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A Comparative Study of Communicative Behavior in Grasshoppers
Abstract
http://deepblue.lib.umich.edu/bitstream/2027.42/56385/1/MP141.pdf
... Sound is produced by many insects as a defense against predators (Low et al. 2021). The sound produced by many species of grasshoppers when they escape by flying (Willey & Willey 1969;Otte 1970) is also considered to be an antipredator defense. However, no empirical studies have examined the antipredator function of the sounds produced by grasshoppers during flight (e.g. ...
... The sound produced by grasshoppers during flight is called crepitation (Otte 1970), regardless of the context of sound occurrence. Crepitation is observed in various species of three subfamilies within Acrididae: Oedipodinae, Gomphocerinae and Acridinae (Song et al. 2020). ...
... The crepitation sound is considered to be produced by the hindwings because the sound is heard after removing the forewings (Isely 1936;Riede 1987), and it is not heard after injuring the hindwings (Isely 1936). Previous studies have suggested that grasshoppers could produce the crepitation sound when the membrane of hindwings becomes taut (Otte 1970;Lorier et al. 2002). Studies supporting this idea are based on the sound being heard when an observer moved the hindwings of Circotettix rabula Rehn & Hebard (Orthoptera: Acrididae: Oedipodinae) and ...
Many species of grasshoppers in subfamilies Oedipodinae, Gomphocerinae and Acridinae make sounds when they escape by flying. We carried out four kinds of experiments with the Chinese grasshopper, Acrida cinerea (Orthoptera: Acrididae: Acridinae), whose males often make sounds while flying, to examine the mechanism of sound production during flight by grasshoppers. First, we recorded high‐speed video and audio of the tethered flight of the males and found that they produced sounds when they clapped their hindwings together at the end of the upstroke. Second, we recorded the tethered flights of the males when we prevented them from clapping their hindwings to clarify whether the hindwing clapping produced the sounds, and we found that the obstruction of clapping hindwings prevented sound production by the males. Third, we recorded the free flight of the males and determined whether the sound produced by clapping hindwings was not specific to the tethered condition. The males clapped their hindwings during free flight, and sounds were produced when they clapped their hindwings. Finally, we observed the tethered and free flights of the females because the possibility of crepitation flight by the females existed, and we found that sound was produced by females clapping their hindwings during flight. These results showed that male A. cinerea make sounds during flight by clapping their hindwings, and females also have the ability to make sounds by using the same mechanism. This study provides the first experimental evidence on the mechanism of sound production during flight through observing the behaviors of grasshoppers.
... Within Caelifera, stridulatory organs evolved at least 10 times across the phylogeny based on our current taxon sampling, involving many different body parts (Fig. 3). However, the ability of these structures to produce sound remains largely unconfirmed, except for a few species that use acoustic signalling for mating or defence 11,35,36 , and thus, we must consider them putative for now. Definitive sound-producing organs used for ...
... "Left-over-right" stridulation "Right-over-left" stridulation "Ambidextrous" stridulation mating evolved at least three times in Caelifera (Fig. 3), once in the common ancestor of Pneumoridae using abdomino-femoral stridulation, once in the common ancestor of Pamphagidae using Krauss's organ-femoral stridulation, and once in the common ancestor of acridid subfamilies Acridinae, Gomphocerinae and Oedipodinae using hind femora and tegmina, although the location of the stridulatory file varies within these insects 36 . Our analysis showed that abdominal tympana likely evolved at least three times (Fig. 3), once in the common ancestor of Pyrgomorphidae, once in the common ancestor of Pamphagidae, and once in the common ancestor of Romaleidae, Ommexechidae and Acrididae. ...
... However, even within this lineage, the sound-producing organs followed different evolutionary trajectories in terms of specific modifications of the stridulatory apparatus. For example, in Gomphocerinae, stridulatory pegs are located on the hind femora, which rub against the thick veins in the forewings, whereas in Oedipodinae, a row of stridulatory files on the intercalary veins in the forewing rubs against the scrapers in the hind femora 36 . In addition to the stridulatory signalling, Oedipodinae and some members of Acridinae evolved an alternative and nonstridulatory acoustic mechanism, called crepitation, which produces sound by snapping wings when they fold and unfold 36 . ...
Acoustic communication is enabled by the evolution of specialised hearing and sound producing organs. In this study, we performed a large-scale macroevolutionary study to understand how both hearing and sound production evolved and affected diversification in the insect order Orthoptera, which includes many familiar singing insects, such as crickets, katydids, and grasshoppers. Using phylogenomic data, we firmly establish phylogenetic relationships among the major lineages and divergence time estimates within Orthoptera, as well as the lineage-specific and dynamic patterns of evolution for hearing and sound producing organs. In the suborder Ensifera, we infer that forewing-based stridulation and tibial tympanal ears co-evolved, but in the suborder Caelifera, abdominal tympanal ears first evolved in a non-sexual context, and later co-opted for sexual signalling when sound producing organs evolved. However, we find little evidence that the evolution of hearing and sound producing organs increased diversification rates in those lineages with known acoustic communication.
... Field studies of S. gregaria have indicated that males outnumber females at the pairing and oviposition sites, and that there are struggles between males for the possession of females (Ellis and Ashall 1957;Popov 1958). This results in males displaying a highly active, dominant role in intersexual interactions (Golov et al. 2018), with very basic or limited courtship, as also shown for the migratory locust (Oberlin 1973), and which is assumed to be a characteristic of most Catantopinae (Otte 1970). Strong and Amerasinghe (1977) also suggested that limited male courtship is a characteristic of gregarious populations that experience strong intra-male competition. ...
... The gregarious females' pre-copulatory behavior is mainly expressed by behavioral patterns aimed at rejecting males, e.g. jumping, kicking and bending the abdomen: (Popov 1958;Loher 1959;Strong and Amerasinghe 1977;Golov et al. 2018), as known for most Catantopinae (Otte 1970;Golov et al. 2018). Fig. 6 The probability of successful copulations in intra-and interphase pairs. ...
... Short and long stridulations were also demonstrated significantly more often by the solitarious compared to the gregarious females. Interestingly, these patterns of male courtship prior to mounting are not common among Catantopinae (Otte 1970). Consistent with our hypothesis of a need for continuous reinforcement of the contact between the male and female, high level stridulation and leg vibration in the solitarious pairs also occurred in the mounting stage. ...
Mating and reproduction behaviors and strategies are fundamental aspects of an organism’s evolutionary and ecological success. In locusts, intra- as well as inter-phase reproductive interactions among gregarious and solitarious locust populations have a major impact on the locust population dynamics. However, practically all previous work on locust sexual behavior has been limited to the gregarious phase. Here we provide a first detailed description of pre-copulatory behavior of solitarious desert locusts. We compare our findings with those of previous reports of pre-copulatory behavior of gregarious locusts, focusing on the behavioral elements that serve in inter-sex signaling and communication. We also studied inter-phase (mixed pairs) reproductive interactions. Solitarious males were found to invest more in pre-copulatory courtship and signaling compared to their gregarious counterparts; and solitarious females played a more dominant role in the inter-sex communication than gregarious females. The solitarious females were also less likely to demonstrate rejection-related behavioral patterns than gregarious females. As a consequence of the particular characteristic behavior of each phase, the most successful among intra- and inter-phase pairs were gregarious males with solitary females. Least successful were solitary males paired with gregarious females, indicating a strong asymmetry in inter-phase reproductive interactions.
... Field studies of S. gregaria have 242 indicated that males outnumber females at the pairing and oviposition sites, and that 243 there are severe struggles between males for the possession of females (Ellis & Ashall, 244 1957; Popov, 1958). This results in males displaying a highly active, dominant role in 245 inter-sexual interactions (Golov et al., 2018), with very basic or limited courtship, as also 246shown for the migratory locust(Oberlin, 1973), and which is assumed to be a 247 characteristic of most Catantopinae(Otte, 1970).Strong et al. (1977) also suggested 248 that limited male courtship is a characteristic of gregarious populations that experience 249 strong intra-male competition. The gregarious females' pre-copulatory behavior is mainly 250 expressed by behavioral patterns aimed at rejecting the males (e.g. ...
... The gregarious females' pre-copulatory behavior is mainly 250 expressed by behavioral patterns aimed at rejecting the males (e.g. jumping and kicking, 251 bending the abdomen:(Popov, 1958;Loher, 1959;Strong & Amerasinghe, 1977; Golov 252 et al., 2018), as known for most Catantopinae(Otte, 1970;Golov et al., 2018). 253In marked contrast, the solitarious locusts' life history involves very different and major 254 challenges in the path to successful mating and reproduction. ...
... Short and long 274 stridulations were also demonstrated significantly more often by the solitarious 275 compared to the gregarious females. Interestingly, these patterns of male courtship prior 276to mounting are not common among Catantopinae(Otte, 1970). Consistent with our 277 hypothesis of a need for continuous reinforcement of the contact between the male and 278 female, high level stridulation and leg vibration in the solitarious pairs were also 279 extended into the mounting stage. ...
Mating and reproduction behaviors and strategies are fundamental aspects of an organism evolutionary and ecological success. In locusts, intra- as well as inter-phase reproductive interactions among gregarious and solitarious locust populations have a major impact on the locust population dynamics. However, practically all previous work on locust sexual behavior has been limited to the gregarious phase. Here we provide a first detailed description of pre-copulatory behavior of solitarious desert locusts. We compare our findings with those of previous reports of pre-copulatory behavior of gregarious locusts, focusing on the behavioral elements that serve in inter-sex signaling and communication. We also studied inter-phase (mixed pairs) reproductive interactions. Solitarious males were found to invest more in pre-copulatory courtship and signaling compared to their gregarious counterparts; and the solitarious females played a comparatively more dominant role in the inter-sex communication. The solitarious females were also less prone to demonstrate the typical rejection-related behavioral patterns displayed by the gregarious females. As a consequence of the particular characteristic behavior of each phase, the most successful among intra- and inter-phase pairs were gregarious males with solitary females. Least successful were solitary males encountered with gregarious females, indicating a strong asymmetry in inter-phase reproductive interactions. We discuss these results in the context of non-random or assortative mating in locust mixed or sympatric solitarious-gregarious populations.
... We have shown that the original definition of Gomphocerinae based on the presence of stridulatory file is no longer valid. Nevertheless, the species of typical Gomphocerinae are among the best-studied grasshoppers in terms of mating behavior (Faber 1929, Jacobs 1953, Otte 1970, Eisner 1974. Many species show elaborate pre-copulatory courtship behaviors using a combination of acoustic, vibrational, and even visual signals sometimes. ...
... However, not all species have this characteristic and there are many other species outside of Oedipodinae that have convergently evolved this trait. Many oedipodine grasshoppers exhibit acoustic communication by crepitation, but a more prominent mode of communication is visual, which involves hind wing color display during flight and various types of hind leg signaling (Otte 1970), although the function of hind wing color display might also be related to defense against visual predators. Many oedipodines are cryptically colored to mimic substrate coloration and prefer sandy habitats. ...
... Melanoplines do not exhibit any pre-copulatory courtship behavior, such as visual or acoustic. Instead, mating behavior appears to be coercive, in which males stealthily approach and jump onto females to initiate copulation (Otte 1970). A recent study by Woller and Song (2017) examined the internal morphology of copulation in Melanoplus rotundipennis (Scudder, 1878) using micro-CT technology and clarified the function of many genital components in both males and females. ...
The grasshopper family Acrididae is one of the most diverse lineages within Orthoptera, including more than 6,700 valid species distributed worldwide. Grasshoppers are dominant herbivores, which have diversified into grassland, desert, semi-aquatic, alpine, and tropical forest habitats, and exhibit a wide array of morphological, ecological, and behavioral diversity. Nevertheless, the phylogeny of Acrididae as a whole has never been proposed. In this study, we present the first comprehensive phylogeny of Acrididae based on mitochondrial genomes and nuclear genes to test monophyly of the family and different subfamilies as well as to understand the evolutionary relationships among them. We recovered the monophyletic Acrididae and identified four major clades as well as several well-characterized subfamilies, but we also found that paraphyly is rampant across many subfamilies, highlighting the need for a taxonomic revision of the family. We found that Acrididae originated in the Paleocene of the Cenozoic period (59.3 million years ago) and, because the separation of South America and Africa predates the origin of the family, we hypothesize that the current cosmopolitan distribution of Acrididae was largely achieved by dispersal. We also inferred that the common ancestor of modern grasshoppers originated in South America, contrary to a popular belief that they originated in Africa, based on a biogeographical analysis. We estimate that there have been a number of colonization and recolonization events between the New World and the Old World throughout the diversification of Acrididae, and, thus, the current diversity in any given region is a reflection of this complex history.
