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Auditory behaviour of a parasitoid fly (Emblemasoma auditrix, Sarcophagidae, Diptera).

Females of the parasitoid fly Emblemasoma auditrix find their host cicada (Okanagana rimosa) by its acoustic signals. In laboratory experiments, fly phonotaxis had a mean threshold of about 66 dB SPL when tested with the cicada calling song. Flies exhibited a frequency dependent phonotaxis when testing to song models with different carrier frequencies (pulses of 6 ms duration and a repetition rate of 80 pulses s(-1)). However, the phonotactic threshold was rather broadly tuned in the range from 5 kHz to 11 kHz. Phonotaxis was also dependent on the temporal parameters of the song models: repetition rates of 60 pulses s(-1) and 80 pulses s and pulse durations of 5-7 ms resulted in the highest percentages of phonotaxis performing animals coupled with the lowest threshold values. Thus, parasitoid phonotaxis is adapted especially to the temporal parameters of the calling song of the host. Choice experiments revealed a preference of a song model with 9 kHz carrier frequency (peak energy of the host song) compared with 5 kHz carrier frequency (electrophysiologically determined best hearing frequency). However, this preference changed with the relative sound pressure level of both signals. When presented simultaneously, E. auditrix preferred 5-kHz signals, if they were 5 dB SPL louder than the 9-kHz signal.

Animal Communication↗

Temporally patterned sound pulse trains affect intensity and frequency sensitivity of inferior collicular neurons of the big brown bat, Eptesicus fuscus.

This study examined the effect of temporally patterned pulse trains on intensity and frequency sensitivity of inferior collicular neurons of the big brown bat, Eptesicus fuscus. Intensity sensitivity of inferior collicular neurons was expressed by the dynamic range and slope of rate-intensity functions. Inferior collicular neurons with non-monotonic rate-intensity functions have smaller dynamic ranges and larger slopes than neurons with monotonic or saturated rate-intensity functions. Intensity sensitivity of all inferior collicular neurons improved by increasing the number of non-monotonic rate-intensity functions when the pulse repetition rate of pulse trains increased from 10 to 30 pulses per second. Intensity sensitivity of 43% inferior collicular neurons further improved when the pulse repetition rate of pulse trains increased still from 30 to 90 pulses per second. Frequency sensitivity of inferior collicular neurons was expressed by the Q10, Q20, and Q30 values of threshold frequency tuning curves and bandwidths of isointensity frequency tuning curves. Threshold frequency tuning curves of all inferior collicular neurons were V-shape and mirror-images of their counterpart isointensity frequency tuning curves. The Q10, Q20, and Q30 values of threshold frequency tuning curves of all inferior collicular neurons progressively increased and bandwidths of isointensity frequency tuning curves decreased with increasing pulse repetition rate in temporally patterned pulse trains. Biological relevance of these findings to bat echolocation is discussed.

Animals↗

Freshwater to seawater acclimation of juvenile bull sharks (Carcharhinus leucas): plasma osmolytes and Na+/K+-ATPase activity in gill, rectal gland, kidney and intestine.

This study examined the osmoregulatory status of the euryhaline elasmobranch Carcharhinus leucas acclimated to freshwater (FW) and seawater (SW). Juvenile C. leucas captured in FW (3 mOsm l(-1) kg(-1)) were acclimated to SW (980-1,000 mOsm l(-1) kg(-1)) over 16 days. A FW group was maintained in captivity over a similar time period. In FW, bull sharks were hyper-osmotic regulators, having a plasma osmolarity of 595 mOsm l(-1) kg(-1). In SW, bull sharks had significantly higher plasma osmolarities (940 mOsm l(-1) kg(-1)) than FW-acclimated animals and were slightly hypo-osmotic to the environment. Plasma Na(+), Cl(-), K(+), Mg(2+), Ca(2+), urea and trimethylamine oxide (TMAO) concentrations were all significantly higher in bull sharks acclimated to SW, with urea and TMAO showing the greatest increase. Gill, rectal gland, kidney and intestinal tissue were taken from animals acclimated to FW and SW and analysed for maximal Na(+)/K(+)-ATPase activity. Na(+)/K(+)-ATPase activity in the gills and intestine was less than 1 mmol Pi mg(-1) protein h(-1) and there was no difference in activity between FW- and SW-acclimated animals. In contrast Na(+)/K(+)-ATPase activity in the rectal gland and kidney were significantly higher than gill and intestine and showed significant differences between the FW- and SW-acclimated groups. In FW and SW, rectal gland Na(+)/K(+)-ATPase activity was 5.6+/-0.8 and 9.2+/-0.6 mmol Pi mg(-1) protein h(-1), respectively. Na(+)/K(+)-ATPase activity in the kidney of FW and SW acclimated animals was 8.4+/-1.1 and 3.3+/-1.1 Pi mg(-1) protein h(-1), respectively. Thus juvenile bull sharks have the osmoregulatory plasticity to acclimate to SW; their preference for the upper reaches of rivers where salinity is low is therefore likely to be for predator avoidance and/or increased food abundance rather than because of a physiological constraint.

