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448! ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 104, no. 3
G F F A ? ? ? 13-year species 11 12 13 14 15 16 17 18 19 20 21 22
17-year species 11 12 13 14 15 16 17 18 19 20 21 22
B C D E ! Fig. 2. Evidence for life-cycle length plasticity in 13- and 17-yr cicadas from experiments or from observation of cicadas emerging off-schedule! in numbers too large to be plausibly accounted for by mutation. Evidence is limited because most historical records do not include data on emergence density. Overall, the evidence is strongest for 1-yr premature and/or delayed emergences<$#-+3'= and,!+D)$/!'%0I*)4%15!/D#!I*)0/%(%/4!&.!HQ for jumps of 4 yr. Supporting data_ for!)1'!HR each_!4#)$!*%.#!(4(*#P!!?&3$(#,! capital-letter-labeled life-cycle variant are as follows: (A) Data from this study, including speciÞcinformationformultiple13-yrspecies.(B)Manypublishedexamples(Dybas 1969, Kritsky 1988,L)$0D)**! Kritsky3*''"68 andA!!cFGHHdP! Simon 1996Ñ! M. cassini,Marshalletal.2003citingG.KritskyÑallthreespecies,Zyla2004), including speciÞcevidenceforlargenumbersofbothM. septendecim and M. cassini in the 1969 example according to Simon and Lloyd (1982).! (C) Maier (1985) and multiple unpublished observations (see text). (D) Published observations (Marlatt 1907, p. 24; Boyd 1952) and experimental data (Karban et al. 2000). (E) Published observations including White and Lloyd (1975, 1979), but! see Kritsky (2004). Indirect evidence suggesting 1- and/or 4-yr shifts in emergence timing (including F) is found in the biogeography of the various broods of both life cycles, which are generally assumed to have originated via mass shifts in life-cycle! timing (Marlatt 1907, Alexander and Moore 1962, Lloyd and Dybas 1966a, Lloyd and White 1976). Eight-year-delayed emergence of 13-yr species (G) has been suggested by Marshall (2001) based on historical data, and it is weakly suggested by our observations of adult cicadas in 2006 within the range of brood XIX. of brood XIX in 2006, 8 yr after an emergence of that Different Magicicada species with the same life cycle brood, suggests that additional search efforts aimed at seem to respond similarly to environmental triggers af- uncovering such developmental plasticity would be fecting emergence timing. All three 17-yr species have worthwhile. Multiple-year premature emergence of been observed emerging together after 13 yr, and we 13-yr cicadas also has not yet been documented. observed all four 13-yr species during our search for Straggling events involving large numbers of cicadas delayed cicadas in 2006 (see references in Fig. 2 legend). are not well established for differences other than plus or M. tredecula was observed only as singletons in 2006, but minus 4 yr, for either life-cycle type. White and Lloyd this may be attributed to the fact that the -decula siblings (1979) reported an example of millions of cicadas from are usually rare compared with the other Magicicada brood XIV emerging 1 yr late, but Kritsky (2004) (p. 105) species (Alexander and Moore 1962, Lloyd and White has since argued that the cicadas belonged to a previ- 1983). The fact that we heard more M. tredecassini strag- ously unrecognized, disjunct population of 13-yr brood glers may be due to the louder, higher pitched, and more XXII. A review of historical records in Indiana (Kritsky obvious song of this species. Approximately equal num- 1987) showed 1- and 4-yr accelerations