Surface structures of the antenna of Mantophasmatodea (Insecta)

The cuticular fine structure of the antenna in Mantophasmatodea (Austrophasmatidae: Hemilobophasma montaguense, Austrophasma gansbaaiense) is described based on SEM images. The armature of sensilla (=s.) on the basiflagellum (14 basal flagellomeres, the distal ones of which are subdivided) and distiflagellum (7 distal flagellomeres) differs strongly. A basiflagellomere has a sporadic vestiture of prominent s. chaetica B as well as distal s. scolopidia and scattered s. coeloconica. The distiflagellum also bears scattered s. chaetica B and s. coeloconica but is also densely covered with sensilla of various other types: numerous s. trichodea and s. basiconica, sporadic s. coelocapitula, and a probably new type of branched sensilla; the distiflagellum is probably the main sensory region, including gustatory and olfactory chemoreception, mechanoreception, and hygroreception. The scape and pedicel also bear s. chaetica B, and s. chaetica A arranged in 4 (scape) or 2 (pedicel) hair-plates. Terminal s. campaniformia are only found on the pedicel. A ‘dark spot’ is present apically on distiflagellomeres 6 (as reported previously) and 1. The two spots are of similar structure and are possibly associated with glands; each comprises a complicated external aperture and a larger internal pouch enclosing a cavity; projections protrude from the pouch walls into the cavity. The surface structure of the basiflagellomeres shows differences that are possibly species specific. An overview is given on antennal apomorphies in Mantophasmatodea.     

Evolution and Diversity of Vibrational Signals in Mantophasmatodea (Insecta)

Vibrational communication for species identification and mate location is widespread among insects. We investigated the vibrational communication signals of 13 species of the insect order Mantophasmatodea (Heelwalkers). Males and females produce percussive signals by tapping their abdomens on the substrate to locate conspecific mates. We show that male and female calls are of similar general structure but differ in temporal characteristics. Using a principal component analysis, we demonstrate that most species can be distinguished by their calls only. Mapping the calls onto an existing molecular phylogenetic tree reveals a slow diverging drift of male call pattern but no specific trend. For females, a trend from faster towards slower pulse repetition times is indicated. Two sympatric species, Karoophasma biedouwense and Viridiphasma clanwilliamense (Austrophasmatidae), exhibit very different call parameters. The latter species produces calls rather different from all other investigated species, which might hint at reproductive character displacement.     

Evolutionary scenarios for unusual attachment devices of Phasmatodea and Mantophasmatodea (Insecta)

Abstract.  The distal parts of the legs of representatives of Phasmatodea and Mantophasmatodea were examined. The condition found in Mantophasma zephyra and Timema nevadense is described in detail. In both species the arolium is highly modified, i.e. strongly enlarged and pan-shaped and densely covered with acanthae. The presence of acanthae on the euplantulae is another very unusual feature shared by the two taxa. A cladistic analysis based mainly on a data matrix from an earlier study of the authors was carried out, with the inclusion of three new characters derived from attachment devices. The results suggest three possible evolutionary scenarios for the features in question. If Phasmatodea are the sister group of Mantophasmatodea, the apomorphic features of the attachment devices may be synapomorphies of both groups, with different degrees of reversal within the suborder Euphasmatodea. A branching pattern Phasmatodea + (Mantophasmatodea + Grylloblattodea) is consistent with the presence of an enlarged pan-shaped arolium and euplantulae with acanthae in the common ancestor of this lineage, with reversal in Grylloblattodea and within Euphasmatodea. The acanthae on the surface of the arolium may or may not have evolved independently in Timema . A placement of Phasmatodea as sister taxon of Orthoptera, Dictyoptera, or a clade comprising both groups implies that the features in question have evolved independently in phasmids and Mantophasmatodea.     

The tentorium and anterior head sulci in Dictyoptera and Mantophasmatodea (Insecta)

The tentorium, the anterior sulci of the head capsule (epistomal, subgenal, subantennal, circumantennal, and circumocular sulci), and the extension of the anterior tentorial pit were studied in 26 species of Blattaria (representing most principal lineages), 4 species of Mantodea (including the basal Mantoida schraderi), and 1 species each of Isoptera (the basal Mastotermes darwiniensis) and Mantophasmatodea (Austrophasma caledonense). The morphology of these head structures is compared with literature data on other insect orders, mainly Phasmatodea, Orthoptera, Dermaptera, Embioptera, and Plecoptera, and partly Odonata and Zygentoma. Characters are defined, presented in a matrix, and evaluated with regard to phylogenetic implications and homoplastic evolution. The structural relationships of the subantennal sulcus to the subgenal, circumocular, and circumantennal sulci, which are highly variable and strongly homoplastic (depending much on the size of the compound eyes) are a focal issue; several types of subantennal sulci are defined. The presence of an anterior transverse bridge in the tentorium (“perforated tentorium”) of all Dictyoptera here studied confirms the monophyly of this group. Mantophasmatodea lacks this element.     