... Nonetheless, the rejection of courting males has been reported, including the female's jumping (before and during mounting), kicking, and lateral movements of her abdomen in the attempt to prevent copulation (Loher, 1959;Strong & Amerasinghe, 1977;Uvarov, 1977). Hind leg vibration and wing stridulation have been reported to be displayed during the pre-copulatory behavior (Morse, 1896;Norris, 1954;Laub-Drost, 1959cited in Uvarov, 1977Loher, 1959;Loher, 1961;Otte, 1970;Uvarov, 1966;Uvarov, 1977), as in other acridid grasshoppers (Haskell, 1957;Haskell, 1958;Otte, 1970). Unlike wing stridulation (displayed by both sexes), the vibration of the hind legs is soundless and much more common in the female (Loher, 1959). ...
... Nonetheless, the rejection of courting males has been reported, including the female's jumping (before and during mounting), kicking, and lateral movements of her abdomen in the attempt to prevent copulation (Loher, 1959;Strong & Amerasinghe, 1977;Uvarov, 1977). Hind leg vibration and wing stridulation have been reported to be displayed during the pre-copulatory behavior (Morse, 1896;Norris, 1954;Laub-Drost, 1959cited in Uvarov, 1977Loher, 1959;Loher, 1961;Otte, 1970;Uvarov, 1966;Uvarov, 1977), as in other acridid grasshoppers (Haskell, 1957;Haskell, 1958;Otte, 1970). Unlike wing stridulation (displayed by both sexes), the vibration of the hind legs is soundless and much more common in the female (Loher, 1959). ...
... Unlike wing stridulation (displayed by both sexes), the vibration of the hind legs is soundless and much more common in the female (Loher, 1959). The role of both behavioral elements in the sexual interaction has remained uncertain (Loher, 1959;Uvarov, 1966;Otte, 1970). ...
Studies of mating and reproductive behavior have contributed much to our understanding of various animals’ ecological success. The desert locust, Schistocerca gregaria , is an important agricultural pest. However, knowledge of locust courtship and precopulatory behavior is surprisingly limited. Here we provide a comprehensive study of the precopulatory behavior of both sexes of the desert locust in the gregarious phase, with particular emphasis on the conflict between the sexes. Detailed HD-video monitoring of courtship and mating of 20 locust pairs, in a controlled environment, enabled both qualitative and quantitative descriptions of the behavior. A comprehensive list of behavioral elements was used to generate an eight-step ethogram, depicting from first encounter between the sexes to actual copulation. Further analyses included the probability of each element occurring, and a kinematic diagram based on a transitional matrix. Eleven novel behavioral elements are described in this study, and two potential points of conflict between the sexes are identified. Locust sexual interaction was characterized by the dominance of the males during the pre-mounting stage, and an overall stereotypic male courtship behavior. In contrast, females displayed no clear courtship-related behavior and an overall less organized behavioral sequence. Central elements in the sexual behavior of the females were low-amplitude hind-leg vibration, as well as rejecting males by jumping and kicking. Intricate reciprocal interactions between the sexes were evident mostly at the mounting stage. The reported findings contribute important insights to our knowledge of locust mating and reproductive behavior, and may assist in confronting this devastating agricultural pest.
... Grapples and mating attempts only occur when males mount each other. Because female grasshoppers can show aggression towards males [53,54], we also recorded any instance of female-male aggression. Female aggressive behaviours are limited to kicks and grapples (Table 1). ...
... Female aggression occurs in other grasshopper species (e.g. Gomphocerus rufus, Arphia pseudonietana, Shistocerca lineata), [53] and [54] reported pre-copulatory female aggression towards males. Because many grasshopper species do not have a distinguishable courtship phase, communication between males and females typically occurs only once the male has mounted a female. ...
... Because many grasshopper species do not have a distinguishable courtship phase, communication between males and females typically occurs only once the male has mounted a female. The 'struggling' observed between a female and a mounted male in species like Schistocerca sp. and G. rufus occurs when a female is unreceptive or partially unreceptive [53,54]. A similar process may occur in K. tristis, where a lack of obvious courtship combined with the high likelihood that late-season females have already mated, may consequently lead to more aggression towards males. ...
Throughout the breeding season, changing environmental and biological conditions can lead to variation in the reproductive landscape of many species. In alpine environments temperature is a key driver of behaviour for small ectotherms such as insects, but variable biotic factors such as mate quality and availability can also influence behaviour. Kosicuscola tristis is a small semelparous grasshopper of the Australian alpine region. In a rare behaviour among grasshoppers, K. tristis males engage in vigorous fights over access to females, involving mandible displays, kicking, biting and grappling. In this study we describe the variation in fighting behaviour of K. tristis throughout the breeding season and test several hypotheses related to temperature, body size, mating behaviour, and female quality. We show that K. tristis males are more aggressive toward each other at the end of the breeding season than at the beginning. This increased aggression is associated with decreased daily average temperatures (from ~20°C to ~9°C), decreased mating activity, increased female fecundity, and an unexpected trend toward an increase in female-to-male aggression. These results suggest that K. tristis is likely under increased selective pressure to time key life cycle events with favourable biological and climatic conditions. The stochastic nature of alpine environments combined with a relatively short life span and breeding season, as well as limited mating opportunities toward the end of the season may have contributed to the evolution of this extraordinary mating system.
... This is the reason why acoustic signals are often used in taxonomy, when sibling species are similar in morphology, but quite different in songs. Among Acrididae subfamilies, acoustic communication in Gomphocerinae is most developed in terms of structure of acoustic apparatus, temporal pattern of the song, and mating strategies (e.g., Otte, 1970;Helversen & Helversen, 1994;Ragge & Reynolds, 1998). The song is produced by stroking a stridulatory file of each hind femur across a raised vein on the fore wing. ...
... A solitary male produces a calling song, listening for the response song of a female which is ready to mate. When a male finds a female, in many species the male begins a special courtship song, which may reach a high complexity and may be accompanied by conspicuous movements of different parts of the body such as abdomen, head, antennae or palps (Faber, 1953, Otte, 1970, Helversen & Helversen, 1994. ...
Male courtship songs of 9 grasshopper species of Gomphocerinae from Kazakhstan and Orenburg region and Altai Republic of Russia were studied. We analyzed not only the sound, but also the stridulatory movements of the hind legs to more entirely describe the songs. We also analyze the frequency spectra of the songs and the whole visual display during courtship. The courtship songs of three species, Stenobothrus miramae, Chorthippus dubius and Ch. angulatus, were studied for the first time. In four species, Omocestus haemorrhoidalis, O. petraeus, Myrmeleotettix pallidus, Ch.karelini, we found certain differences in temporal pattern of the courtship songs in comparison with the previous data on the respective species from other regions. In five species, O. viridulus, S. miramae, M. pallidus, Ch. dubius and Ch. karelini, various parts of elaborate courtship songs differed in the carrier frequency. In four species, O. haemorrhoidalis, O. petraeus, M. pallidus and Ch. dubius, the dominant frequencies of the courtship song were shown to lie in the range higher than 20 kHz. The conspicuous movements of antennae and legs during courtship were studied in M. pallidus, S. miramae and Gomphocerus sibiricus.
... This is the reason why acoustic signals are often used in taxonomy to solve the problems at the species level, when sibling species are similar in morphology, but quite different in songs. Among Acrididae subfamilies, acoustic communication in Gomphocerinae is most developed in terms of structure of acoustic apparatus, temporal pattern of the song, and mating strategies (e.g., Otte, 1970;Helversen & Helversen, 1994;Ragge & Reynolds, 1998). The song is produced by stroking a stridulatory file of each hind femur across a raised vein on the fore wing. ...
... A solitary male produces a calling song, listening for the response song of a female that is ready to mate. When a male finds a female, in many species the male begins a special courtship song, which may reach a high complexity and may be accompanied by conspicuous movements of different parts of the body such as abdomen, head, antennae or palps (Faber, 1953, Otte, 1970, Helversen & Helversen 1994. ...
On the basis of the song and morphological analyses, we revised the status of the subspecies Stenobothrus eurasius eurasius Zubowsky, 1898, and S. eurasius hyalosuperficies Vorontsovskii, 1927. The status of the subspecies S. eurasius hyalosuperficies Vor. has been changed to the species level. The most striking difference between S. eurasius and S. hyalosuperficies lies in the song parameters. The calling songs differ not only in temporal parameters but are also produced by different mechanisms. S. eurasius generates calling songs by common leg stridulation, whereas S. hyalosuperficies produces sound by wing clapping. The courtship songs of both species are complex (contain several elements) and very different in temporal structure. The morphological differences between the two species are not as striking as the differences in bioacoustics: we found the only differences in the hind wing venation. At the same time, we suggest these differences to be important since they might be due to different mechanisms of sound production. We revised the ranges of the two species on the territory of Ukraine, Russia and Kazakhstan. We also reviewed the type localities of S. eurasius Zub. and designated lectotype and paralectotype of this species.
... This is the reason why acoustic signals are often used in taxonomy, when sibling species are similar in morphology, but quite different in songs. Among Acrididae subfamilies, acoustic communication in Gomphocerinae is most developed in terms of structure of acoustic apparatus, temporal pattern of the song, and mating strategies (e.g., Otte, 1970;Helversen & Helversen, 1994;Ragge & Reynolds, 1998). The song is produced by stroking a stridulatory file of each hind femur across a raised vein on the fore wing. ...
... A solitary male produces a calling song, listening for the response song of a female which is ready to mate. When a male finds a female, in many species the male begins a special courtship song, which may reach a high complexity and may be accompanied by conspicuous movements of different parts of the body such as abdomen, head, antennae or palps (Faber, 1953, Otte, 1970, Helversen & Helversen, 1994. ...
Male courtship songs of 9 grasshopper species of Gomphocerinae from Kazakhstan and Orenburg region and Altai Republic of Russia were studied. We analyzed not only the sound, but also the stridulatory movements of the hind legs to more entirely describe the songs. We also analyze the frequency spectra of the songs and the whole visual display during courtship. The courtship songs of three species, Stenobothrus miramae, Chorthippus dubius and Ch. angulatus, were studied for the first time. In four species, Omocestus haemorrhoidalis, O. petraeus, Myrmeleotettix pallidus, Ch.karelini, we found certain differences in temporal pattern of the courtship songs in comparison with the previous data on the respective species from other regions. In five species, O. viridulus, S. miramae, M. pallidus, Ch. dubius and Ch. karelini, various parts of elaborate courtship songs differed in the carrier frequency. In four species, O. haemorrhoidalis, O. petraeus, M. pallidus and Ch. dubius, the dominant frequencies of the courtship song were shown to lie in the range higher than 20 kHz. The conspicuous movements of antennae and legs during courtship were studied in M. pallidus, S. miramae and Gomphocerus sibiricus.
... This is the reason why acoustic signals are often used in taxonomy to solve the problems at the species level, when sibling species are similar in morphology, but quite different in songs. Among Acrididae subfamilies, acoustic communication in Gomphocerinae is most developed in terms of structure of acoustic apparatus, temporal pattern of the song, and mating strategies (e.g., Otte, 1970;Helversen & Helversen, 1994;Ragge & Reynolds, 1998). The song is produced by stroking a stridulatory file of each hind femur across a raised vein on the fore wing. ...
... A solitary male produces a calling song, listening for the response song of a female that is ready to mate. When a male finds a female, in many species the male begins a special courtship song, which may reach a high complexity and may be accompanied by conspicuous movements of different parts of the body such as abdomen, head, antennae or palps (Faber, 1953, Otte, 1970, Helversen & Helversen 1994. ...
