Pygmy Grasshoppers Research Articles

Oriental macropterous leaf-mimic pygmy grasshoppers-genera Oxyphyllum and Paraphyllum (Orthoptera: Tetrigidae) and their taxonomic assignment.

There are numerous pygmy grasshoppers (Tetrigidae) that exhibit leaf-like appearance. Leaf-mimic species can be found mainly in the subfamily Cladonotinae (tribes Cladonotini, Xerophyllini). Two leaf-mimic pygmy grasshopper species found in India, Pakistan (Oxyphyllum pennatum Hancock, 1909) and Borneo (Paraphyllum antennatum Hancock, 1913) are the only macropterous leaf-like species of Asia and were traditionally assigned to subfamily Cladonotinae. Our study present new records of these species, as well as updated descriptions. Oxyphyllum pennatum is recorded for the first time from Chhattisgarh (Central India), and furthermore, records from Pakistan were reviewed and confirmed. Paraphyllum antennatum is reported from a few localities in the mountains of Borneo (East Malaysia). New records found in online social media (Flickr, iNaturalist) were implemented. Brachypronotal specimens of P. antennatum are reported. After comparison with Cladonotini and Xerophyllini members, we conclude that Oxyphyllum and Paraphyllum should not be regarded Cladonotinae members. Traditional assignment was based on superficial resemblance to leaf-like Cladonotinae genera (e.g. Phyllotettix, Hymenotes, Holoarcus, Xerophyllum). However, comparison of morphological characters reveals that Oxyphyllum shares morphology with Tetriginae genera, while Paraphyllum shares morphology with Asian Metrodorinae. Here, we assign the genus Oxyphyllum to Tetriginae and the genus Paraphyllum to Metrodorinae.

New taxa of pygmy grasshoppers from Australia with notes on classification of the subfamily Batrachideinae (Orthoptera: Tetrigidae)

The pygmy grasshopper subfamily Batrachideinae consists of three tribes: Batrachideini Bolívar, 1887; Cassitettigini Yin, 1984, nom. resurr.; Bufonidini Hancock, 1907, nom. resurr., stat. nov. The Australian pygmy grasshoppers of the tribe Bufonidini are reviewed. Three new genera are described, namely Paraselina gen. nov. (type species Vingselina multifora Rehn, 1952), Anaselina gen. nov. (type species Vingselina minor Sjöstedt, 1921), and Selivinga gen. nov. (type species Selivinga tribulata sp. nov.). Four new combinations are proposed: Paraselina multifora (Rehn, 1952), comb. nov.; P. trituberculata (Sjöstedt, 1932), comb. nov.; P. brunneri (Bolívar, 1887), comb. nov.; Anaselina minor (Sjöstedt, 1921), comb. nov. A key to Australian genera of Batrachideinae is provided.

Contrasting patterns of neutral and functional genetic diversity in stable and disturbed environments.

Genetic structure among and diversity within natural populations is influenced by a combination of ecological and evolutionary processes. These processes can differently influence neutral and functional genetic diversity and also vary according to environmental settings. To investigate the roles of interacting processes as drivers of population‐level genetic diversity in the wild, we compared neutral and functional structure and diversity between 20 Tetrix undulata pygmy grasshopper populations in disturbed and stable habitats. Genetic differentiation was evident among the different populations, but there was no genetic separation between stable and disturbed environments. The incidence of long‐winged phenotypes was higher in disturbed habitats, indicating that these populations were recently established by flight‐capable colonizers. Color morph diversity and dispersion of outlier genetic diversity, estimated using AFLP markers, were higher in disturbed than in stable environments, likely reflecting that color polymorphism and variation in other functionally important traits increase establishment success. Neutral genetic diversity estimated using AFLP markers was lower in disturbed habitats, indicating stronger eroding effects on neutral diversity of genetic drift associated with founding events in disturbed compared to stable habitats. Functional diversity and neutral diversity were negatively correlated across populations, highlighting the utility of outlier loci in genetics studies and reinforcing that estimates of genetic diversity based on neutral markers do not infer evolutionary potential and the ability of populations and species to cope with environmental change.

An annotated catalogue of the pygmy grasshoppers of the tribe Scelimenini Bolívar, 1887 (Orthoptera: Tetrigidae) with two new Scelimena species fromthe Malay Peninsula and Sumatra.

