Population dynamics of the yew gall midge Taxomyia taxi (Inchbald) (Diptera: Cecidomyiidae)

Abstract. 1. The causes and incidence of mortality in populations of the yew gall midge Taxomyia taxi at Kingley Vale, West Sussex, are described in relation to the various life cycles of the midge.
2. Life table analysis of populations studied for 10 years indicates that the key factor in mortality is failure to achieve maximum fecundity, although in 1-year life cycles parasitism by Mesopolobus diffinis is equally important. Densities and mortality vary considerably between different life cycles, and no regulation has been firmly established.
3. Interactions between the host populations and their parasites are complex and as yet not fully analysable, but the pattern of parasitism seen may explain some of the complex life history pattern of the host. The existence of long-term cycles of host and parasite densities are predicted.     

Population dynamics of the yew gall midge Taxomyia taxi and its chalcid parasitoids: a 24-year study

Abstract.

• 1 The dynamics of three populations of Taxomyia taxi (Inchbald) (Diptera: Cecidomyiidae) and its chalcid parasitoids have been studied over a 24-year period. Most individuals have a 2-year life cycle but some develop in 1 year. Details of within-generation mortalities in T.taxi are used for life table analyses.
• 2 Mortality in the period between emergence of adult T.taxi and larval infestation of buds appears to be density-dependent and is the largest component of overall mortality. In 2-year life cycles, this mortality and that caused by Torymus nigritarsus (Walker) contribute equally to variance in overall mortality. In 1-year cycles, mortality caused by Mesopolobus diffinis (Walker) is density-dependent and accounts for most within-gall losses.
• 3 T. nigritarsus, which attacks only 2-year galls, is absent from all populations for a number of years in the middle of the study period. Its searching efficiency declines as its density and that of its host increase.
• 4 Densities of M.diffinis are strongly correlated with those of available hosts. Percentage parasitism of 2-year galls is less than that of 1-year galls, suggesting the occurrence of enemy-free space.
• 5 Although there are some correlations in densities and mortality between trees, the dynamics of populations on each are frequently different.
• 6 An earlier analysis of 10 years' data failed to demonstrate density-dependent effects. The extension to a 24-year run has revealed such effects, but also demonstrates long-term fluctuations in population densities, with corresponding changes in the balance of advantage between 1- and 2-year life cycles of T.taxi.

     

Parasite species richness in carnivores: effects of host body mass, latitude, geographical range and population density

Aim  Comparative studies have revealed strong links between ecological factors and the number of parasite species harboured by different hosts, but studies of different taxonomic host groups have produced inconsistent results. As a step towards understanding the general patterns of parasite species richness, we present results from a new comprehensive data base of over 7000 host–parasite combinations representing 146 species of carnivores (Mammalia: Carnivora) and 980 species of parasites.
Methods  We used both phylogenetic and non-phylogenetic comparative methods while controlling for unequal sampling effort within a multivariate framework to ascertain the main determinants of parasite species richness in carnivores.
Results  We found that body mass, population density, geographical range size and distance from the equator are correlated with overall parasite species richness in fissiped carnivores. When parasites are classified by transmission mode, body mass and home range area are the main determinants of the richness of parasites spread by close contact between hosts, and population density, geographical range size and distance from the equator account for the diversity of parasites that are not dependent on close contact. For generalist parasites, population density, geographical range size and latitude are the primary predictors of parasite species richness. We found no significant ecological correlates for the richness of specialist or vector-borne parasites.
Main conclusions  Although we found that parasite species richness increases instead of decreases with distance from the equator, other comparative patterns in carnivores support previous findings in primates, suggesting that similar ecological factors operate in both these independent evolutionary lineages.     

Ant mutualists alter the composition and attack rate of the parasitoid community for the gall wasp Disholcaspis eldoradensis (Cynipidae)

Abstract.  1. The strength or density dependence of pairwise species interactions can depend on the presence or absence of other species, especially potential mutualists.
2. The gall wasp Disholcaspis eldoradensis induces plant galls that secrete a sweet honeydew from their top surfaces while the wasp larvae are active. These galls are actively tended by Argentine ants, which collect the honeydew and drive off parasitoids attempting to attack the gall wasp.
3. When ants were excluded, the total rate of parasitism by seven species of parasitoids increased by 36%, and the rate of gall-wasp emergence decreased by 54%.
4. The total percentage parasitism was affected by gall density when ants were excluded but not when ants were unmanipulated, suggesting a change in parasitoid functional responses due to ant tending.
5. In addition, excluding ants significantly altered the proportions of different parasitoid species that emerged from galls; one parasitoid species increased from 1% to 34%, and another decreased from 46% to 19%.
6. The invasive Argentine ants studied are capable of maintaining the mutualism with the gall wasps that evolved in the presence of different ant species and also act as a selective filter for the local community of generalist parasitoids trying to attack this gall species.     

