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.     

Spatial variability in oviposition damage by periodical cicadas in a fragmented landscape

Effects of the periodical cicada (Magicicada spp.) on forest dynamics are poorly documented. A 1998 emergence of M. cassini in eastern Kansas led to colonization of a fragmented experimental landscape undergoing secondary succession. We hypothesized that per-tree rates of oviposition damage by cicadas would reflect: (1) distance from the source of the emergence, (2) patch size, and (3) local tree density. Ovipositing females displayed clear preferences for host species and damage incidence showed predictable spatial patterns. Two species (smooth sumac, Rhus glabra, and eastern red cedar, Juniperus virginiana) were rarely attacked, whereas others (rough-leaved dogwood, Cornus drummondii; slippery elm, Ulmus rubra; box elder, Acer negundo, and honey locust, Gleditsia triacanthos) were strongly attacked. The dominant early successional tree, dogwood, received on average the most attacks. As predicted, attacks per stem declined strongly with distance from the emergence source, and with local stem density (a "dilution" effect). Contrary to expectations, there were more attacks per stem on larger patches. Because ovipositing cicadas cut damaging slits in host tree branches, potentially affecting tree growth rate, competitive ability, and capacity to reproduce, cicada damage could potentially influence spatial variation in secondary succession.     

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.     

Does the periodical cicada,Magicicada septendecim,prefer to oviposit on native or exotic plant species?

Abstract 1. In 2004, Brood X of the periodical cicada (Magicicada spp.) emerged in Delaware. Extensive suburban development and concomitant planting of exotic species has occurred since the previous emergence of Brood X in 1987. 2. Exotic species could suffer extensive damage during years of cicada emergences if they are preferred for oviposition. Alternately, a shortage of suitable plant hosts may negatively affect remaining cicada populations. 3. We determined if the periodical cicada, Magicicada septendecim, preferred to oviposit on native or exotic woody plant species. Potential hosts were divided into three groups and planted in a randomised design near a likely source of cicadas. The first group, Natives, included 15 species native to Delaware. The second group, Non‐natives, included 15 exotic species that had a native congener represented in the Native group. The final group, Aliens, included 13 exotic species that did not have a native congener. 4. Based on observations of 428 plants, cicadas were more likely to oviposit on Natives or Non‐natives, which did not differ from each other in this measure, than on Aliens. Non‐natives had more oviposition holes per metre than Natives, which had more holes per metre than Aliens. The likelihood of stem flagging was lowest on Alien species; the percentage of the total plant that flagged was greatest for Natives. Plant morphology also influenced host preference and likelihood of flagging. 5. Taking plant morphology into consideration, we speculate that the evolutionary history between periodical cicadas and potential hosts is an important component of host preference.     

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.     

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.     

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.

     

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.     

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.     

Resilience of tropical rain forests: tree community reassembly in secondary forests

Understanding the recovery dynamics of ecosystems presents a major challenge in the human-impacted tropics. We tested whether secondary forests follow equilibrium or non-equilibrium dynamics by evaluating community reassembly over time, across different successional stages, and among multiple life stages. Based on long-term and static data from six 1-ha plots in NE Costa Rica, we show that secondary forests are undergoing reassembly of canopy tree and palm species composition through the successful recruitment of seedlings, saplings, and young trees of mature forest species. Such patterns were observed over time within sites and across successional stages. Floristic reassembly in secondary forests showed a clear convergence with mature forest community composition, supporting an equilibrium model. This resilience stems from three key factors co-occurring locally: high abundance of generalist species in the regional flora, high levels of seed dispersal, and local presence of old-growth forest remnants.     

On the dynamics,floristic subdivision and geographical distribution of várzea forests in Central Amazonia

Abstract. Within different stands of the white-water inundation forest (várzea forest) in the Central Amazon region, composition, abundance, frequency and basal area of tree species were recorded. Determinations of age and radial growth rates were conducted using dendrochronological methods. Results show significant differences in age, history and species composition between stands as well as different growth strategies among dominant species. Assignment of tree species to growth strategies by means of anatomical and morphological features together with quantitative aspects of vegetational analysis permit the further differentiation of successional stages of várzea forests. General features of successional stages were quantitatively described and compared with forest types from outside the várzea. Many tree species of the várzea forests are widespread in South America, and not limited to floodplains. Their occurrence on sites with distinct dry seasons suggests that they are not specifically adapted to flooding but are tolerant to seasonality in general.     

