Density-dependent resistance of the gypsy moth <Emphasis Type="Italic">Lymantria dispar</Emphasis> to its nucleopolyhedrovirus,and the consequences for population dynamics

The processes controlling disease resistance can strongly influence the population dynamics of insect outbreaks. Evidence that disease resistance is density-dependent is accumulating, but the exact form of this relationship is highly variable from species to species. It has been hypothesized that insects experiencing high population densities might allocate more energy to disease resistance than those at lower densities, because they are more likely to encounter density-dependent pathogens. In contrast, the increased stress of high-density conditions might leave insects more vulnerable to disease. Both scenarios have been reported for various outbreak Lepidoptera in the literature. We tested the relationship between larval density and disease resistance with the gypsy moth (Lymantria dispar) and one of its most important density-dependent mortality factors, the nucleopolyhedrovirus (NPV) LdMNPV, in a series of bioassays. Larvae were reared in groups at different densities, fed the virus individually, and then reared individually to evaluate response to infection. In this system, resistance to the virus decreased with increasing larval density. Similarly, time to death was faster at high densities than at lower densities. Implications of density–resistance relationships for insect–pathogen population dynamics were explored in a mathematical model. In general, an inverse relationship between rearing density and disease resistance has a stabilizing effect on population dynamics.     

Two's a Crowd: Phenotypic Adjustments and Prophylaxis in Anticarsia gemmatalis Larvae Are Triggered by the Presence of Conspecifics

Defence from parasites and pathogens involves a cost. Thus, it is expected that organisms use this only at high population densities, where the risk of pathogen transmission may be high, as proposed by the "density-dependent prophylaxis" (DDP) hypothesis. These predictions have been tested in a wide range of insects, both in comparative and experimental studies. We think it pertinent to consider a continuum between solitarious and gregarious living insects, wherein: (1) solitarious insects are those that are constitutively solitary and do not express any phenotypic plasticity, (2) the middle of the continuum is represented by insects that are subject to fluctuations in local density and show a range of facultative and plastic changes; and (3) constitutively gregarious forms live gregariously and show the gregarious phenotype even in the absence of crowding stimuli. We aimed to chart some of the intermediary continuum with an insect that presents solitarious aspects, but that is subject to fluctuations in density. Thus, Anticarsia gemmatalis (Lepidoptera: Noctuidae) larvae reared at higher densities showed changes in coloration, a greater degree of encapsulation, had higher hemocyte densities and were more resistant to Baculovirus anticarsia, but not to Bacillus thuringiensis. Meanwhile, with increased rearing density there was reduced capsule melanization. Hemocyte density was the only variable that did not vary according to larval phenotype. The observed responses were not a continuous function of larval density, but an all-or-nothing response to the presence of a conspecific. As A. gemmatalis is not known for gregarious living, yet shows these density-dependent changes, it thus seems that this plastic phenotypic adjustment may be a broader phenomenon than previously thought.     

Isolation from a marching band increases haemocyte density in wild locusts (Chortoicetes terminifera)

1. The density‐dependent prophylaxis hypothesis predicts that individuals in high‐density populations will invest more resources in immune defence than individuals at lower densities. 2. However, recent work suggests that this prediction may not apply to all situations; solitarious species may paradoxically have higher scores than crowded counterparts in certain immune assays. 3. To investigate the relationship between a key immune parameter and field population densities, the total haemocyte counts (THCs) of Australian plague locusts (Chortoicetes terminifera) from three population densities in Western Australia were compared. 4. THCs were negatively correlated with field population densities, and locusts removed from a marching band and kept in isolation had increased THCs relative to group‐housed controls. 5. These results demonstrate that immune investment can inversely relate to population density in field conditions. 6. We suggest that isolated locusts increase their haemocyte densities relative to crowded conspecifics in response to potentially greater exposure to parasitoids and nematodes.     

Does immune function influence population fluctuations and level of parasitism in the cyclic geometrid moth?