Host resources govern the specificity of swiftlet lice: size matters

1. An important component of parasite diversity is the specificity for particular host taxa shown by many parasites. Specificity is often assumed to imply adaptive specialization by the parasite to its host, such that parasites are incapable of surviving and reproducing on 'foreign' hosts.
2. Specificity, however, need not be due to adaptation to particular hosts. Some parasites may be specific simply because they are incapable of dispersing among host taxa. For example, 'permanent' parasites like chewing lice spend their entire lifecycle on the body of the host and require direct contact between hosts for dispersal.
3. The role of adaptive constraints in parasite host-specificity has seldom been tested in natural populations. We conducted such a test by comparing the relative fitness of host-specific lice experimentally transferred among closely related species of cave swiftlets in northern Borneo.
4. The survival of lice in most of these transfers was significantly reduced in proportion to the mean difference in feather barb size between the donor and recipient species of hosts. Thus, adaptation to a particular resource on the body of the host does appear to govern the specificity of swiftlet lice.
5. In transfers where lice survived, microhabitat shifting on the body of the host was observed, whereby the mean barb diameter of the feathers on which the lice occurred was held 'constant'.     

On understanding seasonal increases in damselfly defence and resistance against ectoparasitic mites

Abstract.  1. Defence against parasites and pathogens can be essential, yet not all hosts respond similarly to parasitic challenge. Environmental conditions are thought to explain variation in host responses to parasites.
2.  Lestes forcipatus damselflies emerging later in the season have shown higher resistance to the mite, Arrenurus planus , than hosts emerging earlier. This study was undertaken to determine whether variation in environmental temperatures characteristic of early vs. late emergence times, degree or costs of mite parasitism, and/or size of newly emerged adults could explain seasonal variation in defence and resistance to ectoparasitic mites.
3. In this study damselflies from early vs. late emergence groups differed in size at emergence and mite intensity. In general, early hosts were larger and had more mites than later hosts. However only experimental temperatures experienced by damselflies at emergence influenced defence and resistance against mites and not host size or degree of parasitism.
4. More specifically, hosts from early and late emergence groups did not differ in defence and resistance when held at the same temperatures in incubators. Housing at a high temperature, indicative of later in the season, was associated with higher defence and resistance for damselflies from both early and late emergence groups.
5. These results indicate that daily temperatures in relation to emergence timing can account for seasonal increases in resistance for this temperate insect. Seasonal increases in resistance may be expected for other temperate insect–parasite associations and should have important implications for the phenology of parasites and for seasonal variation in parasite-mediated selection.     

Host gall size and oviposition success by the parasitoid Eurytoma gigantea

Abstract. 1. Eurytoma gigantea Walsh is a specialist parasitoid of the tephritid gallmaker Eurosta solidaginis (Fitch).
2. In the natural environment the incidence of parasitism by Eurytoma is greater in small galls than in large ones.
3. Laboratory experiments demonstrated that small galls are not more frequently discovered; however, oviposition attempts on small galls were more likely to be successful.
4. Eurytoma spends much time probing galls too big to penetrate; this leads to a decrease in foraging efficiency when many large galls are present.
5. The chance of successfully penetrating a gall depends on the thickness of the gall wall and the length of the parasitoid's ovipositor.
6. A simulation model was constructed which shows that a gallmak-er's chance of being parasitized depends on gall size, the number of parasitoids that discover the gall, and their ovipositor lengths.     

The origin and genetic differentiation of the socially parasitic aphid Tamalia inquilinus