On the basis of the song and morphological analyses, we revised the status of the subspecies Stenobothrus eurasius eurasius Zubowsky, 1898, and S. eurasius hyalosuperficies Vorontsovskii, 1927. The status of the subspecies S. eurasius hyalosuperficies Vor. has been changed to the species level. The most striking difference between S. eurasius and S. hyalosuperficies lies in the song parameters. The calling songs differ not only in temporal parameters but are also produced by different mechanisms. S. eurasius generates calling songs by common leg stridulation, whereas S. hyalosuperficies produces sound by wing clapping. The courtship songs of both species are complex (contain several elements) and very different in temporal structure. The morphological differences between the two species are not as striking as the differences in bioacoustics: we found the only differences in the hind wing venation. At the same time, we suggest these differences to be important since they might be due to different mechanisms of sound production. We revised the ranges of the two species on the territory of Ukraine, Russia and Kazakhstan. We also reviewed the type localities of S. eurasius Zub. and designated lectotype and paralectotype of this species.
... This is the reason why acoustic signals are often used in taxonomy, when sibling species are similar in morphology, but have quite different songs. Among the Acrididae subfamilies, acoustic communication in Gomphocerinae is most developed in terms of structure of acoustic apparatus, temporal pattern of the song, and mating strategies (e.g., Otte 1970;von Helversen and von Helversen 1994;Ragge and Reynolds 1998). The song is produced by ZooKeys 1200: 1-26 (2024), DOI: 10.3897/zookeys.1200.118422 ...
... Several males sitting in a close vicinity can produce rival songs. When a male finds a female, in many species the male begins a special courtship song, which may reach a high complexity and may be accompanied by conspicuous movements of different parts of the body such as the abdomen, head, antennae, or palps (Faber 1953;Otte 1970;von Helversen and von Helversen 1994). ...
The songs of seven grasshopper species of subfamily Gomphocerinae from Russia, Ukraine, Georgia, and Kazakhstan were studied. We analysed not only the sound, but also the stridulatory movements of the hind legs to more entirely describe the songs. In Mesasippus kozhevnikovi, Chorthippus macrocerus, and C. hammarstroemi, the legs are moved in a relatively simple pattern; four other species, Myrmeleotettix palpalis, Stenobothrus newskii, C. pullus, and Megaulacobothrus aethalinus demonstrate more complex leg movements. In six of the seven species studied, the courtship songs contain more sound elements than the calling songs. The highest number of courtship sound elements was found in M. palpalis and M. aethalinus. The different parts of a remarkably long stridulatory file in M. aethalinus are thought to participate in the production of different sound elements. The songs in S. newskii are shown for the first time. This species produces sound not only by common stridulation but also by wing beats. A relationship of S. newskii to some other species of the genus Stenobothrus, which are able to crepitate, is discussed. We also analyse the frequency spectra of the songs. A maximum energy of the song power spectra in six species studied lies in ultrasound range (higher than 20 kHz). In only M. aethalinus, the main peaks in the song power spectra lie lower than 20 kHz. The courtship behaviour in M. palpalis, C. macrocerus, and C. hammarstroemi was shown to include conspicuous visual display (movements of antennae, palps and the whole body).
... Among Orthoptera, crickets (Gryllidae) have been used extensively to study malemale combat (Hack 1997;Hofmann and Schildberger 2001), however aggressive behaviour is extremely rare among other Orthoptera such as grasshoppers (Acrididae) (Otte 1970;Greenfield and Minckley 1993;Umbers et al. 2012Umbers et al. , 2013. The thermocolour skyhopper or 'chameleon' skyhopper, Kosciuscola tristis, presents an interesting exception, and males frequently engage in fierce fights over access to females (Umbers et al. 2012Muschett et al. 2017Muschett et al. , 2018. ...
... These fights involve males flaring their mandibles, biting, kicking, and grappling with opponents, often leading to injury (Umbers et al. 2012) (Fig. 1). Male combat in this species raises questions relating to the evolution and maintenance of aggression in an otherwise relatively non-combative taxon (Otte 1970). ...
The evolution of male-male aggression is of interest because at its extreme it can be very energetically costly, leave males vulnerable to preadtors, and give rise to weaponry such as exaggerated traits. In grasshoppers (Acrididae), one group stands out as exceptionally aggressive, the skyhoppers ( Kosciuscola ) in which males bite, kick, mandible flare, and wrestle each other for access to females or when females are laying eggs. In this study we asked whether there is variation in aggressive behaviour among four skyhopper species and aimed to determine whether the traits used in fighting bear signatures of sexual selection in their size, variability, and allometric scaling. We found clear differences in the numbers and types of aggressive behaviours among species. Kosciuscola tristis and K. usitatus were the most aggressive, K. cognatus was the least aggressive, and K. tristis was the only species that performed the ‘mandible flare’ behaviour. Mandible size was larger among the three species that showed aggressive behaviour, all except K. cognatus , and was negatively allometric for all species possibly suggesting a functional size constraint. Pronotum size was different among most species and K. tristis ’ pronotum was the largest and borderline positively allometric perhaps suggesting that pronotum size is related to aggressive behaviour but the nature of that relationship remains obscured. Our study suggests that further work investigates skyhoppers’ aggressive behaviour and how it varies with ecology, and paves the way for establishing them as a model system in the evolution of aggressive behaviour.
... Male imagines typically have a brown background color combined with some white and black characteristic patterns (i.e., brown morph); whereas females show green, brown, and sometimes intermediate color morphs (Otte, 1981;Otte & Williams, 1972). This species exhibits one of the most complex courtship behaviors in insects, in which males display acoustic signals and a diverse set of body movements (Otte, 1970(Otte, , 1972. Once mating occurs, it normally lasts a few hours. ...
... First, expanding our understanding of the development after final ecdysis (i.e., imago sexual maturation process) in both males and females, and its implications for the reproductive biology of the admirable grasshopper, might be relevant for disentangling the conditions that promoted the evolution of its highly complex courtship behavior (Otte, 1970(Otte, , 1972. Second, our results call for a more detailed analysis of the relative contribution of environmental and genetic components to the large between-and within-individual variation in coloration we described for this species (see for example Schielzeth & Dieker, 2020). ...
Understanding how water limitation affects the performance of herbivorous insects is relevant in a context of worldwide alterations of rainfall regimes due to climate change. Apart from its influence on insect growth and development, water availability has also been considered a cue for color polyphenism in some species. This is the case of the grassland‐inhabiting admirable grasshopper (Syrbula admirabilis), for which water content in plant tissues, a potentially reliable cue to predict future vegetation conditions, has been proposed to determine the frequency of green–brown color morphs. We performed an experiment that manipulated water content of the diet (dry/wet treatment) in a group of individually monitored admirable grasshoppers to explore the effects on imago size, timing of female first mating, and coloration. Our results showed that grasshoppers in the wet treatment achieved larger sizes, and females mated comparatively earlier, than their counterparts in the dry treatment. We also found that hue (the dominant color) in imagines was significantly lower (browner) than in nymphs, but the magnitude of this color shift did not differ between treatments. This study supports the idea that water limitation has a negative impact on grasshopper growth and development, but rules out dietary water content as a cue for color polyphenism. This result encourages additional research to assess the actual contribution of environmental and genetic factors to color determination in admirable grasshoppers. Such information could help to better understand how color polymorphism is maintained in natural populations and to predict how it could evolve in a future in which environmental conditions will become more unpredictable.
... This is the reason why acoustic signals are often used in taxonomy to solve the problems at the species level, when sibling species are similar in morphology, but quite different in songs. Among Acrididae subfamilies, acoustic communication in Gomphocerinae is most developed in terms of structure of acoustic apparatus, temporal pattern of the song, and mating strategies (e.g., Otte, 1970;Helversen & Helversen, 1994;Ragge & Reynolds, 1998). The song is produced by stroking a stridulatory file of each hind femur across a raised vein on the fore wing. ...
... A solitary male produces a calling song, listening for the response song of a female that is ready to mate. When a male finds a female, in many species the male begins a special courtship song, which may reach a high complexity and may be accompanied by conspicuous movements of different parts of the body such as abdomen, head, antennae or palps (Faber, 1953, Otte, 1970, Helversen & Helversen 1994. ...
On the basis of the song and morphological analyses, we revised the status of the subspecies Stenobothrus eurasius eurasius Zubowsky, 1898, and S. eurasius hyalosuperficies Vorontsovskii, 1927. The status of the subspecies S. eurasius hyalosuperficies Vor. has been changed to the species level. The most striking difference between S. eurasius and S. hyalosuperficies lies in the song parameters. The calling songs differ not only in temporal parameters but are also produced by different mechanisms. S. eurasius generates calling songs by common leg stridulation, whereas S. hyalosuperficies produces sound by wing clapping. The courtship songs of both species are complex (contain several elements) and very different in temporal structure. The morphological differences between the two species are not as striking as the differences in bioacoustics: we found the only differences in the hind wing venation. At the same time, we suggest these differences to be important since they might be due to different mechanisms of sound production. We revised the ranges of the two species on the territory of Ukraine, Russia and Kazakhstan. We also reviewed the type localities of S. eurasius Zub. and designated lectotype and paralectotype of this species.
... The North American band-winged grasshoppers are an ideal animal group for the exploration of female-biased VA because 1) females are often larger than males, and 2) their mating behaviors (although variable) often involve female discrimination of fine visual signals (Otte, 1970;Otte, 1984). Band-winged grasshoppers (subfamily Oedipodinae) are a morphologically diverse globally distributed subfamily of ~200 diurnal species known for their colorful hindwing patterns. ...
... One possibility is that females could benefit by more accurately interpreting visual signals. Although mating systems and behaviors are variable across band-winged grasshoppers, many species use a variety of potential visual signals including those involving either their colorful hindwings or leg movements (Kerr, 1974;Otte, 1970;Otte, 1984;Willey and Willey, 1969). Females may therefore benefit by being better able to perceive and interpret these potential signals of species identity and mate quality, and the smaller ∆Φy values could be especially useful for detecting leg motion signals with a strong vertical component. ...
Visual acuity (VA) — a measurement of the fineness or coarseness of vision — may vary within a species including between the biological sexes. Although numerous studies have found males with finer VA than females, relatively few have shown the opposite with females having finer vision. This is surprising because our understanding of between species differences in VA suggests that females may have finer vision than males if they 1) are larger than males, or 2) need finer vision to detect and/or discriminate between males. Here, we estimate the interommatidial angle (ΔΦ, an anatomical measurement of VA) in three species of band-winged grasshoppers in which females are both the larger sex and likely interpret visual signals (Arphia pseudonietana, Dissosteira carolina, and Spharagemon equale; total n = 98). Using a radius of curvature estimation method, we find that females have ∼19% finer estimated ΔΦ than males in the most acute region and axis of the eye, but that this dimorphism varies between species. Further visual explorations of the species showing the greatest body size dimorphism (D. carolina) suggest that this ΔΦ dimorphism is driven by females having larger eyes with more ommatidia. In contrast to many diurnal flying insects where males have finer vision to acquire mates, our study is one of the first to demonstrate a female-biased sexual dimorphism in acuity. Given 1) the number of species in which females are larger than males, and 2) the variability of mating behaviors across taxa, our results suggest that differences in VA between the sexes may be more common than currently appreciated.
... For instance, orthopterans and certain cicadas engage in "crepitation," a loud, sudden, crackling sound produced by the hind wings. The source mechanism is reported to be stiffened veins in the hindwing that mechanically snap back and forth between stable states (Otte 1970). This sound production mechanism appears to have received little research. ...
... In grasshoppers, crepitation is under facultative control (Willey and Willey 1970;Niedzlek-Feaver 1995). For instance, many species produce crepitation when jumping to escape an approaching human, apparently as a startle; and many species also produce it as a sexual signal in flight (Otte 1970). ...
