Potential for climate effects on the size-structure of host–parasitoid indirect interaction networks

Communities of insect herbivores are thought to be structured mainly by indirect processes mediated by shared natural enemies, such as apparent competition. In host–parasitoid interaction networks, overlap in natural enemy communities between any pair of host species depends on the realized niches of parasitoids, which ultimately depend on the foraging decisions of individuals. Optimal foraging theory predicts that egg-limited parasitoid females should reject small hosts in favour of future opportunities to oviposit in larger hosts, while time-limited parasitoids are expected to optimize oviposition rate regardless of host size. The degree to which parasitoids are time- or egg-limited depends in part on weather conditions, as this determines the proportion of an individual''s lifespan that is available to foraging. Using a 10-year time series of monthly quantitative host–parasitoid webs, we present evidence for host-size-based electivity and sex allocation in the common secondary parasitoid Asaphes vulgaris. We argue that this electivity leads to body-size-dependent asymmetry in apparent competition among hosts and we discuss how changing weather patterns, as a result of climate change, may impact foraging behaviour and thereby the size-structure and dynamics of host–parasitoid indirect interaction networks.     

Do nine-primaried passerines have nine or ten primary feathers? The evolution of a concept

The number of primary feathers in a birds wing has been used as a systematic character since the first half of the nineteenth century. During the years, though, the definition of which feathers to count as a primary has changed and today the species historically denoted as having only nine primaries are instead said to have, for example, nine functional primaries. In this study, I investigated the borderline between nine-primaried and ten-primaried birds to search for a proper definition of the term nine-primaried. A total of 161 specimens of 104 bird species, mainly passerines, were examined. All species examined had ten primaries although the nine-primaried species had primary ten more or less concealed under primary covert nine. The number of primary coverts has decreased over time, with ten primary coverts as the ancestral state within Passeriformes and nine primary coverts among most oscine species. In conclusion, a proper definition of nine-primaried might be with primary ten concealed by primary covert nine. This definition includes all taxa historically denoted nine-primaried, i.e. systematically it is a definition of a paraphyletic group. The term nine-primaried is thus too inclusive to be of more than very limited systemtic value and, consequently, the New World nine-primaried oscines group might gain from a new denotation.Electronic Supplementary Material Supplementary material is available for this article at     

Refuge evolution and the population dynamics of coupled host—parasitoid associations

Summary We have investigated the theoretical consequences of character evolution for the population dynamics of a host—parasitoid interaction, assuming a monophagous parasitoid. In the purely ecological model it is assumed that hosts can escape parasitism by being in absolute refuges. A striking property of this model is a threshold effect in control of the host by the parasitoid, when host density dependence is weak. The approximate criteria for the parasitoid to regulate the host to low densities are (1) that the parasitoid's maximum population growth rate should exceed the host's and (2) that the maximum growth rate of the host in the refuge should be less than unity. We then use this ecological framework as a basis for a model which considers evolutionary changes in quantitative characters influencing the size of the absolute refuge. For each species, an increase in its refuge-determining character comes at a cost to maximum population growth rate. We show that refuge evolution can substantially alter the population dynamics of the purely ecological model, resulting in a number of emergent and sometimes counter-intuitive properties. In general, when the host has a high carrying capacity, systems are polarized either with low or minor refuge and top-down control of the host by the parasitoid or with a refuge and bottom-up control of the host by a combination of its own density dependence and the parasitoid. A particularly tantalizing result is that co-evolutionary dynamics can modify ecologically unstable systems into ones which are either stable or quasi-stable (with bouts of unstable dynamics, punctuating long-term periods of quasi-stable behaviour). We present five quantitative criteria which must all be met for the parasitoid to be the agent responsible for control of the host at a co-evolutionary equilibrium. The apparent stringency of this full set of requirements supports the empirically-based suggestion that monophagous parasitoid-driven systems should be less common in nature than those driven by multiple forms of density dependence. Further, we apply our theory to the question of whether exploiters may harvest their victims at maximum sustainable yields and to the evolutionary stability of biological control. Finally, we present a series of testable predictions of our theory and methods useful for testing them.     

