A study of the abundance and patchiness of cicada nymphs (Homoptera: Tibicinidae) in a New Zealand subalpine shrub-grassland

Abstract

Cicada emergence skins in a subalpine shrub grassland have been sampled during 1969–75 to determine the abundance and spatial distributions of nymphs feeding on plant roots. A guild of six cicada species is primarily associated with two forms of vegetation: shrubs (Dracophyllum and Cassinia) and tall tussock (Chionochloa). Skin locations were mapped relative to dominant vegetation species, litter zones, and soil and rock pavements over a range of aspects, altitudes, and vegetation types, and sampling methods were scaled at four levels: the locality, plot, quadrat, and individual plant. There were significant differences in skin counts over four years, and different measures of mean skin densities are given for the four sampling scales. The two primary vegetation types had cumulative 1969–72 mean densities of 5.2 ± 4.0 and 12.9 ± 10.0 skins/quadrat (2.3 m²) , and the 1969–72 mean productivities of the upper 25% of quadrats (adjusted for percent ground cover) were, respectively, 5.5 and 35.5 skins/m². These productivities are believed to be conservative estimates of the maturing nymph numbers per individual host plant over the span of one cicada generation. Over a 17-year span, such productivities lie within the upper range of mean densities recorded for 17-year periodical cicadas in the United States. As the dominant subalpine vegetation species are very slow-growing,it is suggested that high densities of nymphs feeding on root sap may affect plant vitality, although 1971/1987 comparisons of vitality in 52 Chionochloa tussocks could not positively demonstrate a correlation across all data. Skin dispersion analyses indicated significant levels of patchiness, in agreement with other nymphal studies and with known cicada oviposition behaviour. No single dispersion model fitted the data comprehensively, and it is suggested that a gradual shifting of the centres of cicada aggregation may occur over a cumulative period of several generations.     

Red-winged blackbird predation on periodical cicadas (Cicadidae: Magicicada spp.): bird behavior and cicada responses

Summary Predation by red-winged blackbirds Agelaius phoeniceus L. on 13-year periodical cicadas (Magicicada spp.) and reactions by periodical cicadas to predators were studied during emergence of Brood XIX during summer of 1985 in northwestern Arkansas (USA). Emergences of periodical cicadas are classic examples of predator satiation due to high local densities of cicadas and birds are the major predators of adult periodical cicadas. Reactions of periodical cicadas to predators were assessed by recording behaviors exhibited by cicadas when approached in trees by a human hand during the 3-week period of peak adult densitics. Most male cicadas made a noise in association with escape behaviors when approached, and 50% of the females, which are silent, attempted escape behaviors. Observations of predation attempts on cicadas by red-winged blackbirds were made during the period of peak predation pressure. Red-winged blackbirds spent less time searching for cicadas over that 2-week period of increasing predation pressure, and became increasingly efficient at capturing cicadas. Handling time of cicadas by red-winged blackbirds increased by about 20 seconds over that period, as blackbirds spent more time consuming female cicadas. The flysquawk response, used only by male cicadas, was effective in deterring red-winged blackbirds; only 5% of the attacks by blackbirds were successful when that behavior was exhibited. All cicadas that remained motionless and silent when approached by blackbirds were captured and consumed. Because females remained inactive when approached more often than did males, blackbirds may have consumed more female cicadas. Changes that appeared in reactions of preiodical cicadas to the model predator and to the attacks of blackbirds reflect both changes in the sex ratio of the cicada populations and changes in behaviors of cicadas associated with mating and egg laying. The loud noise made by male periodical cicadas at mating centers did not appear to deter predation by blackbirds. Changes in the behavior of blackbirds that appeared to be in association with greater predation on female periodical cicadas relates directly to aspects of foraging theory, particularly predictions concerning more selective foraging during periods of abundant food resources.     

Energetics of emergence in the cicadas, Cyclochila australasiae and Abricta curvicosta (Homoptera: Cicadidae)

The final moult in cicadas marks a major transition in lifestyle and is a behaviour that makes the cicada vulnerable to predation. Consequently, emergence times are short and, we predict, therefore the rate of energy consumption would be high. Hence, we measured the energetic cost of emergence in Cyclochila australasiae (green grocer) and Abricta curvicosta (floury baker) cicadas during the final moult from nymph to adult cicada. Maximum energy expended whilst emerging was compared between the sexes and species. Even though C. australasiae take longer to emerge than A. curvicosta, the mass-specific cost of emergence is not different between the two species (C. australasiae: 11.34+/-2.55 J g(-1); A. curvicosta: 12.91+/-1.90 J g(-1)). The mass-specific metabolic rates of fully emerged adults of both species are approximately twice those of the nymphs and the maximum metabolic rate during emergence is about 1.5 times higher than the resting metabolic rate of emerged adults. Emergence times, as indicated by rates of oxygen consumption, are longer than expected and probably reflect limitations in the oxygen capacity of the cicadas during moulting.     

