Functional forewing morphology and stridulation in crickets (Orthoptera, Grylloidea)

Forewing morphology and infrastructure are analysed in three cricket species ( Gryllus campestris, Oecanthus pellucens and Lerneca fuscipennis ) and checked in 20 other genera. A strong dorsoventral asymmetry is demonstrated for the first time. The upper side of the tegmina is covered with hexagonal, more or less high crests forming reliefs. On the lower side, annulate, flat and thick veins exist. The distribution of the hexagons and that of the different vein types is uneven on the forewing surfaces. Their significance in forewing functional morphology is discussed in relation to stridulation and to current knowledge on cricket sound production mechanisms. Their extent and distribution, together with the number and strength of the veins and the surface of wing cells, could account for the vibrational characteristics of the cricket forewing and explain the diverse results obtained by several authors on Gryllus and Oecanthus .     

Stridulatory activity of Eugaster (Orthoptera, Ensifera)

Under natural conditions Eugaster stridulates in the afternoon until after sunset and in the morning; it has been shown that the threshold for activity is higher in darkness than in dim light.The concept of a continuously increasing sensitization was found also to cover the observation of the stridulation of Eugaster.In continuous darkness and in dim light an endogenous rhythm does appear at high temperature; at lower temperature and at higher illumination the stridulation is irregularly distributed through the diel.The typical round of song may be recognized at constant temperature and 12 hours light and 12 hours darkness; however, it becomes more distinct and more like the natural pattern at high temperature in the daytime and low in the night.

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Résumé Des mâles isolés d'Orthoptères Ensifères stridulent chaque jour à une certaine heure, commencant généralement au crépuscule.Dans des travaux précédents (Nielsen et Dreisig 1970, Dreisig et Nielsen 1970) il a été démontré que cette forme d'activité rhythmique et d'autres analogues sont dues à une restauration progressive d'un état de sensibilité spécial, pendant la période de repos. La sensibilisation se produit plus vite à une température élevée qu'à une température basse. L'activité se déclenche lorsqu'un certain seuil de sensibilité est atteint, ce qui dépend des conditions extérieures actuelles, par exemple l'éclairage et la température. Eugaster diffère des Ensifères déjà examinés en ce que la stridulation, dans le cas de l'alternance naturelle entre le jour et la nuit, est divisée en deux périodes, la première au cours de l'après-midi et se prolongeant jusqu'au delà du crépuscule, la seconde se déclenchant vers le lever du soleil.L'existence d'un «sensibilisateur» chez Eugaster est en accord avec le fait que l'ampleur d'activité des deux périodes de chant dépend de la température au cours de la matinée précédant la reprise de la l'activité de l'après-midi (t.max. dans les figures 1 et 2).Dans les expériences de laboratoire pendant lesquelles l'éclairage et la température furent réglés, il a été constaté que le seuil de déclenchement était plus bas (donc atteint plus vite) à une lumière faible dans l'obscurité absolue. Cela confirme les suppositions tirées des expériences faites à la lumière naturelle, à savoir que l'obscurité inhibe la stridulation et que la pause entre les deux périodes d'activité est l'effet d'une faible illumination.Dans le laboratoire la répartition normale se produisit à une température constante et avec 12 heures de lumière et 12 heures d'obscurité (ou d'éclairage très faible). La répartition fut bien plus distincte en cas de température élevée pendant la période d'éclairage et de température basse pendant la période d'obscurité.A l'obscurité (ou en éclairement faible) constante, l'activité s'est manifestée en «libre cours» (free-running rhythm) avec une période inférieure à 24 heures, si la température était constamment au-dessus de 23°. En cas de température plus basse (au-dessous de 20°) la répartition de la stridulation fut irrégulière. C'est aussi le cas avec un fort éclairage constant.

     

Diversification of New Zealand weta (Orthoptera: Ensifera: Anostostomatidae) and their relationships in Australasia