to be most com- bers of M. 13-yr -decim and M. tredecassini or M. tredecula mon in 17-yr cicadas, but most of the records do not were observed in the exuviae counts from Giant City include data on emergence density. State Park. Although we did not measure the number of 13-yr Parallel 13- and 17-Yr Developmental Plasticity and cicadas that emerged at our study sites 1, 2, and 3 yr Magicicada Life-Cycle Evolution. Many evolutionary late, a numerical exercise suggests that our 2006 ob- mechanisms and historical scenarios have been of- servations probably did not represent the tail end of fered to explain the origin of the seven periodical normally distributed life-cycle variation. Given a cicada species and their long, prime-numbered life mean life-cycle length of 13 yr, an assumption of nor- cycles. All begin with a nonperiodical ancestor with a mally distributed life-cycle variation, and an assumed shorter cycle length more like those observed today in population density of 200,000 per ha (toward the high other temperate zone cicadas (i.e., up to 11 yr, Karban end of the published range), an SD of 1.505 is neces- 1997). Some scenarios derive the Þrst fully periodical sary to account for 1,724/200,000 ϭ 0.86% of the pop- 13- and 17-yr forms independently (Cox and Carlton ulation emerging 4 yr late. But under these assump- 1988, Yoshimura 1997), whereas others propose the tions tens of thousands of cicadas (21% of the evolution of periodicity Þrst in one form (e.g., a 13-yr population) would emerge 1 yr late, and 11% would ancestor), followed by a life-cycle shift to derive the emerge 2 yr late, etc. White and Lloyd (1975) found other form (Marlatt 1907, Lloyd and Dybas 1966a). just 0.7% of 17-yr cicadas emerging in the 18th year in Developmental plasticity plays an important role in astudyplotbeneathanappletree,inanabandoned hypotheses of both types, but its generality has been orchard where they believed delayed development limited by the absence, until now, of evidence that the was especially likely. SigniÞcantly, when they dug up extant 13-yr species possess latent potential to emerge their study plot in the 19th year they found evidence in 17 yr. For example, Yoshimura (1997) proposed a suggesting a bimodal distribution of emergence theory deriving long, periodical life cycles indepen- timesÑÞfth-instar nymphs, amounting to 1.7% of the dently during Pleistocene cold shocks that would have population, still in the ground. nonadaptively extended life length, increased juvenile ! [!
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I$#')/&$0!D)'!.%**#'!/D#%$!0/&:)(D0!)1'!(&3*'!)..&$'!/&!N#!:&$#!0#*#(/%9#!%1!/D#%$! I$#')/%&1P!!6D#0#!'#.#10#!(D)$)(/#$%0/%(0!@#$#!.&31'!)*:&0/!#T(*30%9#*4!%1!:)*#0A! @D%(D!:)4!D)9#!)00%0/#'!/D#!N*)( &1*4!/%:#!)!.#:)*#!(%()')!@)0!&N0#$9#'!/&!N#!'#.#10%9#!%0!@D#1!0D#!@)0!)//#:I/%15! /&!'#I&0%/!D#$!#550!%1!)!4&315!/@%5P!!O.!)!I$#')/&$!@#$#!/D$#)/#1%15!/&!(&:#!(*&0#A! /D#!.#:)*#!@&3*'!%::#'%)/#*4!/)<#!.*%5D/!c?/#@)$'!3*'"68!!H]]^dP!!L)*#!(%()')0!@#$#! &N0#$9#'!/&!'#:&10/$)/#!#%5D/!'%..#$#1/!$#)(/%&10!@D#1!)II$&)(D#'!N4!)!/D$#)/P!! 6D&0#!$#)(/%&10!@#$#,!!@)* 083)@ /)(/%(0A!/D#!*#)0/!#..#(/%9#!:)1#39#$!'%0I*)4#'!@)0!/&!$#:)%1!%1)(/%9#!5%9%15!/D#! I$#')/&$!)1!#)04!/)$5#/!.&$!(&103:I/%&1P!!-9#$4!%10#(/!/D)/!0/)4#'!:&/%&1*#00!%1! .$&1/!&.!I$#4!@)0!()I/3$#'A!@D%*#!&9#$!][l!&.!/D#!%10#(/0!/D)/!'%0I*)4#'!/D#!.*4!)1'! 083)@ /D#%$!I$#')/&$0!@#$#!)N*#!/&!03$9%9#!)1'!I)00!&1!/D#%$!5#1#0!/&!/D#!1#T/!5#1#$)/%&1! &.!