There are at least eight ways that wings potentially produce sound. Five mechanisms are aerodynamic sounds, created by airflow, and three are structural sound created by interactions of solid surfaces. Animal flight is low Mach (M), meaning all animals move at less than 30% of the speed of sound. Thus in aerodynamic mechanisms the effects of air compressibility can be ignored, except in mechanism #1. Mechanism #1 is trapped air, in which air approaches or exceeds Mach 1 as it escapes a constriction. This mechanism is hypothetical but likely. #2 is Gutin sound, the aerodynamic reaction to lift and drag. This mechanism is ubiquitous in flight, and generates low frequency sound such as the humming of hummingbirds or insect wing tones. #3 is turbulence-generated atonal whooshing sounds, which are also widespread in animal flight. #4 are whistles, tonal sounds generated by geometry-induced flow feedback. This mechanism is hypothetical. #5 is aeroelastic flutter, sound generated by elasticity-induced feedback that is usually but not always tonal. This is widespread in birds (feathers are predisposed to flutter) but apparently not bats or insects. Mechanism #6 is rubbing sound (including stridulation), created when bird feathers or insect wings slide past each other. Atonal rubbing sounds are widespread in bird flight and insects; tonal stridulation is widespread in insects. #7 is percussion, created when two stiff elements collide and vibrate, and is present in some birds and insects. Mechanism #8 are tymbals and other bistable conformations. These are stiff elements that snap back and forth between two conformations, producing impulsive, atonal sound. Tymbals are widespread in insects but not birds or bats; insect cuticle appears predisposed to form tymbals. There are few examples of bat wing sounds: are bats intrinsically quiet, or just under-studied? These mechanisms, especially Gutin sound, whooshes, and rubbing (#2, #3, and #6) are prominent cues in ordinary flight of all flying animals, and are the ‘acoustic substrate’ available to be converted from an adventitious sound (cue) into a communication signal. For instance, wing sounds have many times evolved into signals that are incorporated into courtship displays. Conversely, these are the sounds selected to be suppressed if quiet flight is selected for. The physical mechanisms that underlie animal sounds provides context for understanding the ways in which signals and cues may evolve.
... Por un lado, los caracteres morfológicos propios para la identificación son la forma de la placa subgenital, la carina pronotal lateral, los detalles del fastigio, la presencia o ausencia de un aparato estridulador y, en algunos casos, la nervadura del ala posterior [46]. Seguido a esto, muchos taxones de esta tribu muestran reducción o pérdida del aparato estridulador femoral, asimismo, a pesar de esta reducción en el comportamiento estridulatorio, tienden a tener una región engrosada y esclerotizada en el borde del ala posterior y células más o menos agrandadas en el área media [46], [48]. (Otte, 1979) [1], [49]. ...
... El género actualmente incluye 18 especies, seis presentes en Suramérica y doce en Centroamérica y sur de México. Las especies en este grupo generalmente están asociadas con regiones montañosas [1], [48]. ...
Traditionally, the subfamilies Acridinae, Gomphocerinae and Oedipodinae belon to the family Acrididae are part of those insects that are taken as a reference when thinking of grasshoppers. They are part of an important link as a food resource in the trophic chain and are also indicators of ecosystem changes. Acrididae is one of the most diverse Orthoptera groups in the world, however, the records to Neotropical region show that much remains to be described and reported. In the last decades, the discovery of species and genera for the South American continent has had an exponential increase and the different descriptions have been gradually clarified due to the implementation of revision characters to delimit species such as: sexual dimorphism, genitalia characters, presence or absence of specific morphological characters and the geographic distribution of the species as factors to take into account for a more accurate group review. On the other hand, the status of these subfamilies are controversial. Acridinae over the decades has become a group that gathers isolated organisms with morphological similarity and requires prompt revision. Gomphocerinae is the most diverse subfamily in relation to the others, its distribution and morphological variation make it difficult to separate species, highlighting the scarce information available for the groups; Oedipodinae is a phylogenetically group further away from those already mentioned, the ecosystems it inhabits contrast with the characteristics presents in Neotropics, however, Colombia has the presence of two species. This study sought to make a synopsis of the subfamilies Acridinae, Gomphocerinae and Oedipodinae (Orthoptera: Acrididae) for Colombia, based on a search for documents in different information sources (Database, journals, entomological collections, among others) including generalities in the morphology and natural history of the group, current distribution of the species and the design of identification keys. A total of 3,115 specimens deposited in entomological collections of Museums and Institutes in the most representative cities of Colombia were reviewed, of which a total of 14 species were identified, grouped into: Acridinae three species, Gomphocerinae nine species and Oedipodinae two
species respectively. The distribution analysis showed that most of the species found have a widespread distribution in the Colombian territory, with the exception of Silvitettix malasmonotus with restricted distribution in the departments: Boyacá, Cundinamarca and Tolima. The results allow us to conclude that the information that exists for these groups is incomplete and makes it difficult to manage it for the review of certain groups. Likewise, no new species records were found for the country and the role of entomological collections as repositories of information for studies like this one is highlighted.
... This is the reason why acoustic signals are often used in taxonomy, when sibling species are similar in morphology, but quite different in songs. Among Acrididae subfamilies, acoustic communication in Gomphocerinae is most developed in terms of structure of acoustic apparatus, temporal pattern of the song, and mating strategies (e.g., Otte, 1970;Helversen & Helversen, 1994;Ragge & Reynolds, 1998). The song is produced by stroking a stridulatory file of each hind femur across a raised vein on the fore wing. ...
... A solitary male produces a calling song, listening for the response song of a female which is ready to mate. When a male finds a female, in many species the male begins a special courtship song, which may reach a high complexity and may be accompanied by conspicuous movements of different parts of the body such as abdomen, head, antennae or palps (Faber, 1953, Otte, 1970, Helversen & Helversen, 1994. ...
Male courtship songs of 9 grasshopper species of Gomphocerinae from Kazakhstan and Orenburg region and Altai Republic of Russia were studied. We analyzed not only the sound, but also the stridulatory movements of the hind legs to more entirely describe the songs. We also analyze the frequency spectra of the songs and the whole visual display during courtship. The courtship songs of three species, Stenobothrus miramae, Chorthippus dubius and Ch. angulatus, were studied for the first time. In four species, Omocestus haemorrhoidalis, O. petraeus, Myrmeleotettix pallidus, Ch.karelini, we found certain differences in temporal pattern of the courtship songs in comparison with the previous data on the respective species from other regions. In five species, O. viridulus, S. miramae, M. pallidus, Ch. dubius and Ch. karelini, various parts of elaborate courtship songs differed in the carrier frequency. In four species, O. haemorrhoidalis, O. petraeus, M. pallidus and Ch. dubius, the dominant frequencies of the courtship song were shown to lie in the range higher than 20 kHz. The conspicuous movements of antennae and legs during courtship were studied in M. pallidus, S. miramae and Gomphocerus sibiricus.
... Onset of the rather limited courtship behavior can be recognized initially by the 320 display of "orienting", in which the male points his antennae towards the female. This behavior is 321 common in the subfamily Catantopinae (Otte 1970). 322 ...
... have been reported to feature during both pre-mounting and mounting (see also Norris 1954;Laub-342 Drost cited in Uvarov 1977;Otte 1970). While their role is still not fully resolved, in our current 343 observations they were more frequent during mounting (as also mentioned by Loher, 1959). ...
Studies of mating and reproductive behavior have contributed much to our understanding of various animals’ ecological success. The desert locust, Schistocerca gregaria , is an important agricultural pest. However, knowledge of locust courtship and precopulatory behavior is surprisingly limited. Here we provide a comprehensive study of the precopulatory behavior of both sexes of the desert locust in the gregarious phase, with particular emphasis on the conflict between the sexes. Detailed HD-video monitoring of courtship and mating of 20 locust pairs, in a controlled environment, enabled both qualitative and quantitative descriptions of the behavior. A comprehensive list of behavioral elements was used to generate an eight-step ethogram, depicting from first encounter between the sexes to actual copulation. Further analyses included the probability of each element occurring, and a kinematic diagram based on a transitional matrix. Eleven novel behavioral elements are described in this study, and two potential points of conflict between the sexes are identified. Locust sexual interaction was characterized by the dominance of the males during the pre-mounting stage, and an overall stereotypic male courtship behavior. In contrast, females displayed no clear courtship-related behavior and an overall less organized behavioral sequence. Central elements in the sexual behavior of the females were low-amplitude hind-leg vibration, as well as rejecting males by jumping and kicking. Intricate reciprocal interactions between the sexes were evident mostly at the mounting stage. The reported findings contribute important insights to our knowledge of locust mating and reproductive behavior, and may assist in confronting this devastating agricultural pest.
... At the same time, most researchers consider acoustic behavior to be the most important and, occasionally, the only mechanism of reproductive isolation in the subfamily Gomphocerinae. Therefore, it is commonly recognized that studies of the acoustic signals of these grasshoppers must have taxonomic implications (Otte, 1970;Ragge and Reynolds, 1998). Bukhvalova (1998) stated even more explicitly that the difference in the male signal structure was the principal premating reproductive barrier, and, therefore, the amplitude-temporal structure of calling signals is the most important diagnostic character in Gomphocerinae. ...
... Thus, the amplitude-temporal parameters of signals, like any other taxonomic character taken separately, are of limited value in the taxonomy of Gomphocerinae. The bioacoustic data may prove to be more useful when applied to particular taxonomic problems at the species level (Otte, 1970;Helversen, 1986;Ingrisch, 1993;Ragge and Reynolds, 1998). However, it should always be borne in mind that closely related species may have identical calling songs, and in such cases serious taxonomic conclusions cannot be based solely on an analysis of acoustic signals. ...
The acoustic communication and specific features of ecology and distribution of Ramburiella turco-mana and R. bolivari in Russia and the Transcaucasia were studied. The calling song of R. bolivari and the complete acoustic repertoire of R. turcomana are described for the first time. The acoustic repertoire of R. turcomana includes calling, rivalry, courtship, and jumping-on songs. Pallasiella is treated as a subgenus of the genus Ram-buriella. The genus Caucasippus is shown to belong to the tribe Hypernephiini. The importance of amplitude-temporal patterns of acoustic signals in the taxonomy of Gomphocerinae is discussed. The genus Ramburiella Bol. belongs to the tribe Arcypterini and includes five species. Two of them, R. turcomana F.-W. and R. bolivari Kuthy, are typical elements of the fauna of semi-desert landscapes in Russia and the Transcaucasia. The acoustic behavior of Ramburiella species has been studied poorly. Calling songs of R. hispanica Ramb. from France and Spain (Ragge and Reynolds, 1998) and those of R. tur-comana from southeastern Kazakhstan (Bukhvalova and Vedenina, 1998) have been described. We studied the specific features of acoustic communication of R. turcomana and R. bolivari from various localities of the Lower Volga region, Daghestan, and Armenia. Calling signals of R. bolivari and the complete acoustic repertoire of R. turcomana, including four types of signals, are described in the present communication. In addition, it summarizes the data on distribution of Ramburiella species in Russia and the Transcaucasia and considers some ecological features of R. turco-mana and R. bolivari. This communication also includes taxonomic notes concerning the composition of the tribe Arcypterini and the genus Ramburiella. Finally , based on a comparison of amplitude-temporal patterns of acoustic signals emitted by representatives of the genus Ramburiella and related genera and on other data on the acoustic communication in Gom-phocerinae, we discuss the possible use of bioacoustic data in the taxonomy of this subfamily. MATERIALS AND METHODS This study is based primarily on the signal records and collections of grasshoppers, made in 1995-2000 in the following localities: (1) northeastern part of Astra-khan Province, near Lake Baskunchak and Bolshoi Bogdo Mt., 17.VII-7.VIII.1995 and 1-7.VII.1996 (be-low referred to as Baskunchak); (2) southwestern part of Astrakhan Province, near Basinskaya Station, on elongated sandy hills, 4-10.VII.1998 (Basinskaya); (3) Daghestan, 20 km W of Makhachkala, near Sarykum sand-dune, NE slopes of Karatebe Range, 100-200 m above sea level, 11-16.VII.1997 (Sary-kum); (4) Armenia, Vedinskii District, E of Gorovan Vill., 900-1000 m above sea level, rocky semi-desert, 13.VII.2000 (Armenia). A total of 95 acoustic signals emitted by 14 individuals of Ramburiella spp. were analyzed. Most signals were recorded by the author; signals of several individuals of R. turcomana from Baskunchak were recorded by A.Yu. Lekarev. The methods for rearing of the grasshoppers in laboratory and studying their acoustic behavior were described in detail in a previous communication (Savitsky, 2000). The signals were recorded with MD 382 and MKE-15A microphones (frequency range 80 Hz-12.5 kHz and 50 Hz-15 kHz, respectively), connected to "Elektronika-302-1" and "Elektronika M-401C" tape recorders (frequency range 63 Hz-10 kHz and 63 Hz-12.5 kHz, respectively). The amplitude-temporal parameters were analyzed using an IBM AT computer equipped with an analog-digital converter. Data on the phenology and distribution of grasshoppers within the area studied were obtained from the author's own collections, published data, and examina
... All other aspects of mating behavior and structure of courtship are similar to A. thalassinus. The researchers (Larrosa et all, 2007) assume the courtship behavior is being of «Acrolophitus-type» (Otte, 1970). ...
