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.     

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.     

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.     

Permanence induced by life-cycle resonances: the periodical cicada problem

Periodical cicadas are known for their unusually long life cycle for insects and their prime periodicity of either 13 or 17 years. One of the explanations for the prime periodicity is that the prime periods are selected to prevent cicadas from resonating with predators with submultiple periods. This paper considers this hypothesis by investigating a population model for periodical predator and prey. The study shows that if the periods of the two periodical species are not coprime, then the predator cannot resist the invasion of the prey. On the other hand, if the periods are coprime, then the predator can resist the invasion of the prey. It is also shown that if the periods are not coprime, then the life-cycle resonance can induce a permanent system, in which no cohorts are missing in both populations. On the other hand, if the periods are coprime, then the system cannot be permanent.     

Permanence induced by life-cycle resonances: the periodical cicada problem

Periodical cicadas are known for their unusually long life cycle for insects and their prime periodicity of either 13 or 17 years. One of the explanations for the prime periodicity is that the prime periods are selected to prevent cicadas from resonating with predators with submultiple periods. This paper considers this hypothesis by investigating a population model for periodical predator and prey. The study shows that if the periods of the two periodical species are not coprime, then the predator cannot resist the invasion of the prey. On the other hand, if the periods are coprime, then the predator can resist the invasion of the prey. It is also shown that if the periods are not coprime, then the life-cycle resonance can induce a permanent system, in which no cohorts are missing in both populations. On the other hand, if the periods are coprime, then the system cannot be permanent.     

Reconstructing asymmetrical reproductive character displacement in a periodical cicada contact zone

Selection against costly reproductive interactions can lead to reproductive character displacement (RCD). We use information from patterns of displacement and inferences about predisplacement character states to investigate causes of RCD in periodical cicadas. The 13-year periodical cicada Magicicada neotredecim exhibits RCD and strong reproductive isolation in sympatry with a closely related 13-year species, Magicicada tredecim. Displacement is asymmetrical, because no corresponding pattern of character displacement exists within M. tredecim. Results from playback and hybridization experiments strongly suggest that sexual interactions between members of these species were possible at initial contact. Given these patterns, we evaluate potential sources of selection for displacement. One possible source is 'acoustical interference', or mate-location inefficiencies caused by the presence of heterospecifics. Acoustical interference combined with the species-specificity of song pitch and preference appears to predict the observed asymmetrical pattern of RCD in Magicicada. However, acoustical interference does not appear to be a complete explanation for displacement in Magicicada, because our experiments suggest a significant potential for direct sexual interactions between these species before displacement. Another possible source of selection for displacement is hybrid failure. We evaluate the attractiveness of inferred hybrid mating signals, and we examine the viability of hybrid eggs. Neither of these shows strong evidence of hybrid inferiority. We conclude by presenting a model of hybrid failure related to life cycle differences in Magicicada.     

Dynamics of periodical cicada chorus centers (Homoptera: Cicadidae:Magicicada)

Mating aggregations of three species of periodical cicadas were monitored during the emergence of Brood XIX at a 16-ha study site in northwest Arkansas, May–June 1985. Magicicada tredecassiniappeared first and formed the most choruses. M. tredecimand M. tredeculachoruses formed next, and M. tredeculachoruses outnumbered those of M. tredecim.Of the 268 choruses seen, 84% were composed of M. tredecassini. M. tredecassiniwere often found chorusing in the same trees with the other two species. Such multispecies mating aggregations apparently are unique to periodical cicadas. Choruses were dynamic with respect to their locations and durations. Initially, choruses were located near areas of high cicada emergence densities. One week later, cicadas chorused in trees throughout the forest and at the forest edge. Many choruses were seen only once at a location. Although cicadas chorused for almost 4 weeks, individual choruses persisted only approximately 8 days, on average. Sound intensities under chorus centers ranged from 50 to 80 dh and were correlated with arena sizes during times of peak chorus activity. No distinct habitat preferences of the three species were observed, however, the tree species used by chorusing cicadas differed among the species.     

