Gomphocerine Grasshoppers Research Articles

Ontogeny of color development in two green-brown polymorphic grasshopper species.

Many insects, including several orthopterans, undergo dramatic changes in body coloration during ontogeny. This variation is particularly intriguing in gomphocerine grasshoppers, where the green and brown morphs appear to be genetically determined (Schielzeth & Dieker, 2020, BMC Evolutionary Biology, 20, 63; Winter etal., 2021, Heredity, 127, 66). A better understanding of how these color morphs develop during ontogeny can provide valuable insights into the evolution and ecology of such a widespread color polymorphism. Here, we focus on the color development of two green-brown polymorphic species, the club-legged grasshopper Gomphocerus sibiricus and the steppe grasshopper Chorthippus dorsatus. By following the color development of individuals from hatching to adulthood, we found that color morph differences begin to develop during the second nymphal stage, are clearly defined by the third nymphal stage, and remain stable throughout the life of an individual. Interestingly, we also observed that shed skins of late nymphal stages are identifiable by color morphs based on their yellowish coloration, rather than the green that marks green body parts. Furthermore, by assessing how these colors are perceived by different visual systems, we found that certain potential predators can chromatically discriminate between morphs, while others may not. These results suggest that the putative genes controlling color morph are active during the early stages of ontogeny, and that green color is likely composed of two components, one present in the cuticle and one not. In addition, the effectiveness of camouflage appears to vary depending on the specific predator involved.

Comparative analysis of the multivariate genetic architecture of morphological traits in three species of Gomphocerine grasshoppers.

Evolutionary change is the change in trait values across generations, and usually occurs in multidimensional trait space rather than along isolated traits. Genetic covariation influences the magnitude and direction of evolutionary change and can be statistically summarized by the additive genetic (co)variance matrix, G. While G can affect the response to selection, it is exposed to evolutionary change by selection and genetic drift, but the magnitude and speed of these changes are poorly understood. We use comparative G matrix analyses to assess evolution of the shape and orientation of G over longer timescales in three species of Gomphocerine grasshoppers. We estimate 10 × 10 G matrices for five morphological traits expressed in both sexes. We find low-to-moderate heritabilities (average 0.36), mostly large cross-sex correlations (average 0.54) and moderate between-trait correlations (average 0.34). G matrices differ significantly among species with wing length contributing most to these differences. Wing length is the trait that is most divergent among species, suggesting it has been under selection during species divergence. The more distantly related species, Pseudochorthippus parallelus, was the most different in the shape of G. Projection of contemporary genetic variation into the divergence space D illustrates that the major axis of genetic variation in Gomphocerippus rufus is aligned with divergence from Chorthippus biguttulus, while the major axis of genetic variation in neither of the species is aligned with the divergence between Pseudochorthippus parallelus and the other two species. Our results demonstrate significant differences in G matrices with a phylogenetic signal in the differentiation.

Contributions to the study of gomphocerine grasshoppers calling songs (Orthoptera: Acrididae: Gomphocerinae) with notes on taxonomic status and distribution of some forms from Kyrgyzstan

Male calling songs of 10 widely distributed species of Gomphocerinae from localities in Kyrgyzstan were investigated. In all species songs of males from the Central Asiatic and European populations are almost identical in spite of the fact, that in some species localities in Kyrgyzstan are isolated from the main parts of their ranges by the deserts of the South Kazakhstan and Central Asia. Glyptobothrus maritimus is the only species from G. biguttulus group widely distributed over all the territory of Kyrgyzstan. G. mollis was found only in the North and Central Tien Shan Mts.; no other species of this group were found in Kyrgyzstan. Chorthippus apricarius from the Central and West Tien Shan Mts. does not differ from the nominotypical subspecies in the calling song pattern and demonstrates variability of diagnostic characters used earlier for distinguishing between Ch. apricarius apricarius and Ch. apricarius asiaticus. This gives grounds for establishing a synonymy Ch. apricarius apricarius (Linnaeus, 1758) = Ch. apricarius asiaticus Mistshenko, 1951, syn. n.

Chemical cues from females trigger male courtship behaviour in grasshoppers.