Two new genera and two new species of Mantophasmatodea (Insecta, Polyneoptera) from Namibia

Two new species and two new genera (Pachyphasma, Striatophasma) of Mantophasmatodea are described from Namibia. Pachyphasma brandbergense is endemic to the Brandberg massif; Striatophasma occupies an extensive area south of the region inhabited by Mantophasma. Phylogenetic analyses (see Predel et al. in press) suggest a sistergroup relationship of Striatophasma and the South African Austrophasmatidae.     

Postembryonic development of the spider Enoplognatha ovata (Clerck) (Araneae:Theridiidae)

The postembryonic development of the three colour varieties of Enoplognatha ovata (CI.) was studied in the field and in the laboratory for two generations. Native and European spiders of this species were compared. Estimates of the number and duration of each stage were obtained from laboratory mating studies and verified by field studies. Males and females matured after five instars. Males matured faster than females but did not live as long. Each instar was characterized by several measurable parameters. All varieties over-winter in the second instar. Sexes could be easily separated at the fourth instar by comparing the ratio of the tarsal to patellar width of the pedipalps. Sexual dimorphism occurs at maturation ecdysis. Filial and mating studies indicate that the three colour varieties belong to a single species.     

Phylogeny of the Heelwalkers (Insecta: Mantophasmatodea) based on mtDNA sequences, with evidence for additional taxa in South Africa

We examined the phylogeny of Mantophasmatodea from southern Africa (South Africa, Namibia) using approx. 1300 bp of mitochondrial DNA sequence data from the genes encoding COI and 16S. The taxon sample comprised multiple specimens from eight described species (Namaquaphasma ookiepense, Austrophasma rawsonvillense, A. caledonense, A. gansbaaiense, Lobatophasma redelinghuysense, Hemilobophasma montaguense, Karoophasma botterkloofense, K. biedouwense) and four undescribed species of Austrophasmatidae; three specimens of Sclerophasma paresisense (Mantophasmatidae); and two specimens of Praedatophasma maraisi and one of Tyrannophasma gladiator (not yet convincingly assigned to any family). For outgroup comparison a broad selection from hemimetabolous insect orders was included. Equally weighted parsimony analyses of the combined COI+16S data sets with gaps in 16S scored as a fifth character state supported Austrophasmatidae and all species and genera of Mantophasmatodea as being monophyletic. Most species were highly supported with 98-100% bootstrap/7-39 Bremer support (BS), but K. biedouwense had moderate support (87/4) and A. caledonense low support (70/1). Mantophasmatodea, Austrophasmatidae, and a clade Tyrannophasma gladiator+Praedatophasma maraisi were all strongly supported (99-100/12-25), while relationships among the two latter clades and Mantophasmatidae remain ambiguous. Concerning the relationships among genera of Austrophasmatidae, support values are moderately high for some nodes, but not significant for others. We additionally calculated the partitioned BS values of COI and 16S for all nodes in the strict consensus of the combined tree. COI and 16S are highly congruent at the species level as well as at the base of Mantophasmatodea, but congruence is poor for most intergeneric relationships. In forthcoming studies, deeper relationships in the order should be additionally explored by nuclear genes, such as 18S and 28S, for a reduced sample of specimens.     

Postembryonic development of the cervicothoracic skeleton of Euborellia annulipes (Lucas) (Dermaptera: Labiduridae)

The nymphal and adult cervicothoracic skeleton of Euborellia annulipes (Lucas) is described and discussed.     

Postembryonic development of Tisbe gracilis (T. Scott) (Copepoda, Harpacticoida)

All six naupliar and five copepodite stages of Tisbe gracilis (T. Scott, 1895) are described. A key for the identification of the nauplius stages is given. The oral appendages of all copepodite stages are described. Sexual dimorphism is visible from the copepodite IV stage on.     

Postembryonic development of Paracentrophyes praedictus (Homalorhagida): neoteny questionable among the Kinorhyncha

The postembryonic ontogeny of the kinorhynch Paracentrophyes praedictus is described by light microscopy and scanning electron microscopy. The development of head scalids and trunk Characteristics has been studied. The first of 6 juvenile stages possesses 11 segments already. One segment each is added both in stages 2 and 5. The number of head scalids increases during development. Every scalid becomes visible as a spinose anlage in one stage and differentiates in the next stage. All head scalids in stage 1 represent anlagen of scalids. Late juvenile stages J-5 and J-6 express two different scalid patterns which do not seem to depend on the gender. Scalids and trunk segments develop in a subfrontal and a subcaudal growing zone, respectively. The presumed neotenic status of P. praedictus is rejected because this species exhibits the same number of juvenile stages as observed in all other Kinorhyncha studied. It is suggested that a weakly sclerotized, juvenile-like trunk cuticle as found, e.g. in adult P. praedictus represents the plesiomorphic characteristic within the Kinorhyncha; a thickened cuticle has probably evolved independently within both the cyclorhagid and the homalorhagid Kinorhyncha.     