... Song terminology is accepted after Zhantiev (1981) and Otte (1970): the song consists of pulses (result of friction of the femur of hind legs against the tegmina) (mono-or bisyllabic echemes sensu: Larrosa et al., 2007) and chirps (result of sliding hitting of the tibia of hind legs against the tegmina). ...
Впервые описаны и проиллюстрированы осциллограммами акустические прекопуляционные сигналы самцов трех видов: Epacromius pulverulentus, E. tergistinus и Platypygius crassus из России (Южная Сибирь) и Украины (Приазовье). Их звуки продуцируются двумя способами: тегмина-феморальной (у всех видов) и тибиа- тегминальной (у E. tergistinus) стридуляцией. Призывные звуки у этих видов не обнаружены.
... Basking may last approximately two to three hours before they become active, start walking about and feeding. Grasshopper have been observed to mate and oviposit later in the day (3,92,103). As soil temperatures rise, individuals will climb vegetation to help thermo-regulate. ...
... As soil temperatures rise, individuals will climb vegetation to help thermo-regulate. When temperatures began to decline in late afternoon, grasshoppers will bask for a second time and then become active, pottering, feeding, mating, and ovipositing, before taking their nighttime positions (92,98,99,117). ...
Grasshoppers have caused significant damage to crops of farmers and rangeland forage of ranchers throughout history. Populations of grasshoppers can build and explode with exponential growth under the right climatic factors and habitat. Melanoplus sanguinipes is the most economic species of grasshopper in the United States. The migratory flight capabilities are very similar to true locusts. This species has been known to travel distances of over 500 miles. Millions of dollars in damage occurs to crops and rangeland annually in the United States due to this species and other economic grasshopper species. The life history of this species and timing of control measures are critical in the suppression of populations to safeguard crops and rangeland. Various techniques are employed to survey, detect, and treat grasshoppers. One of the most cost-effective methods of treatment is to use Reduced Agent-Area treatments versus blanket coverage treatments.
... At the same time, most researchers consider acoustic behavior to be the most important and, occasionally, the only mechanism of reproductive isolation in the subfamily Gomphocerinae. Therefore, it is commonly recognized that studies of the acoustic signals of these grasshoppers must have taxonomic implications (Otte, 1970;Ragge and Reynolds, 1998). Bukhvalova (1998) stated even more explicitly that the difference in the male signal structure was the principal premating reproductive barrier, and, therefore, the amplitude-temporal structure of calling signals is the most important diagnostic character in Gomphocerinae. ...
... Thus, the amplitude-temporal parameters of signals, like any other taxonomic character taken separately, are of limited value in the taxonomy of Gomphocerinae. The bioacoustic data may prove to be more useful when applied to particular taxonomic problems at the species level (Otte, 1970;Helversen, 1986;Ingrisch, 1993;Ragge and Reynolds, 1998). However, it should always be borne in mind that closely related species may have identical calling songs, and in such cases serious taxonomic conclusions cannot be based solely on an analysis of acoustic signals. ...
The acoustic communication and specific features of ecology and distribution of Ramburiella turcomana and R. bolivari were investigated in Russia and Transcaucasia. The calling song of A. bolivari and complete acoustic repertoire of A. turcomanaare described for the first time. The acoustic repertoire of R. turcomana includes calling, rivalry, courtship, and jumping on songs. Pallasiella is treated as a subgenus of the genus Ramburiella. The genus Caucasippus is shown to be a member of the tribe Hypemephiini. The importance of amplitude-temporal patterns of acoustic signals in taxonomy of Gomphocerinae family is discussed.
... The bibliography at the end of the paper includes all recent papers and phonograph records dealing with Michigan singing insects. Because no keys to the sounds and associated behavior of locusts (Acrididae) are included in the present paper, discussions of these characteristics by Cantrall (1943Cantrall ( , 1968 and Otte (1971) will be especially useful. ...
... Such groups, in which there is no pre-copulation courtship mechanism, are characterized by the absence of specificity of the female genitalia (Cigliano & Lange 2007). In other groups of Acrididae (e.g., Acridinae and Oedipodinae), the genitalia are uniform at the specific level (Dirsh 1956;Otte 1984), but they show visual and acoustic specific pre-reproductive behavior (Otte 1970). Song & Bucheli (2010) compared the phylogenetic signal between genitalia and non-genitalia characters based on 41 cladistic analyses of different groups of insects and showed that the phylogenetic signal is similar in both sources of characters, concluding that although male genitalia diverge rapidly by being under sexual selection, it is not necessarily equated with the lack of phylogenetic signal. ...
Diponthus Stål, one of the most diversified and widely distributed Romaleinae genera, is endemic to southern South America. This colorful grasshopper genus is placed within Romaleini and currently includes 16 valid species from a total of 22 nominal ones, most of them only known from their original descriptions. The aims of this study were to propose a morphology-based phylogenetic hypothesis of relationships among the species of Diponthus, and to conduct a taxonomic revision, including the description of four previously unknown species. Parsimony analysis of 79 morphological characters strongly recovered the monophyly of genus Diponthus, and Gurneyacris as its sister group. Internal clades within Diponthus were supported by external morphology and coloration patterns, while characters from male genitalia were shown to be more useful for species differentiation. Based on the results of the taxonomic review, Diponthus is constituted by 16 valid species, with the following nomenclatural changes proposed: Diponthus nigroconspersus (Stål) is considered a valid name; D. invidus Carl and D. bilineatus Rehn are synonymized under D. virgatus (Gerstaecker); Diponthus clarazianus Pictet & Saussure is synonymized under D. cribratus (Serville); D. paulista Rehn is synonymized under D. porphyreus (Gerstaecker); D. maculiferus (Walker) is synonymized under D. electus (Serville). New names are proposed for four undescribed species from Bolivia (D. colorbellus sp. nov. and D. dilatatus sp. nov.), Argentina (D. paranaensis sp. nov.) and Brazil (D. salvadorii sp. nov.).
... At the evolutionary level, the hindwings of band-winged grasshoppers present an interesting case study of how a single structure may be under multiple different selective pressures. In addition to their anti-predator purposes and inherent biomechanical function, hindwings may also serve in visual conspecific signaling (Otte, 1970(Otte, , 1985 although behavioral data is lacking). Hindwing patterning often varies. ...
Some brightly colored structures are only visible when organisms are moving, such as parts of wings that are only visible in flight. For example, the primarily brown Carolina grasshopper (Dissosteira carolina) has contrasting black-and-cream hindwings that appear suddenly when it takes off, then oscillate unpredictably throughout the main flight before disappearing rapidly upon landing. However, the temporal dynamics of hindwing coloration in motion have not previously been investigated, particularly for animals that differ from humans in their temporal vision. To examine how quickly this coloration appears to a variety of non-human observers, we took high-speed videos of D. carolina flights in the field. For each of the best-quality takeoffs and landings, we performed a frame-by-frame analysis on how the relative sizes of the different-colored body parts changed over time. We found that in the first 7.6 ± 1.5 ms of takeoff, the hindwings unfurled to encompass 50% of the visible grasshopper, causing it to roughly double in size. During the main flight, the hindwings transitioned 6.4 ± 0.4 times per second between pauses and periods of active wing-beating (31.4 ± 0.5 Hz), creating an unstable, confusing image. Finally, during landings, the hindwings disappeared in 11.3 ± 3.0 ms, shrinking the grasshopper to 69 ± 9% of its main flight size. Notably, these takeoffs and landings occurred faster than most recorded species are able to sample images, which suggests that they would be near-instantaneous to a variety of different viewers. We therefore suggest that D. carolina uses its hindwings to initially startle predators (deimatic defense) and then confuse them and disrupt their search images (protean defense) before rapidly returning to crypsis.
... Crepitation. Crepitation is defined as the production of sounds by a weak wing membrane between veins that suddenly expands and vibrates due to a fast airflow 22 . Some extant Acridinae, Gomphocerinae, and Oedipodinae grasshoppers crepitate with fenestrated zones of their hind wings when they fly away, even for a short distance 19,23,24 . ...
Acoustic communication is well-known in insects since the Mesozoic, but earlier evidence of this behavior is rare. Titanoptera, an ‘orthopteroid’ Permian-Triassic order, is one of the few candidates for Paleozoic intersex calling interactions: some specimens had highly specialized broadened zones on the forewings, which are currently considered—despite inconclusive evidence—as ‘resonators’ of a stridulatory apparatus. Here we argue that the stridulatory apparatus hypothesis is unlikely because the Titanoptera lack a stridulatory file on their bodies, legs or wings. Instead, comparing these broadened zones with similar structures in extant locusts, flies, and fossil damselflies, we find evidence that the Titanoptera used their wings to produce flashes of light and/or crepitated sounds. Moreover, we describe the first Carboniferous (~310 Mya) Titanoptera, which exhibits such specialized zones, thus corresponding to the oldest record of wing communication in insects. Whether these communication systems were used to attract sexual partners and/or escape predators remain to be demonstrated. Schubnel, Nel and colleagues present the first Carboniferous representative of Titanoptera, Theiatitan azari. Specially modified wing zones suggest that Titanoptera were capable of wing-based communication using light or sound, as seen in modern species.
... Some of the most complex signal patterns and motor programs in invertebrates are displayed in the context of intraspecific communication. The elaborate acoustic and/ or vibratory signals of drosophilids (Mazzoni et al. 2013;Herrnández and Fabre 2016), hemiptera such as cicadas and bugs (Claridge 1985;Fonseca 1991;Virant-Doberlet and Cokl 2004), and orthopteran insects such as acridid grasshoppers (Otte 1970;Bull 1979;Vedenina and von Helversen 2003) and crickets (Alexander 1962;Otte 1992) are crucial for species-specific recognition during mating behaviour. Their evolution has been shaped by sexual selection (Zahavi 1980;Huber and Gerhardt 2002;Ronacher 2019) to produce reliable and species-specific communication signals, which can be stereotypic signal sequences like in a Morse code. ...
Chirping male crickets combine a 30 Hz pulse pattern with a 3 Hz chirp pattern to drive the rhythmic opening-closing movements of the front wings for sound production. Lesion experiments suggest two coupled modular timer-networks located along the chain of abdominal ganglia, a network in A3 and A4 generating the pulse pattern, and a network organized along with ganglia A4–A6 controlling the generation of the chirp rhythm. We analyzed neurons of the timer-networks and their synaptic connections by intracellular recordings and staining. We identified neurons spiking in phase with the chirps and pulses, or that are inhibited during the chirps. Neurons share a similar “gestalt”, regarding the position of the cell body, the dendritic arborizations and the contralateral ascending axon. Activating neurons of the pulse-timer network elicits ongoing motor activity driving the generation of pulses; this activity is not structured in the chirp pattern. Activating neurons of the chirp-timer network excites pulse-timer neurons; it drives the generation of chirps and during the chirps the pulse pattern is produced. Our results support the hypothesis that two modular networks along the abdominal ganglion chain control the cricket calling song, a pattern generating network in the mesothoracic ganglion may not be required.
