Cicada Species Research Articles

Two new genera and five new species of Mugadina-like small grass cicadas (Hemiptera: Cicadidae: Cicadettini) from Central and Eastern Australia: comparative morphology, songs, behaviour and distributions.

Moulds (2012) established the genus Mugadina for two small cicadas, M. marshalli (Distant) and M. emma (Goding and Froggatt), both grass inhabiting species known from Queensland and New South Wales. Both species are notable for their relatively simple 'ticking' songs. Moulds further noted that there were at least two superficially similar genera of cicadas, but each with different genitalia. This paper describes two new genera of small (9-15 mm body lengths) and distinctive grass cicadas with genitalia that are very similar to those of Mugadina, but possess clear morphological, colour and calling song differences. The new genera are: Heremusina n. gen. with two known species namely H. udeoecetes n. sp. and H. pipatio n. sp.; the second new genus is Xeropsalta n. gen., containing four known species, X. thomsoni n. sp., X. aridula n. sp., X. rattrayi n. sp., and X. festiva n. comb. Heremusina n. gen. species are described from the Alice Springs area of Northern Territory and the Cloncurry area of northwest Queensland, from arid to semi arid habitats. The Xeropsalta n. gen. species are described from western, southwest and central Queensland, and from the Simpson and Strzelecki Deserts in northeastern South Australia and northwestern New South Wales, respectively, all locations in very arid to arid habitats, but close to seasonal (often irregular) rivers and lakes. X. festiva n. comb. occurs in semi arid habitats in southern and southeastern Australia. Detailed taxonomic descriptions are provided of the new species, together with distributions, habitats, and the calling songs. The Heremusina species emit songs with short repetitive buzzing echemes, the echeme durations differing between each species. The Xeropsalta songs are notable for their complexity, containing multiple elements with rapid changes of amplitudes and temporal structures, rather atypical of the songs of most small grass dwelling cicadas. Detailed song structures distinguishing each of the species are illustrated and interpreted in each case in light of their respective taxonomic status.

Cicada parasitic moths from China (Lepidoptera: Epipyropidae): morphology, identity, biology, and biogeography

Species of the family Epipyropidae are infrequently collected due to their ectoparasitic lifestyle and have therefore received little study. Based on our morphological and molecular analyses, all specimens found parasitizing different species of cicadas in China belong to one species, Epipomponia nawai (Dyar). The species Epipomponia oncotympana Yang (nom. invalid.) was found to be identical with E. nawai. The morphology of the egg, larva, pupa, and adult of E. nawai is described, including the ultrastructure of the egg and larva. Seven types of sensilla and their distribution pattern on the antennae of both females and males of E. nawai are examined using scanning electron microscopy, resulting in a better understanding of sexual dimorphism in this species. Population differentiation and phylogenetic relationships amongst populations of E. nawai occurring in China, the Korean Peninsula, and Japan are analysed. The intraspecific genetic distances suggest a low genetic differentiation which is also consistent with the minor morphological variations. The biogeographic analysis reveals that the Qinling Mountain Range, which is geographically the border of the subtropical and temperate zones, is presumably one of the most important geographic barriers to the gene flow of E. nawai. Our molecular phylogenetic analysis based on the COI gene of E. nawai indicates that frequent gene flow might have occurred in the China mainland, Taiwan and Japan islands, and possibly also the Ryukyu Islands after the Quaternary ice ages. In addition, the biology and behaviour of both larvae and adults of E. nawai were also observed, and results indicate the possibility of bisexual reproduction in this moth. Our results increase the understanding of the morphology, diversity, distribution, and biology of this unusual moth, and should aid future studies on patterns of population differentiation, biogeography, and reproductive behaviour in other epipyropids.

Urban heat island effect on cicada densities in metropolitan Seoul.