Nature and fate of the nutrient reserves of the periodical (17 year) cicada

The metabolic reserves of Magicicada cassini, a periodical cicada with a 17 year life cycle, were examined. Analysis of males and females which were collected shortly after emergence and just before death provided information about the nature and fate of the reserves which had been accumulated during the attenuated larval period. Results showed that young females had a mean weight of 565 mg compared with only 351 mg for equivalent males. Lipids and soluble proteins were the principal reserve materials. Glycogen was a relatively minor reserve of both sexes. While young females contained about twice as much soluble protein (110 mg) as the males (51 mg), they contained three times as much total lipid material (58–19 mg). Lipids were fractionated into neutral and phospholipids and the absolute amounts of each class were determined; triglycerides were the main component.Further analyses of old females provided information about the utilization of reserves for oögenesis. The data showed that even though the adults feed, lipids and soluble proteins stored during the larval stages are essential for egg maturation. The results are discussed in relation to the special requirements of an insect which has a protracted immature stage, yet a fleeting adult existence.     

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.     

Plant architecture affects periodical cicada oviposition behavior on native and non‐native hosts

Variation in plant quality provides a basis for oviposition site selection for a variety of insects. Of the plant traits that influence plant–insect interactions, plant architecture has received little attention despite its putative role in modulating oviposition behavior. In a common garden comprised of native and non‐native plant species, we assessed how host plant architecture and identity influenced the oviposition behavior of 17‐year periodical cicadas (Homoptera: Cicadidae: Magicicada). On each host, we quantified the availability of branches suitable for oviposition and compared those measures with the branches used by ovipositing cicadas. Using this approach, we determined how the structural attributes of plants (i.e. branch diameter, length and incline) affected oviposition site selection. We then related cicada oviposition preferences to offspring performance by quantifying egg hatching success. On each host species, cicadas selectively used broader and longer branches for oviposition, suggesting that branch architecture provides a basis for oviposition behavior irrespective of plant identity. Broader and longer branches were more abundant on native than on non‐native hosts in our study, contributing to greater oviposition loads among the native species. Egg hatching success was similar among native and non‐native hosts. However, it is possible that the use of native plants for oviposition could enhance offspring output because native hosts generally contained more viable eggs per egg nest and more egg nests per plant. While previous accounts of cicada oviposition preferences have focused on differences in oviposition loads among hosts, our evaluation of within‐host branch selection by ovipositing cicadas helps to clarify oviposition preferences at a higher resolution and demonstrates that plant architecture provides an important basis for oviposition behavior. Furthermore, because branch structure can differ substantially among host species, our results suggest that periodical cicadas may be sensitive to the changes in plant composition that often result from non‐native plant invasions.     

Synrhabdosome life cycles

Synrhabdosomes are monospecific colonies of colonies of graptolites. Their mode of formation is unknown. We draw attention to a type of colony formation found in two genera of colonial rotifers and suggest that these represent plausible models for the life cycle of graptolite synrhabdosomes.     

Evaporative cooling and endothermy in the 13-year periodical cicada,Magicicada tredecem (Homoptera: Cicadidae)

Summary The thermobiology of a cicada, Magicicada tredecem, from a warm, high humidity environment was investigated. Thoracic temperature (T_th) of M. tredecem in the field was strongly dependent on, and consistently higher than, ambient temperature (T_am), averaging 33.0±0.19°C on warm sunny days (T_am=28–29°C, rh=60–75%). Laboratory studies documented cuticle water fluxes high enough ( 5mg · cm^–2 · h^–1 in dry air at 40°C) to result in a significant degree of passive evaporative cooling, but the ability of M. tredecem to actively facilitate evaporative water loss during thermal stress is comparatively limited: water loss rates (WLR) of live M. tredecem at 40°C (dry air) were only 35–45% greater than those of dead cicadas. The limited ability of M. tredecem to facilitate transcuticular WLR is associated with limited surface distribution of the cuticular ducts through which water is actively extruded during evaporative cooling. In the laboratory, active extrusion of water had no appreciable effect on T_th, demonstrating that evaporative cooling was due largely to passive water flux through the highly permeable cuticle. The location of the abdominal pore tracts is such that extrusion of water through the ducts may preferentially cool the heart and perhaps other abdominal tissues. Long-term climatological data indicate that M. tredecem rarely encounters T_am levels high enough (i.e., above its apparent T_th setpoint of 34–35°C) to require evaporative cooling. Inactive M. tredecem can endothermically increase T_th. An hypothesis is proposed to account for the diversity of body temperature setpoints in cicadas.Abbreviations rh relative humidity - SOT standard operating temperature - T _am ambient temperature - T _b body temperature - T _sp body temperature setpoint - T _th thoracic temperature - TWF transcuticular water flux - WLR water loss rate     

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.     