Populations of the autumnal moth, Epirrita autumnata, exhibit cycles with high amplitudes in northernmost Europe, culminating in devastating outbreak densities at favourable sites. Parasitism by hymenopteran parasitoids has been hypothesised to operate with a delayed density dependence capable of producing the observed dynamics. It has also been hypothesised that insects in crowded conditions invest greatly in their immunity as a counter-measure to increased risk of parasitism and pathogen infections. Furthermore, inducible plant defences consequent to grazing by herbivorous insects may be linked to the performance of parasitoids and pathogens through increased immunocompetence of the herbivore feeding on the foliage, in which the defence induction has taken place. At ten sampling sites, we quantified larval abundance, outbreak status and percentage larval parasitism during an extended peak phase of a population cycle. These population level covariates, together with an individual pupal mass, were used to explain differences in the immune defence, measured as an encapsulation reaction to artificial antigen. We also conducted a field study for an investigation of the susceptibility of autumnal moth pupae to naturally occurring pupal parasitoids. We did not find obvious differences between the encapsulation rate of autumnal moths originating from the sites with different past and current larval densities and risks for parasitism. The best ranked statistical models included pupal mass and outbreak status as explanatory variables, although both showed only slight effects on the encapsulation rate. The host resistance test revealed positive relationships between the encapsulation rate, body size and percentage parasitism of the exposed pupae, indicating that pupal parasitoids chose, and/or survived better, in large host individuals irrespective of their encapsulation ability. Thus, our results do not provide support for the hypothesis that variation in the immune function drives or modulates population cycles of autumnal moths. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorised users.     

Strong density-dependent survival and recruitment regulate the abundance of a coral reef fish

Debate on the control of population dynamics in reef fishes has centred on whether patterns in abundance are determined by the supply of planktonic recruits, or by post-recruitment processes. Recruitment limitation implies little or no regulation of the reef-associated population, and is supported by several experimental studies that failed to detect density dependence. Previous manipulations of population density have, however, focused on juveniles, and there have been no tests for density-dependent interactions among adult reef fishes. I tested for population regulation in Coryphopterus glaucofraenum, a small, short-lived goby that is common in the Caribbean. Adult density was manipulated on artificial reefs and adults were also monitored on reefs where they varied in density naturally. Survival of adult gobies showed a strong inverse relationship with their initial density across a realistic range of densities. Individually marked gobies, however, grew at similar rates across all densities, suggesting that density-dependent survival was not associated with depressed growth, and so may result from predation or parasitism rather than from food shortage. Like adult survival, the accumulation of new recruits on reefs was also much lower at high adult densities than at low densities. Suppression of recruitment by adults may occur because adults cause either reduced larval settlement or reduced early post-settlement survival. In summary, this study has documented a previously unrecorded regulatory mechanism for reef fish populations (density-dependent adult mortality) and provided a particularly strong example of a well-established mechanism (density-dependent recruitment). In combination, these two compensatory mechanisms have the potential to strongly regulate the abundance of this species, and rule out the control of abundance by the supply of recruits.     

Population dynamics of the beech caterpillar, <Emphasis Type="Italic">Syntypistis punctatella</Emphasis>, and biotic and abiotic factors

The beech caterpillar, Syntypistis punctatella (Motschulsky) (Lepidoptera: Notodontidae), often causes extensive defoliation of beech forests in Japan. Outbreaks have often occurred synchronously among different areas at intervals of 8–11 years. Synchrony of outbreaks was considered to be caused by synchrony of weather. Populations of this insect exhibit periodical dynamics in both outbreak and nonoutbreak areas. Factors that might influence the population dynamics of the beech caterpillar were classified from the point of view of the natural bioregulation com-plex, which includes a coleopteran predator, Calosoma maximowiczi, avian predators, parasitoids, entomopathogenic fungi, and delayed induced defensive response (DIR) of beech trees. Because such periodic population dynamics are believed to be caused by one or more delayed density-dependent factors, delayed density-dependent mortality has been identified as a likely source of population cycles. The DIR and pathogenic diseases showed a high order of density dependence. An infectious pathogen, Cordyceps militaris, was considered to be the most plausible agent responsible for periodic dynamics of the beech caterpillar population because insect diseases were effective in cases in which the S. punctatella population started to decrease without reaching outbreak densities, but DIR was not. Conspicuous defoliation caused by this insect tends to occur at certain elevations, where forests are composed of pure stands of beech trees. I propose three different hypotheses to explain this phenomenon: the diversity–stability hypothesis, the resource concentration hypothesis, and the altitudinal soil nutrient hypothesis. Received: November 20, 1999 / Accepted: August 3, 2000     