(%()')0P!!6D$&35D&3/!/D#!#:#$5#1(#!I#$%&'!$#'_@%15#'!N*)( 0#*#(/%9#!/D)/!/D#4!@&3*'!0#*#(/!/D#!(%()')0A!@D%(D!'#:&10/$)/#'!/D#!.#@#0/!'#.#10#! /)(/%(0P!!! ! ! ! ! ! ! H]! I"C63!F,!?3((#00.3*!)1'!3103((#00.3*!)//)( !Table 4. Percent attack success of red-winged blackbirds on period- tion), the high percentage of inactive females in mid-May ical cicadas as influenced by cicada behavior may be due to the large number of young adult females in the population. Also, female periodical cicadas signal Predator Cicada behavior exhibited - sexual receptivity by remaining motionless when ap- attack (%) proached by a male (Dunning et al. 1979), so that the higher Inactive Fly Fly-Fall Fly-Squawk of motionless females in mid- to late (N=55) (JV=111) (N=53) (JV=22) percentage May may be due to a large number of unmated females in the popula- Successful 100 44.1 56.6 4.6 tion. By the end of May, all females had emerged on our study site and both the number and sizes of chorus centers, Unsuccessful 0 55.9 43.4 95.4 ! where mating occurs, were decreasing rapidly in early June ! (Williams and Smith, unpublished work). At the end of the model predator experiments, relatively few females re- (9;"%*',)'G6")*:'")&'L-*+$3)*'45%6$)#Table 5. Percent periodical cicadas' exhibiting predator-avoidance mained inactive since most had mated and were in the pro- behavior (as described in text) to attacking red-winged blackbirds cess of ovipositing eggs. Ovipositing female cicadas are very ! different during time-periods skittish : on 20 occasions, we attempted to approach a fe- ! 70!:#1/%&1#'!#)$*%#$A!/D#!N#*&@5$&31'!*%.#!&.!)!(%()')!I$&9%'#0!)!31%83#! male that had her ovipositor embedded in a twig, but never Time-period ? Inactive Fly Fly-Fall Fly- got closer than 1 m before she flew away. Squawk &II&$/31%/4!.&$!)!(4(*%15!&.!/D#!N#*&@!5$&31'!13/$%#1/0!/&!/D#!0&%*!03$.)(#P!!+%()')0! Males also remained inactive less often in early June, due to their and the that 0I#1'!1#)$*4!/D#%$!#1/%$#!*%9#0!.##'%15!&1!/D#!Tlow bird 62 27.4 30.74*#:!&.!/$##!$&&/0P!!`D%*#! 19.3 22.6 probably age dynamic changes were in the location of chorus centers at that pr?dation occurring 31'#$5$&31'!/D#!(%()')!5�!/D$&35D![!(28-30 May) ():*"+!0/)5#0!&.!5$&@/D!c`D%/#!H]R[dP!!6D#!time (Williams and Smith, unpublished work), and squawks were associated with 65% of their avoidance behav- moderate bird 105 19.0 52.4 21.9 6.7 nearly .%./D!0/)5#!&.!'#9#*&I:#1/!()1!*)0/!)0!*&15!)0![!4#)$0P!!O1!/D#!.%./D!0/)5#!/D#!(%()')! iors at that time. cicadas from pr?dation Squawking very rarely escape birds once are communi- @%**!$#)(D!%/0!.%1)*!14:ID!0/)5#!N#.&$#!%/!#:#$5#0!.$&:!/D#!5(31 May to 3 June) $&31'!)1'!:&*/0!%1/&!)1! they captured (M. Lloyd, personal cation; V.B. Steward, personal observation), suggesting high bird 74 24.3 50.0 24.3 1.4 )'3*/!(%()')P!!! that squawking apparently does not startle avian predators pr?dation (but see Smith and Langley 1978). Rather, squawking more !(7-9 June)O1!/D#!.%./D! $):*"+!0/)5#A!/D#!(%()')!@%**!$#)(D!.3**!5$&@/D!)1'!@%**!D)9#! likely alerts a predator that it is about to prey upon a male )//)%1#'!1#)$*4!)**!&.!%/0!13/$%/%&1!.$&:!/D#!$&&/!T4*#:!)1'!/D#!0&%*!)$&31'!%/P!!`%/D! cicada. Since males contain little more than air in their abdomens, birds probably learn to identify females, /D#!(%()')!*%9%15!%/0!*%.#A!&1!)9#$)5#A!I$%&$!/&!#:#$Avoidance behavior used by cicadas5#1(#!)N&3/!HG!/&!MG!(:!N#*&@!