Gomphocerine grasshoppers use species-specific calling songs for sex recognition and mate attraction. In two closely related species, Chorthippus biguttulus and C. mollis, acoustic communication is the only experimentally characterized communication channel that elicits male courtship behaviour. However, courtship in these species involves extensive close-range interactions that are likely to be mediated by other signalling modalities, in particular chemical cues. We developed a bioassay to determine if female cuticular hydrocarbons (CHCs) act as chemical cues that induce courtship behaviour, and if males assess variation in CHCs to determine whether or not to court a female. The results of this approach provide evidence that grasshopper males use species- and sex-specific information from CHC signals and respond with a courtship song to the CHC profile of conspecific females but not to the CHC profile of heterospecific females and conspecific males. We conclude that males of C. biguttulus and C. mollis use multimodal channels for mating decisions, based on both acoustic and olfactory cues. We discuss various factors that might favour the evolution of male choosiness in grasshoppers.

Physical mapping of 18S and 5S rDNA loci and histone H3 gene in grasshopper species of the subfamily Gomphocerinae (Acrididae)

In this study, fluorescence in situ hybridization (FISH) analysis was used to determine and compare the numbers and chromosomal locations of two multigene families (rDNA and histone H3) in four Neotropical species of gomphocerine grasshoppers. FISH using the 18S rDNA probe identified a single site on the S9 chromosome of Amblytropidia sp and Cauratettix borelli, a single site on chromosome M6 of Compsacris pulcher, and two sites (chromosomes L1 and L2) in Orphulella punctata. By contrast, FISH with a 5S rDNA probe identified dispersion of this sequence in the genomes of the four species, with evidence of intraspecific variations. Amblytropidia sp had six to eight FISH signals on autosomal chromosomes, while C. pulcher exhibited a signal only on the M5 bivalent. The histone H3 gene was less variable and was restricted to a single pair in all species. The conservation of the numbers and locations of 18S rDNA and H3 genes in conjunction with data from the literature was useful for evaluating karyotype evolution in this subfamily. The variation in the number and sizes of 5S rDNA sites indicates a process of recent dispersion that might have been mediated by transposition.

The Elaborate Courtship Behavior ofStenobothrus clavatusWillemse, 1979 (Acrididae: Gomphocerinae)*

The elaborate courtship behavior of the gomphocerine grasshopper Stenobothrus clavatus, endemic to a few mountains in Greece, incorporates acoustic signalling and visual display elements to prime the conspecific female. The courtship song, produced by rubbing hind legs against an elytral vein, can be divided into four consecutive phases. The basic movements of phases I, II and IV consist of simple up and interrupted downstrokes, with sound produced during the downstroke only. In contrast, phase III is characterized by a fast sound-producing elevation of the femora, a kick of the tibiae and other visual display elements, such as a fast raising of the antennae with their dark spatulate tips during each movement cycle. The latter is strongly pronounced and therefore a characteristic feature of this species. The stridulatory file of each hind leg consists of two parts. The proximal part bears a double row of little pegs, whereas the pegs in the distal part are regularly aligned in a single row. The proximal part of the stridulatory file is incorporated in sound production only during phase III. However, frequency spectra from different sound elements produced during phase III were similar. Nevertheless, differences are found between phase I/II and phase III/IV, which are ascribed to variations in contact pressure of the stridulatory file against the forewing, rather than to different portions of the stridulatory file being involved.

Comparative analysis of rDNA location in five Neotropical gomphocerine grasshopper species

We report here, for the first time, the chromosome complement, number and location of the nucleolar organizer regions (NORs) revealed by silver staining (AgNO(3)) and fluorescent in situ hybridization (FISH) in five Neotropical gomphocerine species: Rhammatocerus brasiliensis, R. brunneri, R. palustris, R. pictus and Amblytropidia sp. The objective of this study was to summarize available data and propose a model of chromosome evolution in Neotropical gomphocerines. All five species studied showed chromosome numbers consisting of 2n = 23,X0 in males and 2n = 24,XX in females. Amblytropidia sp. was the only species showing a bivalent (M(8)) with megameric behavior during meiosis. The rDNA sites were restricted to autosomal pairs, i.e. the pericentromeric region of the S(9) chromosome, the consensus NOR location in all five species. R. brasiliensis was the only species showing additional NORs on M(4) and M(6) pairs which, likewise the S(9) NOR, were active in all cells analyzed. Comparison of these results with those reported previously in Palearctic gomphocerine species suggests higher resemblance of Neotropical species with the Old World species also possessing 23/24 chromosomes. Evolutionary mechanisms responsible for the observed interspecific variation in NOR location in this group are discussed.