Adaptation, Physiological↗

Dynamics of blood chylomicron fatty acids in a marine carnivore: implications for lipid metabolism and quantitative estimation of predator diets.

Blubber fatty acid(s) (FA) signatures can provide accurate estimates of predator diets using quantitative FA signature analysis, provided that aspects of predator FA metabolism are taken into account. Because the intestinal absorption of dietary FA and their incorporation into chylomicrons (the primary transport lipoproteins for dietary FA in the blood) may influence the relationship between FA composition in the diet and adipose tissue, we investigated the metabolism of individual FA at these early stages of assimilation. We also investigated the capacity of chylomicron signatures to provide quantitative estimates of prey composition of an experimental meal. Six captive juvenile grey seals (Halichoerus grypus) were fed either 2.3 kg (n = 3) or 4.6 kg (n = 3) of Atlantic herring (Clupea harengus). Although chylomicron FA signatures resembled diet signatures at all samplings, absolute differences were smallest at 3-h post-feeding, when chylomicrons were likely largest and had the greatest ratio of triacylglycerol to phospholipid FA. Specific FA that differed significantly between diet and chylomicron signatures reflected either input from endogenous sources or loss through peroxisomal beta-oxidation. When these aspects of metabolism were accounted for, the quantitative predictions of diet composition generated using chylomicron signatures were extremely accurate, even when tested against 28 other prey items.

Adipose Tissue↗

The aerobic physiology of the air-breathing blue gourami, Trichogaster trichopterus, necessitates behavioural regulation of breath-hold limits during hypoxic stress and predatory challenge.

Physiological characteristics of the blood oxygen transport system and muscle metabolism indicate a high dependence on aerobic pathways in the blue gourami, Trichogaster trichopterus. Haemoglobin concentration and haematocrit were modest and the blood oxygen affinity (P50=2.31 kPa at pH 7.4 and 28 degrees C) and its sensitivity to pH (Bohr factor, phi=-0.34) favour oxygen unloading at a relatively high oxygen pressure (PO2). The intracellular buffering capacity (44.0 slykes) and lactate dehydrogenase (LDH) activity (154.3 iu g(-1)) do not support exceptional anaerobic capabilities. Air-breathing frequency in the blue gourami is expected to increase when aquatic oxygen tensions decline. Under threat of predation, however, this behaviour must be modified at a potential cost to aerobic metabolism. We therefore tested the hypothesis that metabolic responses to predatory challenge and aquatic hypoxia are subject to behavioural modulation. Computer-generated visual stimuli consistently reduced air-breathing frequency at 19.95, 6.65 and 3.33 kPa PO2. Bi-directional rates of spontaneous activity were similarly reduced. The metabolic cost of this behaviour was estimated and positively correlated with PO2 but not with visual stimulation thus indicating down-regulation of spontaneous activity rather than breath-holding behaviour. Neither PO2 nor visual stimulation resulted in significant change to muscle lactate and ATP concentrations and confirm that aerobic breath-hold limits were maintained following behavioural modulation of metabolic demands.

Adaptation, Physiological↗

The blowfly Chrysomya albiceps (Wiedemann) (Diptera: Calliphoridae) as a new forensic indicator in Central Europe.