/D#! prior to attacks which are more nutritious than males (Brown and Chippen- from red-winged blackbirds also changed over the study dale 1973), a conclusion also reached by Karban (1983). 03$.)(#A!/D#!13/$%#1/0!/D)/!/D#!(%()')!N$%150!/&!/D#!03$.)(#!)$#!/4I%()**4!1&/!)9)%*)N*#! period in early June (Table 5). More cicadas flew silently Possible cues that birds could use to discriminate sexes in- and fewer cicadas when flew /&!0D)**&@!$&&/#'!I*)1/0P!significantly ! squawked they clude squawks made by males and the larger size and less = over time (^ 23.25, df=2, P<0.001). Numbers of cica- aggressive behaviors (Karban 1983) of females. das remaining inactive and making fly-fall responses re- During the period of increasingly heavy bird pr?dation mained relatively constant over the study period. in June, which occurred after peak cicada abundance, more cicadas that were attacked by blackbirds were silent (Ta- ble Discussion 5). This could reflect a shift in the sex ratio in favor of females. Since males emerged before females, we assume In contrast to the conclusions of Lloyd and Dybas (1966 b), that males started to die before females. Alternatively, the periodical cicadas appear to exhibit some predator-avoid- great decrease in June in the number of cicadas that ance behaviors. When approached by a model predator, squawked when attacked by blackbirds could be the result most males produced a squawk and > 50% of the females of blackbirds preferentially preying on females and not at- demonstrated some type of predator-avoidance behavior. tacking male cicadas. Since nearly all males cicadas that However, it would appear that, even though periodical cica- squawked when attacked were not captured and all silent, do das possess some predator-avoidance behaviors, they motionless cicadas that were attacked were successfully cap- still are relatively easy to capture. Red-winged blackbirds tured, it would appear that blackbirds were able to discrimi- quickly learned to catch periodical cicadas, a prey item that nate between male and female cicadas and were preferen- they had never encountered, and had relatively high capture tially eating females. rates for a species that does not ordinarily forage in trees. The significant increase in handling times of cicadas by There also seemed to be no evidence that sound produced birds also reflects the change to more efficient consumption by male cicadas in the chorus centers deterred pr?dation of female cicadas. Initially, blackbirds were observed re- by blackbirds. moving the wings from cicadas and consuming the entire Changes that appeared in reactions of periodical cicadas insect. During the period of heaviest avian pr?dation, birds to the model predator and to the attacks of blackbirds re- were seen removing the wings, slitting the abdomen with flect both changes in the sex ratio of the cicada populations their bills, eating the contents of the abdomen, and discard- and changes in behaviors of cicadas associated with mating ing the exoskeleton. These results are consistent with the and egg laying. At the start of the model predator experi- prediction made by Sih (1980) that a lower proportion of ments, the sex ratio of the cicada population was biased each prey item maybe consumed during periods of prey towards males. Nearly all male cicadas had emerged by abundance. He also predicted that handling time would mid-May on our study site, but females continued to emerge decrease, whereas, our results demonstrate that avian han- until late May. Since young adult cicadas tend to be less dling times increased when periodical cicadas were partially active than older ones (M. Lloyd, personal communica- consumed. Birds, however, were probably still maximizing This content downloaded from 128.143.23.241 on Mon, 12 Aug 2013 19:50:30 PM All use subject to JSTOR Terms and Conditions ! FG! ! O1!)