BackgroundUrban heat island (UHI) effect, the ubiquitous consequence of urbanization, is considered to play a major role in population expansion of numerous insects. Cryptotympana atrata and Hyalessa fuscata are the most abundant cicada species in the Korean Peninsula, where their population densities are higher in urban than in rural areas. We predicted a positive relationship between the UHI intensities and population densities of these two cicada species in metropolitan Seoul.MethodsTo test this prediction, enumeration surveys of cicada exuviae densities were conducted in 36 localities located within and in the vicinity of metropolitan Seoul. Samples were collected in two consecutive periods from July to August 2015. The abundance of each species was estimated by two resource-weighted densities, one based on the total geographic area, and the other on the total number of trees. Multiple linear regression analyses were performed to identify factors critical for the prevalence of cicada species in the urban habitat.ResultsC. atrata and H. fuscata were major constituents of cicada species composition collected across all localities. Minimum temperature and sampling period were significant factors contributing to the variation in densities of both species, whereas other environmental factors related to urbanization were not significant. More cicada exuviae were collected in the second rather than in the first samplings, which matched the phenological pattern of cicadas in metropolitan Seoul. Cicada population densities increased measurably with the increase in temperature. Age of residential complex also exhibited a significantly positive correlation to H. fuscata densities, but not to C. atrata densities.DiscussionEffects of temperature on cicada densities have been discerned from other environmental factors, as cicada densities increased measurably in tandem with elevated temperature. Several mechanisms may contribute to the abundance of cicadas in urban environments, such as higher fecundity of females, lower mortality rate of instars, decline in host plant quality, and local adaptation of organisms, but none of them were tested in the current study.ConclusionsIn sum, results of the enumeration surveys of cicada exuviae support the hypothesis that the UHI effect underlies the population expansion of cicadas in metropolitan Seoul. Nevertheless, the underlying mechanisms for this remain untested.

A New Species of Carineta Amyot & Audinet-Serville, 1843 (Hemiptera: Cicadoidea: Cicadidae) from Martinique

Abstract. A new Caribbean cicada Carineta acclivis Sanborn, new species is described and illustrated. This is the second species of Carineta Amyot & Audinet-Serville, 1843 described from Martinique and the Lesser Antilles. Carineta martiniquensis Davis, 1934 is illustrated and distinguished from the new species. The adults of both cicada species on Martinique are active in forest habitats and have extended emergence periods during the year but appear to be separated by an altitudinal gradient.

Idiosyncratic Genome Degradation in a Bacterial Endosymbiont of Periodical Cicadas

When a free-living bacterium transitions to a host-beneficial endosymbiotic lifestyle, it almost invariably loses a large fraction of its genome [1, 2]. Theresulting small genomes often become stable in size, structure, and coding capacity [3-5], as exemplified by Sulcia muelleri, a nutritional endosymbiont of cicadas. Sulcia's partner endosymbiont, Hodgkinia cicadicola, similarly remains co-linear in some cicadas diverged by millions of years [6, 7]. But in the long-lived periodical cicada Magicicada tredecim, the Hodgkinia genome has split into dozens of tiny, gene-sparse circles that sometimes reside in distinct Hodgkinia cells [8]. Previous datasuggested that all other Magicicada species harbor complex Hodgkinia populations, but the timing, number of origins, and outcomes of the splitting process were unknown. Here, by sequencing Hodgkinia metagenomes from the remaining six Magicicada and two sister species, we show that each Magicicada species harbors Hodgkinia populations of at least 20 genomic circles. We find little synteny among the 256 Hodgkinia circles analyzed except between the most closely related cicada species. Gene phylogenies show multiple Hodgkinia lineages in the common ancestor of Magicicada and its closest known relatives but that most splitting has occurred within Magicicada and has given rise to highly variable Hodgkinia gene dosages among species. These data show that Hodgkinia genome degradation has proceeded down different paths in different Magicicada species and support a model of genomic degradation that is stochastic in outcome and nonadaptive for the host. These patterns mirror the genomic instability seen in some mitochondria.

A new species of Platypleura Amyot & Audinet-Serville, 1843 (Hemiptera: Cicadidae: Cicadinae) from the Eastern Ghats of Andhra Pradesh, India

We here describe a new cicada species, Platypleura poorvachala sp. nov., (Hemiptera: Cicadidae: Cicadinae) from the Eastern Ghats of India. The combination of following characters distinguishes this species from its platypleurine relatives: (a) transverse continuous infuscation along the nodal line and in the central area of the forewings, and (b) opaque hindwings, which are indochine-coloured with two prominent sub-distal black bands. We illustrate the male and female types and the male genitalia, and present a photographic plate for comparison of this species with other Indian and Southeast Asian Platypleura species with respect to its morphology and distribution.

Exploring the Role of Habitat on the Wettability of Cicada Wings.