Chromosomal location of esterase,peroxidase and phosphoglucomutase isozyme structural genes in cultivated rye (Secale cereale L.)

Summary Isoelectric focusing of esterase (EST), peroxidase (PRX), and phosphoglucomutase (PGM) isozymes in Chinese Spring wheat, Imperial rye and several Chinese Spring/Imperial and Holdfast/King II addition, translocation and substitution lines revealed the chromosomal location of nine Est loci previously described and of one Prx and Pgm locus. Loci Est1, Est2, Est3, Est5, Est6 and Est7 were found on chromosome arm 5RL, Est8 and Est9 on chromosome 6R in Imperial rye, and the Est10 locus on chromosome arm 4RL in Imperial rye and King II rye. A discrepancy was found between the chromosomal location of the Prx locus in Imperial where chromosome 2R was responsible for the expression of the peroxidase enzyme, and King II with chromosome 1R carrying the Prx gene. As a possible explanation, the occurrence of translocation events during the production of wheat/rye aneuploid lines is discussed. The rye Pgm locus could be associated with chromosome 4RS in Imperial and King II rye. Except for the location of Est loci on chromosome 5RL, the results reported in this paper lend further evidence for the assumed homoeology relationships between the chromosomes of Triticinae and for the conservation of gene synteny groups during the evolution of the Triticeae tribe.     

Who is the top dog in ant communities? Resources,parasitoids, and multiple competitive hierarchies

A wide variety of animal communities are organized into interspecific dominance hierarchies associated with the control and harvest of food resources. Interspecific dominance relationships are commonly found to be linear. However, dominance relations within communities can form a continuum ranging from intransitive networks to transitive, linear dominance hierarchies. How interference competition affects community structure depends on the configuration of the dominance interactions among the species. This study explores how resource size and the trait-mediated indirect effect (TMIE) specialist phorid fly parasitoids exert on interference competition, affect the transitive nature of competitive interactions in an assemblage of woodland ants. I quantify the linearity of networks of interactions associated with large and small food resources in the presence and absence of phorid parasitoids. Two distinct, significantly linear dominance hierarchies exist within the ant assemblage depending on the size of the disputed resource. However, the presence of phorid fly parasitoids eliminates the linearity of both dominance hierarchies. The hosts phorid defense behaviors reduce the competitive asymmetries between the host and its subdominant competitors, increasing the indeterminacy in the outcome of competitive interactions. Thus, both resource size variation and phorid-induced TMIEs appear to facilitate coexistence in assemblages of scavenging ants.     

Reflections on the phytosociological approach and the epistemological basis of vegetation science

Summary It is tried to identify the logical basis of the phytosociological method. Both the inductive and the deductive approach are discussed. The starting point can be summarized with the axiom vegetation is organized in communities. Some problems related to the use of numerical methods and computers are discussed.This paper was written under the auspices of the Programma finalizzato Qualità dell'Ambiente of the C.N.R. (Roma) Subprogetto 5 Metodologie matematiche e basi di dati. I thank the following people for helpful comments: E. Feoli, D. Lausi, P. Nimis, E. Pignatti Wikus (all at the Istituto Botanico, Trieste), D. Dal Maso (Istituto di Matematica, Trieste), E. van der Maarel (Botanisch Laboratorium, Nijmegen).     

Reversible and irreversible damage in reoxygenated ‘ischemic’ ventricular myocytes in culture

The LDH release pattern from cardiomyocytes under ischemia-like conditions shows two phases. In the initial slow phase, reoxygenation immediately stops further enzyme release. Accelerated LDH release, which occurs concomitantly with Iysosomal enzyme release, characterizes the second phase of ischemia. Reoxygenation at this stage does not put a stop to further enzyme release.Reoxygenation during the first phase of ischemia rapidly restored ATP level, while in the second phase, ATP levels remained low even after 6 h of reoxygenation.This study as well as previous data seem to suggest that irreversible cellular damage leading to cell death, occurs by synergistic action of many effectors, each of which does not necessarily cause irreversible damage.     