During the summer months of the year 2001, six forensic cases (one is reported in the present paper), a pig carrion study in the city of Vienna (latitude 48 degrees 12'N, longitude 16 degrees 22'E) and several liver-baited traps north of Vienna, yielded large numbers of maggots of the blowfly Chrysomya albiceps (Wiedemann) (Diptera: Calliphoridae). Apart from some records from France, reports of C. albiceps from the palearctic region north of the Alps (i.e. north of a latitude of 48 degrees N) have been scarce. Our findings provided an opportunity to derive developmental schedules for C. albiceps at five different constant temperatures (15, 20, 25, 30, 35 degrees C). The minimal duration of development from oviposition to adult was inversely related to temperature, ranging from 8.3 +/- 0.5 days at 35 degrees C to 19.2 +/- 0.92 days at 20 degrees C. Although eggs hatched after 1.9 +/- 0.16 days at 15 degrees C, larvae did not complete development and frequently died during the first instar stage. We also found a high mortality rate (up to 99%) of native L. sericata larvae caused by predation of C. albiceps larvae under laboratory conditions, indicating a high susceptibility of L. sericata to attack by C. albiceps. Apart from this, the current and possible future distribution of C. albicepsin Europe is discussed. The northward expansion of its range beyond southern Europe obviously decreases the value of C. albiceps in estimating the site of death, in that it is no longer exclusive to southern European regions. Moreover, the aggressive feeding behaviour of second and third instar larve of C. albiceps could reset the post-mortem insect clock by clearing a corpse of all earlier arrivers.

Animals↗

Slippery surfaces of carnivorous plants: composition of epicuticular wax crystals in Nepenthes alata Blanco pitchers.

Plants in the genus Nepenthes obtain a substantial nutrient supply by trapping insects in highly modified leaves. A broad zone of the inner surface of these pitchers is densely covered with wax crystals on which most insects lose their footing. This slippery wax surface, capturing prey and preventing its escape from the trap, plays a pivotal role in the carnivorous syndrome. To understand the mechanism of slipperiness, the present investigation aimed at an ultrastructural and physico-chemical characterization of the wax crystals in pitchers of N. alata Blanco. Scanning electron microscopy revealed that entire platelets protruded perpendicularly from the surface. Methods were developed that allowed the mechanical removal of wax crystals from the pitcher surface. It could be shown that the sampling was selective for the epicuticular wax, relevant for plant-insect interactions. The crystals consisted of a mixture of aliphatic compounds dominated by very-long-chain aldehydes. Triacontanal, at 43% the most abundant constituent, was largely responsible for crystal formation. Solubility data indicate that the Nepenthes crystals contained polymeric forms of this aldehyde. The resulting mechanical properties of the polymer crystals and the mechanism of slipperiness are discussed.

Animals↗

Eggs and embryos in Xenoturbella (phylum uncertain) are not ingested prey.

Xenoturbella is an enigmatic animal that has puzzled science for almost a century. The eggs and embryos found in Xenoturbella have recently been interpreted as ingested prey. However, PCR on individual eggs as well as in situ hybridisation and in situ PCR unambiguously show that they are Xenoturbella's own. The eggs and embryos are individually enclosed within follicles with the same ultrastructure. The cortical granules in oocytes and eggs from Xenoturbella but not Nucula stained positively with an antiserum against Reissner's substance. The embryos incorporated 5-bromodeoxyuridine in vivo, i.e. they replicate their genome and are living.

Animals↗

Seasonal occurrence of the tapeworm Proteocephalus longicollis and its transmission from copepod intermediate host to fish.

Seasonal occurrence in terms of prevalence, intensity of infection, abundance and density of the tapeworm Proteocephalus longicollis (Zeder, 1800) and its transmission between its intermediate host (Cyclops abyssorum prealpinus) and definitive host (common whitefish, Coregonus lavaretus) in Lake Annecy, an oligotrophic lake in the western part of the Alps, France, were studied in the period of 1998-2000. A copepod Cyclops abyssorum prealpinus (Kiefer, 1939), the dominant species among planktonic crustaceans in the lake, served as the only intermediate host for this parasite. Infection with plerocercoids was higher in adult copepods (predominantly females) than in copepodite stages IV and V. The prevalence rate of 25% found in C. abyssorum prealpinus females in June 1998 represents a unique infection rate of intermediate hosts with fish tapeworms in natural conditions. The final host, the common whitefish Coregonus lavaretus (L.), was heavily infected with P. longicollis throughout the year (prevalence 90%; mean abundance 40.3; maximum intensity of infection more than 500 tapeworms per fish); immature tapeworms predominated in all samples (P<0.01). Transmission of tapeworm larvae from copepods to the common whitefish took place most intensively in summer and autumn, and depended on seasonal changes in the density of the C. abyssorum prealpinus population, infection of this copepod with plerocercoids and their density in the lake. In addition, transmission efficiency also seems to be determined by the longevity of tapeworm larvae in the intermediate host, timing of predation of the fish host and rapid development of the parasite within this host during the summer period. Overall transmission potential of P. longicollis was low, with only about 9% of juvenile specimens reaching maturity in common whitefish.