Social and brood parasitisms are nonconsumptive forms of parasitism involving the exploitation of the colonies or nests of a host. Such parasites are often related to their hosts and may evolve in various ecological contexts, causing evolutionary constraints and opportunities for both parasites and their hosts. In extreme cases, patterns of diversification between social parasites and their hosts can be coupled, such that diversity of one is correlated with or even shapes the diversity of the other. Aphids in the genus Tamalia induce galls on North American manzanita (Arctostaphylos) and related shrubs (Arbutoideae) and are parasitized by nongalling social parasites or inquilines in the same genus. We used RNA sequencing to identify and generate new gene sequences for Tamalia and performed maximum‐likelihood, Bayesian and phylogeographic analyses to reconstruct the origins and patterns of diversity and host‐associated differentiation in the genus. Our results indicate that the Tamalia inquilines are monophyletic and closely related to their gall‐forming hosts on Arctostaphylos, supporting a previously proposed scenario for origins of these parasitic aphids. Unexpectedly, population structure and host‐plant‐associated differentiation were greater in the non‐gall‐inducing parasites than in their gall‐inducing hosts. RNA‐seq indicated contrasting patterns of gene expression between host aphids and parasites, and perhaps functional differences in host‐plant relationships. Our results suggest a mode of speciation in which host plants drive within‐guild diversification in insect hosts and their parasites. Shared host plants may be sufficient to promote the ecological diversification of a network of phytophagous insects and their parasites, as exemplified by Tamalia aphids.     

Plant sex alters Allee effects in aggregating plant parasites

Species interactions are central to our understanding of population dynamics. While density typically strengthens competition, reducing absolute fitness, Allee effects can reverse this pattern, increasing fitness with density. Allee effects emerge in host–parasite systems when higher parasite densities dilute immune responses or increase resource‐mobilization. The optimal density of individuals in these systems should be influenced by how host quality alters the rates at which facilitative and competitive effects change across densities. We tested these ideas using sumac Rhus typhina and a gall‐forming parasite Melaphis rhois that attacks sumac leaves. Fitness peaked at intermediate densities, indicating an Allee effect, but the fitness peak depended on host sex. Patterns of abundance mirrored fitness patterns, with galls clustered on leaves and female hosts supporting greater numbers of galls. Within leaves, galls near the stem were more fit, and gall‐makers preferentially oviposited near to the stem. The patterns of fitness and abundance are consistent with Allee effects caused by increased resource mobilization at higher gall‐maker densities rather than diluted immune responses. Our results suggest that Allee effects in parasites can be described as the summative effects of competitive and facilitative processes and, because both are common, Allee effects are likely common in host–parasite systems.     

Patterns of kleptoparasitism and inquilinism in social and non-social Dunatothrips on Australian Acacia

Abstract.  1. Australian Acacia thrips and their parasitic exploiters provide a novel system for analysing fundamental questions about host–parasite evolution and ecology.
2. In this study, colony collections were used to examine interactions between members of the genus Dunatothrips and the diverse assemblage of parasitic thrips that invade domiciles constructed by the host on various species of Acacia .
3.  Dunatothrips parasites adopt a range of strategies for exploiting their hosts, including lethal fighting, host eviction, and cohabitation.
4. Social Dunatothrips ( D. aneurae and D. skene ) were more frequently parasitised than non-social D. gloius , which could be related to differences in social organisation or the fact that D. gloius has shifted onto a novel host that is distantly related to other Dunatothrips ' hosts.
5. Parasitism rates of D. aneurae and D. skene domiciles were high (46% and 51% respectively), and were positively density dependent.     

The fecundity, development and host relationships of Ceratobaeus spp. (Hymenoptera: Scelionidae), parasites of spider eggs

Abstract. 1. Ceratobaeus spp. enter the nest of their host spider and oviposit into eggs through the thin silk eggsac.
2. Temperatures below 15° C limit oviposition by slowing parasites down.
3. Males emerge prior to females and mate with their sibs. Sex ratios of 6.6–6.0:1 in favour of females were observed in the field and laboratory for two species.
4. Even though females can oviposit almost immediately after emergence their full complement of eggs (= 65) is not reached until several days later.
5. Superparasitism is low and appears to occur from subsequent accidental ovipositions. No marking of the external surface of hosts was observed.
6. Parasites overwinter as adults under bark. They do not feed as adults nor do they reabsorb their eggs, but rather oviposit as soon as host eggs become available in spring.
7. Successful oviposition occurs in later stages of host eggs reared at 15° C and 20° C than it does at 25°C. Different rates of development between host and parasite is proposed as an explanation for this phenomenon.
8. The species studied show varying degrees of specificity but each has a dominant host. Location of hosts involves cues from the habitat (bark), silk nests of spiders, and some factor associated with host eggs.     