Food-web models that generate constant predator-prey ratios

Summary An approximately constant ratio of number of predator species/number of prey species is observed in several natural communities, although the exact value of the ratio may vary with habitat and the types of organisms in the food web. We test the hypothesis that a constant predator/prey ratio can be generated by what Holt (1977) terms apparent competition and what Jeffries and Lawton (1984) call competition for enemy-free space. We create simple, two trophic-level communities by drawing species of predators and prey at random from a species pool, simulating their interactions using Lotka-Volterra models. The simulated food webs converge over successive periods of invasion and extinction to locally stable systems with approximately constant ratios of number of predator species/number of prey species, despite varying initial conditions. As expected, predator/prey ratios take different values depending upon the biology of the simulated species. We conclude that apparent competition between prey species via shared enemies may be one mechanism whereby approximately constant predator/prey ratios are generated in natural communities.     

Establishment and dominance of an introduced herbivore has limited impact on native host-parasitoid food webs

The gypsy moth is considered one of the most harmful invasive forest insects in North America. It has been suggested that gypsy moth may indirectly impact native caterpillar communities via shared parasitoids. However, the impact of gypsy moth on forest insect food webs in general remains unstudied. Here we assess such potential impacts by surveying forest insect food webs in Ontario, Canada. We systematically collected caterpillars using burlap bands at sites with and without histories of gypsy moth outbreak, and then reared these caterpillars until potential parasitoid emergence. This procedure allowed us to generate quantitative food webs describing caterpillar-parasitoid interactions. We estimated the degree of parasitoid sharing between gypsy moth and native caterpillars. We also statistically modeled the effect of gypsy moth outbreak history and current gypsy moth abundance on standard indices of quantitative food web structure and the diversity of parasitoid communities. Rates of gypsy moth parasitism were very low and gypsy moth shared very few parasitoids with native caterpillars, suggesting limited potential for indirect interactions. We did not detect any significant effects of gypsy moth on either food web structure or parasitoid diversity, and the small amount of parasitoid sharing strongly implies that this lack of significance is not merely due to low statistical power. Our study suggests that gypsy moth has limited impact on native host-parasitoid food webs, at least for species that use burlap bands. Our results emphasize that extrapolations of theoretical and experimental conclusions on the impacts of invasive species should be tested in natural settings.     

Learning to discriminate between infochemicals from different plant-host complexes by the parasitoids Cotesia glomerata and C. rubecula

Comparison of closely related species can elucidate adaptive differences in species characteristics. The present study compares the effect of experience on the host-finding behaviour of two Cotesia (Hymenoptera: Braconidae) parasitoid species that differ in their degree of specialization. After multiple experiences with host larvae, females of the generalist parasitoid Cotesia glomerata showed a clear preference for volatiles from Pieris brassicae-infested Brussels sprouts leaves over P. rapae-infested Brussels sprouts leaves in two-choice tests (preference learning at herbivore level). A single experience with a host did not lead to such preferences.Experience of adult C. glomerata with different P. brassicae-infested cabbage varieties or nasturtium (Tropaeolum majus L.) led to preferences for the experienced plant-host complex in most cases (preference learning at plant level). No effect of rearing plant (early adult learning) on plant preference was found.In contrast to the generalist C. glomerata, females of the specialist C. rubecula did not show preference learning at the herbivore level. At the plant level, experience with different P. rapae-infested cabbage varieties in no case resulted in a difference in preference between treatments. The results support the hypothesis that learning plays a more important role in searching in generalists than in specialist parasitoids. The behaviour of the generalist C. glomerata was more easily changed by experience than that of the specialist C. rubecula.     

Searching for Food or Hosts: The Influence of Parasitoids Behavior on Host–Parasitoid Dynamics

A host–parasitoid system with overlapping generations is considered. The dynamics of the system is described by differential equations with a control parameter describing the behavior of the parasitoids. The control parameter models how the parasitoids split their time between searching for hosts and searching for non-host food. The choice of the control parameter is based on the assumption that each parasitoid maximizes the instantaneous growth rate of the number of copies of its genotype. It is shown that optimal individual behavior of parasitoids, with respect to time sharing between hosts and food searching, may have a stabilizing effect on the host–parasitoid dynamics.     