Baculovirus resistance in the noctuid Spodoptera exempta is phenotypically plastic and responds to population density

Parasite resistance mechanisms can be costly to maintain. We would therefore predict that organisms should invest in resistance only when it is likely to be required. Insects that show density-dependent phase polyphenism, developing different phenotypes at high and low population densities, have the opportunity to match their levels of investment in resistance with the likelihood of exposure to pathogens. As high population densities often precipitate disease epidemics, the high-density form should be selected to invest relatively more in resistance. We tested this prediction in larvae of the noctuid Spodoptera exempta. Larvae reared at a high density were found to be considerably more resistant to a nuclear polyhedrosis virus than those reared in isolation. A conspicuous feature of the high-density phase of S. exempta and other phase-polyphenic Lepidoptera is cuticular melanization. As melanization is controlled by the phenoloxidase enzyme system, which is also involved in the immune response, this suggests a possible mechanism for increased resistance at high population densities. We demonstrated that melanized S. exempta larvae were more resistant than non-melanized forms, independent of rearing density. We also found that haemolymph phenoloxidase activity was correlated with cuticular melanization, providing further evidence for a link between melanization and immunity. These results suggest that pathogen resistance in S. exempta is phenotypically plastic, and that the melanized cuticles characteristic of the high-density form may be indicative of a more active immune system.     

Population dynamics ofMalacosoma neustria testacea (Lepidoptera: Lasiocampidae): Stabilizing process in a field population

Stabilizing mechanism in population of Malacosoma neustria testacea was investigated in a central part of Japan based on eight year survey. Population fluctuation in each developmental stage in the experimental field was rather small, i.e., 5.9 times in egg and 85.0 times in female adult. Pupal weight negatively correlated with the densities of 5th instar larvae and prepupae (cocoons) and correlation coefficient was highly significant in females. Population density was stabilized by density-dependent dispersal of female moths in preovipositional period. Comparison between fecundity of emerged moths and that of actually oviposited ones in the experimental field suggested that density-dependent dispersal took place as the result of density-dependent size variation, i.e., small-sized female months have higher flying ability. This hypothesis was supported by the experiment in which flying ability of newly emerged female moth was measured. Similar stabilizing mechanism is expected to occur in semelparous or pro-ovigenic insects.     

Density-dependent prophylaxis in the mealworm beetle Tenebrio molitor L. (Coleoptera: Tenebrionidae): cuticular melanization is an indicator of investment in immunity

If there are costs involved with the maintenance of pathogen resistance, then higher investment in this trait is expected when the risk of pathogenesis is high. One situation in which the risk of pathogenesis is elevated is at increased conspecific density. This paper reports the results of a study of density-dependent polyphenism in pathogen resistance and immune function in the mealworm beetle Tenebrio molitor. Beetles reared at high larval densities showed lower mortality when exposed to a generalist entomopathogenic fungus and a higher degree of cuticular melanization than those reared solitarily. The degree of cuticular melanization was a strong indicator of resistance, with darker beetles being more resistant than lighter ones regardless of rearing density. No differences were found between rearing densities in the levels of phenoloxidase, an enzyme key to the insect immune response. The results show that pathogen resistance is phenotypically plastic in T. molitor, suggesting that the maintenance of this trait is costly.     

Interactions ofMonoporeia affinis (Lindström) (Amphipoda) with sedentary Chironomidae

Interannual population oscillations ofMonoporeia affinis and sedentary chironomids are negatively correlated in Lake Mälaren. Sedentary chironomid abundance regressed againstM. affinis density was highly significant at a lag response of one year (adj. R²=0.54, P=0.0001). The inverse correlation between Tanytarsini (Micropsectra sp. andTanytarsus sp.) andM. affinis densities supports a recruitment limitation hypothesis,viz. that the growth or survival of early instar chironomids is low at high amphipod densities. Microcosm studies showed density-dependent effects on the growth and survival of second instarCh. riparius larvae across an amphipod density gradient. No significant effects were found on growth or survival of third instars. These findings support the chironomid recruitment limitation hypothesis. Amphipod predation on early instar chironomid larvae is suggested as a population controlling mechanism.     