How to analyse host discrimination

Abstract. 1. Two usual definitions of host discrimination are presented. The first is 'the ability of a parasite to distinguish unparasitized from parasitized hosts and to lay eggs in the former'. This definition is not useful and even confusing since it does not include the ability of a parasite to distinguish hosts containing different numbers of parasite eggs and to lay preferably only in those with the lowest numbers.
2. The second definition is 'the ability of a parasite to distribute its eggs in a non-random, regular way among its hosts'. It is argued that most field data are insufficient to permit any conclusions about host discriminative ability on the basis of this definition. An example is given of an apparent random distribution by parasites that are perfectly able to discriminate.
3. Arguments are given for studying the behaviour of the parasite in order to answer the question of host discrimination. Five examples of parasites are presented that would erroneously have been classified as non-discriminators on the basis of the first definition, since they all superparasitize.
4. It is suggested that the meaning of the term 'host discrimination' be extended to include the ability of a parasite to distinguish hosts with different numbers of eggs.     

Associational resistance mediated by natural enemies

Abstract.  1. Associational resistance theory suggests that the association of herbivore-susceptible plant species with herbivore-resistant plant species can reduce herbivore density on the susceptible plant species. Several casual mechanisms are possible but none has so far invoked natural enemies. Associational resistance mediated by natural enemies was tested for by examining densities of a gall fly, Asphondylia borrichiae (Diptera: Cecidomyiidae), and levels of parasitism on two closely related seaside plants, Borrichia frutescens and Iva frutescens , when alone and when co-occurring.
2. Both Borrichia and Iva grow alone or together on small offshore islands in Florida. Each host plant species has its own associated race of fly, but both races of fly are attacked by the same four species of parasitoids. Borrichia normally has a higher density of galls than Iva , and galls are larger on Borrichia than on Iva .
3. Gall size, gall abundance, parasitism levels, and parasitoid community composition were quantified on both Borrichia and Iva on islands where each species grew alone or together. Some islands were then manipulated by adding Borrichia to islands supporting only Iva , and by adding Iva to islands supporting only Borrichia . Subsequent gall densities and gall parasitism levels on the original native species were then examined.
4. On both natural and experimentally manipulated islands, gall densities on Iva were significantly lowered by the presence of Borrichia . This is because bigger parasitoid species that were common on Borrichia galls, which are bigger, spilled over and attacked the smaller Iva galls. Thus, parasitism rates on Iva were higher on islands where Borrichia co-occurred than on islands where Borrichia were absent. Most parasitoids from Iva were too small to successfully attack the large Borrichia galls and so gall density on Borrichia was unaffected by the presence of Iva .     

On the capacity of macroparasites to control insect populations

A graphical model of the population dynamics of macroparasites and their hosts is developed. Three principal means by which the parasites can be regulated are considered: reduction in host density as a result of parasite-induced host mortality, reduction in host density as a result of parasite-induced host sterility, and competition among parasites within multiply-infected hosts. The means by which parasites are regulated has a major effect on the degree to which they can depress host population densities. In particular, a parasite that sterilizes its host is expected to reduce host density more than one that causes an equivalent decline in host fitness through increased mortality. A special case of the model is developed for herbivorous insects that, in the absence of parasites, are limited by larval food resources. Parasites that are regulated via parasite-induced host sterility will control the insect populations below the level set by larval resources if the threshold host density for the parasites (N(T)) is less than the ratio of carrying capacity to net reproductive rate of the insects (K/R). Data are presented showing that all three means of parasite regulation, but especially parasite-induced host sterility, can operate in Howardula aoronymphium, a nematode parasite of mycophagous Drosophila flies. Data from a field cage experiment show that, if these nematodes are regulated primarily via reductions in host density due to this sterility, the parameters N(T), K, and R are such that Howardula is likely to play an important role in controlling Drosophila populations. However, this conclusion must be tempered by the fact that these nematodes also cause increased host mortality and experience within-host competition, making the conditions for parasite control of the flies more stringent.     

Plant resistance, plant traits, and host plant choice of the leaf-folding sawfly on the arroyo willow

Abstract. 1. The hypotheses that genetic variation in host plant resistance of the arroyo willow affected leaf folder ( Phyllocolpa sp.) (Hymenoptera: Tenthredinidae) density and that genetic variation in shoot length and leaf length was correlated with resistance were tested.
2. Willows grown in pots and exposed to ovipojsition by the leaf folding sawfly in cages had significantly different densities among clones, indicating variation in resistance caused by genetic differences among conspecific host plants.
3. There was a general correspondence between leaf folder density on potted cuttings and on the plants in the field that were the sources of cuttings.
4. In behavioural choice experiments, susceptible clones (with highest leaf fold densities) had the highest oviposition activity of female leaf folders compared to clones that were resistant to the leaf folder.
5. Clones differed significantly in shoot length and leaf length among clones grown in pots, among clones in the field, and between shoots with galls and shoots without galls on clones in the field.
6. Leaf folder density was significantly positively correlated with mean shoot length on field clones in 1985 and 1986, but was not correlated with leaf length, although leaf length and shoot length were correlated.
7. Leaf length variation among willow clones accounted for a significant portion of the variation in resistance of potted willows, but shoot length was unimportant.     