Adding physiological realism to dynamic state variable models of parasitoid host feeding

Summary The behavioural decisions of insect parasitoids have been the subject of dynamic state variable models, which explicitly incorporate the physiological state of the parasitoid, e.g. her age and egg load (number of mature eggs). Such models have been most recently applied to parasitoid host feeding, the consumption of host body fluids or tissues by the adult female wasp. The models developed to date, and recent empirical work withAphytis melinus, have highlighted the importance of the physiology associated with host feeding to the behavioural decision whether to host feed or oviposit. Below, I develop a new dynamic state variable model which incorporates the physiological features associated with host feeding inAphytis. My intention is to make qualitative predictions about the effects of various physiological featues on host-feeding behaviour and to make quantitative predictions about the relationship between egg load and host feeding inAphytis.     

Spontaneous chlorophyll mutants of Pennisetum americanum: Genetics and chlorophyll quantities

Summary Thirteen spontaneously occurring chlorophyll deficient phenotypes have been described and their genetic basis was established. Ten of these — white, white tipped green, patchy white, white virescent, white striping 1, white striping 2, white striping 4, fine striping, chlorina and yellow virescent showed monogenic recessive inheritance and the remaining three — yellow striping, yellow green and light green seedling phenotypes showed digenic recessive inheritance. The genes for (i) white tipped green (wr) and yellow virescent (yv) and (ii) patchy white (pw) and white striping 1 (wst 1) showed independent assortment. Further, the genes for white (w), white tipped green (wr) and yellow virescent (yv) were inherited independently of the gene for hairy leaf margin (Hm).In the mutants — white tipped green, patchy white, white striping 1, white striping 2, fine striping, chlorina, yellow virescent, yellow striping, yellow green and light green phenotypes total quantity of chlorophyll was significantly less than that in the corresponding controls, while in white virescent there was no reduction in the mature stage. For nine of the mutants the quantity of chlorophyll was also estimated in F₁'s (mutant x control green). In F₁'s of six of the mutants — white tip, patchy white, chlorina, yellow virescent, fine striping and yellow striping the quantity of chlorophyll was almost equal to the wild type. In the F₁'s of three of the mutants — white striping 1, white striping 2 and light green an intermediate value between the mutant and wild types was observed. In yellow virescent retarded synthesis of chlorophyll, particularly chlorophyll a was observed in the juvenile stage. Reduced quantity of chlorophyll was associated with defective chloroplasts. In the mutants — white tipped green, white virescent, fine striping, chlorina, yellow striping, yellow green and light green defective plastids were also observed. In patchy white secondary destruction of chlorophylls and the presence of defective plastids were found to be associated with reduced chlorophyll quantity at maturity.Paper chromatographic studies of leaf flavonoids revealed some variation between the inbreds, but there were three common spots, 7, 8 and 9, except for PDP in which the spot 8 was absent. Chlorophyll deficient mutants differed from their respective controls in the absence of one or more of the spots present in the controls and in the presence of new spots in some of the mutants.Most of the chlorophyll mutants showed higher survival rate in the Kharif season than in Rabi season which was attributed to the higher mean day temperature and longer day light period in the Kharif season than in Rabi season.     

Oviposition behavior of Anastrepha fraterculus in apple and diel pattern of activities in an apple orchard in Brazil

The oviposition behavior and diel pattern of activities of Anastrepha fraterculus (Wied.) (Diptera: Tephritidae) were observed in an apple orchard and fruit characteristics involved in oviposition preferences were investigated in field cage tests. Fruit size influenced fruit acceptability as an oviposition site by females which did not discriminate among the cultivars Gala, Fuji, and Golden Delicious when same-size fruits were presented simultaneously. Oviposition behavior in apples was basically the same as that described for primary hosts. Hourly census of fly activity indicated that adults did not overnight in the orchard and that they entered the orchard around 1100 h when temperature reached about 21 °C . Bird droppings were an important food item for adults. Behavioral differences between males and females might account for a significant biased sex ratio both in the orchard and at the edge of the native surrounding vegetation. Implications for fruit fly management in Brazilian apple orchards are discussed.     