Diversity and abundance of insect herbivores foraging on seedlings in a rainforest in Guyana

1. Free-living insect herbivores foraging on 10 000 tagged seedlings representing five species of common rainforest trees were surveyed monthly for more than 1 year in an unlogged forest plot of 1 km² in Guyana. 2. Overall, 9056 insect specimens were collected. Most were sap-sucking insects, which represented at least 244 species belonging to 25 families. Leaf-chewing insects included at least 101 species belonging to 16 families. Herbivore densities were among the lowest densities reported in tropical rainforests to date: 2.4 individuals per square metre of foliage. 3. Insect host specificity was assessed by calculating Lloyd’s index of patchiness from distributional records and considering feeding records in captivity and in situ. Generalists represented 84 and 78% of sap-sucking species and individuals, and 75 and 42% of leaf-chewing species and individuals. In particular, several species of polyphagous xylem-feeding Cicadellinae were strikingly abundant on all hosts. 4. The high incidence of generalist insects suggests that the Janzen–Connell model, explaining rates of attack on seedlings as a density-dependent process resulting from contagion of specialist insects from parent trees, is unlikely to be valid in this study system. 5. Given the rarity of flushing events for the seedlings during the study period, the low insect densities, and the high proportion of generalists, the data also suggest that seedlings may represent a poor resource for free-living insect herbivores in rainforests.     

Population dynamics ofMalacosoma neustria testacea (Lepidoptera: Lasiocampidae): Stabilizing process in a field population

Summary Stabilizing mechanism in population ofMalacosoma neustria testacea was investigated in a central part of Japan based on eight year survey. Population fluctuation in each developmental stage in the experimental field was rather small, i.e., 5.9 times in egg and 85.0 times in female adult. Pupal weight negatively correlated with the densities of 5th instar larvae and prepupae (cocoons) and correlation coefficient was highly significant in females. Population density was stabilized by density-dependent dispersal of female moths in preovipositional period. Comparison between fecundity of emerged moths and that of actually oviposited ones in the experimental field suggested that density-dependent dispersal took place as the result of density-dependent size variation, i. e., small-sized female moths have higher flying ability. This hypothesis was supported by the experiment in which flying ability of newly emerged female moth was measured. Similar stabilizing mechanism is expected to occur in semelparous or pro-ovigenic insects. Contribution Ser. A, No. 64 from Fruit Tree Research Station, Ministry of Agriculture and Forestry, Japan     

High larval density does not induce a prophylactic immune response in a butterfly

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Greater cuticular melanism is not associated with greater immunogenic response in adults of the polymorphic mountain stone weta, Hemideina maori

Abstract.  1. Greater immune function is associated with the high-density melanic phase of polyphenic insects, appearing to compensate for density-dependent increases in susceptibility to parasites and/or pathogens. Other types of discrete variation in cuticular colour occur in insects (which may or may not be associated with melanin pigmentation), but whether this variation is predictive of immune ability has not been investigated.
2. In the mountain stone weta Hemideina maori , a black morph and yellow banded morph occur. These morphs are not seasonally polyphenic and have discrete haplotype genetic markers. Black individuals are typically found at lower local densities than yellow individuals, contrary to relations between cuticular melanism and density seen in polyphenic insects.
3. Yellow males and females had greater melanotic encapsulation responses upon immune challenge than did black males and females, but these differences were not associated with differences in temperature selection between morphs. Morph differences in melanotic encapsulation responses were somewhat related to differences between morphs in haemocyte concentrations.
4. These results indicate that a common form of immune expression is not heightened with dark coloration in the mountain stone weta. Thus, earlier findings of greater immunity associated with darker cuticles in phase polyphenic insects cannot be extended to insects with other forms of discrete colour variation. These findings will help in elucidating causes and consequences of such colour polymorphism, which is widespread in several insect orders.     