Evolutionary pressure has pushed many extant species to develop micro/nanostructures that can significantly affect wettability and enable functionalities such as droplet jumping, self-cleaning, antifogging, antimicrobial, and antireflectivity. In particular, significant effort is underway to understand the insect wing surface structure to establish rational design tools for the development of novel engineered materials. Most studies, however, have focused on superhydrophobic wings obtained from a single insect species, in particular, the Psaltoda claripennis cicada. Here, we investigate the relationship between the spatially dependent wing wettability, topology, and droplet jumping behavior of multiple cicada species and their habitat, lifecycle, and interspecies relatedness. We focus on cicada wings of four different species: Neotibicen pruinosus, N. tibicen, Megatibicen dorsatus, and Magicicada septendecim and take a comparative approach. Using spatially resolved microgoniometry, scanning electron microscopy, atomic force microscopy, and high speed optical microscopy, we show that within cicada species, the wettability of wings is spatially homogeneous across wing cells. All four species were shown to have truncated conical pillars with widely varying length scales ranging from 50 to 400 nm in height. Comparison of the wettability revealed three cicada species with wings that are superhydrophobic (>150°) with low contact angle hysteresis (<5°), resulting in stable droplet jumping behavior. The fourth, more distantly related species (Ma. septendecim) showed only moderate hydrophobic behavior, eliminating some of the beneficial surface functional aspects for this cicada. Correlation between cicada habitat and wing wettability yielded little connection as wetter, swampy environments do not necessarily equate to higher measured wing hydrophobicity. The results, however, do point to species relatedness and reproductive strategy as a closer proxy for predicting wettability and surface structure and resultant enhanced wing surface functionality. This work not only elucidates the differences between inter- and intraspecies cicada wing topology, wettability, and water shedding behavior but also enables the development of rational design tools for the manufacture of artificial surfaces for energy and water applications.

Bacterial diversity of bacteriomes and organs of reproductive, digestive and excretory systems in two cicada species (Hemiptera: Cicadidae).

Cicadas form intimate symbioses with bacteria to obtain nutrients that are scarce in the xylem fluid they feed on. The obligate symbionts in cicadas are purportedly confined to specialized bacteriomes, but knowledge of bacterial communities associated with cicadas is limited. Bacterial communities in the bacteriomes and organs of reproductive, digestive and excretory systems of two cicada species (Platypleura kaempferi and Meimuna mongolica) were investigated using different methods, and the bacterial diversity and distribution patterns of dominant bacteria in different tissues were compared. Within each species, the bacterial communities of testes are significantly different from those of bacteriomes and ovaries. The dominant endosymbiont Candidatus Sulcia muelleri is found not only in the bacteriomes and reproductive organs, but also in the “filter chamber + conical segment” of both species. The transmission mode of this endosymbiont in the alimentary canal and its effect on physiological processes merits further study. A novel bacterium of Rhizobiales, showing ~80% similarity to Candidatus Hodgkinia cicadicola, is dominant in the bacteriomes and ovaries of P. kaempferi. Given that the genome of H. cicadicola exhibits rapid sequence evolution, it is possible that this novel bacterium is a related endosymbiont with beneficial trophic functions similar to that of H. cicadicola in some other cicadas. Failure to detect H. cicadicola in M. mongolica suggests that it has been subsequently replaced by another bacterium, a yeast or gut microbiota which compensates for the loss of H. cicadicola. The distribution of this novel Rhizobiales species in other cicadas and its identification require further investigation to help establish the definition of the bacterial genus Candidatus Hodgkinia and to provide more information on sequence divergence of related endosymbionts of cicadas. Our results highlight the complex bacterial communities of cicadas, and are informative for further studies of the interactions and co-evolution of insect-microbial symbioses in Cicadoidea.

A new genus and new species of Cicadettini (Hemiptera: Cicadidae: Cicadettinae) from Pakistan.

Pakistan contains parts of the Indomalayan and Palaearctic ecozones. As a result, there is a diversity of habitats that has led to diversity in the cicada fauna. We reviewed the Pakistani cicada fauna (Ahmed and Sanborn 2010) but new records and new species continue to be found (Ahmed et al. 2010; 2012; 2013; 2014). This work continues to expand the knowledge of the Pakistani cicada fauna with the description of a new genus and new species of cicada provided to the second author from his contacts.

Morphology, songs and genetics identify two new cicada species from Morocco: Tettigettalna afroamissa sp. nov. and Berberigetta dimelodica gen. nov. & sp. nov. (Hemiptera: Cicadettini).