Social mutilation in the Ponerine ant <Emphasis Type="Italic">Diacamma</Emphasis>: cues originate in the victims

Summary. In the queenless ponerine ant genus Diacamma, all workers eclose with a pair of innervated thoracic appendages termed gemmae. The gamergate (= mated egg laying worker) maintains reproductive monopoly by mutilating the gemmae of all eclosing individuals. Such mutilation leads to irreversible behavioural and neurological changes such that the individual lacking gemmae becomes incapable of appropriate sexual calling and mating. In one population related to Diacamma ceylonense from India, Diacamma sp. from Nilgiri (hereafter referred to as nilgiri), gamergates do not mutilate their nestmates and yet maintain reproductive monopoly. To understand what triggers mutilation, we exchanged cocoons between the mutilating D. ceylonense colonies and the non mutilating nilgiri colonies. nilgiri callows were not mutilated even in D. ceylonense colonies while D. ceylonense callows were mutilated even in nilgiri colonies, suggesting that the cues for mutilation originate in the victims (callows), presumably in the gemmae themselves. This finding should facilitate understanding the proximate mechanism and evolutionary significance of mutilation of gemmae as a method of resolution of reproductive conflicts in the genus Diacamma.Received 27 August 2003; revised 26 April 2004; accepted 27 April 2004.     

Food web structure of aphids and their parasitoids in Belgian fruit agroecosystems

Community structures of aphids and their parasitoids were studied in fruit crop habitats of eastern Belgium in 2014 and 2015. Quantitative food webs of these insects were constructed separately for each year, and divided into subwebs on three host‐plant categories, fruit crop plants, non‐crop woody and shrub plants and non‐crop herbaceous plants. The webs were analyzed using the standard food web statistics designed for binary data. During the whole study period, 78 plant species were recorded as host plants of 71 aphid species, from which 48 parasitoid species emerged. The community structure, aphid / parasitoid species‐richness ratio and trophic link number varied between the two years, whereas the realized connectance between parasitoids and aphids was relatively constant. A new plant–aphid–parasitoid association for Europe was recorded. Dominant parasitoid species in the study sites were Ephedrus persicae, Binodoxys angelicae and Praon volucre: the first species was frequently observed on non‐crop trees and shrubs, but the other two on non‐crop herbaceous plants. The potential influence, through indirect interactions, of parasitoids on aphid communities was assessed with quantitative parasitoid‐overlap diagrams. Symmetrical links were uncommon, and abundant aphid species seemed to have large indirect effects on less abundant species. These results show that trophic indirect interactions through parasitoids may govern aphid populations in fruit crop habitats with various non‐crop plants, implying the importance for landscape management and biological control of aphid pests in fruit agroecosystems.     

Dynamic heterogeneous spatio-temporal pattern formation in host-parasitoid systems with synchronised generations

In this paper we develop a general mathematical model describing the spatio-temporal dynamics of host-parasitoid systems with forced generational synchronisation, for example seasonally induced diapause. The model itself may be described as an individual-based stochastic model with the individual movement rules derived from an underlying continuum PDE model. This approach permits direct comparison between the discrete model and the continuum model. The model includes both within-generation and between-generation mechanisms for population regulation and focuses on the interactions between immobile juvenile hosts, adult hosts and adult parasitoids in a two-dimensional domain. These interactions are mediated, as they are in many such host-parasitoid systems, by the presence of a volatile semio-chemical (kairomone) emitted by the hosts or the hosts food plant. The model investigates the effects on population dynamics for different host versus parasitoid movement strategies as well as the transient dynamics leading to steady states. Despite some agreement between the individual and continuum models for certain motility parameter ranges, the model dynamics diverge when host and parasitoid motilities are unequal. The individual-based model maintains spatially heterogeneous oscillatory dynamics when the continuum model predicts a homogeneous steady state. We discuss the implications of these results for mechanistic models of phenotype evolution.P. Schofield gratefully acknowledges the financial support of the BBSRC and The Wellcome Trust.     