Pygmy grasshoppers (Tetrigidae) are a speciose group of complicated taxonomy, with many species requiring clarification on their species boundaries, and more still awaiting discovery. Two new species of Scelimena Serville, 1838 are described: (1) from S. discalis species group S. gombakensis sp. nov. and (2) from S. hexodon species group Scelimena marta sp. nov. Catalogue of Scelimenini genera (15) and species (100) is presented and taxonomy and biogeography of the tribe are discussed. New and resurrected combinations are: Falconius becvari (Buzzetti Devriese, 2008) comb. nov. (of Gavialidium becvari), Gavialidium carli Hebard, 1930 comb. resurr. (of Bidentatettix carli), Indoscelimena india (Hancock, 1907) comb. nov. (of Scelimena india), Paragavialidium nodiferum (Walker, 1871) comb. nov. (of Platygavialidium nodiferum), Platygavialidium productum (Walker, 1871) comb. nov. (of Gavialidium productum), Scelimena hexodon (Haan, 1843) comb. resurr. (of Hexocera hexodon), Scelimena rosacea (Hancock, 1915) comb. resurr. (of Amphibotettix rosaceus), Tegotettix bufocrocodil (Storozhenko Dawwrueng, 2015) comb. nov. (of Gavialidium bufocrocodil). New synonyms are: Gavialidium phangensum Mahmood, Idris Salmah, 2007 syn. nov. (of Eufalconius pendleburyi), Gavialidium philippinum Bolívar, 1887 syn. nov. (of Platygavialidium productum comb. nov.), Hexocera Hancock, 1915 syn. nov. (of Scelimena), Paracriotettix Liang, 2002 syn. nov. (of Scelimena), Paracriotettix zhengi Liang, 2002 syn. nov. (of Scelimena melli), Scelimena mellioides Deng, 2016 syn. nov. (of Scelimena melli), Scelimena wuyishanensis Deng, 2016 syn. nov. (of Platygavialidium sinicum). The genus Scelimena is divided into six species groups. Eucriotettix neesoon Tan Storozhenko nom. nov. is new name for homonym Eucriotettix guentheri Tan Storozhenko, 2017. Finally, a tabular key to 15 Scelimenini genera, based on 16 morphological characters, is presented.

On the role of sex differences for evolution in heterogeneous and changing fitness landscapes: insights from pygmy grasshoppers.

Much research has been devoted to study evolution of local adaptations by natural selection, and to explore the roles of neutral processes and developmental plasticity for patterns of diversity among individuals, populations and species. Some aspects, such as evolution of adaptive variation in phenotypic traits in stable environments, and the role of plasticity in predictable changing environments, are well understood. Other aspects, such as the role of sex differences for evolution in spatially heterogeneous and temporally changing environments and dynamic fitness landscapes, remain elusive. An increased understanding of evolution requires that sex differences in development, physiology, morphology, life-history and behaviours are more broadly considered. Studies of selection should take into consideration that the relationships linking phenotypes to fitness may vary not only according to environmental conditions but also differ between males and females. Such opposing selection, sex-by-environment interaction effects of selection and sex-specific developmental plasticity can have consequences for population differentiation, local adaptations and for the dynamics of polymorphisms. Integrating sex differences in analytical frameworks and population comparisons can therefore illuminate neglected evolutionary drivers and reconcile unexpected patterns. Here, I illustrate these issues using empirical examples from over 20 years of research on colour polymorphic Tetrix subulata and Tetrix undulata pygmy grasshoppers, and summarize findings from observational field studies, manipulation experiments, common garden breeding experiments and population genetics studies.This article is part of the theme issue ‘Linking local adaptation with the evolution of sex differences’.

To the knowledge of pygmy grasshoppers (Orthoptera: Tetrigidae) from Cambodia

To the knowledge of pygmy grasshoppers (Orthoptera: Tetrigidae) from Cambodia

Seasonal differences in body mass and circulating metabolites in a wing‐dimorphic pygmy grasshopper: implications for life history?

1. Functional wing polymorphism in insects is an intriguing topic, especially with respect to the adaptive advantage of each wing morph. The common pygmy grasshopper in Germany, Tetrix subulata, displays wing polymorphism skewed towards macropterous (LW) individuals capable of flight. Furthermore, T. subulata is known to undergo adult diapause in winter and reproduce in spring.2. Morphometric and biochemical parameters were examined in field‐collected grasshoppers during autumn and spring to obtain a ‘snapshot’ from the same/one cohort of grasshoppers in the wild.3. Flight muscles are largely reduced in brachypterous (SW) specimens, whereas they are well developed in LW individuals. Body mass measurements indicated gain in female T. subulata in spring, especially in LW morphs, which could be attributed to increased reproductive activity (egg production).4. Metabolic fuel in haemolymph is differentially distributed in autumn: the concentration of lipids is highest in males, while carbohydrates are most abundant in LW specimens. The metabolic data imply that dispersal in T. subulata is predominantly in autumn, by flight in the case of LW specimens and by hopping/walking in males.5. The season seems to be an important factor for the reproductive versus dispersal trade‐off in this species. Moreover, this study shows that morphological differences in T. subulata individuals are reflected in physiological differences that may ultimately affect behaviour and ecology.