... 50-51, 215) фразы, образованные более или менее однотипными пульсами или парами пульсов. При этом крыловые сигналы разных видов хорошо различаются периодичностью повторения фраз, продолжительностью фраз и пульсов, числом пульсов в составе фраз и частотой их повторения [ см. также : Otte, 1970;Бенедиктов, 1998]. ...
ABSTRACT: Amplitude-temporal patterns of
acoustic signals of Asiotmethis muricatus (Pall.), A.
zacharjini (B.-Bien.), Dericorys tibialis (Pall.), Truxalis
eximia Eich., Ochrilidia hebetata kazaka (Tarb.),
Mecostethus alliaceus (Germ.) and Hyalorrhipis clausi
(Kitt.) and also acoustic repertoires of Stenobothrus
eurasius hyalosuperficies Vor. и Chorthippus
macrocerus purpuratus Vor. are described for the first
time. Acoustic signals of Stenobothrus fischeri (Ev.),
Chorthippus maritimus Mistsh., Ch. mollis (Ch.), Ch.
parallelus (Zett.), Ch. dichrous (Ev.), Ch. karelini (Uv.),
Euchorthippus pulvinatus (F.-W.), Stethophyma
grossum (L.), Aiolopus thalassinus (F.) and Bryodema
gebleri (F.-W.) from before nonstudied populations are
described. Some features of acoustic and sexual
behaviour of Oedaleus decorus (Germ.), Oedipoda
caerulescens (L.), Sphingonotus rubescens (Walk.) and
some other species are noted. A comparative analysis
of the acoustic and sexual behaviour of studied species
of grasshoppers was performed.
РЕЗЮМЕ: Впервые описаны амплитудно-времен-
ные параметры акустических сигналов Asiotmethis
muricatus (Pall.), A. zacharjini (B.-Bien.), Dericorys
tibialis (Pall.), Truxalis eximia Eich., Ochrilidia hebetata
kazaka (Tarb.), Mecostethus alliaceus (Germ.) и
Hyalorrhipis clausi (Kitt.), а также акустические ре-
пертуары Stenobothrus eurasius hyalosuperficies Vor.
и Chorthippus macrocerus purpuratus Vor. Описаны
акустические сигналы Stenobothrus fischeri (Ev.),
Chorthippus maritimus Mistsh., Ch. mollis (Ch.), Ch.
parallelus (Zett.), Ch. dichrous (Ev.), Ch. karelini (Uv.),
Euchorthippus pulvinatus (F.-W.), Stethophyma
grossum (L.), Aiolopus thalassinus (F.) и Bryodema
gebleri (F.-W.) из ранее не изучавшихся популяций. Описаны некоторые особенности акустического и
полового поведения Oedaleus decorus (Germ.),
Oedipoda caerulescens (L.), Sphingonotus rubescens
(Walk.) и ряда других видов. Проведен сравнитель-
ный анализ акустического и полового поведения изу-
ченных видов саранчовых.
... Ôàêò ñîâìåñòíîãî îáèòàíèÿ A. muricatus è D. tibialis â íåêîòîðûõ áèîòîïàõ òàêaeå ïîäòâåðaeäàåò ýòîò âûâîä.  ýòîé ñâÿçè óìåñòíî îòìåòèòü, ÷òî â Ñåâåðíîé Àìåðèêå êðûëîâûå ñèãíàëû âèäîâ ïîäñåìåéñòâà Oedipodinae, íàñåëÿþùèõ îäèí áèîòîï, âñåãäà ðàçëè÷àþòñÿ ÷àñòîòîé ïîâòîðåíèÿ ïóëüñîâ è õàðàêòåðîì èõ ãðóïïèðîâàíèÿ âî ôðàçû [Otte, 1970]. ...
... Primary defences include camouflage, hiding, roosting, akinesis and aggregation. Secondary defences include running, jumping, flying, dodging, dropping, climbing, moving into the centres of thick bushes (hiding), remaining motionless, kicking, biting, gouging, regurgitation, defecation, autotomy, threat-signalling, startle displays, flash-thenhide behaviour and group scatter/confusion behaviour (Edmunds, 1973;Lagos, 2017;Maeno et al., 2013Otte, 1970;Uvarov, 1977). Given their abundance, large size, wealth of defences, daily cyclical movements among different microhabitats, highly variable daily temperatures and diversity of possible predators, locusts are ideal animals to study how temperature, photoperiod and microhabitat interact to influence antipredator defence. ...
We examined environmental factors influencing plasticity in antipredator defences of adult gregarious desert locusts, Schistocerca gregaria, including daily cycles in temperature, light, microhabitat occupied and predator threat. In the Sahara Desert in Mauritania, West Africa, daily temperature fluctuated widely from below locusts’ cold thermal limits for jump‐ and flight‐defence, to above their preferred body temperature. Locusts changed microhabitats throughout the 24‐hr period in synchrony with the daily thermo‐photocycle. They roosted in tall trees and large bushes at night, moved to the ground in the morning, shaded under or in small bushes and annuals at midday, moved back to the ground in the afternoon and then returned to night roosting sites around dusk. Locust antipredator defences varied throughout the 24‐hr period, and these changes were correlated with temperature, photocycle and habitat. Flight escape was associated with daytime, high temperatures and the ground habitat. Dropping escape (= releasing hold of vegetation and dropping to the ground or into vegetation) was associated with cool temperatures and low‐to‐medium sized bushes. Stationary behaviour was associated with the tree microhabitat and height off the ground. Roosting (a primary defence) was associated with cool temperatures at night and early morning, tree habitats and nocturnal ground‐foraging times of endothermic mammals. In summary, we propose that temperature is the key factor in determining both changing microhabitat choice and changing antipredator defence, due to thermal constraints on locust muscle for this ectothermic insect. The thermocycle also influences temporal predator loads, which influence the evolution of locust diel defence strategies. These various environmental factors not only influence one another, they also interact to influence antipredator defence expression in locusts. Overall, our study suggests that plasticity in locust antipredator defences is a complicated matter mediated by the interactions of multiple environmental factors and physiological and ecological constraints and trade‐offs.
... Wing communication sounds are produced in flight by birds that include flappet larks (Mirafra spp.) (Norberg, 1991), manakins (Bostwick and Prum, 2003) and guans (Delacour and Amadon, 1973); fruit bats (Boonman et al., 2014); and insects such as butterflies ( Yack et al., 2000), grasshoppers (Otte, 1970) and mosquitoes (Cator et al., 2009). In many of these examples, the wing sounds are produced only during displays, such that the communication sound is not also present in ordinary flight. ...
Wing trills are pulsed sounds produced by modified wing feathers at one or more specific points in time during a wingbeat. Male Allen's Hummingbird (Selasphorus sasin) produce a sexually dimorphic 9 kHz wing trill in flight. Here we investigate the kinematic basis for trill production. The wingtip velocity hypothesis posits that trill production is modulated by the airspeed of the wingtip at some point during the wingbeat, whereas the wing rotation hypothesis posits that trill production is instead modulated by wing rotation kinematics. To test these hypotheses, we flew six male Allen's Hummingbirds in an open jet wind tunnel at flight speeds of 0, 3, 6, 9, 12 and 14 m s-1, and recorded their flight with two 'acoustic cameras' placed below and behind, or below and lateral to the flying bird. The acoustic cameras are phased arrays of 40 microphones that used beamforming to spatially locate sound sources within a camera image. Trill Sound Pressure Level (SPL) exhibited a U-shaped relationship with flight speed in all three camera positions. SPL was greatest perpendicular to the stroke plane. Acoustic camera videos suggest that the trill is produced during supination. The trill was up to 20 dB louder during maneuvers than it was during steady state flight in the wind tunnel, across all airspeeds tested. These data provide partial support for the wing rotation hypothesis. Altered wing rotation kinematics could allow male Allen's Hummingbird to modulate trill production in social contexts such as courtship displays.
... In Drosophila , where species differ in several temporal characters of male song, evolutionary trends are not apparent within the groups that have been analyzed (Hoikkala and Mazzi 2009 ). Similarly, analyses of gomphocerine grasshoppers, which are noted for complex, multimodal signaling (Otte 1970 ), have not revealed trends toward increasing complexity in song (Nattier et al. 2011 ). In summary, temporal song elements, being controlled by neuromuscular factors, may be particularly labile characters (cf. ...
Tympanal organs for hearing in the far field have evolved on multiple occasions among insects and are currently found in seven orders. Many, if not most, cases of insect hearing probably originated as a means for detecting and avoiding predators. In particular, sensitivity to ultrasound appears to have coevolved with echolocation signaling by insectivorous bats. However, on an overall scale, hearing is relatively rare among insects in comparison with other modalities of perception, including detection of substrate vibration. Sound signaling in insects, which typically occurs in the context of mating communication, is rarer still and is known in only five orders. Phylogenetic analyses suggest that acoustic communication in the Lepidoptera and in the suborder Caelifera (grasshoppers) of the Orthoptera originated via a “sensory bias” mechanism. Hearing was ancestral and sound signaling by males subsequently arose on multiple, independent occasions. On the other hand, acoustic communication in the Cicadidae and in the suborder Ensifera (crickets, katydids) of the Orthoptera may have originated via coevolution between female perception and male signaling. The diversity of songs among acoustic insects may reflect genetic drift and reproductive character displacement. There is little evidence, however, that insect songs are adapted to specific physical environments. In one clade of acoustic insects, the diversification of song is associated with an unusually high rate of population differentiation and speciation, which may be facilitated by a genomic co-localization of loci influencing female response/preference and male signaling. The extent to which co-localization is a general factor in speciation remains to be explored.
... Furthermore, the first author observed a male perched on the back of a female on June 11 and July 3, 2015, although it is unknown if the male and female were the same pair on both dates as the majority of the 15 specimens (from the second visit to Seminole State Park (Field #PG191-2-A)) were still alive at the time and almost all were adults. This behavior is typical of Melanoplus mating behavior (Otte 1970), usually before attempts at copulation, although no copulation was ever observed. ...
Melanoplus foxi Hebard, 1923 (Orthoptera: Acrididae: Melanoplinae) is a flightless and morphologicallydistinct member of the Puer Group of grasshoppers that is endemic to sandhills in the state of Georgia, U.S.A. This species had not been collected in almost 60 years despite our combined eight years of independently searching for it in and around all known localities. Thus, the species was thought to possibly be extinct until our rediscovery of a combined total of 33 specimens of M. foxi in May of 2015 in three locations: Seminole State Park and Little Ocmulgee State Park and Lodge, which are protected areas, and along the road just outside of the latter. The discovery of a large number of specimens of M. foxi among the unidentified grasshoppers in the collection of the University of Michigan’s Museum of Zoology’s Insect Division and the subsequent examination of the collectors’ digitized field notes (linked to the specimens via a code system) led directly to the discovery of these extant populations. This rediscovery highlights the need for detailed field notes and retaining their link to the specimens they describe, often an arduous task, but one which some institutions are embracing by converting field notebooks into digital form either via
wholesale scanning, transcription, or a combination of both, and allowing the public to access them on the internet. Such data are invaluable and should be recorded by field explorers and made available to others whenever possible. In addition, absence or negative data, largely unreported in natural history publications, also played a role in the rediscovery of this unique species because it gave us the ability to keep track of investigated sites, and these are reported here to assist in planning future field trips to discover new locations harboring M. foxi.
Many grasshopper species produce conspicuous sounds while escaping from approaching predators; however, they occasionally escape without producing sounds. The Chinese grasshopper, Acrida cinerea , often exhibits noisy escape behavior. Therefore, a field experiment was conducted using A. cinerea to identify factors related to the production of sound during escape. This study utilized a predator model with an investigator approaching A. cinerea three times. We examined the relationship between the production of sound during escape and the following factors: ambient temperature and relative humidity as environmental factors; sex, body length, body weight, and limb autotomy as prey traits; and the repeated approach as a predator trait. The relationships between noisy escape and flight initiation distance (i.e., predator-prey distance when the prey initiates the escape), distance fled (i.e., distance the prey covered during the escape), and the mode of locomotion during escape (i.e., flying or jumping) were also examined. Noisy escape was observed only in males that escaped by flying, whereas the females and males that escaped by jumping invariably escaped silently. Among males that flew, noisy escape was related to ambient temperature, limb autotomy, and distance fled. The proportion that produced sound increased in parallel with the ambient temperature and distance fled. This proportion was lower among individuals that had autotomized one of their hind legs. These results indicate that noisy escape behavior is most frequent in healthy male A. cinerea under warm conditions.