Induced responses of cherry trees to periodical cicada oviposition

Cherry trees (Prunus serotina) responded to oviposition by periodical cicadas (Magicicada spp.) by depositing gum at the egg nest. The proportion of cicada eggs that hatched successfully was significantly reduced at egg nests with visible gum compared to non-gummed egg nests. The number of egg nests with gum increased in proportion to the total number of egg nests on a tree. The probability of an egg nest having visible gum increased as the total number of egg nests increased. Mortality at hatching due to gum deposition increased as a direct density-dependent function of the number of cicada eggs laid in the tree. Although statistically significant, this relationship was weak and appeared to hold only at densities above 100 egg nests per tree. Gum deposition is discussed as an induced plant response to cicada attack. A cherry may reduce the number of cicada nymphs that will parasitize it up to the next oviposition period (17 or 13 years later) by reducing cicada hatching through gum deposition at the site of oviposition.     

SELECTION FOR PRIME-NUMBER INTERVALS IN A NUMERICAL MODEL OF PERIODICAL CICADA EVOLUTION

Periodical cicadas are known for unusually long and prime-numbered life cycles (13 and 17 years) for insects. To explain the evolution of prime-numbered reproductive intervals (life cycles), the hybridization hypothesis claims that prime numbers greatly reduce the chance of hybridization with other life cycles. We investigate the hybridization hypothesis using a simulation model. This model is a deterministic, discrete population model with three parameters: larval survival per year, clutch size, and emergence success. Reproductive intervals from 10 years to 20 years compete for survival in the simulations. The model makes three key assumptions: a Mendelian genetic system, random mating among broods of different life-cycle lengths, and integer population sizes. Longer life cycles have larger clutch sizes but suffer higher total mortality than shorter life cycles. Our results show that (1) nonprime-numbered reproductive intervals disappear rapidly in comparison to the selection among the various prime-numbered life cycles, (2) the selection of prime-numbered intervals happens only when populations are at the verge of extinction, and (3) the 13- and 17-year prime phenotypes evolve under certain conditions of the model and may coexist. The hybridization hypothesis is discussed in light of other hypotheses for the evolution of periodical cicada life cycles.     

Self‐organizing cicada choruses respond to the local sound and light environment

1. Periodical cicadas exhibit an extraordinary capacity for self‐organizing spatially synchronous breeding behavior. The regular emergence of periodical cicada broods across the United States is a phenomenon of longstanding public and scientific interest, as the cicadas of each brood emerge in huge numbers and briefly dominate their ecosystem. During the emergence, the 17‐year periodical cicada species Magicicada cassini is found to form synchronized choruses, and we investigated their chorusing behavior from the standpoint of spatial synchrony.
2. Cicada choruses were observed to form in trees, calling regularly every five seconds. In order to determine the limits of this self‐organizing behavior, we set out to quantify the spatial synchronization between cicada call choruses in different trees, and how and why this varies in space and time.
3. We performed 20 simultaneous recordings in Clinton State Park, Kansas, in June 2015 (Brood IV), with a team of citizen‐science volunteers using consumer equipment (smartphones). We use a wavelet approach to show in detail how spatially synchronous, self‐organized chorusing varies across the forest.
4. We show how conditions that increase the strength of audio interactions between cicadas also increase the spatial synchrony of their chorusing. Higher forest canopy light levels increase cicada activity, corresponding to faster and higher‐amplitude chorus cycling and to greater synchrony of cycles across space. We implemented a relaxation‐oscillator‐ensemble model of interacting cicadas, finding that a tendency to call more often, driven by light levels, results in all these effects.
5. Results demonstrate how the capacity to self‐organize in ecology depends sensitively on environmental conditions. Spatially correlated modulation of cycling rate by an external driver can also promote self‐organization of phase synchrony.