Molecular phylogenetic evidence for multiple dispersal events in gomphocerine grasshoppers

The gomphocerine grasshoppers, comprising over 1000 species, occur on all continents excepting Australia and Madagascar. This work provides an independent examination of previous accounts of their taxonomic relationships and intercontinental connections, focusing on selected North American and Eurasian taxa. Our study is based on portions of four mitochondrial genes (coding for cytochrome b, cytochrome oxidase subunits I and II, and NADH dehydrogenase subunit V) which were sequenced and phylogenetically analyzed using weighted and unweighted maximum parsimony, maximum likelihood and Bayesian methods. Maximum resolution was achieved using weighted parsimony (counting transversions at third codon positions only) and Bayesian methods, and treating all four sequences, totalling 1892 bp, as a unit. The subfamily is provisionally accepted as monophyletic. The tribe Chrysochraontini is monophyletic, whereas the monophyletic status of Aulocarini and Dociostaurini is unclear. Tribes, Arcypterini, Chorthippini and Gomphocerini, are not monophyletic and require further scrutiny. Regarding biogeographic origins of the subfamily, our molecular data generally support Vickery's assertion that there were multiple periods of dispersal, most likely from Eurasia to North America. Assigning the range 50 to 70 Mya to the time of gomphocerine divergence, we provide estimates of the times of these biogeographic events.

Grasshopper populations across 2000 m of altitude: is there life history adaptation?

Life history differentiation along climatic gradients may have allowed a species to extend its geographic range. To explore this hypothesis, we compared eleven Omocestus viridulus (Orthoptera: Acrididae) populations along an altitudinal gradient from 410 to 2440 m in Switzerland, both in the field and laboratory. In situ temperature records indicated a striking decline in available heat sums along the gradient, and field populations at high altitudes reached egg hatching and adulthood much later in the year than at low elevation. The reproductive period at high altitude is thus severely limited by season length, especially during a cool year. However, controlled environment experiments revealed that intrinsic rates of embryonic and juvenile development increased with the populations’ altitude of origin. This countergradient variation is largely genetic and conforms to predictions of life history theory. No corresponding differentiation in the overwintering egg stage, a pivotal determinant of phenology, was found. This trait seems conserved within the gomphocerine grasshopper subfamily. Although we found evidence for altitudinal adaptation in development, the potential of O. viridulus to adapt to cool alpine climates appears restricted by a phylogenetic constraint.

Stridulatory pattern generation in acridid grasshoppers: metathoracic interneurons in Stenobothrus rubicundus (Germar 1817).

Stridulation was elicited in tethered gomphocerine grasshoppers of the species Stenobothrus rubicundus in order to identify interneurons of the stridulation pattern generator, and describe their morphological and physiological properties. Nine types of such neurons could be found and characterized; eight of those could additionally be compared to corresponding neuron types previously known from other species. As shown in detail for one selected type, the neurons of the stridulation pattern generator are very similar in their anatomical appearance, and possess similar physiological qualities at least in two species with similar stridulation patterns. Stridulation interneurons of species with largely different stridulatory motor patterns have a similar morphology, but show a different activation timing throughout the stridulation. Nevertheless, special properties such as resetting or initiation capability of certain stridulation interneurons seem to be conserved throughout the species. The results suggest that the stridulation pattern generator of different species consists of a uniform set of interneurons that change their activity pattern to produce species-specific song movements.

Pharmacological brain stimulation releases elaborate stridulatory behaviour in gomphocerine grasshoppers - conclusions for the organization of the central nervous control

Grasshoppers produce a variety of sounds generated by complex movements of the hindlegs. Stridulation, performed in the context of partner finding, mating and rivalry, can be released by pressure injection of cholinergic agonists into the protocerebrum. Particularly stimulation with muscarinic agonists induced long-lasting stridulation that resembled the natural behaviour to an astonishing degree, not only with respect to their temporal structure and right/left coordination, but also to changes in the song sequences according to the progress of courtship stridulation, even including accessory movements of other parts of the body. According to the complexity of their stridulatory behaviour ten gomphocerine species were chosen for this comparative study. The results indicate that the protocerebrum fulfils two important tasks in the control of stridulation: (1) it integrates sensory input relevant to stridulation that represents a certain behavioural situation and internal state of arousal, and (2) it selectively activates and deactivates the thoracic networks that generate the appropriate movement and sound patterns. With the knowledge of the natural behaviour and the accessibility to pharmacological and electrophysiological studies, the cephalic control system for stridulation in grasshoppers appears to be a suitable model for how the brain selects and controls appropriate behaviours for a given situation.