The evolution of the calling songs in Gomphocerinae was evaluated via estimating a phylogenetic signal of the song characters and an ancestral character state reconstruction. Analyses of the calling songs in 80 palearctic gomphocerine species allowed us to define 24 characters describing the temporal pattern of the sound and the stridulatory leg‐movement pattern. The ancestral song of Gomphocerinae was shown to consist of numerous short echemes lasting on average 0.9 s; each echeme comprised only one syllable produced by movements of only one leg. The next step of the song evolution could be producing longer echemes or longer echeme‐sequence. Later, echeme duration again decreased, but this was accompanied by increasing of echeme or syllable complexity. The characters describing the echeme structure were found to be conservative in their evolution. By contrast, most characters of the syllable temporal structure were shown to be relatively labile and more likely under natural or sexual selection. Our study shows that the song evolution in Gomphocerinae implied not only increasing but also decreasing complexity of the syllable temporal structure.
The genus Chilecicada Sanborn, 2014 is shown to be a complex of closely related species rather than a monospecific genus. Chilecicada citatatemporaria Sanborn & Cole n. sp., C. culenesensis Sanborn & Cole n. sp., C. curacaviensis Sanborn & Cole n. sp., C. impartemporaria Sanborn & Cole n. sp., C. magna Sanborn & Cole n. sp., C. mapuchensis Sanborn n. sp., C. oraria Sanborn & Cole n. sp., C. parrajaraorum Sanborn n. sp., C. partemporaria Sanborn & Cole n. sp., C. pehuenchesensis Sanborn & Cole n. sp., C. trifascia Sanborn n. sp., C. trifasciunca Sanborn & Cole n. sp., and C. viridicitata Sanborn & Cole n. sp. are described as new. Chilecicada occidentis Walker, 1850 is re-described to facilitate separation of the new species from the only previously known species. Song and cytochrome oxidase I analysis available for most species support the separation of the new taxa from the type species of the genus. Known species distributions and a key to the species of the genus are also provided. The new species increases the known cicada diversity 61.9% to 34 species, 91.2% of which are endemic to Chile.
Inventories of Amazon invertebrates are relatively incipient and fragmented. The state of Amapá is one of the Amazonian states with a large knowledge gap regarding invertebrate biodiversity. Also, there is no record in the literature of systematic studies that focus mainly on the Acridofauna. Therefore the goal of this study was to understand the diversity and abundance of grasshoppers (Gomphocerinae) of the Environmental Protection Area of the Curiaú river, Macapá - AP. Twelve samples were collected from October 2011 to September 2012 using the active search technique with sweep nets. A total of 508 Gomphocerinae individuals were sampled and classified into five genera and twelve species. The floristic composition of sites A1 and A3, and sites A5 and A6, are considered more similar since the locusts are closely related to the vegetation. This is the first study assessing the diversity of orthopteroides in the state of Amapá. However, this pioneering study is expected to be the basis of future studies.
In many species, males can increase their fitness by mating with the highest quality females. Female quality can be indicated by cues, such as body size, age and mating status. In the alpine grasshopper Kosciuscola tristis, males can be found riding on subadult females early in the season, and as the season progresses, males engage in fights over ovipositing females. These observations suggest that males may be competing for females that are either unmated (early season) or sperm‐depleted (late season). We thus hypothesised that male K. tristis may be choosy in relation to female mating status, and specifically, we predicted that males prefer females that are unmated. We conducted behavioural experiments in which males were given the choice of two females, one mated and one unmated. Contrary to our prediction, males did not mate preferentially with unmated females. However, copulation duration with unmated females was, on average, 24 times the length of copulation with mated females. While female K. tristis can reject mates, we did not observe any evidence of overt female choice during our trials. Females may gain additional benefits from mating multiply and may therefore not readily reject males. While our experiment cannot definitively disentangle female from male control over copulation duration, we suggest that males choose to invest more time in copula with unmated females, perhaps for paternity assurance, and that male mate assessment occurs during copulation rather than beforehand.
The Orthoptera are the most species‐rich group among the lower neopterans, and there is a tremendous amount of diversity in biology, ecology, and morphology in the order. This chapter presents a general overview of the systematics of the Orthoptera, as well as diversity in form and function in the order, and provides a brief synopsis of each of the known families. The most notable study was by Flook, who produced the first modern phylogeny of the Orthoptera, based on 31 in‐group taxa representing all major lineages and few ribosomal loci; they also reclassified some superfamilies. The monophyly of the Orthoptera has been supported by morphological and molecular data. The taxonomic classification of the Orthoptera has a tumultuous and complex history, as different taxonomists proposed conflicting classification schemes based on different character sets, such as fossil wing venation, internal organs, external morphology, and the male phallic complex.
Two species of the genus Sinipta, S. dalmani and S. hectorisperonii, exist in Uruguay, the latter reported here for the first time. Both species have quite similar morphological features and can only be distinguished by slight details, which have led to misidentifications in the past. Specimens from both taxa were studied in order to detect suitable characters to distinguish them and to clarify their taxonomic status, by integrating previously unstudied morphological, biological and molecular characteristics. Detailed structures of the stridulatory file of both sexes were studied and described for the first time, and the sounds produced by both species were studied and described on the basis of recordings made in captivity with an analog recorder. The signal was digitized and analyzed using Avisoft software. Three different types of sound were described for both species: calling song, courtship song and disturbance song. Oscillograms and frequency spectra were provided, and the physical characteristics of the sounds of interest for species identification were established. Finally, molecular studies showed that the two taxa are reciprocally monophyletic groups. The combined results of the analyses suggest that the two studied taxa are distinct species, and point to features suitable for their identification.
We investigated probable functions of the interacting genitalic components of a male and a female of the flightless grasshopper species Melanoplus rotundipennis (Scudder, 1878) (frozen rapidly during copulation) via correlative microscopy; in this case, by synergizing micro-computed tomography (micro-CT) with digital single lens reflex camera photography with focal stacking, and scanning electron microscopy. To assign probable functions, we combined imaging results with observations of live and museum specimens, and function hypotheses from previous studies, the majority of which focused on museum specimens with few investigating hypotheses in a physical framework of copulation. For both sexes, detailed descriptions are given for each of the observed genitalic and other reproductive system components, the majority of which are involved in copulation, and we assigned probable functions to these latter components. The correlative microscopy approach is effective for examining functional morphology in grasshoppers, so we suggest its use for other animals as well, especially when investigating body regions or events that are difficult to access and understand otherwise, as shown here with genitalia and copulation. J. Morphol., 2017.
Sounds of different kinds and intensities are produced by a number of species in all the main orders of insects; for general reviews see Alexander (1967), Dumortier (1963a, b and c), Frings and Frings (1958, 1960), Haskell (1961), Michelsen and Nocke (1974) and Bennet-Clark (1975). In some cases, the sound-producing organs are similar in the two sexes (as in many Coleoptera) but often they are confined to, or more strongly developed in, the male (e.g. most Orthoptera and Cicadidae). The biological significance of the sounds which are produced is not always clear but in some species they facilitate mating by attracting the sexes or stimulating the female and in other instances they may express sexual rivalry between males, subserve species-recognition (thus helping to keep members of the same species together), communicate warnings of danger or have a defensive function. A single species may make several sounds, each with its own function.
During the past decades considerable information has been gathered on the neuronal mechanisms underlying animal behaviour. Such progress has become possible since new stimulation and recording techniques have been developed, which often permit neurophysiological studies on unrestrained and normally behaving animals, therefore justifying the introduction of a new term — Neuroethology — for this field of research (reviews: Hoyle 1970, Usherwood and Newth 1975, Ewert 1976, Fentress 1976, Kandel 1976). The main goal of neuroethology is to reveal the functions and interactions of receptors, the central nervous system and effector organs which lead to the performance of behaviour. However, this attempt will remain incomplete if it does not include investigations on the genetics, the development and the phylogenetic origin of the neuronal mechanisms underlying behaviour.
The toothpick grasshopper, Achurum sumichrasti (Saussure), was collected in 12 pitcher plant bogs in long-leaf pine savannah associations in southeast Texas, from four south-central Texas coastal habitats and three west Texas localities. Comparisons of morphology, karyotypes and song of east Texas and west Texas populations of Achurum sumichrasti (Saussure) were made by the junior author. Food preferences and life histories were determined by the senior author from caged specimens brought back from their native habitats.
Acoustically conspicuous animals offer special opportunities for field biologists and students of reproductive behavior. Cicadas, crickets, katydids, and some grasshoppers are the birds and frogs of the insect world. Males loudly advertise their species, sexual readiness, and location. Biologists can use these acoustical displays to detect sibling species, to find local populations and individuals for study, and to determine seasonal, ecological, and geographical distributions. The “songs” can be recorded on magnetic tape and played back with high fidelity and analyzed and synthesized with devices that are simple and inexpensive compared to those needed to do research of equivalent quality with visual and olfactory displays. For ease of study only animals using long-range bioluminescent signals compete with acoustically displaying ones, and a much more limited array of species is involved (Lloyd, 1978).
The primary thermoregulatory function of body feathers and their movements is discussed and four general feather postures are distinguished : 1. S l e e k e d : The feathers are fully depressed against the body. This is characteristic of active birds, or birds ready for action, and streamlines them in flight and also reduces the insulating effect of the plumage. 2. R e l a x e d: The feathers are neither depressed nor erected. This is the usual state of birds when only moderately active, or inactive. 3. F l u f f e d : The feathers are partially erected giving the body a rounded appearance with smooth outlines. This increases the thickness of the insulating layer and helps to keep the bird warm. 4. R u f f l e d : The feathers are fully erected giving the bird a round but ragged appearance. The separating out of the individual feathers tends to destroy the insulating layer and a cooling effect results. It is observed in very hot birds which are inactive. The secondary signal function of body feathers is investigated. It is shown that in non-thwarting situations the mood of a bird may under certain conditions be conveyed to its fellows by its unspecialised feather postures, which therefore act as simple signals. In thwarting situations, the autonomic discharge, which accompanies the primary somatic response, is known to produce dramatic pilomotoric effects. It is suggested that this physiological relationship between thwarting and feather erection is the basis of all feather-posture displays (with the exception of the simple ones mentioned above which occur in non-thwarting situations). Not only generalised body fluffing or ruffling, but also the highly specialised body-feather displays are explicable in this way. Examples given include the evolution of crests, ruffs, chin-growths, throat-plumes, eartufts, flank-plumes, etc. It is stressed that a detailed study of the physio-ethology of thwarting would greatly assist in advancing the study of social signals in animals. The general problem of the evolution as signals of responses to thwarting situations is next considered. Three conditions of thwarting are recognised, namely: The absence of indispensible stimuli, following arousal; simple physical obstruction; and simultaneous arousal of incompatible tendencies. The Primary Response to thwarting is subdivided into a Somatic Response, which tends to adjust the animal to its external environment, and an Autonomic Response, which tends to adjust the internal environment of the animal to the requirements of this Somatic Response. Five types of Somatic Response are conceived:
1.The stridulation of the grasshoppers Stenobothrus lineatus, Omocestus virudulus, Chorthippus brunneus and Chorthippus parallelus was recorded on magnetic tape and analysed oscillographically and with an audio-spectrometer.2.The various types of songs are described and inter- and intra-specific differences as revealed by physical analysis are shown to reside almost completely in differing pulse repetition frequencies.3.The behaviour of adult mature males on stimulation with the normal song of their species is described. Normally this behaviour leads to group singing, with no locomotor movement, but males of all species except O. viridulus when segregated from their fellows for 24 hours or more show taxic locomotor reactions to the source of sound. The locomotor movements cease when a male joins a singing group.4.It is suggested that this reaction may be related to the distributional ecology of the insects.