     

Feeding ecology and evidence for amino acid synthesis in the periodical cicada (Magicicada)

The periodical cicadas of the genus Magicicada (including M. septendecim, M. cassini, and M. septendecula) have the longest juvenile life span of any insect, living underground for 13 or 17 years and feeding exclusively on root xylem fluids. Due to their inaccessible life cycles very little is known about cicada nutrition, despite the fact that members of Magicicada can achieve a very large biomass in woodland habitats east of the Mississippi and hence constitute a major part of the ecosystem where they occur in high densities. Live cicadas were collected at two sites in early June of 2004, during the emergence of Brood X (both M. septendecim and M. cassini were recovered). We used a combination of stable isotopic measurements (δ¹⁵N and δ¹³C) and multivariate statistical techniques to test for differences in resource acquisition among the cicada species and sexes collected at two locations within the 17-year periodical Brood X range. The amino acid constituents of cicada chitin and organs, plus xylem extracted from a deciduous sapling, were also analyzed. The data show that male and female cicadas have different carbon fractionations, which could reflect differential resource utilization due to oviposition in females. Several essential amino acids for the cicada were absent in xylem. Carbon-isotopic composition of all amino acids in the cicadas was distinctly different from the limited set measured in the xylem. Because of the differences in isotopic composition, we conclude that amino acids were synthesized de novo rather than incorporated directly, most likely produced by endosymbiotic bacteria.     

Why is one of the periodical cicadas (Magicicada septendecula) a comparatively rare species?

ABSTRACT.

• 1 The same woodland rarely supports two different broods of 17-year cicadas, and never two that appear in successive years. Theoretically, this is because a build-up of predators, generated by the leading brood, would eliminate the lagging one, or because nymphs of the lagging brood would suffer a competitive disadvantage, or both.
• 2 Nevertheless, substantial numbers of 17-year cicadas of all three species (Magicicada septendecim, M.cassini and M.septendecula) did appear in Grant County, Kentucky, in 1974 (Brood XIV, expected) and 1975 (not expected). This is interpreted to mean that many individuals delayed their emergence until the eighteenth year.
• 3 Dissected eggnests show that the expected build-up of hymenopterous egg parasitoids failed to occur, although there may have been a build-up of dipteran parasitoids, which may leave no trace in the eggnest.
• 4 Excavated young nymphs show that establishment rate was poorer in 1975 than in 1974, which is attributed to nymphal competition.
• 5 Identification of eggnests from the two years shows that one of the species (M.septendecula, which is usually the rarest in mixed-species populations) is much more susceptible to delaying its development by a year than are the other two species.
• 6 It is predicted that the 1975 emergence will produce no adult progeny, i.e. that there will be no emergence in 1992.
• 7 The sporadic occurrence of this phenomenon, other factors being equal, would be sufficient to explain the comparative rarity of M. septendecula, or even its local extinction.
• 8 Historical records from northern Illinois show frequent emergences one year behind schedule. These delayed individuals never produce progeny that survive to the next generation. M.septendecula is absent from northern Illinois and indeed absent from the northern areas of all periodical cicada broods in the northern part of the range.

     

Xylem feeding by periodical cicada nymphs on tree roots

Abstract. 1. Histological sections of tree roots on which periodical cicada nymphs of both the 17-year and the 13-year race arid various instars had fed showed that salivary sheaths made by the nymphs ended in xylem vessels.
2. Because no salivary sheaths were found in phloem cells, this observation indicates that cicada nymphs are xylem feeders throughout their developmental period.
3. The habit of feeding on xylem fluid, which is extremely dilute, may explain why periodical cicada nymphs require so many years (13 or 17) to mature. It may also explain why they excrete amino acids rather than sugars, as phloem feeders do.     

Body temperature and the acoustic behavior of the cicadaTibicen winnemanna (Homoptera: Cicadidae)

Tibicen winnemanna (Davis) produces a broad-band frequency- and amplitude-modulated advertisement call. Observations of the animals in the field show that the cicadas produce varied acoustic output. This work investigates the connection between the acoustic behavior of the cicada and the body temperature of the calling animal. The results show there is strong evidence to support the hypothesis that both acoustic output and song intensity are highly dependent on body temperature.     

Seasonal cues in tropical organisms. Rainfall? Not necessarily!