Fitness in invasive social wasps: the role of variation in viral load,immune response and paternity in predicting nest size and reproductive output

Within any one habitat, the relative fitness of organisms in a population can vary substantially. Social insects like the common wasp are among the most successful invasive animals, but show enormous variation in nest size and other fitness‐related traits. Some of this variation may be caused by pathogens such as viruses that can have serious consequences in social insects, which range from reduced productivity to colony death. Both individual immune responses and colony‐level traits such as genetic diversity are likely to influence effects of pathogen infections on colony fitness. Here we investigate how infections with Kashmir Bee Virus (KBV), immune response and intracolony genetic diversity (due to queen polyandry) affect nest size in the invasive common wasp Vespula vulgaris. We show that KBV is highly prevalent in wasps and expression of antiviral immune genes is significantly increased with higher viral loads across individuals. Patriline membership within a nest did not influence KBV susceptibility or immune response. A permutational MANCOVA revealed that polyandry, viral load and expression of the immune gene Dicer were significant predictors of variation in nest size. High intracolony genetic diversity due to polyandry has previously been hypothesized to improve colony‐level resistance to parasites and pathogens. Consistent with this hypothesis, we observed genetically diverse colonies to be significantly larger and to produce more queens, although this effect was not driven by the pathogen we investigated. Invasive wasps clearly suffer from pathogens and expend resources, as indicated here by elevated immune gene expression, toward reducing pathogen‐impact on colony fitness.     

Investment in Constitutive Immune Function by North American Elk Experimentally Maintained at Two Different Population Densities

Natural selection favors individuals that respond with effective and appropriate immune responses to macro or microparasites. Animals living in populations close to ecological carrying capacity experience increased intraspecific competition, and as a result are often in poor nutritional condition. Nutritional condition, in turn, affects the amount of endogenous resources that are available for investment in immune function. Our objective was to understand the relationship between immune function and density dependence mediated by trade-offs between immune function, nutritional condition, and reproduction. To determine how immune function relates to density-dependent processes, we quantified bacteria killing ability, hemolytic-complement activity, and nutritional condition of North American elk (Cervus elaphus) from populations maintained at experimentally high- and low-population densities. When compared with elk from the low-density population, those from the high-density population had higher bacteria killing ability and hemolytic-complement activity despite their lower nutritional condition. Similarly, when compared with adults, yearlings had higher bacteria killing ability, higher hemolytic-complement activity, and lower nutritional condition. Pregnancy status and lactational status did not change either measure of constitutive immunity. Density-dependent processes affected both nutritional condition and investment in constitutive immune function. Although the mechanism for how density affects immunity is ambiguous, we hypothesize two possibilities: (i) individuals in higher population densities and in poorer nutritional condition invested more into constitutive immune defenses, or (ii) had higher parasite loads causing higher induced immune responses. Those explanations are not mutually exclusive, and might be synergistic, but overall our results provide stronger support for the hypothesis that animals in poorer nutritional condition invest more in constitutive immune defenses then animals in better nutritional condition. This intriguing hypothesis should be investigated further within the larger framework of the cost and benefit structure of immune responses.     

An analysis of distribution and abundance of populations of the high-shore limpet,Notoacmea petterdi (tenison-woods)

Summary Notoacmea petterdi occurs only on vertical surfaces at the highest levels on the shore. On a given surface, density of limpets decreases with increasing height. Mean and maximum sizes of limpets increase with increasing height, due to faster rates of growth at higher levels where densities are lower. Rates of mortality are also negatively correlated with height on the shore, and are therefore generally less at higher levels. Both adult and juvenile limpets exhibit well-developed homing behaviour. There is no evidence of seasonal migrations, density-dependent migrations, nor gradual upward migrations as limpets grow older.Hypotheses are formed on the basis of these preliminary observations and measurements. Results of subsequent experimental manipulations of densities support the hypothesis that density-dependent mortality, due to intraspecific competition, plays a major role in regulating population densities of this limpet. It is also suggested that the pattern of settlement of juvenile limpets is the major factor that determines the limits to the vertical distribution of this species.     