Mean latitudinal range sizes of bird assemblages in six Neotropical forest chronosequences

Aim The geographical range size frequency distributions of animal and plant assemblages are among the most important factors affecting large‐scale patterns of diversity. Nonetheless, the relationship between habitat type and the range size distributions of species forming assemblages remains poorly understood. We examined how the mean latitudinal range sizes of species in Neotropical bird species assemblages shift during forest clearance and subsequent regeneration. We tested the hypothesis that bird species assemblages in early successional habitats tend to have larger latitudinal ranges than those in more mature forests. Location We considered breeding bird chronosequence data from six Neotropical forests. Results Breeding bird assemblages were found to have the species with the largest average latitudinal range sizes in cleared areas, intermediate in young secondary forests and smallest in old secondary and mature forests. Similar differences were also found when we compared congeners differing in their successional preferences. Sizes of regional ranges (within the Neotropics) did not, however, differ consistently among successional stages. The larger latitudinal (but not regional) ranges of early successional species was as a result in part of the tendency of early successional species to have ranges that extend beyond the Neotropical forest biome. Conclusions Our analysis of chronosequences suggests that as early successional habitats mature, a consistent shift from large‐ranged species towards more small ranged species occurs. Even relatively old secondary forests have bird species with larger average ranges than mature forests. As a consequence, conservation of secondary forests alone will miss many of the species most at risk of extinction and most unlikely to be conserved in other locations or biomes.     

Feeding ecology and emergence production of annual cicadas (Homoptera: Cicadidae) in tallgrass prairie

The emergence phenology and feeding ecology of annual cicadas in tallgrass prairie are poorly documented. However, these large insects are abundant, and their annual emergence represents a potentially important flux of energy and nutrients from belowground to aboveground. We conducted a study at Konza Prairie Research Natural Area in eastern Kansas to characterize and quantify cicada emergence and associated energy and nutrient fluxes. We established emergence trap transects in three habitat types (upland prairie, lowland prairie, and riparian forest), and collected cicadas every 3 days from May to September. A subset of trapped cicadas was used for species- and sex-specific mass, nutrient, and stable isotope analyses. Five species were trapped during the study, of which three were dominant. Cicadetta calliope and Tibicen aurifera exhibited significantly higher emergence production in upland prairie than in lowland prairie, and were not captured in forested sites at all. T. dorsata emerged from all three habitat types, and though not significant, showed a trend of greater abundance in lowland grasslands. Two less abundant species, T. pruinosa and T. lyricen, emerged exclusively from forested habitats. Nitrogen fluxes associated with total cicada emergence were estimated to be ∼4 kg N ha^–1 year^–1 in both grassland habitats, and 1.01 kg N ha^–1 year^–1 in forested sites. Results of stable isotope analyses showed clear patterns of resource partitioning among dominant cicada species emerging from grassland sites. T. aurifera and C. calliope had δ¹³C and δ¹⁵N signatures indicative of feeding on shallowly rooted C₄ plants such as the warm-season grasses dominant in tallgrass prairie ecosystems, whereas T. dorsata signatures suggested preferential feeding on more deeply rooted C₃ plants. Received: 20 September 1999 / Accepted: 9 December 1999     

Alternate pathways of succession on the Lake Erie Islands

In order to determine the effect of prior land use on secondary succession, woody plant species abundances were determined on the Lake Erie Islands in 21 sites 20–90 yr post-abandonment: 11 former vineyards, 8 former orchards, and 2 former quarries. Detrended Correspondence Analysis ordination was used to compare the relative abundances of woody plants in the 21 successional sites to each other and to 5 mature upland forests and 3 mature lowland forests. Vineyards 20–45 yr post-abandonment had closed canopies 2–4 m tall and were dominated by the same mix of species dominating mature forests at similar elevations. Abandoned orchards 20–45 yr post-abandonment supported species common in mature forest floras as well as many shade-intolerant, pioneer tree and shrub species not found in mature stands. These differences in successional patterns seem to be related to differences in the suite of colonization sites presented upon abandonment, with orchards being dominated by open, bare ground, and vineyards by shaded ground receiving ca. 20% of full sunlight. Newly abandoned vineyards had appreciable ‘advanced regeneration’ of saplings of shade-tolerant species 3–7 yr old growing among the grapes. While former quarries remained open, marginal habitats with sparse, open canopies dominated by species typical of rocky cliff edges, the community structure of former vineyards and orchards converged toward each other and toward the composition of mature forests at similar elevations. Thus patterns of secondary succession varied with prior land use and were determined by the suite of colonization sites available at the time of abandonment.     