Nymph inadvertently described as new species for a fourth time? On the identity of Euscelimena hardi (Tetrigidae: Scelimeninae) with general remarks on the identification of pygmy grasshopper nymphs.

Euscelimena hardi Gupta et Chandra, 2018 was described earlier this year from Chhattisgarh (India) as a unique species within the genus, characterized in small size, long pronotal projections, and a lack of wings. In this commentary we provide evidence that E. hardi is a nymph of E. harpago (Serville, 1838), hence a new synonymy is proposed: Euscelimena harpago (Serville, 1838) = Euscelimena hardi Gupta et Chandra, 2018 syn. nov. Since this is the fourth time a nymph of Euscelimena Günther, 1938 has been described as a new species, we present general remarks on how to distinguish pygmy grasshopper nymphs and adults. Presence of a groove on the dorsal margin of the hind femora, directly before the knee (separating antegenicular and genicular teeth) is the only definite feature to separate nymphs from adults in brachypronotal or micropronotal wingless species. In macropterous species, morphology of tegmina and alae is helpful to separate nymphs from adults, nymphs having alae rich in radial venation in front of tegmina, while in adults tegmina are in front of developed alae. When dealing with a potential new species, researchers should firstly take into account known pygmy grasshoppers fauna of the studied area via Orthoptera Species File Online database Complex search (with imput Scope of search: Tetrigidae, Place name from geographic hierarchy: continent or country name).

A new tribe of tarsonemid mites (Trombidiformes: Heterostigmatina) parasitic on tetrigid grasshoppers (Orthoptera).

Podotarsonemini tribe nov. (Tarsonemidae: Acarapinae), with one nominate genus Podotarsonemus gen. nov., is proposed for seven new species of tarsonemid mites collected from the hindwings of pygmy grasshoppers of the family Tetrigidae (Orthoptera). The new tribe is placed within the tarsonemid subfamily Acarapinae on the basis of several morphological synapomorphies, as well as on parasitism of insects. In light of this new tribe, a revised description of the Acarapinae is provided, as are full tribal, generic and species descriptions for the Podotarsonemini, and a key to species. In view of Podapolipidae, the sister family of Tarsonemidae, consisting entirely of obligatory parasitic mites, the distinction between the two constituent sister families of Tarsonemoidea as well as the ancestral feeding habits of that superfamily are considered. These mites and their host grasshoppers were collected from Bolivia, Ecuador, Honduras, southern India, Japan, Papua New Guinea, South Africa and two localities in north-eastern Australia. This distribution suggests that the Podotarsonemini are an ancient lineage of Tarsonemidae that radiated on the Gondwanan Tetrigidae during the Jurassic.

Morphometrics reveal correlation between morphology and bioclimatic factors and population mixture in Tetrix japonica (Orthoptera: Tetrigidae)

AbstractThe morphology of an organism is limited by genetic and environmental factors, and the precise mechanism is inconsistent between species. Tetrix japonica Bolivar, 1987, is a widely distributed pygmy grasshopper in East Asia. However, the population clustering and relationships between the morphology and bioclimatic factors have not been previously investigated. Here, 32 geographic populations were sampled from China, and morphometrics and multiple statistical analyses were applied to detect the population clustering and relationship between the morphology and bioclimatic factors. The results suggested that T. japonica with females are significantly bigger than males in the eight morphological traits. The 32 populations do not obviously cluster according to the natural geographic area. The body sizes of females are mainly related to the lowest temperature and precipitation; by contrast, males only have a significant relationship with the lowest temperature. The forewing size is significantly related to the maximum precipitation. Furthermore, the Mantel test showed that the morphological size variation of females has a weak positive correlation to geographic distance, but is insignificant in males. It was concluded that Chinese geographic populations of T. japonica mixed and that the size of the morphological structure is limited by bioclimatic factors.

