cape Floristic Region (CFR) of South Africa. In addition, we tested the ability of mites to feed on the associated fungi. Four species of scolytine beetles were collected from various Virgilia hosts and from across the CFR. All were consistently associated with various Geosmithia species, fungi known from phloeophagous beetles in many parts of the world, but not yet reported as Scolytinae associates in South Africa. Four beetle species, a single mite species and five Geosmithia species were recovered. The beetles, Hapalogenius fuscipennis, Cryphalini sp. 1, and Scolytoplatypus fasciatus were associated with a single species of Elattoma phoretic mite that commonly carried spores of Geosmithia species. Liparthrum sp. 1 did not carry phoretic mites. Similar to European studies, Geosmithia associates of beetles from Virgilia were constant over extended geographic ranges, and species that share the same host plant individual had similar Geosmithia communities. Phoretic mites were unable to feed on their Geosmithia associates, but were observed to feed on bark beetle larvae within tunnels. This study forms the first African-centred base for ongoing global studies on the associations between arthropods and Geosmithia species. It strengthens hypotheses that the association between Scolytinae beetles and dry-spored Geosmithia species may be more ubiquitous than commonly recognised.     

Ultrastructure of the spermatozoa ofCorystes cassivelaunus (Corystidae),Platepistoma nanum (Cancridae) andCancer pagurus (Cancridae) supports recognition of the Corystoidea (Crustacea, Brachyura, Heterotremata)

A combination of characters, not individually unique, possessed by the corystid,Corystes cassivelaunus, and the two cancrids,Platepistoma nanum andCancer pagurus, defines a corystoid-type of spermatozoon: the basally bulbous, anteriorly narrowing perforatorium, the extent of this almost to the plasma membrane through a widely perforate operculum, and the simple inner acrosome zone, lacking an acrosome ray zone. The sperm of the two cancrids are closely similar, that of the corystid differing, for instance, in the less pointed, and less tapered, form of the perforatorium. This relative uniformity of spermatozoal ultrastructure in the cancrid+corystid assemblage so far investigated supports inclusion of the two families in the superfamily Corystoidea by Guinot (1978). The combination of perforation of the operculum and absence of an acrosome ray zone (at least in a clearly recognizable form) are features of the Potamidae which possibly indicate that the latter family, modified for a freshwater existence, is related to the cancrid+corystid assemblage. Some elongation of the centrioles, apparent at least inCorystes, may be a further link with potamids in which they are greatly elongated. The coenospermial spermatophores of cancridoids are a notable difference from the cleistospermia of potamids; but the latter is probably an apomorphic modification for fertilization biology.