Diallel analysis of submergence tolerance in rice,Oryza sativa L.

Summary The genetics of submergence tolerance in rice was studied in a 10 × 10 half-diallel cross set involving 10 lowland rice varieties, four of which were tolerant (FR13A, FR43B, Kurkaruppan, and Goda Heenati) and the remaining six were nontolerant (RD19, IR42, IR17494-32-1, IR19672-24-3, Jagannath, and CR1009). Estimates of genetic parameters following Hayman's method showed significant additive and nonadditive gene action and the latter appeared to be solely due to dominance. Narrow sense heritability (0.70) indicated that additive gene effects were more important in the inheritance of the trait. Tolerance was dominant over nontolerance and the average dominance was within the range of incomplete dominance. Dominant alleles were more concentrated in the three tolerant parents, FR13A, Kurkaruppan, and FR43B in that order. Wr/Vr graphic analysis suggested the involvement of both major and minor genes. Combining ability analysis by Griffing's method also indicated significance of both additive and nonadditive effects, and the former appeared to be more important than the latter. The hybrids involving FR13A with RD19, IR42, and IR17494-32-1, and those of Kurkaruppan with RD19 and CR1009 appeared to be promising for incorporating an adequate level of tolerance to submergence into lowland rice cultivars.     

Cascading trophic interactions: Uncoupling at the zooplanktonphytoplankton link

Four limnocorrals (8 m dia. by 15 m deep) located at Lake St. George, Ontario, Canada; were used to examine the interactions between planktivorous fish, crustacean zooplankton (notably Daphnia galeata mendotae), and phytoplankton. During the spring, in all of the limnocorrals, high Daphnia biomasses were correlated with increased water transparency and a spring clear-water phase. However, as the summer progressed, the relationship between Daphnia biomass and phytoplankton abundance became more complex and less predictable. Investigation of these interactions suggested four conclusions. (1) During late July and throughout August and September, water transparency decreased and algal cell counts increased. In 3 of 4 limnocorrals, deterioration in water quality occurred 3–5 weeks before zooplankton (and Daphnia populations) declined. In all cases decreased transparency was associated with increased concentrations of algal cells (Gloeococcus) that were poor food sources for Daphnia. These results suggested that decreased water transparency was not caused by decreases in Daphnia biomass. (2) Taken together, data from all of the limnocorrals showed no correlation between the magnitude of July Daphnia biomasses and the percentage of poor food source algae that were observed in August. This suggested that grazer effects were not necessary for the onset of summer poor food source algal blooms. (3) In two limnocorrals, there was a positive correlation between increased Daphnia mortality and the onset of poor food source blooms. In the other two limnocorrals there was no correlation. In all limnocorrals there was no correlation between decreased Daphnia reproductive capacity and poor food source blooms. These data suggested that blooms of poor food source algae were not necessary for the collapse of Daphnia populations. (4) In all 4 limnocorrals there was a strong correlation between the time that 0+ yellow perch planktivores reached biomasses of 30–50 kg ha^–1 and the collapse of Daphnia populations Species and size selection was also observed. These results suggested that for this set of limnocorral experiments, fish biomasses in the 30–50 kg ha^–1 were responsible for the collapse of Daphnia populations in the summer.     

Effect of giving insulin and adrenalin alternately on changes in the level of blood glucose in chickens of different breeds

Summary Differences in reaction to exogenous insulin and adrenalin were studied among chickens of Leghorn, White Rock and Rhode Island Red breeds, using level of blood glucose as indicator of the reaction to hormones. It was found that: 1.) The physiological quantity of glucose in blood in chickens of the three breeds ranged from 157 to 194 mg %. 2.) The reaction of chickens to insulin and adrenalin given alternately, measured by changes of blood glucose, differed according to the breed. The smallest reaction to hormones was in Leghorn, the biggest in White Rock. 3.) The chickens of Leghorn and Rhode Island Red breeds, as well as Leghorn and White Rock breeds, differed significantly in maximal glucose level after adrenalin, 4.) The correlation between the normal quantity of blood glucose and the body weight of birds appeared highly significant in cockerels of all three races combined.