Animals↗

Impact of a microphallid trematode on the behaviour and survival of its isopod intermediate host: phylogenetic inheritance?

The extent to which the ability of parasites to alter host behaviour is phylogenetically inherited as opposed to independently evolved has received little attention. We investigated the impact of an undescribed species of Microphallus on the behaviour and survival of its host, the freshwater isopod Austridotea annectens, to determine if it produced effects comparable to those induced by other trematodes of this genus. There was no difference between the vertical distribution and responses to light of infected isopods and those of uninfected isopods. In contrast, we found that infected isopods were more active swimmers than uninfected isopods, and that they failed to show the evasive responses shown by uninfected isopods when exposed to a simulated predator. There was no detectable effect of Microphallus infection on isopod survival, however. Overall, our results provide no strong evidence that the behavioural manipulation shown by our Microphallus species and that displayed by the well-studied Microphallus papillorobustus are inherited from a common ancestor rather than independently derived.

Animals↗

Ultrastructure of a novel eurytele nematocyst of Carybdea alata Reynaud (Cubozoa, Cnidaria).

The ultrastructural characteristics of nematocysts from the cubozoan Carybdea alata Reynaud, 1830 (Hawaiian box jellyfish) were examined using light, scanning and transmission electron microscopy. We reclassified the predominant nematocyst in C. alata tentacles as a heterotrichous microbasic eurytele, based on spine, tubule and capsule measurements. These nematocysts exhibited a prominent and singular stylet, herein referred to as the lancet. Discharged nematocysts from fixed tentacle preparations displayed the following structures: a smooth shaft base, lamellae, a hemicircumferential fissure demarking the proximal end of a stratified lancet, and a gradually tapering tubule densely covered with large triangularly shaped spines. The lancet remained partially adjoined to the shaft base in a hinge-like fashion in rapidly fixed, whole-tentacle preparations. In contrast, this structure was not observed in discharged nematocyst preparations which involved multiple transfer steps prior to fixation. Various approaches were designed to detect this structure in the absence of fixative. Detached lancets were located in proximity to discharged tubules in undisturbed coverslip preparations of fresh tentacles. In addition, examination of embedded nematocysts from fresh tentacles laid on polyacrylamide gels revealed still-attached lancets. To examine the function of this structure in prey capture, Artemia sp. laden tentacles were prepared for scanning electron microscopy. While carapace exteriors exhibited structures proximal to the lancet, i.e., the nematocyst capsule and shaft base, neither tubule nor lancet structures were visible. Taken together, the morphological data suggested a series of events involved in the discharge of a novel eurytele from C. alata.

Animals↗

Intraguild interference and biocontrol effects of generalist predators in a winter wheat field.

Arable land typically harbours communities of polyphagous invertebrate natural enemies, among them numerous soil-surface dwelling predators such as ground beetles (Carabidae) and spiders (Lycosidae, Linyphiidae). Numbers of these predators were experimentally manipulated in a winter wheat field in order to study the predation impact of a generalist predator assemblage on herbivorous insects, the possible interferences among the predators concerned, and subsequent effects on wheat plant parameters. Removing ground beetles doubled numbers of Lycosidae indicative of intraguild interference between these two predator groups. Aphid densities were highest in carabid removal plots implying a substantial predation impact of ground beetles on the pest population. The predation impact of ground beetles was strongest earlier and disappeared later in the season. In mid-season, at intermediate aphid densities, the combined impact of carabid beetles and spiders appeared to be responsible for the reduction in aphid abundance. This result was probably due to a biomass effect rather than to a synergistic effect of the predator community. Thysanoptera decreased when spiders were removed (perhaps because spiders were preying on a predator of thrips), while Cicadellidae and Delphacidae showed no effect at all. The rise of aphid numbers in carabid removal plots corresponded to an increase in protein content of the wheat grains, while other plant parameters such as plant numbers and grain mass were not affected. In conclusion, this study provided field evidence for intraguild interference among generalist ground predators in arable land. Despite this interference the polyphagous predator community was able to depress numbers of aphids in winter wheat, a result cascading down to plant quality parameters.

Animals↗

Why does the larval integument of some sawfly species disrupt so easily? The harmful hemolymph hypothesis.