Morocco has been the subject of very few expeditions on the last century with the objective of studying small cicadas. In the summer of 2014 an expedition was carried out to Morocco to update our knowledge with acoustic recordings and genetic data of these poorly known species. We describe here two new small-sized cicadas that could not be directly assigned to any species of North African cicadas: Tettigettalna afroamissa sp. nov. and Berberigetta dimelodica gen. nov. & sp. nov. In respect to T. afroamissa it is the first species of the genus to be found outside Europe and we frame this taxon within the evolutionary history of the genus. Acoustic analysis of this species allows us to confidently separate T. afroamissa from its congeners. With B. dimelodica, a small species showing a remarkable calling song characterized by an abrupt frequency modulation, a new genus had to be erected. Bayesian inference and maximum likelihood phylogenetic analyses with DNA-barcode sequences of Cytochrome C Oxidase 1 support the monophyly of both species, their distinctness and revealed genetic structure within B. dimelodica. Alongside the descriptions we also provide GPS coordinates of collection points, distributions and habitat preferences.

How the Aridification of Australia Structured the Biogeography and Influenced the Diversification of a Large Lineage of Australian Cicadas.

Over the last 30 million years, Australia's landscape has undergone dramatic cooling and drying due to the establishment of the Antarctic Circumpolar Current and change in global CO$_{2}$ levels. Studies have shown that many Australian organisms went extinct during these major cooling events, while others experienced adaptive radiations and increases in diversification rates as a result of exploiting new niches in the arid zone. Despite the many studies on diversification and biogeography in Australia, few have been continent-wide and none have focused on a group of organisms adapted to feeding on plants. We studied 162 species of cicadas in the Australian Pauropsalta complex, a large generic lineage within the tribe Cicadettini. We asked whether there were changes in the diversification rate of Pauropsalta over time and if so: 1) which clades were associated with the rate change? 2) did timing of rate shifts correspond to known periods of dramatic historical climate change, 3) did increases in diversification rate along select lineages correspond to adaptive radiations with movement into the arid zone? To address these questions, we estimated a molecular phylogeny of the Pauropsalta complex using ${\sim}$5300 bp of nucleotide sequence data distributed among five loci (one mtDNA locus and four nDNA loci). We found that this large group of cicadas did not diversify at a constant rate as they spread through Australia; instead the signature of decreasing diversification rate changed roughly around the time of the expansion of the east Antarctic ice sheets ${\sim}$16 Ma and the glaciation of the northern hemisphere ${\sim}$3 Ma. Unlike other Australian taxa, the Pauropsalta complex did not explosively radiate in response to an early invasion of the arid zone. Instead multiple groups invaded the arid zone and experienced rates of diversification similar to mesic-distributed taxa. We found evidence for relictual groups, located in pre-Mesozoic habitat, that have not diversified and continue to reside on mesic hosts in isolated "habitat islands". Future work should focus on groups of similar ages with similar distribution patterns to determine whether this tempo and pattern of diversification and biogeography is consistent with evidence from other phytophagous insects.

Genomic divergence and lack of introgressive hybridization between two 13-year periodical cicadas support life cycle switching in the face of climate change.

Life history evolution spurred by post-Pleistocene climatic change is hypothesized to be responsible for the present diversity in periodical cicadas (Magicicada), but the mechanism of life cycle change has been controversial. To understand the divergence process of 13-year and 17-year cicada life cycles, we studied genetic relationships between two synchronously emerging, parapatric 13-year periodical cicada species in the Decim group, Magicicada tredecim and M.neotredecim. The latter was hypothesized to be of hybrid origin or to have switched from a 17-year cycle via developmental plasticity. Phylogenetic analysis using restriction-site-associated DNA sequences for all Decim species and broods revealed that the 13-year M.tredecim lineage is genomically distinct from 17-year Magicicada septendecim but that 13-year M.neotredecim is not. We detected no significant introgression between M.tredecim and M.neotredecim/M.septendecim thus refuting the hypothesis that M.neotredecim are products of hybridization between M.tredecim and M.septendecim. Further, we found that introgressive hybridization is very rare or absent in the contact zone between the two 13-year species evidenced by segregation patterns in single nucleotide polymorphisms, mitochondrial lineage identity and head width and abdominal sternite colour phenotypes. Our study demonstrates that the two 13-year Decim species are of independent origin and nearly completely reproductively isolated. Combining our data with increasing observations of occasional life cycle change in part of a cohort (e.g. 4-year acceleration of emergence in 17-year species), we suggest a pivotal role for developmental plasticity in Magicicada life cycle evolution.