THE ADULT MALE OF THE LITTLE-KNOWN PYGMY GRASSHOPPER ARMASIUS IBERIANUS PEREZ-GELABERT & YONG, 2014 (ORTHOPTERA: TETRIGIDAE: CLADONOTINAE)

An adult male of the Cuban endemic pygmy grasshopper Armasius iberianus Perez-Gelabert & Yong, 2014 is herein described. It was collected on the El Toldo High Plateau and represents the first record of this monotypic genus from Holguín Province. As a result, diagnosis of the species is updated and further data on its morphology and morphometry are given. The description is supported by a thorough illustrative complement, which includes a precise map and color photographs of habitus, main morphological diagnostic characters and habitat.

Ground dwelling pygmy grasshoppers (Orthoptera: Tetrigidae) in Southeast Asian tropical freshwater swamp forest prefer wet microhabitats

Ground dwelling pygmy grasshoppers (Orthoptera: Tetrigidae) in Southeast Asian tropical freshwater swamp forest prefer wet microhabitats

The importance of validated alpha taxonomy for phylogenetic and DNA barcoding studies: a comment on species identification of pygmy grasshoppers (Orthoptera, Tetrigidae).

In a recently published paper on colour polymorphism in a Pygmy grasshopper from China (Zhao et al 2016) an unidentified Paratettix sp. was misidentified as Tetrix bolivari. This case highlights the need for correct species identification and provides an opportunity to recommend some aspects of Good Taxonomic Practice (GTP) in Tetrigidae to reduce the number of erroneous identifications.

The role of dispersal for genetic and phenotypic variation: insights from comparisons of sympatric pygmy grasshoppers

Patterns of genetic and phenotypic variation within and among populations are influenced by a complex interplay of ecological and evolutionary processes. Theory posits that gene flow should increas ...

Integrative taxonomy reveals two sympatric species of the genus Eucriotettix Hebard, 1930 (Orthoptera: Tetrigidae).

Limno-terrestrial pygmy grasshoppers are mostly recognised using morphology but characters used are not always clear-cut for species identification. From a faunistic survey in Singapore, Eucriotettix were collected and two new species are identified and described here: E. guentheri sp. nov. and E. simulans sp. nov. This was done by iteratively incorporating morphological, molecular, morphometric, and ecological data. Mitochondrial COI trees, COI cluster analysis, microhabitat preference, and morphometric analyses provide strong enough indirect evidence that the specimens comprise a pair of species. A key to species of Eucriotettix from Singapore is also provided.

Population genetic structure, differentiation, and diversity in Tetrix subulata pygmy grasshoppers: roles of population size and immigration.

Genetic diversity within and among populations and species is influenced by complex demographic and evolutionary processes. Despite extensive research, there is no consensus regarding how landscape structure, spatial distribution, gene flow, and population dynamics impact genetic composition of natural populations. Here, we used amplified fragment length polymorphisms (AFLPs) to investigate effects of population size, geographic isolation, immigration, and gene flow on genetic structure, divergence, and diversity in populations of Tetrix subulata pygmy grasshoppers (Orthoptera: Tetrigidae) from 20 sampling locations in southern Sweden. Analyses of 1564 AFLP markers revealed low to moderate levels of genetic diversity (PPL = 59.5–90.1; Hj = 0.23–0.32) within and significant divergence among sampling localities. This suggests that evolution of functional traits in response to divergent selection is possible and that gene flow is restricted. Genetic diversity increased with population size and with increasing proportion of long‐winged phenotypes (a proxy of recent immigration) across populations on the island of Öland, but not on the mainland. Our data further suggested that the open water separating Öland from the mainland acts as a dispersal barrier that restricts migration and leads to genetic divergence among regions. Isolation by distance was evident for short interpopulation distances on the mainland, but gradually disappeared as populations separated by longer distances were included. Results illustrate that integrating ecological and molecular data is key to identifying drivers of population genetic structure in natural populations. Our findings also underscore the importance of landscape structure and spatial sampling scheme for conclusions regarding the role of gene flow and isolation by distance.

Characterization and analysis of a de novo transcriptome from the pygmy grasshopper Tetrix japonica.