The larvae of several sawfly species belonging to the Tenthredinidae (Hymenoptera) have such a low mechanical resistance in the integument that slight mechanical damage to the integument is enough to provoke the release of hemolymph at a given spot. We quantified this phenomenon, which we call "easy bleeding", by measuring the pressure needed to pierce dissected sawfly integument. We also investigated the feeding deterrance of ethanolic extracts of the hemolymph by laboratory bioassays using Myrmica rubra ant workers. These traits, integument resistance and hemolymph deterrence, were inversely related, considering 22 tenthredinid species. A negative correlation was obtained by only taking into account the species of one tenthredinid tribe, namely the Phymatocerini (nine species studied). Our results support the "harmful hemolymph hypothesis" that we present here and that assumes a functional link between these morphological and chemical traits, jointly acting as a chemical defense strategy. We suspect hemolymph deterrence to be often due to sequestration of plant secondary metabolites. We discuss the role of easy bleeding, considering the fact that sawfly larvae are frequently the prey of invertebrate and vertebrate predators. It is suggested that invertebrates such as ants were more important than vertebrates in the evolution of easy bleeding.

Animals↗

Morphological variation of the predatory cladoceran Leptodora kindtii in relation to prey characteristics.

The responses of invertebrate predators to changes in the morphology of their prey, and especially the responses for induced defences, are largely unexplored, compared with the vast amount of data on predator-induced defences. This study demonstrates that the size of the feeding basket, the anatomical structure used to capture prey by the predaceous freshwater cladoceran Leptodora kindtii, shows significant allometric changes with the average body size of the prey (herbivorous cladocerans) in six lakes of northeastern Poland. Prey density influences the basket size only in adult Leptodora individuals, whereas induced defences such as helmets of Daphniasp. seem to have no effect on Leptodora's morphology. In a feeding experiment it is shown that the increase in the feeding basket enables Leptodora to capture larger prey. The seasonal pattern of morphological change, and its measurable benefit suggest that the observed morphological variability of Leptodora is phenotypic plasticity.

Adaptation, Physiological↗

Effects of a simple plant morphological mutation on the arthropod community and the impacts of predators on a principal insect herbivore.

Plant features that enhance predator effectiveness can be considered extrinsic-resistance factors because they result in reduced insect herbivory. In this paper we test the hypothesis that reduced epicuticular wax (EW) in Pisum sativum L. is an extrinsic-resistance factor contributing to field resistance to Acyrthosiphon pisum (Harris). We monitored pea aphid populations in the field on reduced EW and normal EW near isolines of peas for two seasons and confirmed that aphid populations are lower on reduced EW peas than on normal EW peas. We also monitored predators within the canopies of the two pea lines to discover community level patterns in response to differences in EW. We found that while predator numbers were similar between the two lines, there were more syrphids on the normal EW peas, and a trend towards more coccinellids on reduced EW peas. We tested the impact of predators on pea aphids on the two EW lines by monitoring their population levels in cages that excluded predators, and in cages that allowed predators to enter. We found that pea aphid populations were similar on the two EW lines when predators were excluded. When predators were allowed access to the plants, pea aphid populations were reduced more on reduced EW peas than on normal EW peas. We also examined the intrinsic resistance to aphids in reduced EW peas with laboratory dual-choice tests comparing aphid response to reduced EW and normal EW peas, and found that walking, apterous aphids displayed no preference for one pea line over the other. Bioassays to measure growth and fecundity of the pea aphid on the two EW types in the greenhouse and in the field showed that intrinsic rate of increase, and other life table parameters, were not different for aphids on the two lines. Together these results support the hypothesis that reduced EW in peas is a predator-dependent extrinsic resistance factor. Genetically reducing EW bloom in peas and other waxy crop plants might improve the effectiveness of arthropod natural enemies of insect pests. More generally, the results show that a subtle change in plant morphology can substantially influence the impact of predators on insect herbivore populations. The benefit of extrinsic resistance to herbivory conferred by reduced EW may balance any benefits of a prominent EW bloom, thereby sustaining EW polymorphisms in some natural plant populations.

Adaptation, Physiological↗

Infochemical-mediated intraguild interactions among three predatory mites on cassava plants.