Heterogeneity and homologies of the repeating and unique DNA of dragonflies (Odonata, Insecta)

A relative content of unique and reiterated nucleotide sequences in DNA of eleven dragonfly species was estimated. The degree of intra- and intergenomic divergence of these DNA sequences was determined by means of DNA-DNA hybridization. Species from different genera share 40-45% of the repetitive sequences and those from different families--from 11 to 20% only. Data on the thermostability of homo- and heteroduplexes suggest that new families of the repetitive sequences have arisen repeatedly during dragonflies evolution. The quality of homologous unique sequences in the DNA compared (20-97%) correlates with the taxonomic relationships of species. Phylogenesis of some dragonfly families is discussed in view of the results obtained.     

Postembryonic development of the parasitic amphipodHyperia galba

Hyperia galba Montagu is associated with gelatinous zooplankton as are many species of the Hyperiidea. The hosts preferred in the European seas are the large scyphomedusaeAurelia aurita, Chrysaora hysoscella, Rhizostoma pulmo, Cyanea capillata andCyanea lamarckii, which harbour the first developmental stages. The anamorphic development produces young that are incapable of swimming at the time of hatching. They are characterized by an embryonic abdomen without extremities and external segmentation; the eyes are not completely developed and the mouth is primitive lacking bristles, molar and incisor. The postembryonic development, described in detail, is subdivided into two phases: the pantochelis phase and the protopleon phase; the former comprises only one stage; the latter can be subdivided into four stages. In the course of postnatal development the larval organs are reduced and characters typical of the adult are gradually differentiated.H. galba plays an important role as obligatory endoparasite of scyphomedusae at least during the first stages of development; without a host this amphipod cannot survive, neither benthically nor in the plankton. The transition from life in the female's marsupium to endoparasitism in the jellyfish generally occurs during stage of the postembryonic development which is the first stage of the protopleon phase. The specific adaptations of its reproductive biology to a parasitic mode of life such as moult inhibition under starvation, development of larval organs and the behavioural patterns of the females as well as the young are described. Further, the influence of external factors such as temperature and food supply on the course of development is examined. Dedicated to Prof. Dr. H. Mergner on the occasion of his 70th birthday.     

Sperm ultrastructure of<Emphasis Type="Italic"> Mantophasma zephyra</Emphasis> (Insecta,Mantophasmatodea)

The ultrastructure of Mantophasma zephyra spermatozoa is described. Sperm cells have a trilayered acrosome with conspicuous extra-acrosomal material which expands along the nucleus. The nucleus is crossed anteriorly by a canal and its posterior end is embedded in the centriole adjunct material. A centriole with microtubular triplets is present. The flagellum has a 9+9+2 axonemal pattern, two partially crystallised mitochondrial derivatives, two membranous sacs and three connecting bands. The accessory microtubules are filled with dense material and have 16 protofilaments in their tubular wall. The intertubular material is not very expanded. In the seminal vesicles spermatozoa are stuck together to form spermatodesms, and their heads are also joined by adherens junctions. A cladistic analysis based on sperm features indicates a close relationship of Mantophasmatodea with Mantodea.     

Morphology of the antenna cleaner in the Hymenoptera with particular reference to non-aculeate families (Insecta)

An electron microscopic survey of antenna cleaner morphology, mainly in non-aculcate Hymenoptera, is presented. Modified, scale- or paddle-shaped setae on the fore basitarsus were found to be widely distributed throughout the order, but were particularly well developed in the Xyeloidea. Megalodontoidea, Blasticotomidae, Siricoidca, Orussoidea, Cephoidea and Chalcidoidea. as well as in the aculeate family Formicidae. A comb of fine setae on the fore basitarsus was present in all Apocrita, with the exception of the Trigonalyoidea, Evaniidae, and some families of Chalcidoidea, but among the symphytan families was present only in the Orussidae. Members of the symphytan family, Anaxyelidae have a distinct line of discrete setae in the same position as the fine comb of the Orussidae and apocritans which we term a protocomb; members of the Cephidae also show an indication of a protocomb, in the form of a line of more widely spaced, erect setae, that could form part of the same transition series. Members of the Trigonalyoidea and of the Evaniidae have no comb of fine setae but do possess one or more rows of highly modified, plate-like structures on the fore basitarsi.