!/#0/!)$#)!/D)/!@)0!I#$.&$:#'!%1!)1!&$(D)$'!)1'!1&1_&$(D)$'! #19%$&1:#1/A!%/!@)0!.&31'!/D)/!/D#!)9#$)5#!)'3*/!(%()')0!N&'4!4%#*'#'!/D#!.&**&@%15! 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N)(/#$%)!)1'!.315%!@%/D%1!F!/&!Q!@##<0!&.!/D#!()$()00#0!*)1'%15!&1!/D#!.&$#0/!.*&&$! ci)15!FGGMdP!!`%/D!/D%0!1%/$&5#1!%1($#)0#A!$#03*/0!0D&@#'!/D)/!0:)**#$!I*)1/0!03(D!)0! /D#!N#**.*&@#$!I$&'3(#'!0##'0!/D)/!@#$#!]l!*)$5#$!/D)1!(&1/$&*!5$&3I0P!! 96 ! L. H. Yang Fig. 1 a Intraspecific competition between neighboring bellflower rosettes decreased plant biomass, while fertilization with dead cicada carcasses increased bellflower biomass. b Cicada fertilization increased foliage nitrogen concentration (i.e., decreased nitrogen use efficiency). The effect of competition on foliage nitrogen concentration was not significant ! ! p = 0.038). Competition did not affect foliage nitrogen declined. Future studies should aim to examine the effects concentrations (Fig. 1b, F1,53 = 0.40, p = 0.53). These of pulsed resources on competition using longitudinal experiments were unable to detect<$#-+3 any!M,!C)$!J$)ID0!&.!C#**.*&@#$!N%&:)00P! effect of cicada designs that quantify consumer!+&:I)$%15!%1/$)0I#(%.%(!(&:I#/%/%&1! responses through time. fertilization on the biomass asymmetry of bellflower pairs The second possibility is that the competitive interac- when grown in competition (p = @%/D!1#%5DN&$0!)1'!.#$/%*%W)/%&1!@%/D!(%()')0P!!?&3$(#,!i)15A!cFGHFdP!0.38). tions among conspecifics could have been undiminished! even if belowground resources were supplied in excess of ! demand, if competition shifted toward some other limiting Discussion resource. In general, if specific resources are supplied in ! 7/!/D#!(D&$30!(#1/#$0!@D#$#!13:N#$0!()1!N#!%1!/D#!/D&30)1'0!&$!#9#1!excess of demand, competitive interactions are generally In this experiment, the addition of cicada carcasses sub- expected to shift to other niche dimensions (e.g., Harpole stantially increased plant biomass, while competition and Tilman 2007). For example, strong resource enrich- decreased plant biomass. The positive:%**%&10A!(%()')0!@%**!(&103:#!0:)**!):&31/0!&.!*#)9#0!)1'!/D#!.#:)*#0!@%**!'# effects of fertiliza- ment in the belowground environment commonly increases I&0%/! tion on plant biomass were counterbalanced by the nega- aboveground competition for light (Weiner 1986; Wilson tive effects of intraspecific competition/D#%$!#550!%1!4&315!N$)1(D#0!)1'!/@%50!/D)/!()1!$#03*/!%1!/D#!'#)/D!&.!/D#!/@%5P!!`%/D! in an additive and Tilman 1993). In this experiment, the cumulative manner. This suggests that the addition of cicada carcasses effects of intraspecific competition could have been undi- increased plant growth, but did not:)00!13:N#$0A!%/!0##:0!*&5%()*!/D)/!/D#$#!@&3*'!N#!)!1&/%(#)N*#!'#($#)0#!