Cicada sound impacts avian dawn chorus in a subtropical forest of Taiwan

Acoustic space is a limited resource for animal communications. To reduce signal competition, animals could differentiate their acoustic frequency range or time to produce sound. Many birds call or sing at dawn for higher signal transmission efficiency. However, in tropical and subtropical forests, the dawn chorus were frequently interferenced by cicadas. In order to examine the effect of cicadas on the avian dawn chorus, we used automated acoustic recorders at subtropical forest of Yangmingshan National Park, Taiwan, to monitor acoustic signals of eight bird species and three cicada species. We found that all bird species postponed their acoustic activities during cicada's breeding season. Meanwhile, we also found frequency partitioning of acoustic signals between birds and cicadas when their acoustic signals overlapped temporally. The results show that cicadas could play a crucial role in the avian dawn chorus. Birds would delay their signal producing time and acoustic frequency to partition the acoustic niche.

Cicada Wing Surface Topography: An Investigation into the Bactericidal Properties of Nanostructural Features.

Recently, the surface of the wings of the Psaltoda claripennis cicada species has been shown to possess bactericidal properties and it has been suggested that the nanostructure present on the wings was responsible for the bacterial death. We have studied the surface-based nanostructure and bactericidal activity of the wings of three different cicadas (Megapomponia intermedia, Ayuthia spectabile and Cryptotympana aguila) in order to correlate the relationship between the observed surface topographical features and their bactericidal properties. Atomic force microscopy and scanning electron microscopy performed in this study revealed that the tested wing species contained a highly uniform, nanopillar structure on the surface. The bactericidal properties of the cicada wings were investigated by assessing the viability of autofluorescent Pseudomonas fluorescens cells following static adhesion assays and targeted dead/live fluorescence staining through direct microscopic counting methods. These experiments revealed a 20-25% bacterial surface coverage on all tested wing species; however, significant bactericidal properties were observed in the M. intermedia and C. aguila species as revealed by the high dead:live cell ratio on their surfaces. The combined results suggest a strong correlation between the bactericidal properties of the wings and the scale of the nanotopography present on the different wing surfaces.

Evolution of periodicity in periodical cicadas

Periodical cicadas (Magicicada spp.) in the USA are famous for their unique prime-numbered life cycles of 13 and 17 years and their nearly perfectly synchronized mass emergences. Because almost all known species of cicada are non-periodical, periodicity is assumed to be a derived state. A leading hypothesis for the evolution of periodicity in Magicicada implicates the decline in average temperature during glacial periods. During the evolution of periodicity, the determinant of maturation in ancestral cicadas is hypothesized to have switched from size dependence to time (period) dependence. The selection for the prime-numbered cycles should have taken place only after the fixation of periodicity. Here, we build an individual-based model of cicadas under conditions of climatic cooling to explore the fixation of periodicity. In our model, under cold environments, extremely long juvenile stages lead to extremely low adult densities, limiting mating opportunities and favouring the evolution of synchronized emergence. Our results indicate that these changes, which were triggered by glacial cooling, could have led to the fixation of periodicity in the non-periodical ancestors.

On the use and importance of Multivariate Receiver Operating Characteristic (MROC) curve in identification of cicada species

ABSTRACTIn real-world situations, problems pertaining to classification are commonly observed. To handle such problems, prominent statistical classification tools are being used. This article focuses on one such classification tool, namely Receiver Operating Characteristic (ROC) curve. The entire framework of the work is to demonstrate the use and importance of the ROC curve under multivariate setup. Real data on cicadas is considered where the objectives are to identify the type of species as well as the gender. The results disclose interesting facts about the species considered.

Phylogeography of six codistributed New Zealand cicadas and their relationship to multiple biogeographical boundaries suggest a re‐evaluation of the Taupo Line