New Zealand taxa from the Orthopteran family Anostostomatidae have been shown to consist of three broad groups, Hemiandrus (ground weta), Anisoura/Motuweta (tusked weta) and Hemideina-Deinacrida (tree-giant weta). The family is also present in Australia and New Caledonia, the nearest large land masses to New Zealand. All genera are endemic to their respective countries except Hemiandrus that occurs in New Zealand and Australia. We used nuclear and mitochondrial DNA sequence data to study within genera and among species-level genetic diversity within New Zealand and to examine phylogenetic relationships of taxa in Australasia. We found the Anostostomatidae to be monophyletic within Ensifera, and justifiably distinguished from the Stenopelmatidae among which they were formerly placed. However, the New Zealand Anostostomatidae are not monophyletic with respect to Australian and New Caledonian species in our analyses. Two of the New Zealand groups have closer allies in Australia and one in New Caledonia. We carried out maximum-likelihood and Bayesian analyses to reveal several well supported subgroupings. Our analysis included the most extensive sampling to date of Hemiandrus species and indicate that Australian and New Zealand Hemiandrus are not monophyletic. We used molecular dating approaches to test the plausibility of alternative biogeographic hypotheses for the origin of the New Zealand anostostomatid fauna and found support for divergence of the main clades at, or shortly after, Gondwanan break-up, and dispersal across the Tasman much more recently.     

Activity patterns of Eugaster (Orthoptera: Ensifera)

In continuation of previous work on Eugaster guyoni (Nielsen 1971) stridulation and locomotor activity has been investigated under various conditions of temperature and illumination.The theory of continuously increasing sensitization was further corroborated, and it is attempted to determine the relative thresholds of release and cessation of stridulation.The subspecies E.g. fernandezi from the Sahara is in its daily stridulation activity adapted to a higher temperature than E.g. berlandi from the Sous valley; however, at the same temperature the number of chirps per minute is the same in the two subspecies.

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Résumé Les deux sous-espèces d'Eugaster guyoni, fernandezi du Sahara et berlandi d'Agadir, diffèrent par leur activité stridulatoire. La sous-espèce du désert semble être adaptée à une température plus élevée de 5° que celle d'Agadir (E. berlandi).Les très jeunes adultes ont une très grande activité locomotrice pendant que la stridulation se développe graduellement pendant une période d'une semaine. Il semble y avoir un certain degré d'inhibition réciproque entre les deux activités locomotrice ou stridulatoire.Dans des conditions semi-naturelles l'activité du matin et du soir dépend de la température maximum atteinte durant la période de repos vers la fin de matinée.En condition d'éclairage permanent, les périodes de stridulation alternent irrégulièrement avec celles de repos. La quantité d'activité dépend de la température.Avec une alternance de 12 heures de lumiére et 12 heures d'obscurité l'activité stridulatoire dépend de la température comme en codition d'éclairage permanent. Cependant cette activité se répartit de façon caractéristique en deux périodes précédant et suivant le changement d'éclairement.Avec 12 heures de pleine lumière et 12 heures de lumière atténuée il n'y a normalement qu'une seule période de stridulation qui se manifeste surtout pendant la phase de lumière atténuée. Pour une température donnée l'activité est plus grande en lumière faible qu'en pleine lumière ou en obscurité.Une tentative est faite, dans la discussion, pour établir les valeurs relatives de la sensibilisation aux différents seuils de déclenchement et d'arrêt de la stridulation.

     

Comparative morphology of mandibular structures in lycid larvae and its phylogenetic implications (Polyphaga, Hexapoda)

The morphology of the larval mandibular structures of the family Lycidae (Polyphaga) is characterized by facultative dicondyly, posterior articulation with long mandibular rods, lateral location of the anterior condyle and its articulation with a paired non‐cranial structure. It is compared to that of Eucrustacea, Chilopoda, Entognatha, Microcoryphia and Zygentoma and found to be more reminiscent of the Entognatha. The phylogenetic implications of this conclusion are discussed, with the Pterygota and Dicondylia hypothesized to be non‐monophyletic.     

Chirp rate variability in male song ofEphippigerida taeniata (Orthoptera: Ensifera)

Variability in the chirp rate of the male song of the ephippigerine speciesEphippigerida taeniata during intraspecific communication was investigated in the laboratory. Conspecific chirps were used as auditory stimuli. The stimulus rate was controlled by computer. Experiments were carried out at 19, 27, and 35°C. Acoustically isolated males ofE. taeniata sang with a relatively constant chirp rate, which depended on the ambient temperature. Chirp rate significantly increased with rising temperature from 19 to 27°C, whereas at 35°C the chirp rate did not differ significantly from that at 27°C. Male chirp rates were affected by stimulus rates. Males significantly increased their chirp rate in response to increasing stimulus rates at temperatures of 19 and 27°C. At 35°C the increase in the chirp rate was not significant. At 27 and 35°C males sang with a higher chirp rate than the stimulus rate within a certain range. Evaluating stimulus and response chirp pattern when the males increased their chirp rate in response to the stimulus rate showed that an alternation pattern was established. More than 50% of the male chirps occurred at a characteristic time range at around 40% of the interstimulus interval, which was slightly affected by temperature.     