Author Institution: Department of Zoology and Entomology, The Ohio State University, Columbus 10
The responses of single sense cells in the tympanal organ of a locust (Schistocerca gregaria Forskål) were examined by means of capillary microelectrodes. The sense cells in three of the four anatomical groups differ as to absolute sensitivity, latency time of maximum response, adaptation, and frequency response. Evidence is presented for the existence of a fifth auditory unit. The so-called ‘on’ response from the entire tympanal nerve is interpreted as an effect of the synchronization of the action potentials at the onset of sound, whereas the ‘tonic’ response is ascribed to a lack in synchronization. There is no evidence for the existence of specific ‘phasic’ and ‘tonic’ receptors in the tympanal organ. Previous arguments for the lack of frequency discrimination in Orthoptera are discussed, and it is argued that none of them are conclusive.
Duration of crepitation: u p to 5 seconds? COMMUNICATIVIC HEHAVlOR I N GRASSHOPPERS Duration of stridulation pulses: 0.08-0.13 secs Femur-shaking rate and # strokes/burst: 17.2-22.0/sec (3-5) n = 6 80°F Morgan Co
- Gillespie Co
Gillespie Co., Texas
Arphia xanthoptera
Crepitation rate:
23.l/sec undisturbed 88°F Hocking Co., Ohio
20.7-23.8 n = 5 undisturbed 70°F Lacledc Co., Mo.
Duration of crepitation: u p to 5 seconds?
COMMUNICATIVIC HEHAVlOR I N GRASSHOPPERS
Duration of stridulation pulses: 0.08-0.13 secs
Femur-shaking rate and # strokes/burst:
17.2-22.0/sec (3-5) n = 6 80°F Morgan Co., Tenn.
16.9-18.0
(5-8) n = 5 84
Chortophaga australior Crepitation rate: 73.3/sec disturbed 94°F Gulf Co
- Ripley Co
- Mo
Ripley Co., Mo.
Duration of pulses of stridulation: 0.05-0.07 secs Washtenaw Co., Mich.
Femur-shaking rate: 70/sec 93°F Washtenaw Co., Mich.
Femur-tipping duration: 0.2-0.3 secs n = 7 93°F Washtenaw Co., Mich.
Chortophaga australior
Crepitation rate:
73.3/sec disturbed 94°F Gulf Co., Fla.
Femur-shaking rate and # strokes/burst: 66.6/sec (11) 78°F Alachua Co
- Co
- Fla
/sec disturbed 94°F Alachua Co., Fla.
Femur-shaking rate and # strokes/burst:
66.6/sec (11) 78°F Alachua Co., Fla.
53.0
(10) 86
Male femur-shaking rate and # strokes/burst: 29.0-35.0/sec (14-19) n = 5 88°F Livingston Co
- Sutton Co
Sutton Co., Texas
Femur-tipping duration: 0.75 seconds 86°F Irnuris, Sonora, Mex.
Pardalophora apiculata
Stridulation rate:
between 6.5 and 20/sec 98°F Livingston Co., Mich.
Duration of vibratory stridulation:
0.55-1.25 secs n = 5 88°F Livingston Co., Mich.
Male femur-shaking rate and # strokes/burst:
29.0-35.0/sec (14-19) n = 5 88°F Livingston Co., Mich.
32.0-36.7/sec (13-25) n = 4 90°F Ozark Co., Mo.
Female femur-shaking Iatc:
40, 90/sec (presenting female) 90°F Ozark Co., Mo.
40.8
(presenting female) 88
San Luis Potosi, Mexico Interval between shaking and stridulation: 0.07-0.41 sccs rrrean = 0.20 n = 15 80°F Anderson Co., Texas Aggressive femur-shaking rate and # strokes/burst: 34.4/sec (15) 80°F Anderson Co
- Andcrson Co
Andcrson Co., Texas
35.1
(I6)
90°F 55 mi. W. Cd. Valles, San Luis Potosi, Mexico
Interval between shaking and stridulation:
0.07-0.41 sccs rrrean = 0.20 n = 15 80°F Anderson Co., Texas
Aggressive femur-shaking rate and # strokes/burst:
34.4/sec (15) 80°F Anderson Co., Texa5
33.3
(20) 90
Trimerotropis palliclipennis Crepitation rate, burst rate, and # snaps/burst: 55.5/sec 5.0/sec (4,5) 88°F Lorclsburg
- Anderson Co
Anderson Co., Texas
Femur-tipping duration:
0.6-1.0 secs n = 7 85°F Gulf Co., Fla.
0.7
87
Berrien Co., Mich.
Trimerotropis palliclipennis
Crepitation rate, burst rate, and # snaps/burst:
55.5/sec 5.0/sec (4,5) 88°F Lorclsburg, New Mexico
58.3-58.8 4.7-4.8 (4-6) 82°F Jeff Davis Co., Texas
57.1
3.8
(4-6)
73°F Lubbock, Texas
60.0
4.9
(4, 5) 72°F Warner Springs, Calif.
4 (4, 5) 86°F Las Aninias Co., Colo. 30.3 3.0 (5-7) 80°F Las Animas Co., Colo
- Colfax Co
Colfax Co., New Mexico
28.5
3.4
(4, 5) 86°F Las Aninias Co., Colo.
30.3
3.0
(5-7) 80°F Las Animas Co., Colo.
Femur-shaking ratc and # strokcs/burst:
23.5/sec (1 1) 90°F Pueblo Co., Colo.
Crepitation rate:
43.0/sec 86°F St. Clair Co., Micll.
44.5
86°F SL. Clair Co., Mich.
Keys for the identification of Colorado Orthoptvra
- C Alexander
ALEXANDER, C. 1941. Keys for the identification of Colorado Orthoptvra. Univ. Colorado
Studies, Series D, 1: 129-164.
Translatrd from Russian for National Science Foundation and the Acadcmy of Natural Sciences of Phila
- G Y Bei-Bienko
- , L Misiiciienko
BEI-BIENKO, G. Y., AND 1,. L. MISIICIIENKO. 1951. Locusts and g r a~~h o p p c r~
of the U.S.S.R.
and adjacent countries. Zool. Inst. U.S.S.R. Acad. Sci., Part 11, No. 40, 291 pp.
(Translatrd from Russian for National Science Foundation and the Acadcmy of
Natural Sciences of Phila. by Israel Program for Scientific Translations).
Orthoptcra of northeastern America
- Blatchi
- W S Ey
BLATCHI.EY, W. S. 1920. Orthoptcra of northeastern America. Nature Publishing CO.,
Indianapolis, 784 pp.
Current Problems in Animal Behaviour
- Bi
- A Fst
BI.FST, A. 1). 1961. T h e concept of 'ritualisation.' In: Thorpe, W. H. and 0. I. Zangwill
(ed.). Current Problems in Animal Behaviour. Cambridge Univ. Press, Cambridge, England, 122 pp.
- L Bruner
BRUNER, L. 1904. Biologia Centrali-Americana. Insccta. Orthoptera. Acrididae. Pp. 19-342.
-1922b. A list of the Orthoptera and Derrnaptera recorded fro111 British Columbia prior to the year 1922, with annotations
- E R Buckell
BUCKELL, E. R. 1920. Life-history notes on some species of Acrididae (Orthoptera) found
in British Columbia. Repi. ent. Soc. Ont. for 1919:53-61.
-----1922a. Notes on the ecological distribution of some Orthoptcra from the
Chilcotin district of British Columbia. Pmc. ent. Soc. Br. Columb., No. 18:
32-38.
-----1922b. A list of the Orthoptera and Derrnaptera recorded fro111 British Columbia prior to the year 1922, with annotations. Proc. ent. Soc. Br. Columb., No. 20:
9-41.
Notes on Orthoptera from Colorado
- A N Caudell
CAUDELL, A. N. 1903. Notes on Orthoptera from Colorado, New Mexico, Arizona, and
Texas, with descriptions of new species. Proc. U.S. natn. Mus., 26:775-809.
T h e grasshoppers of Oklahoma (Orthoptera: Acrididae)
- S Coppock
COPPOCK, S. 1962. T h e grasshoppers of Oklahoma (Orthoptera: Acrididae). P11.D. Thesis,
Oklahoma State University, May 1962, 220 pp.
O n locomotory niovements in birds and the intention rnovements dcrivcd from them
- A Daanje
DAANJE, A. 1950. O n locomotory niovements in birds and the intention rnovements
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1872. T h c expression of the emotions in man and aninials
- C Darwin
DARWIN, C. 1872. T h c expression of the emotions in man and aninials. Phoenix Rooks,
Univ. Chicago Press, Chicago and London, 1965, 372 pp.
African genera of Acridoidea
- T H E
T h e African genera of Acridoidea. Cambridge Univ. Press, Cambridge,
England, 579 pp.
Chorthi$f)t(s longicornis I.atr. (= fiarallrlus Zett.) und Clzorthippus m o n t a n~~s Charp. (hisller nach Finot als "longicornis Latr
- A Fader
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m o n t a n~~s Charp. (hisller nach Finot als "longicornis Latr." bezcichnet). Zool.
Anz., 81:l-24.
-----1953. Laut-und Gcbirdensprachc bei Insekten. Orthoptcra (Geradfliiglcr). I.
Mitt. st. Mus. Naturk. Stuttg., 198 pp.
Data on the ortliopteran farwisticc of eastern Pennsylvania and southern New Jersey
- I-I Fox
Fox, I-I. 1914. Data on the ortliopteran farwisticc of eastern Pennsylvania and southern
New Jersey. Proc. Ac;~tl. nat. Sci. Philad., 1914: 441-534.
T h c grasshoppers ant1 other Orth,optera of Iowa
- R C Fkoesciiner
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J. Sci. 29: 163-354.
Notes on certain genera of North American grasshoppers of tire subfamily Oedipodinae, with the descriptions of a new genus and species
- A R Gurney
GURNEY, A. R. 1940a. Notes on certain genera of North American grasshoppers of tire
subfamily Oedipodinae, with the descriptions of a new genus and species
(Orthoptera: Acrididae). Ent. Soc. Wash., 42:l-15.
-----1940b. A revision of the grasshoppers of the genus Orfih~clelln Giglio-Tos,
from Amcrica north of Mexico (Orthoptera; Acrididac). Enton~ologica am.,
20:85-157.
A synopsis of tlie North A~ncrican Acridinc grasshoppers of the genus-grollp Chrysochraontes (Orthoptera: Acrididae)
- A B Gurney
- H F Strohecker
- J R Hflfer
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A~ncrican Acridinc grasshoppers of the genus-grollp Chrysochraontes (Orthoptera: Acrididae). Trans. Amcr. ent. Soc., 89: 119-139.
Seasonal snrccssion, soil relations, numbers, and regional distribution of northeastern Texas acridians
- F B Isely
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----1936. Flight-Stridulation in Amcriran acridians. (Orthop.: Acritlidae). Ent.
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----A
1937. Seasonal snrccssion, soil relations, numbers, and regional distribution
of northeastern Texas acridians. Ecol. Monogr., 7:317-344.
-----1944. Corrclation bctwccn mandibular morphology and food specificity in
grasshoppers. Ann. ent. Soc. Am., 37:47-67.
Verhalte~lsbiologische Studicn an Feldheuschrccken
- W Jacobs
JACOBS, W. 1953. Verhalte~lsbiologische Studicn an Feldheuschrccken. Z. Tierpsychol.,
Suppl. I, pp. 1-228.
Mkthodes inhabituelles clc production d c son chcz les orthoptbrcs
- D K Kevan
- Mce
KEVAN, D. K. McE. 1954. Mkthodes inhabituelles clc production d c son chcz les orthoptbrcs. In: Busnel, R. G. (cd.). I.'Acoustique des Orthopteres. Inst. Nat. Rech.
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Contribution to the study of the sexual behavior of Schistocerra gregaria
- W Loner
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Vcrgleichende Bewegungsstudien an Anatinen
- K Lorenz
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Drosophila and the evolution of behavior
- A Manning
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Dcvelopmcnts in the study of animal communication
- P Marler
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