%1!I*)1/! change the proportional minished if reduced nutrient limitations belowground effects of competition. In addition, cicada supplementation resulted in increased light limitations aboveground. How- increased foliage nitrogen concentrations in both compet- ever, competition for light is often considered to be fun- itive and non-competitive environments. This result cor- damentally asymmetric (Weiner 1990), such that increased responds to a pattern of decreased nitrogen use efficiency light competition would be associated with increased bio- under cicada fertilization. mass asymmetry between competing plants in the cicada- The observation that competition was not significantly fertilized group. This biomass asymmetry is generally altered by cicada resources suggests at least two possibil- expected to be self-reinforcing because larger plants shade ities. First, this pattern could result if the observed smaller plants, and greater exposure to light allows more increases in plant resource consumption (due to increased biomass accumulation, as has been shown for bellflower growth and reduced nitrogen use efficiency) occurred at a rosettes (Galloway and Etterson 2009). However, these timescale commensurate with the timescale of cicada- current analyses did not find a significant increase in the detrital decomposition. If this were the case, then cicada biomass asymmetry of competing plants in the cicada fertilization (while pulsed relative to the entire biennial life fertilization treatment, suggesting that belowground fertil- history of the plant) did not present nutrient resources in ization may not have caused a significant shift toward excess of consumer demand at the timescale of this increased competition for light, or that the high light con- experiment. This study did not attempt to determine whe- ditions of the experiment may have reduced the importance ther there may have been a transient period during which of light competition. nutrient supply temporarily exceeded consumer demand, or Taken together, these results suggest that the consumers whether plant requirements that increased under pulsed in this system (American bellflowers) responded to the conditions (due to increased biomass or reduced nutrient pulsed increase in belowground resources (decomposing use efficiency) would ultimately have contributed to more cicada carcasses) with relatively rapid increases in nutrient intense competition as the availability of resources uptake through both increased growth and reduced nutrient 123 ! FQ! 5$&@/D!)1'!@&&'!)((3:3*)/%&1!&9#$!/D#!.&**&@%15!4#)$!&.!)!I3*0#'!(%()')! #:#$5#1(#A!N3/!$#0#)$(D!D)0!0D&@1!&/D#$@%0#P!!!O1!/D#!4#)$0!.&**&@%15!)!(%()')! #:#$5#1(#A!/D#$#!%0!)1!%1($#)0#!%1!/D#!):&31/!&.!@&&'!)((3:3*)/%&1!)1'!I$%:)$4! 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F; This content downloaded from 128.143.23.241 on Mon, 12 Aug 2013 19:45:24 PM All use subject to JSTOR Terms and Conditions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`D)/!/4I%()**4!/)<#0!)! :&1/D!%1!/#$$#0/$%)*!#(&040/#:0!&1*4!/)<#0!&1#!/&!/@&!@##<0!%1!/D#!@)/#$P!!6D#! #..#(/0!&.!/D#!)''%/%&1!&.!(%()')0!%1!)83)/%(!#19%$&1:#1/0!%0!#)0%#$!/&!/$)(#!0%1(#!/D#! I$&(#00!&((3$0!&9#$!)!83%(<#$!I#$%&'!&.!/%:#A!)0!0D&@1!%1!K%53$#!\A!)!D%5D#$! I#$(#1/)5#!&.!(%()')!13/$%#1/0!%0!%1'%$#(/*4!*%1<#'!/&!)83)/%(!I$#')/&$0P!!!! ! ! QH! March 2008 THE ECOLOGY OF RESOURSE PULSES 651 ! FIG. 1. Diagram of predicted responses of terrestrial and aquatic systems to a resource pulse event, such as water or dissolved labile nutrients,<$#-+3 that!^,!!`#N!&.!I3*0#!$#0&3$(#!(4(*%15!%1!/#$$#0/$%)*!)1'!)83)/%(!#19%$&1:#1/0P!!! affects the biomass and productivity of primary producers. Pathways of nutrients and energy between trophic levels are illustrated with arrows. The thickness of the arrows indicates the relative rapidity and magnitude of nutrient and energy S flows throughY%1#!