Carnivorous arthropods exhibit complex intraspecific and interspecific behaviour among themselves when they share the same niche or habitat and food resources. They should simultaneously search for adequate food for themselves and their offspring and in the meantime avoid becoming food for other organisms. This behaviour is of great ecological interest in conditions of low prey availability. We examined by means of an olfactometer, how volatile chemicals from prey patches with conspecific or heterospecific predators might contribute to shaping the structure of predator guilds. To test this, we used the exotic predatory mites Typhlodromalus manihoti and T. aripo, and the native predatory mite Euseius fustis, with Mononychellus tanajoa as the common prey species for the three predatory mite species. We used as odour sources M. tanajoa-infested cassava leaves or apices with or without predators. T. manihoti avoided patches inhabited by the heterospecifics T. aripo and E. fustis or by conspecifics when tested against a patch without predators. Similarly, both T. aripo and E. fustis females avoided patches with con- or heterospecifics when tested against a patch without predators. When one patch contained T. aripo and the other T. manihoti, females of the latter preferred the patch with T. aripo. Thus, T. manihoti is able to discriminate between odours from patches with con- and heterospecifics. Our results show that the three predatory mite species are able to assess prey patch profitability using volatiles. Under natural conditions, particularly when their food sources are scarce, the three predatory mite species might be involved in interspecific and/or intraspecific interactions that can substantially affect population dynamics of the predators and their prey.

Animal Communication↗

Helpful habitant or pernicious passenger: interactions between an infaunal bivalve, an epifaunal hydroid and three potential predators.

The importance of positive interspecific interactions within physically stressful habitats has received increased attention from community ecologists. The exposed sandy beach is an example of a physically rigorous environment where biological interactions have long been considered insignificant. We examined the interaction between the infaunal clam, Donax variabilis, and the hydroid, Lovenella gracilis, on exposed sandy beaches in North Carolina. Epibiotic occupation of Donax by hydroids has been repeatedly observed on ocean beaches but rarely investigated. By providing a stable substrate for attachment, the clam facilitates the persistence of the hydroid in the intertidal beach; however, benefits or costs experienced by the host as a result of this association are unknown. By exposing clams with and without hydroid colonies to multiple types of clam predators, we tested the effectiveness of the hydroid, which possesses stinging nematocysts, in defending its host. The hydroid defended the clam against one common predator, the Florida pompano ( Trachinotus carolinus). Against speckled crabs ( Arenaeus cribrarius) and ghost crabs ( Ocypode quadrata), however, the hydroid offered no protection for its host and instead facilitated predation. The epibiotic hydroid, which projects above the surface of the sand, allowed the crabs to more readily detect clams below the surface. In the field, we evaluated the effect of the hydroid on the tidally synchronized migrations and burrowing speed of the clam. The hydroid, which can form large colonies on the posterior end of the clam, had no effect on Donax burrowing speed, but did reduce the speed of transport of clams by wave swash. Depending on relative predation pressure, the occupation of D. variabilis by L. gracilis can alternately be characterized as beneficial or detrimental to the host.

Animals↗

Seasonal shift from bottom-up to top-down impact in phytophagous insect populations.

Although many studies now examine how multiple factors influence the dynamics of herbivore populations, few studies explicitly attempt to document where and when each is important and how they vary and interact. In fact, how temporal variation in top-down (natural enemies) and bottom-up (host plant resources) factors affect herbivore dynamics has been suggested as a particularly important yet poorly understood feature of terrestrial food webs. In this study we examined how temporal changes in predator density (wolf spiders, sheet-web builders, and mirid egg predators) and host-plant resources (plant quality and structural complexity) influence the population dynamics of the dominant phytophagous insects on Atlantic-coast salt marshes, namely Prokelisia planthoppers (Homoptera: Delphacidae). We designed a factorial experiment in meadows of Spartina alterniflora to mimic natural variation in vegetation quality and structure by establishing two levels of plant nutrition (leaf nitrogen content) by fertilization, and two levels of habitat complexity by adding leaf litter (thatch). We then assessed seasonal changes in the strength of bottom-up (plant quality) and top-down (predator) impacts on planthopper populations. Planthopper populations responded positively to increased plant quality treatments in late summer. Despite the greater number of planthopper adults colonizing fertilized Spartina plots compared to unfertilized controls, the offspring of these colonists were much less abundant at the end of the season in fertilized plots, particularly those with thatch. The initial colonization effect was later erased because arthropod predators selectively accumulated in fertilized plots where they inflicted significant mortality on all stages of planthoppers. Predators rapidly colonized fertilized plots and reached high densities well in advance of planthopper colonization, a response we attribute to their rapid aggregation in complex-structured habitats with readily available alternative prey. Our results suggest that plant resources not only mediate the strength of predator impacts on herbivore populations, but they also promote the coupling of predator and prey populations and thus influence when enemy impacts are realized.

Animals↗