The peristomatic structures of Lithobiomorpha (Myriapoda, Chilopoda): comparative morphology and phylogenetic significance

A comparative survey of the epipharynx and hypopharynx of lithobiomorph centipedes by light and scanning electron microscopy examines 18 species that sample the major groups of both families, the Lithobiidae and Henicopidae. Cladistic analysis of 11 characters of the peristomatic structures together with 29 additional morphological characters serves as a basis for interpreting the evolution of the lithobiomorph peristomatic structures. Scutigeromorpha is used for outgroup comparison in the framework of a homology scheme for the basic components of the epi- and hypopharynx. Compared to other chilopods, the monophyly of Lithobiomorpha is supported by a row of distinctive bottle-shaped gland openings at the border between the labral and clypeal parts of the epipharynx, as well as by a distinctive shape of the hypopharynx. Paired rows of elongate spines on the clypeal part of the epipharynx are an apomorphic character of Lithobiidae. The transformation of these spine rows into a few groups of branching spines is characteristic for the Monotarsobius group sensu Verhoeff. Similar groups of branching clypeal spines characterize the Anopsobiinae within Henicopidae, whereas Henicopinae possess a dense cluster of short, simple spines instead. The recently described genus Dzhungaria is resolved closer to Henicopinae than to Anopsobiinae, a hypothesis supported by a field of grooves on the medial labral part of the epipharynx. Monophyly of Henicopidae does not receive unique support from the peristomatic structures although two homoplastic characters contribute to this node; among these, the reduction of a median spine field between clypeal and labral parts of the epipharynx to a narrow transverse band also supports a close relationship between the Ezembius group and Hessebius within Lithobiidae. An Ezembius+Hessebius clade is additionally supported by the absence of a transverse bulge between the clypeal and labral parts of the epipharynx, a character otherwise present in all lithobiomorph species studied so far. Lithobius is resolved as polyphyletic, with different species being most closely related to such genera as Australobius, Hessebius and Pleurolithobius.     

Secondary structure and phylogenetic analysis of the internal transcribed spacers 1 and 2 of bush crickets (Orthoptera: Tettigoniidae: Barbitistini)

We inferred secondary structure models of the internal transcribed spacers (ITS) 1 and 2 of bush crickets using a combined comparative and thermodynamic approach. The inferred secondary structure models were used to account for interdependency of interacting nucleotides in a phylogenetic analysis of the bush cricket genus Poecilimon. Our analysis indicates that the two previously reported conformational structures (i.e., hairpin and ring) of ITS2 are likely to fold in bush crickets as well and that both predicted structures are similar to those proposed for other eukaryotes. Comparing predicted ITS1 secondary structure models proved to be difficult because of substantial variation in their nucleotide sequence length. Our study revealed that the phylogenetic signal of ITS1 and ITS2 is largely congruent with that preserved in the mitochondrial genes 16S rRNA, tRNA‐Val and 12S rRNA. The phylogenetic signal in both the nuclear and the mitochondrial genome question the monophyly of the genus Poecilimon: species of the genera Poecilimonella, Parapoecilimon, Polysarcus and Phonochorion consistently cluster within Poecilimon.     

Variation in nuptial gift quality in bush crickets (Orthoptera: Tettigoniidae)

Males of many insect species provide the female, during courtshipand copulation, with a nuptial gift consisting of a prey itemor synthesized material (e.g., spermatophores). The studiesperformed so far have mainly focused on effects of nuptial giftsize on male and female reproductive success. However, the qualityof the nuptial gift can differ substantially between taxa andmay potentially have a large impact on male and female reproductiveperformance. In this study the effects of. variation in dieton nuptial gift quality is investigated in several bush cricketspecies with different diets. The effect of diet on nuptialgift quality (e.g., protein in the spermatophylax) and femalereproductive output and, in turn, die effect of variation inspermatophylax quality on female reproductive output are investigated.Female reproductive output and male spermatophore size weremainly found to be influenced by differences in diet betweenspecies. Spermatophylax quality (i.e., protein concentration)was also correlated with differences in diet. There was a largevariation in protein content of the spermatophylax widiin aswell as between species. Moreover, larger spermatophylaxes hada lower protein concentration, indicating a possible trade-offbetween spermatophylax size and quality. Consequendy, productionof larger spermatophylaxes, required for protection of the male'ssperm carrying ampulla during insemination, can lead to a reducedprotein concentration, because the total amount of protein maybe limited. This pattern is also consistent with die idea diatthe spermatophylax functions primarily to ensure sperm transfer.Finally, there was no correlation between the amount of proteinin the spermatophylax and female reproductive output eitheracross diets or within each diet category, further supportingthe finding that female reproductive output is mainly affectedby differences in diet.     