The pygmy grasshopper Tetrix japonica is a common insect distributed throughout the world, and it has the potential for use in studies of body colour polymorphism, genomics and the biology of Tetrigoidea (Insecta: Orthoptera). However, limited biological information is available for this insect. Here, we conducted a de novo transcriptome study of adult and larval T.japonica to provide a better understanding of its gene expression and develop genomic resources for future work. We sequenced and explored the characteristics of the de novo transcriptome of T.japonica using Illumina HiSeq 2000 platform. A total of 107608206 paired-end clean reads were assembled into 61141 unigenes using the trinity software; the mean unigene size was 771bp, and the N50 length was 1238bp. A total of 29225 unigenes were functionally annotated to the NCBI nonredundant protein sequences (Nr), NCBI nonredundant nucleotide sequences (Nt), a manually annotated and reviewed protein sequence database (Swiss-Prot), Gene Ontology (GO) and Kyoto Encyclopedia of Genes and Genomes (KEGG) databases. A large number of putative genes that are potentially involved in pigment pathways, juvenile hormone (JH) metabolism and signalling pathways were identified in the T.japonica transcriptome. Additionally, 165769 and 156796 putative single nucleotide polymorphisms occurred in the adult and larvae transcriptomes, respectively, and a total of 3162 simple sequence repeats were detected in this assembly. This comprehensive transcriptomic data for T.japonica will provide a usable resource for gene predictions, signalling pathway investigations and molecular marker development for this species and other pygmy grasshoppers.

Prey Luring Coloration of A Nocturnal Semi‐Aquatic Predator

AbstractBody coloration serves a variety of purposes in animals. Diurnal and nocturnal predators such as spiders may use their body coloration to lure prey. We predicted here that the white patches on the forelegs on females of the nocturnal semi‐aquatic spider Dolomedes raptor lure prey, explaining why they are primarily displayed when the spider forages along the water edge. To test our prediction, we developed a color vision model assessing whether the patches are visible to pygmy grasshoppers, the spider's primary prey. We conducted a field experiment using cardboard dummies that resemble D. raptor in size, shape, and color, but with half of them lacking leg patches, and we staged interactions between pygmy grasshoppers and D. raptor with and without leg patches in a greenhouse. We found the white patches to be visible to grasshoppers. The dummies with white patches attracted more grasshopper prey than the dummies without the patches. Moreover, grasshoppers were more attracted to spiders when their white patches were present. Our results supported the hypothesis that the white patches of D. raptor lure prey. Our findings, nevertheless, could not be explained as the spider's body coloration acting as a sensory trap but it should not be ruled out. More studies on a wider range of predators and prey will give more meaningful insights into the co‐evolution of predatory lures and prey sensory modalities.

DNA barcoding reveals polymorphism in the pygmy grasshopper Tetrix bolivari (Orthoptera, Tetrigidae).

Many pygmy grasshopper species exhibit colour-marking polymorphism. However, this polymorphism in some species, such as Tetrix bolivari, is almost unknown. The aim of this work is to identify using DNA barcoding the colour-marking polymorphic morphs of this pygmy grasshopper species collected from both grass and sand microhabitats. Analysis by NJ clustering and pairwise distances indicated that all specimens collected showing colour-marking polymorphism are species of Tetrix bolivari. Haplotype network construction showed ten different haplotypes from a total of 57 Tetrix bolivari individuals with H1(82.5%) being the most common type and it also displayed low divergence within Tetrix bolivari population. The haplotype analyses were consistent with the NJ clustering. Our field census showed the frequency of Tetrix bolivari morphs differed significantly, with the rank order of morphs (from high to low) typeA1, type B1, type A2, type A3, type A4, type A5, type A6, type A7, type B2, type B3, and type B4. The most common type A morphs were without contrasting markings, while the rarer type B morphs have contrasting white markings. We suggest that type B morphs have greater camouflage effects against natural backgrounds such as grass or sand than type A morphs. Both our field census and haplotype analysis revealed that type A has higher frequency and more haplotypes than type B.

Discovery of a new species and key to all known species of the genus Aalatettix Zheng & Mao (Orthoptera, Tetrigoidea, Tetrigidae) from China.

The genus Aalatettix Zheng & Mao, 2002 (Orthoptera, Tetrigoidea, Tetrigidae) contains 9 valid species in China (Zheng & Mao, 2002; Zheng, Cao & Chen, 2011; Zheng, Lin & Zhang, 2013; Zheng, 2014a; Zheng, 2014b; Zheng & Lin, 2015; Zheng, Shi & Yin, 2015). During the identification of pygmy grasshopper specimens collected from Taiwan, China in 1998, a new species of the genus Aalatettix Zheng & Mao, 2002 was found and described below. A key to all known species of the genus is given in this paper. The type specimens are deposited in the National Museum of Natural Science, Taichung, Taiwan, China.