AbstractAimComparative biogeographers question the extent to which codistributed species respond similarly to environmental change. Such responses should create similar, appropriately timed patterns of cladogenesis among codistributed taxa compared to evolutionary independence, which may limit the predictions that can be made for unstudied species. Here, we compare phylogeographical patterns across ecologically divergent, codistributed taxa in the light of New Zealand's palaeohistory.LocationNorth Island, New Zealand.MethodsMitochondrial DNA from six codistributed cicada species (Kikihia ochrina, K. cutora, K. laneorum, K. cauta, K. scutellaris and K. dugdalei) was analysed using phylogenetic methods and molecular dating techniques. We analysed phylogeographical distributions using analysis of molecular variance (AMOVA) to determine the significance of hypothesized biogeographical boundaries for clade differentiation and spatial distribution of genetic diversity.ResultsFive species (Kikihia ochrina, K. cutora, K. laneorum, K. cauta and K. scutellaris) show various degrees of intraspecific concordance with biogeographical boundaries found in previously studied taxa – the Kauri Line, the Northland Line and the newly identified Cockayne's Line. Clade splits of forest species correlate with the Kauri Line and/or Northland Line, whereas splits of scrub/hill species correlate with Cockayne's Line. Four species (Kikihia ochrina, K. cutora, K. laneorum and K. cauta) diversified before the Last Glacial Maximum (LGM, 20 ka), whereas two species (K. scutellaris and K. dugdalei) show only post‐LGM diversification.Main conclusionsDespite species idiosyncrasies, we see the imprint of shared palaeoclimatic/geological events. We distinguish between (1) the importance of biogeographical lines as the demarcation between older genetically diverse and newer genetically depauperate populations, and (2) the importance of lines as biogeographical boundaries between sister clades. We also stress the importance of dating clade splits to ensure consistency with explanations for the biogeographical lines in question. We suggest that the Taupo Line has been overemphasized as a biogeographical boundary, whereas the importance of the mountain axis running north‐east to south‐west (‘Cockayne's Line’) has been overlooked.

A redescription of Yoyetta landsboroughi (Distant) and Y. tristrigata (Goding and Froggatt) (Hemiptera: Cicadidae) and description of four new related species.

This study provides redescriptions of two small cicada species, Yoyetta landsboroughi (Distant) and Y. tristrigata (Goding and Froggatt), from eastern Australia, based on a detailed morphological examination of available material. The status of Y. toowoombae (Distant) is re-examined and it is now formally recognised to be a junior synonym of Y. landsboroughi. Four new species of Yoyetta are described, also from eastern Australia. These are: Y. cumberlandi sp. nov., Y. fluviatilis sp. nov., Y. nigrimontana sp. nov., and Y. repetens sp. nov.. Within each species (re)description, sections on distinguishing features, distribution, habitat and behaviour, and calling song structures are described and illustrated where appropriate.

A new genus and species of tettigarctid cicada from the early Miocene of New Zealand: Paratettigarctazealandica (Hemiptera, Auchenorrhyncha, Tettigarctidae).

A new genus and species of primitive cicada (Hemiptera: Tettigarctidae) is described from the early Miocene of southern New Zealand. Paratettigarcta zealandica gen. et sp. n. is the first cicada (Cicadoidea) fossil from New Zealand and exhibits wing venation patterns typical for the subfamily Tettigarctinae. It differs from other fossil taxa and the extant genus Tettigarcta in the early divergence of CuA2 from the nodal line in the forewing, its parallel-sided subcostal cell, the early bifurcation of vein M and long apical cells of the hindwing, and in wing pigmentation patterns.

How do "mute" cicadas produce their calling songs?

Insects have evolved a variety of structures and mechanisms to produce sounds, which are used for communication both within and between species. Among acoustic insects, cicada males are particularly known for their loud and diverse sounds which function importantly in communication. The main method of sound production in cicadas is the tymbal mechanism, and a relative small number of cicada species possess both tymbal and stridulatory organs. However, cicadas of the genus Karenia do not have any specialized sound-producing structures, so they are referred to as “mute”. This denomination is quite misleading, as they indeed produce sounds. Here, we investigate the sound-producing mechanism and acoustic communication of the “mute” cicada, Karenia caelatata, and discover a new sound-production mechanism for cicadas: i.e., K. caelatata produces impact sounds by banging the forewing costa against the operculum. The temporal, frequency and amplitude characteristics of the impact sounds are described. Morphological studies and reflectance-based analyses reveal that the structures involved in sound production of K. caelatata (i.e., forewing, operculum, cruciform elevation, and wing-holding groove on scutellum) are all morphologically modified. Acoustic playback experiments and behavioral observations suggest that the impact sounds of K. caelatata are used in intraspecific communication and function as calling songs. The new sound-production mechanism expands our knowledge on the diversity of acoustic signaling behavior in cicadas and further underscores the need for more bioacoustic studies on cicadas which lack tymbal mechanism.