Entomopathogenic fungi detection and avoidance by mole crickets (Orthoptera: Gryllotalpidae)

A chamber to monitor mole cricket behavior was designed using two different soil-filled containers and photosensors constructed from infrared emitters and detectors. Mole crickets (Scapteriscus spp.) were introduced into a center tube that allowed them to choose whether to enter and tunnel in untreated soil or soil treated with Beauveria bassiana (Balsamo) Vuillemin. Each time the cricket passed through the photosensor located near the entrance of soil-filled containers, the infrared light was blocked and the exact moment that this occurred was logged onto a computer using custom-written software. Data examined included the first photosensor trigger, total number of sensor triggers, presence of tunneling, and final location of the cricket after 18 h. These behaviors were analyzed to discern differences in mole cricket behavior in the presence of different treatments and to elucidate the mechanism that mole crickets use to detect fungal pathogens. The first study examined substrate selection and tunneling behavior of the southern mole cricket, Scapteriscus borellii Giglio-Tos, to the presence of five strains of B. bassiana relative to a control. There were no differences between the first sensor trigger and total number of triggers, indicating the mole crickets are not capable of detecting B. bassiana at a distance of 8 cm. Changes in mole cricket tunneling and residence time in treated soil occurred for some strains of B. bassiana but not others. One of the strains associated with behavioral changes in the southern mole cricket was used in a second experiment testing behavioral responses of the tawny mole cricket, S. vicinus Scudder. In addition to the formulated product of this strain, the two separate components of that product (conidia and carrier) and bifenthrin, an insecticide commonly used to control mole crickets, were tested. There were no differences in mole cricket behavior between treatments in this study. The differences in behavioral responses between the two species could suggest a more sensitive chemosensory recognition system for southern mole crickets.     

On recognition of sound communication signals in bush crickets (Orthoptera,Tettigoniidae)

Two hypotheses have been proposed to explain the mechanisms of calling signal recognition in orthopterans: the filtration and resonance ones. To test these hypotheses, conspecific male calling songs and their models with modified temporal parameters were presented to females of bush crickets in ethological experiments. The models with a double pulse rate evoked positive phonotaxis of females while phase shift significantly complicated the recognition process. These data fit the resonance hypothesis.     

Effect of temperature on auditory receptor functions in crickets (Orthoptera,Tettigoniodea)

By electrophysiological methods, effect of temperature on cricket tympanal organ functions was studied. Activity of auditory receptors was recorded intracellularly in the 5th nerve of I thoracic ganglion in Tettigonia cantans, Metrioptera roeselii, M. bicolor, Platycleis albopunctata, Pholidoptera griseoaptera, and Phaneroptera falcata. The temperature was changed in the range from 17 to 34° C. Heating of the tympanal organ to 30–32°C led to a decrease of impulse amplitude, shortening of their duration, an increase of sensitivity, of the burst instantaneous frequency, and of the number of impulses in the responses as well as to a decrease of latent periods (LP) of receptor reaction. The optimal frequency in all studied cells did not change, although the range of perceived frequencies was enlarged. The frequency threshold curve of receptor either was shifted down along the ordinate scale without changes of its shape or the thresholds at various frequencies decreased non-uniformly. Thus, the obtained data indicate the absence of changes in the frequency tuning of the auditory receptors with changes of temperature.     

Tempo and mode of species diversification in Dolichopoda cave crickets (Orthoptera, Rhaphidophoridae)