Inoculum Density and Population Dynamics of Suppressive and PathogenicStreptomycesStrains and Their Relationship to Biological Control of Potato Scab

SeveralStreptomycesstrains are capable of suppressing potato scab caused byStreptomyces scabies.Although these strains have been successful in the biocontrol of potato scab in the field, little is known about how populations of pathogenicStreptomycesin the potato rhizosphere are influenced by inoculation of the suppressive strains. The effects of inoculum densities of pathogenic and suppressiveStreptomycesstrains on their respective populations on roots and in rhizosphere soil were examined during the growing season. The relationships between inoculum density or rhizosphere population densities and disease severity were also investigated. Populations of suppressiveStreptomycesstrain 93 increased significantly on roots with increasing inoculum dose. At its highest inoculum dose, the suppressive strain reached a population density greater than 10⁶CFU/g root 14 weeks after planting. The ability of the suppressive strain to increase its populations with increasing inoculum density was hindered at high inoculum doses of the pathogen, suggesting that density-dependent competitive interactions may be occurring between the two antagonists. Strain 93 was most effective at preventing scab early in the growing season (8 weeks after planting), when tubers were most susceptible to the scab disease. Population densities of the suppressive strain in soil were more highly negatively correlated with scab severity than were populations on roots, suggesting that rhizosphere soil rather than potato roots may be the primary source of inoculum of the suppressive strain for tubers.     

Limits to predator regulation of rabbits in Australia: evidence from predator-removal experiments

Summary Predator-prey studies in semi-arid eastern Australia demonstrated that populations of rabbits (Oryctolagus cuniculus) could be regulated by predators. The functional, numerical and total responses of foxes (Vulpes vulpes) to rabbits and the numerical response of feral cats (Felis catus) to rabbits, are described. Measurement of the rabbit component of foxes' stomach contents indicates a Type III functional response. The size of the fox population in summer was dependent on the availability of rabbits over the immediately preceding rabbit breeding season but there appeared to be no density-dependent aggregation of young foxes in areas of surplus food. The total response of foxes, estimated using the short-term numerical response of dispersing foxes, was directly density-dependent for low rabbit densities and inversely density-dependent for high rabbit densities. Two states are possible with this form of total response: a state with low rabbit densities regulated by predators and a state with high rabbit densities which occurs when rabbits escape predator regulation. The boundary between regulation and non-regulation by predators was demonstrated by a predator-removal experiment. In the treated areas, predators were initially culled and rabbits increased to higher densities than in an untreated area where predators were always present. When predators were allowed back into the treated areas, rabbit populations continued to increase and did not decline to the density in the untreated area. This is the critical evidence for a two-state system. When predators were present, rabbits could be maintained at low densities which were in the density-dependent part of the total response curve for foxes. Exceptionally high rabbit recruitment, or artificially reduced predation, could result in rabbits escaping predator-regulation. Under these circumstances, rabbits could move into the inversely density-dependent region of the total response curve for foxes.     

Population regulation by post-settlement mortality in two temperate reef fishes

 Input of individuals dispersing into open populations can be highly variable, yet the consequences of such variation for subsequent population densities are not well understood. I explored the influence of variable input (”supply”) on subsequent densities of juveniles and adults in open local populations of two temperate reef fishes, the bluebanded goby (Lythrypnus dalli) and the blackeye goby (Coryphopterus nicholsii). Variable recruitment was simulated by stocking a natural range of densities of young fishes on replicate patch reefs. Density and mortality of the stocked cohorts were followed over time, until the fishes reached maturity. Over the first day of the experiments, mortality of both species was significantly density-dependent; however, there was still a very strong relationship between density on day 1 and density on day 0 (i.e., simulated recruitment was still an excellent predictor of population density). At this point in the study, the main effects of density-dependent mortality were to reduce mean densities and variation about the mean. Over the period from the start of the experiments until the time when maturity was reached by each species (about 1 and 3 months for Lythrypnus and Coryphopterus, respectively), mortality was strongly density-dependent. Such strong density-dependent mortality virtually eliminated any linear relationship between adult density and ”recruit” density. However, for both species, the relationship between these two variables was well fit by an asymptotic curve, with the asymptotic density of adults equal to c. 3/m² for Coryphopterus, and c. 10/m² for Lythrypnus. Natural recruitment (via settlement of larvae) to the reefs over the period of the study (9 months) was above the asymptotic densities of adults for the two species, even though the study did not encompass the periods of peak annual recruitment of either species. This suggests that adult populations of these two gobies may often be limited, and regulated, by post-settlement processes, rather than by input of settlers. Other studies have shown that mortality of the two species is density-independent, or only weakly density-dependent, on reefs from which predators have been excluded. Hence, it appears that predators cause density-dependent mortality in these fishes. Received: 26 November 1996 / Accepted: 5 April 1997