Summary Activity seasons of tropical organisms often start, on the average, at or about the beginning of the rainy or dry seasons. The hypothesis that the onset or cessation of the wet season provides the seasonal cues necessary of the initiation of the activity season of some tropical organisms is tested with data on Panamanian cicadas. Seasonal adult activity patterns are described for cicada species in Panama, mostly from Barro Colorado Island (BCI), some from Las Cumbres. In all species the correlation between the timing of the beginning and the end of the cicada season was low and not significant, so that the actual beginning data of a cicada season in a particular year had little or no predictive value for the end date. Seven out of eleven species on BCI started their average activity season at the average beginning of the dry season (one species) or rainy season (six species). Nevertheless, in 13 years, correlations between the start or end of the cicada seasons and that of the meteorological seasons were low and not significant. At best, the beginning and end of the rains played a minor role as seasonal cues governing cicada emergence or the termination of the cicada season. It is speculated that photoperiod might be a major seasonal cue governing emergence, through its effects on the host plants.     

Periodical cicadas use light for oviposition site selection

Organisms use incomplete information from local experience to assess the suitability of potential habitat sites over a wide range of spatial and temporal scales. Although ecologists have long recognized the importance of spatial scales in habitat selection, few studies have investigated the temporal scales of habitat selection. In particular, cues in the immediate environment may commonly provide indirect information about future habitat quality. In periodical cicadas (Magicicada spp.), oviposition site selection represents a very long-term habitat choice. Adult female cicadas insert eggs into tree branches during a few weeks in the summer of emergence, but their oviposition choices determine the underground habitats of root-feeding nymphs over the following 13 or 17 years. Here, field experiments are used to show that female cicadas use the local light environment of host trees during the summer of emergence to select long-term host trees. Light environments may also influence oviposition microsite selection within hosts, suggesting a potential behavioural mechanism for associating solar cues with host trees. In contrast, experimental nutrient enrichment of host trees did not influence cicada oviposition densities. These findings suggest that the light environments around host trees may provide a robust predictor of host tree quality in the near future. This habitat selection may influence the spatial distribution of several cicada-mediated ecological processes in eastern North American forests.     

Response of host plants to periodical cicada oviposition damage

Insect oviposition on plants is widespread across many systems, but studies on the response of host plants to oviposition damage are lacking. Although patterns of oviposition vary spatially and temporally, ovipositing insects that exhibit outbreak characteristics may have strong effects on host plants during peak abundance. Periodical cicadas (Magicicada spp.), in particular, may reduce the performance of host plants when they synchronously emerge in massive numbers to mate and oviposit on host plants. Here we provide the first experimental manipulation of host plant use by periodical cicadas to evaluate the impact of cicada oviposition on plant performance across a diversity of host species within an ecologically relevant setting. Using a randomized block design, we established a plantation of three native and three exotic host plant species common to the successional forests in which cicadas occur. During the emergence of Brood X in 2004, we employed a highly effective cicada exclusion treatment by netting half of the host plants within each block. We assessed multiple measures of host plant performance, including overall plant growth and the growth and reproduction of individual branches, across three growing seasons. Despite our thorough assessment of potential host plant responses to oviposition damage, cicada oviposition did not generally inhibit host plant performance. Oviposition densities on unnetted host plants were comparable to levels documented in other studies, reinforcing the ecological relevance of our results, which indicate that cicada oviposition damage did not generally reduce the performance of native or exotic host plants.     

Acoustic and Temporal Partitioning of Cicada Assemblages in City and Mountain Environments

Comparing adaptations to noisy city environments with those to natural mountain environments on the community level can provide significant insights that allow an understanding of the impact of anthropogenic noise on invertebrates that employ loud calling songs for mate attraction, especially when each species has its distinct song, as in the case of cicadas. In this study, we investigated the partitioning strategy of cicada assemblages in city and mountain environments by comparing the acoustic features and calling activity patterns of each species, recorded using automated digital recording systems. Our comparison of activity patterns of seasonal and diel calling revealed that there was no significant temporal partitioning of cicada assemblages in either environment. In addition, there was no correlation between the acoustic distance based on spectral features and temporal segregation. Heterospecific spectral overlap was low in both city and mountain environments, although city and mountain cicada assemblages were subject to significantly different levels of anthropogenic or interspecific noise. Furthermore, for the common species found in both environments, the calling activity patterns at both seasonal and diel time scales were significantly consistent across sites and across environments. We suggest that the temporal calling activity is constrained by endogenous factors for each species and is less flexible in response to external factors, such as anthropogenic noise. As a result, cicada assemblages in city environments with low species diversity do not demonstrate a more significant temporal partitioning than those in mountain environments with high species diversity.     