This study focuses on the phylogenetic relationships among ninety percent of known Dolichopoda species (44 out of 49); primarily a Mediterranean genus, distributed from eastern Pyrenees to Caucasus. A total of 2490 base pairs were sequenced corresponding to partial sequences of one nuclear (28SrRNA) and three mitochondrial genes (12S, 16S and COI). A relaxed molecular clock, inferred from Bayesian analysis was applied to estimate the divergence times between the lineages using well dated palaeoevents of the study areas. Molecular substitution rates per lineage per million years were also obtained for each analyzed gene. Based on the nearly complete species phylogeny, temporal patterns of diversification were analyzed using Lineage-Through-Time plots and diversification statistics. Alternative hypotheses about the colonization of present range by Dolichopoda species were tested by means of Approximate Bayesian Computation analysis. Results from this analysis carried out on the 90% of known Dolichopoda species confirmed the previous ones based on subgroups of species, suggesting the ABC analysis as a remarkable tool in biogeographic studies. Based on these results, the distribution of Dolichopoda species appears to have been shaped by the palaeogeographic and climatic events that occurred from Late Miocene up to the Plio-Pleistocene. Both vicariance and dispersal events appear to have influenced Dolichopoda species distributions, with many processes occurring in ancestral epigean populations before the invasion of the subterranean environment.     

First song descriptions of some Anatolian species of Tettigoniidae Krauss, 1902 (Orthoptera,Ensifera)

Fourteen endemic and two sub-endemic species belonging to three subfamilies of Tettigoniidae (Tettigoniinae, Bradyporinae and Saginae) were sampled during field trips throughout the different ranges of Anatolia between the years of 2004 and 2013. Acoustic parameters of these 16 species affiliated to 8 genera (Anterastes, Apholidoptera, Gampsocleis, Parapholidoptera, Pezodrymadusa, Psorodonotus, Bradyporus and Saga) have been described for the first time in this study. Acoustical analysis showed that song characters are species-specific in the genera Saga and Psorodonotus. On the other hand, we could not find big differences among species of the genus Pezodrymadusa and Parapholidoptera castaneoviridis species-group.     

AFLP analysis of phylogenetic relationships among myrmecophytic species of Macaranga (Euphorbiaceae) and their allies

The palaeotropic pioneer tree genus Macaranga Thouars (Euphorbiaceae) is characterized by various types of mutualistic interactions with specific ant partners (mainly Crematogaster spp.). About 30 species are obligate ant-plants (myrmecophytes). We used amplified fragment length polymorphism (AFLP) markers to assess phylogenetic relationships among 108 Macaranga specimens from 43 species, including all available taxa from the three sections known to contain myrmecophytes. Eight primer combinations produced 426 bands that were scored as presence/absence characters. Banding patterns were analyzed phenetically, cladistically and by principal coordinates analysis. Monophyly of section Pruinosae is clearly supported. There is also good evidence for a monophyletic section Pachystemon that includes the puncticulata group. The monophyly of section Winklerianae and relationships between the three sections remain ambiguous. Section Pachystemon is subdivided into four well-supported monophyletic subclades that presumably correspond to taxonomic entities.We acknowledge the support by the Deutsche Forschungsgemeinschaft (DFG Fi606/4-1, DFG We1830/2-1, 4-1 and 4-2), which in part was granted in the frame of the DFG-SPP 1127 Radiations: origins of biological diversity. Part of the plant material was kindly supplied by Dr. H. Feldhaar (University of Würzburg), Dr. U. Moog (University of Kassel) and Dr. F. Slik (Leiden University Branch, Nationaal Herbarium Nederland). We thank the University of Malaysia (Dr. Rosli b. Hashim) and Taman Taman Sabah (Datuk Lamri Ali; Dr. J. Nais) for permits and logistic support, and EPU for permission to conduct research in Malaysia.     

The structure and development of a somatotopic map in crickets: The cercal afferent projection

A group of club-shaped sensilla called clavate hairs, located on the cercus of crickets (Acheta domesticus), are part of a specialized sensory system which monitors the orientation of a cricket with respect to the earth's gravitational field. The clavate hairs occur in rows which run proximodistally on the medial aspect of the cercus and each hair can be identified by specifying which row a hair is in and what position it is in within the row. The array of hairs is constant from individual to individual, and thus each hair can be identified in each specimen. The soma of a single bipolar sensory neuron is located in the integument below each hair; its dendrite projects into the hair and its axon projects to a well-defined area of the abdominal ganglion called the cercal glomerulus. All of the neurons within a row project to a particular area of the cercal glomerulus and different rows project to different areas within the glomerulus. Within a row neurons project to slightly different parts of the target area for that row. Thus a highly ordered projection pattern is produced which is tentatively called somatotopic. The development of the first clavate neuron to appear was examined from the first instar to the adult instar. The terminal arborization of this first hair was in no way unusual and its growth paralleled ganglion growth, maintaining a relatively constant position with respect to ganglion coordinates. A second clavate neuron behaved similarly, its arborization was fully formed when the receptor first appeared in the third instar and merely enlarged as the ganglion grew.