Birds,Cattle, and Bracken Ferns: Bird Community Responses to a Neotropical Landscape Shaped by Cattle Grazing Activities

Large areas of tropical moist forests have been converted to cattle pastures, generating complex landscapes where different habitats are represented by small patches with an uneven spatial distribution. Here, we describe how bird communities respond to the different elements present in a livestock landscape that was originally dominated by tropical moist forest. We surveyed six habitats: open pastures, pastures with shrubs, early‐ and middle‐secondary forests, mature forest, and pastures invaded by bracken ferns (Pteridium aquilinum). Bird diversity was high in secondary and mature forests, and low in fern‐invaded sites and open pastures. Fern‐dominated sites had the lowest bird species richness, and trophic guild diversity of all habitats. Habitat structure affected both bird species richness and densities in similar ways. Tree species richness was the habitat attribute that had a bigger positive effect on bird species richness. Bird community structure varied among sampled habitats, separating habitats in two major groups (forests and pastures). Our data indicate that bracken fern‐invaded pastures were the worst habitat condition for avian communities. To increase bird diversity, we recommend to eliminate or manage bracken fern and to increase shrub and tree cover in open pastures to provide food resources and shelter for birds. Finally, we encourage the maintenance of secondary and mature forest remnants as a strategy to conserve resident birds within a landscape dominated by livestock activities.     

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.     

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.     

黑龙江省不同地点蒙古栎林生态特点研究

通过对黑龙江省６个地点的天然蒙古栎林的结构和更新特点的分析,蒙古栎林可划分为不同特点的蒙古栎群落,即纯蒙古栎群落、蒙古栎桦树群落、蒙古栎槭树群落、蒙古栎红松群落和蒙古栎松混交林群落,其演替趋势如下：红蒙古栎群落、蒙古栎桦树群落至蒙古栎槭树落、蒙古栎红松群落,再至蒙古栎红松混交林群落,蒙古栎群落类型的多样性主要反应群落不同的演替阶段,造成蒙古栎群落多样的原因是人类活动和自然因素作用的结果,随着群落的演替,蒙古栎的优势地位逐渐被消弱,乔木种类丰富度增多,草本种类丰富度增多;蒙栎的相对密度下降,林内环境由于干燥逐渐变中性至较湿润,蒙古栎幼苗和幼树在总幼苗和幼树中所占的比例下降,耐荫物中色木槭等的幼苗和幼树所占比例上升;在演替过程中,蒙古栎分布格局－聚集分布的聚集程度逐渐降低,并向随机分布的方向发展。     

On the evolution of cicada X host-tree associations in Central America

An hypothesis tout court is proposed to account for the association of cicadas (Homoptera: Cicadidae) with large-sized trees of the family Leguminosae in both climax forest formations and forest remnants in Central America. Based upon extensive field studies of these insects in Costa Rica, final-instar nymphal skins of many genera and species of Neotropical cicadas appear to be aggregated annually around individual large legume trees in a broad range of habitats in Costa Rica, and much of the reproductive behavior of the adult insects is also centered around these trees. A series of testable subhypotheses are generated based largely upon the assumption that cicadas, primarily through egg-placement behavior and nymphal feeding behavior, have coevolved with legume trees in climax forest formations of Central America, and perhaps elsewhere in the Neotropical Region. The root crowns of such trees, with some exceptions, might be optimal feeding sites for cicada nymphs in tropical forests.     

长白山针阔混交林不同演替阶段的昆虫多样性

昆虫多样性变化对生态系统健康有重要的指示作用, 为研究昆虫群落变化与生境演替之间的关系, 本研究采用网捕、灯诱和诱捕法系统调查了长白山针阔混交林不同演替阶段（次生白桦林、次生针阔混交林、原始阔叶红松林）昆虫群落的组成和多样性, 分析了昆虫在森林演替过程中的规律及与植被群落之间的关系。系统调查共采集昆虫标本8 183头, 隶属于14个目699种, 其中鳞翅目和鞘翅目是主要优势类群。次生针阔混交林昆虫的个体数量最多, 原始阔叶红松林中物种数最多。不同演替阶段昆虫群落的物种数和个体数差异不显著, 但次生针阔混交林、原始阔叶红松林的Fisher’s α指数显著高于次生白桦林。目水平上的昆虫多样性未表现出显著性差异. 昆虫多样性在森林演替过程中和草本植物多样性的变化趋势相同;由于食性和生境选择的不同, 森林演替过程中鳞翅目昆虫多样性逐渐升高, 而鞘翅目多样性逐渐降低。