Periodical Cicada Detritus Impacts Stream Ecosystem Metabolism

The emergence of 17-year periodical cicadas in Maryland, USA, in 2004 provided a unique opportunity to study the effect of a large, but temporally limited, resource pulse of arthropod detritus on stream ecosystem function. Cicada emergence was quantified in the forests adjacent to two small streams with different histories of riparian disturbance (Intact and Disturbed sites). We estimated the input of cicada detritus to the streams, described its retention and breakdown dynamics, and measured whole-stream respiration over the cicada flight season (May–July). Average emergence density was significantly greater at the Intact site, but average cicada detritus input rates were greater at the Disturbed site. Cicada detritus was locally retained within both streams and rapidly broke down. Daily whole-stream respiration (CR₂₄) at both sites responded dramatically to the cicada pulse, with CR₂₄ doubling pre-cicada measurements following the period of greatest cicada input (Intact: 12.82 → 23.78 g O₂ m⁻² d⁻¹; Disturbed: 2.76 → 5.77 g O₂ m⁻² d⁻¹). CR₂₄ returned to baseline levels when cicada input decreased at the Intact site, but more than doubled again at the Disturbed site (13.14 g O₂ m⁻² d⁻¹), despite a decline in cicada input rate. Differences in respiration response may be a function of differences in cicada input rates as well as differences in microbial community activity. The strong effects on stream ecosystem function exerted by a short but intense input of periodical cicada detritus may provide insights regarding the response of streams to other irregular resource pulses. HM, MP, LC, and DR conceived and designed study; HM, LC, and DR performed research; HM, LC, and DR analyzed data; HM, MP, LC, and DR wrote the paper.     

TRACE FOSSILS OF CICADAS IN THE CENOZOIC OF CENTRAL PATAGONIA, ARGENTINA

Abstract:  Hemispherical pan or dish-shaped trace fossils from the Cenozoic of the Central Patagonia are attributed to the burrowing action of cicadas. A new ichnotaxon, Feoichnus challa igen. et isp. nov. is characterized by its hemispherical shape, mostly subvertical orientation, smoothed internal lining showing knobbly surface texture, and rough and irregular external surface devoid of ornamentation. The wall is composed of a lining plus a layer of soil material consolidated passively by cicada excretions. In other specimens the wall shows a repetition of linings and soil layers reflecting changes in the position of the chambers. Grooves, which represent traces of roots originally related to the feeding activities of cicada nymphs, are located in the wall of many specimens of F. challa . These grooves are subvertical to subhorizontal and show smooth surfaces with longitudinal striations. The cicadan origin of these traces is supported by comparison of the fossils to modern cicada nymph chambers and by laboratory experiments. The most significant characters that emerged from the comparison are the differential preservation of the basal part of the chambers, the interior lining with similar surface texture, and the presence of root traces in the wall and in the interior of the chamber. Additional records of F. challa from the Cretaceous–Pliocene of the USA, the Oligocene of Ethiopia, and the Miocene of the United Arab Emirates and Kenya complete the current information yielded by the known body fossil record to help understand the evolutionary history of cicadas.     

Genetic evidence for assortative mating between 13-year cicadas and sympatric "17-year cicadas with 13-year life cycles" provides support for allochronic speciation

Abstract. Thirteen-year cicadas of brood XIX from northern Arkansas, Missouri, and southern Illinois (lineage A) are known to be genetically different at two marker loci (mitochondrial DNA and abdominal color) from 13-year cicadas to the south (lineage B) that emerge in the same year. Because 17-year cicadas from all broods (year classes) are indistinguishable from lineage A at these two marker loci, previous workers suggested that the lineage A cicadas of 13-year brood XIX were derived from 17-year cicadas by life-cycle switching (allochrony). Data presented here show that, over the same northern geographic range, lineage A is also present in 13-year cicadas belonging to brood XXIII (which always emerges four years later than brood XIX). Detailed sampling along the putative life-cycle-switching boundary in 13-year brood XXIII revealed a previously unsuspected broad zone of overlap where populations contained individuals of both lineages A and B. Despite this sympatry, and previous reports of a lack of behavioral barriers to interbreeding, a strong correlation between mitochondrial haplotype and abdominal color suggests that assortative mating has taken place. Lineage A 13-year cicadas from both broods XIX and XXIII are only found within a gap in the spatial distribution of 17-year cicadas. This, in combination with the lack of differentiation between lineage A 13- and 17-year cicadas at the marker loci and new behavioral data for 13-year brood XIX, suggests a recent derivation of all northern 13-year cicadas from the 17-year cicadas via life-cycle switching. We discuss the implications of these allochronic shifts for speciation.     