Long life cycles in insects

Long life cycles covering more than one year are known for all orders of insects. There are different mechanisms of prolongation of the life cycle: (1) slow larval development; (2) prolongation of the adult stage with several reproduction periods; (3) prolongation of diapause; (4) combination of these mechanisms in one life cycle. Lasting suboptimal conditions (such as low temperature, low quality of food or instability of food resources, natural enemies, etc.) tend to prolong life cycles of all individuals in a population. In this case, the larvae feed and develop for longer than a year, and the active periods are interrupted by dormancy periods. The nature of this dormancy is unknown: in some cases it appears to be simple quiescence, in others it has been experimentally shown to be a true diapause. Induction and termination of these repeated dormancy states are controlled by different environmental cues, the day-length being the principal one as in the case of the annual diapause. The long life cycles resulting from prolonged adult lifespan were experimentally studied mainly in beetles and true bugs. The possibility of repeated diapause and several periods of reproductive activity is related to the fact that the adults remain sensitive to day length, which is the main environmental cue controlling their alternative physiological states (reproduction vs. diapause). Habitats with unpredictable environmental changes stimulate some individuals in a population to extend their life cycles by prolonged diapause. The properties of this diapause are poorly understood, but results of studies of a few species suggest that this physiological state differs from the true annual diapause in deeper suppression of metabolism. Induction and intensity of prolonged diapause in some species appear to be genetically controlled, so that the duration of prolonged diapause varies among individuals in a group, even that of sibles reared under identical conditions. Thus, long life cycles are realized due to the ability of insects to interrupt activity repeatedly and enter dormancy. This provides high resistance to various environmental factors. Regardless of the nature of this dormancy (quiescence, annual or prolonged diapause, or other forms) and the life cycle duration, the adults always appear synchronously after dormancy in the nature. The only feasible explanation of this is the presence of a special synchronizing mechanism, most likely both exo- and endogenous, since the adults appear not only synchronously but also in the period best suited for reproduction. As a whole, the long life cycles resulting from various structural modifications of the annual life cycle, are typical of the species living under stable suboptimal conditions when the pressure of individual environmental factors is close to the tolerance limits of the species, even though it represents its norm of existence. Such life cycles are also typical of the insects living in unstable environments with unpredictable variability of conditions, those developing in cones and galls, feeding on flowers, seeds, or fruits with limited periods of availability, those associated with the plant species with irregular patterns of blossoming and fruiting, and those consuming low-quality food or depending on unpredictable food sources (e.g., predators or parasites). Long cycles are more common in: (1) insect species at high latitudes and mountain landscapes where the vegetation season is short and unstable; (2) species living in deserts or arid areas where precipitation is unstable and often insufficient for survival of food plants; (3) inhabitants of cold and temporary water bodies that are not filled with water every year. At the same time, long life cycles sometimes occur in insects from other climatic zones as well. It is also important to note that while there is a large body of literature dealing with the long life cycles in insects, it mostly focuses on external aspects of the phenomenon. Experimental studies are needed to understand this phenomenon, first of all the nature of dormancy and mechanisms of synchronization of adult emergence.     

Morphoprocess and life cycles of organisms

In developing the ideas of V.N. Beklemishev about an organism as a form, existing in a process of determined transformation and matter/energy exchange, we consider different aspects of the term "morphoprocess" and introduce corresponding additional terms. Momentary morphoprocess characterizes an organism in the given moment of time. This term reflects a constancy of the form ("momentary form"), where the existence of an organism can be imagined as a sequence of "momentary forms". "First derivative" of this momentary characteristic is particular morphoprocess--an organism from its origin to fission/division or death. Compound particular morphoprocess is a determined and reiterating sequence of different particular morphoprocesses. And, at last, general morphoprocess--a "second derivative" of momentary morphoprocess--is rhythmical reiteration of a particular morphoprocess on the long-term scale, an ancestors/descendants lineage. To describe consecutive changes in this material system, the terms ontogenesis and life cycle are used. Ontogenesis characterizes a sequence of the morpho-functional changes of an individual organism during its life, whereas life cycle reflects a sequence of changes during one complete segment of the general morphoprocess represented by a single or several particular morphoprocesses. We also discuss morphoprocess uniformity along with the phase nature of morphoprocesses, both particular and compound particular ones.     

Mathematical description of holometabolous life cycles

The deterministic continuous equations are developed for the population levels of the various life stages of holometabolous species. Special attention is paid to the larval stage in which the variable of weight, in addition to the traditional variables of age and time, is included; growth rate and the genetic variability of the growth rate are allowed for. A pair of equations is derived that permits computation of the larval population level without regard to the levels of the other life stages. Finite difference equations are developed, simple analytical forms for vital rates are adopted, and a numerical example is given.