Pharmacological Activation of Stridulation in the Grasshopper Chorthippus albomarginatus (Orthoptera: Gomphocerinae)

Calling and courtship stridulatory behavior of Chorthippus albomarginatus was induced by injections of acetylcholine agonists into the protocerebrum. Pharmacologically induced stridulation, in many parameters, was quite similar to the natural behavior. However, the order of the courtship element alternation was different from that of the natural song. In some cases the pharmacologically induced stridulation included only one or two courtship elements. Based on the exclusive stimulation of a particular element of courtship songs and similarity of its movement pattern with the calling song, both patterns of Ch. albomarginatus stridulation appear to be homologous. The results obtained on this species confirm the idea of a hierarchic organization of the central nervous control of stridulation in gomphocerine grasshoppers and indicate participation of certain protocerebrum structures in this control.

ACOUSTIC COMMUNICATION DISTANCES OF A GOMPHOCERINE GRASSHOPPER

ABSTRACT I studied in the laboratory and in the field the communication distance of the gomphocerine grasshopper Chorthippus biguttulus, which lives close to the ground in stony, short grass habitats. Using sound measurements and behavioural tests, I examined the frequency dependent attenuation and temporal degradation of the song pattern. Sound measurements (2–40 kHz) in the field revealed that excess attenuation increases with increasing frequency. Close to the ground, excess attenuation reaches 18 dB/m. On the basis of (i) excess attenuations, (ii) tympanic receptor threshold curves and (iii) song spectra, perception distances of female tympanic receptors for male songs and vice versa were calculated. Behavioural tests revealed that, especially in short and stony grass vegetation, the typical habitat of Ch. biguttulus, the temporal song pattern is strongly degraded. Thus, in the laboratory and in tall grass vegetation, sound attenuation was found to limit the communication distance. In contrast, in short grass vegetation, degradation of the temporal pattern was found to limit the communication distance of males listening to female songs. I argue that the exploitation of fast amplitude modulations for song recognition restricts the acoustic communication of grasshoppers to short distances (1–2.2 m). In this respect it seems adaptive to stridulate at low intensities.

Recognition of sex in the acoustic communication of the grasshopper Chorthippus biguttulus (Orthoptera, Acrididae)

Many gomphocerine grasshoppers communicate acoustically: a male's calling song is answered by a female which is approached phonotactically by the male. Signals and recognition mechanisms were investigated in Chorthippus biguttulus with regard to the cues which allow sex discrimination. (1) The stridulatory files on the hindfemur of both sexes are homologous in that they are derived from the same row of bristles, but convergent with respect to the “pegs”. In males the pegs are derived from the bristles, and in females from the wall of the bristle's cup. (2) Male and female songs are generated by similar, probably homologous motor programs, but differ in the duration, intensity, “gappyness” of syllables, risetime of pulses, and the frequency spectra. The hindleg co-ordination during stridulation and the resulting temporal song patterns are less variable in males than in females. (3) For both sexes, recognition of a mate's signal depends on species-specific syllable structure. For males it is essential that the female syllables consist of distinct short pulses, whereas females reject “gappy” syllables. Males strongly prefer “ramped” pulses, females respond to syllables irrespective of steeply or slowly rising ramps. Males react only to the low-frequency component, whereas females prefer spectra containing both, low and high frequency components.

Central nervous control of hindleg coordination in stridulating grasshoppers

Stridulation in many gomphocerine grasshoppers is characterized by specific phase shifts between the two hindlegs as well as different movement patterns produced by the left and the right leg. The underlying neuronal excitation patterns are generated by networks on either side of the metathoracic ganglion. The role of the intraganglionic commissures in right-left coordination and the production of differing movement patterns was investigated by transecting the metathoracic ganglion mediosagittally in Omocestus viridulus, Chorthippus biguttulus and Chorthippus mollis. In all three species, after this operation both hindlegs produced the same pattern and no longer different movements. The effects of transection on coordination differed: rapid movement rhythms, like those typical of Ch. biguttulus and the vibratory parts of the song of Ch. mollis, on the two sides drifted with respect to one another. In contrast, the slow rhythms characteristic of O. viridulus and the song subunits of Ch. mollis were completely synchronized. It is inferred that in intact animals the pathways for coordination of the rapid stridulatory rhythms are exclusively intraganglionic, whereas the phase relations of the slow rhythms are additionally influenced by way of anterior right-left connections, perhaps within the suboesophageal ganglion.