/D%(<1#00!$#I$#0#1/0!/D#!#..#(/!/D)/!I$&(#00!D)0P!!?&3$(#,!B&@*%1! food webs between the specific ecosystem type (i.e., the thicker the line, the faster and/or3*'"68 greaterA!!cFGG^dP! the effect).! PECIAL reduced! intensity of top-down consumer effects in populations in response to resource pulses may translate terrestrial systems relative to aquatic systems. into new biomass for herbivores more rapidly than F terrestrial plants. Pelagic food webs are also highly size EATURE PredictingM)%3+*"$)*$3: the speed' and persistence of aquatic structured; in general, body size increases with trophic and terrestrial responses to resource pulses level (Brose et al. 2006, Shurin et al. 2006) and this size Ostfeld' and Keesing (2000) suggest that community- structure may contribute to more efficient energy level responses to resource pulses depend on three transfer in pelagic systems (Hairston and Hairston interrelated! characteristics70!:3(D!/D)/!%0!<1&@1!)N&3/!/D#!31%83#!*%.#!)1'!#..#(/0!&.!/D#! of consumers in food webs 1993). In contrast, terrestrial!"#$%$%"&": herbivores can! be smaller receiving the resource pulse: (1) the degree of speciali- than plants (e.g., phytophagous insects vs. trees) or zation* on%.#!)1'!#19%$&1:#1/)*!%:I)(/A!/D#$#!%0!#9#1!:&$#!31(#$/)%1/%#0!/D)/!*&&:!&9#$!/D#! the pulsed resource, (2) the rate of consumer larger than plants (e.g., ungulate grazers vs. grasses and population increase in response to the pulse, and (3) the forbs) (Shurin et al. 2006). Benthic primary producers in mobility of consumers in response to the pulse. Are these aquatic systems typically exhibit traits intermediate of features%10#(/P!!e)9%15!03(D!)!31%83#!*%.#!(4(*#A!$#0#)$(D!&1!/D#!%10#(/!%0!*%:%/#'!/&!&1*4!)!.#@! also important in mediating the response of those in pelagic and terrestrial systems, in terms of size, aquatic food webs to resource pulses? Here, we consider growth, and stoichiometry (Shurin et al. 2006). how differences:&1/D0!#9#$4!HQ!&$!HR!4#)$0P!!6D#!31'#$5$&31'!*%.#!0I)1!&.!/D#!%10#(/!%0!&1#!&.!/D#! in the size structure and growth rates of Due to these energetic, stoichiometric, and size-related organisms, nutrient stoichiometry and chemical compo- differences, the proportion of primary production sition of*)$5#0/!31(#$/)%1/%#0!&.!/D#!%10#(/P!!70%'#!.$&:!0I#1'%15!:&0/!&.!%/0!*%.#!)$&31'!/$##! primary producers, and the sensitivity of each consumed by herbivores is approximately three times system type to bottom-up and top-down forces may greater in pelagic than in terrestrial and macrophyte- mediate$&&/0!)1'!*%9%15!&..!&.!T4*#:!/D#$#!%0!*%//*#!/D)/!%0!<1&@1!)N&3/!/D%0!I#$%&'!&.!)! community responses to pulsed resources. dominated systems (Cyr and Pace 1993, Cebrian and Generation times and tissue turnover rates of aquatic Lartigue 2004). Thus, it is likely that primary producers organisms are more rapid than their terrestrial counter- in pelagic and terrestrial food webs respond differently parts (Persson(%()')0!*%.#!0%1(#!/D#4!)$#!D)$'!/&!/$)( ! 7N&9#!5$&31'!/D#$#!)$#!)*0&!:)14!31(#$/)%1/%#0!)1'!I$&N*#:0!@D%(D!&((3$! .$&:!/D#0#!#:#$5#1(#0P!!E1'#$0/)1'%15!/D#!N#D)9%&$0!&.!'%..#$#1/!I$#')/&$0A!'3$%15! /D%0!#:#$5#1(#!I&0#0!I$&N*#:P!!O1!$#0#)$(D!I#$.&$:#'!N4!?/&$:!3*'"687!)1%:)*0!03(D! )0!/D#!5$)4!083%$$#*!@#$#!&N0#$9#'!#)/%15!(%()')0!%1!/D#!.%#*'A!N3/!/D#!5$)4!083%$$#*0! 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