Ecophysiological responses to climate change in cicadas

Cicadas are large hemipteran insects characterized by unique life‐history traits, such as extraordinarily long life cycles, a subterranean/terrestrial habitat transition, xylem sap‐feeding and melodious sound production. These fascinating features of cicadas have attracted much attention in the research fields of physiology and ecology, resulting in an accumulation of knowledge about the underlying mechanisms and their adaptive significance. Although community‐level responses to recent climate change have already been documented for cicada fauna, an understanding of their causal relationships is still at the initial stages. In this review, we summarize current knowledge about environmental adaptations of cicadas to facilitate a deeper understanding of the ecophysiological consequences of climate change. We first outline the diverse responses of cicadas to environmental factors, mainly temperature, and their strategies to cope with naturally fluctuating environments. Then, we discuss the consequence of upcoming climate change by consolidating the current findings. This review highlights the fact that fitness‐relevant activities are fine‐tuned to a species‐specific temperature optimum to achieve habitat segregation among coexisting species, implying that cicada diversity is highly susceptible to climate warming. As a result of their conspicuous large bodies and species‐specific calling songs, cicadas are promising candidates for use as bioindicator species to monitor ecological impacts of climate change. We encourage future works that continuously quantify population‐ and community‐level responses to upcoming climate change, as well as unveil mechanistic links between physiological traits and ecological consequences.     

Cellular and potential molecular mechanisms underlying transovarial transmission of the obligate symbiont Sulcia in cicadas

Vertical transmission of symbionts in insects is critical to persistence of symbioses across host generations. The key time point and related cellular/molecular mechanisms underlying the transmission in most insects remain unclear. Here, we reveal that in the bacteriome–endosymbiont system of the cicada Meimuna mongolica, the obligate symbiont Candidatus Sulcia muelleri (hereafter Sulcia) proliferates and migrates to the ovaries mainly after the adult emergence of cicadas. Sulcia cells swell to approximately twice their previous size with the outer membrane changed to be more irregular during this process. Almost all the Sulcia genes involved in biosynthesis of essential amino acids, heat shock protein, energy metabolism, DNA replication and repair and protein export were highly expressed in all life stages of cicadas. Among which, genes involved in DNA replication and synthesis of leucine and arginine were upregulated in the newly emerged adults relative to fifth-instar nymphs. Signal transduction is the pronounced function exhibited in both Sulcia and the cicada bacteriomes in newly emerged adults. The results suggest host sensing of arginine and leucine integrate Sulcia's output of host-EAAs into mTORC1 signalling. This study highlights the importance of signalling pathways in regulating the host/symbiont interaction and symbiont transmission in sap-feeding auchenorrhynchous insects.     

Phylogeography of the New Zealand cicada Maoricicada campbelli based on mitochondrial DNA sequences: ancient clades associated with cenozoic environmental change

New Zealand's isolation, its well-studied rapidly changing landscape, and its many examples of rampant speciation make it an excellent location for studying the process of genetic differentiation. Using 1520 base pairs of mitochondrial DNA from the cytochrome oxidase subunit I, ATPase subunits 6 and 8 and tRNA(Asp) genes, we detected two well-differentiated, parapatrically distributed clades within the widespread New Zealand cicada species Maoricicada campbelli that may prove to represent two species. The situation that we uncovered is unusual in that an ancient lineage with low genetic diversity is surrounded on three sides by two recently diverged lineages. Using a relaxed molecular clock model coupled with Bayesian statistics, we dated the earliest divergence within M. campbelli at 2.3 +/- 0.55 million years. Our data suggest that geological and climatological events of the late Pliocene divided a once-widespread species into northern and southern components and that near the middle of the Pleistocene the northern lineage began moving south eventually reaching the southern clade. The southern clade seems to have moved northward to only a limited extent. We discovered five potential zones of secondary contact through mountain passes that will be examined in future work. We predict that, as in North American periodical cicadas, contact between these highly differentiated lineages will exist but will not involve gene flow.