Auditory information processing in stridulating grasshoppers: tympanic membrane vibrations and neurophysiology

During stridulation in the gomphocerine grasshopper Omocestus viridulus the leg movements, sound pattern and either summed auditory nerve activity or single interneuron activity were recorded. Simultaneous laser interferometric and vibrometric measurements of the displacement and velocity of the tympanic membrane were performed at the pyriform vesicle (d-receptor group). Slow displacements of the tympanic membrane occur in phase with the ventilatory and stridulatory rhythm and reach 10 μmpeak-peak and 1–3 μmpeak-peak in amplitude, respectively. Additionally, the tympanic membrane oscillates maximally in the range 5–10 kHz. These high-frequency oscillations are due to sound production and motor activity and correspond in amplitude to oscillations evoked by sound pressures of 90-dB SPL. They activate the auditory receptors during most of the stridulatory cycle even during mute stridulation. Only at the lower reversal point of the leg movement are membrane vibrations and receptor activity at a minimum. As a consequence the response of receptors and interneurons to auditory stimuli are generally impaired and an auditory response of receptors and interneurons can be elicited only during a short period at the lower reversal point. Although in this phase of the stridulatory cycle auditory sensitivity is present, males do not show phonotactic responses towards female songs during ongoing own stridulation.

Olfactory sensitivity of gomphocerine grasshoppers to the odours of host and non-host plants

Antennal responses to host and non-host odours were recorded, using the electroantennogram technique, from four gomphocerine grasshoppers with different feeding habits: Bootettix argentatus (monophagous on Larrea, a dicotyledonous plant), Ligurotettix coquilletti (oligophagous on dicotyledonous plants), Cibolacris parviceps (polyphagous) and Chorthippus curtipennis (graminivorous). The patterns of responses across the plant odours were similar for all four grasshopper species, but there was a relatively greater response to the odour of Larrea in B. argentatus and L. coquilletti which specialise on this plant. This suggests that there is considerable overall similarity in the make up of the olfactory receptor populations of these four grasshoppers species, but that some specificity for hostplant odour does exist at the antennal level. Resume Sensibilite olfactive d'orthopteres Gomphocerinae a l'odeur de plantes hotes et non hotes Les reponses antennaires ont ete examinees par electroantennogramme aux odeurs de plantes hotes et non hotes d'orthopteres Gomphocerinae de comportements alimentaires differents: Bootettix argentatus, monophage sur une dicotyledone, Larrea; Ligurotettix coquilletti, oligophage de dicotyledones; Cibolacris parviceps, polyphage; Chorthippus curtipennis, graminivore. Les types de reponse des quatre especes etaient tres voisins, mais les reponses aux odeurs de Larrea de B. argentatus et L. coquilletti, specialistes de cette plante, etaient relativement plus fortes. Ces resultats suggerent une similarite globale considerable dans la composition des populations de recepteurs olfactifs de ces quatre especes d'orthopteres mais aussi une certaine specificite pour les odeurs de la plante hote au niveau antennaire.

Chiasma frequencies and distributions in gomphocerine grasshoppers: a comparative study between sexes

Chiasma frequencies and distributions in gomphocerine grasshoppers: a comparative study between sexes

The chemosensory system of the monophagus grasshopper, Bootettix argentatus bruner (Orthoptera : Acrididae)

Bootettix argentatus (Orthoptera : Acrididae) is a monophagous grasshopper in the subfamily Gomphocerinae. The numbers of chemoreceptors in most groups on the mouthparts increase with each molt and are generally similar in the 2 sexes. There are fewer sensilla in the A3 groups than in other gomphocerine grasshoppers, and the possibility that this is related to host-plant specificity is discussed. Only very small numbers of sensilla are present on the pulvillar pads of the tarsi and on the arolia. The number of sensilla on the antennae is proportional to the length of the flagellum and coeloconic sensilla comprise 30–35%. Most of the remainder are multiporous basiconic sensilla. Adult males have more antennal sensilla than adult females.

The Structure of Grasshopper Song in Relation To Mating Success

Males of the gomphocerine grasshopper Chorthippus brunneus produce a conspicuous calling song consisting of groups of echemes each composed of syllables. Males with intermediate mean syllable lengths have higher mating success than extreme males. The songs of 21 males previously ranked for mating success have been further analysed using fourier transform analysis of sound envelopes. This analysis confirms the importance of intermediate syllable length and indicates other characters of behavioural signifiance. In particular, a previously undescribed grouping of syllables into triplets is associated with high mating success. This demonstrates that female grasshoppers are sensitive to quite complex features of amplitude modulation in male songs. The technique of fourier transform analysis of sound envelopes has considerable potential in the study of acoustic signals.