Sex ratio in relation to season and host plant quality in a monogenous stem-galling midge (Diptera: Cecidomyiidae)

Abstract.  1. The sex determination mechanism in gall midges is little understood, although it is known that the females of several species primarily or exclusively produce unisexual broods throughout their lifetime.
2. The gall midge Izeniola obesula Dorchin is a multivoltine species, inducing multi-chambered stem galls on the salt-marsh plant Suaeda monoica . Each gall contains 5–70 individuals, all being the progeny of a single female. Sampling of more than 450 galls, from which adult midges were reared, suggested that I .  obesula exhibits strict monogeny, resulting in galls that contain either all female or all male progeny.
3. Characterisation of the growth pattern of young S .  monoica shoots revealed that shoots in apical positions grew more rapidly than shoots in more basal positions. Galls that were induced on such shoots were larger and yielded more adult midges.
4. No difference in the site of gall induction was found between male and female galls, with galls of either sex being mainly induced on more rapidly growing shoots. It is concluded that I .  obesula females cannot control the sex of their progeny, and that both female-producers and male-producers strive to maximise their reproductive success by choosing the faster-growing shoots for gall induction.
5. Female galls were larger and more abundant than male galls at almost all times. The sex ratio among galls fluctuated throughout the year, ranging from 4:1 in spring to 1:1 in winter. The skewed sex ratio among galls possibly results from greater mortality rates among male galls than among female galls, due to either primary or secondary factors. Alternatively, it is possible that the number or fitness of male-producers in the population is reduced relative to female-producers.     

Opposing survivorship and fecundity effects of host phenology on the gall-forming aphid Hormaphis hamamelidis

Abstract  1. This study examined the synchronisation and roles of host and insect phenology for fundatrices of the gall-forming aphid Hormaphis hamamelidis , which forms galls on the leaves of the understorey tree Hamamelis virginiana .
2. Host and insect phenologies varied widely among years and individual host trees.
3. Most winter eggs (≈ 85%) hatched before budbreak and galls could be initiated but in 2 years there was no inherent risk for earlier hatching fundatrices in terms of greater mortality. Egg hatch was not synchronised tightly with host trees but instead seemed to occur well in advance of budbreak as a conservative, bet-hedging strategy to ensure the opportunity to form galls.
4. Fundatrices that formed galls later had higher fecundity, contrary to typical patterns for insects that feed on the first flush of leaves in temperate forests, where later herbivores have lower fecundity. Also, unlike many other systems, trees that had later budbreak had greater densities of galls.     

A review of the species of Mesopolobus (Chalcidoidea: Pteromalidae) associated with Ceutorhynchus (Coleoptera: Curculionidae) host-species of European origin

Four species of Mesopolobus Westwood were reared as parasitoids of Ceutorhynchinae hosts in Europe during surveys in 2000-2004. An illustrated key is given to differentiate the four species, M. gemellus Baur & Muller sp. n., M. incultus (Walker), M. morys (Walker) and M. trasullus (Walker), plus M. moryoides Gibson, a parasitoid of the cabbage seedpod weevil, Ceutorhynchus obstrictus (Marsham), in North America. Pteromalus clavicornis Walker is recognized as a junior synonym of M. incultus syn. n., and Pteromalus berecynthos Walker (also a junior synonym of M. incultus) is considered a correct original spelling. For Disema pallipes F?rster (a junior synonym of Mesopolobus morys), a lectotype is designated. Mesopolobus morys is for the first time accurately associated with the seed weevil Ceutorhynchus turbatus (Schultze), a potential agent for classical biological control, of hoary cress, Lepidium draba L. (Brassicaceae), in North America. Mesopolobus gemellus is associated with another seed weevil, Ceutorhynchus typhae (=C. floralis) (Herbst), in pods of shepherd's purse, Capsella bursa-pastoris (L.) Medik. (Brassicaceae). Implications of the host-parasitoid associations are discussed relative to the introduction of species to North America for classical biological control of the cabbage seedpod weevil.     

Gall aphids do not select optimal galling sites (Smynthurodes betae; Pemphigidae)

Abstract. 1. In studies of insect-host plant interaction it is often suggested that insects preferentially colonize host plants (or sites within plants) on which their fitness is maximized (a positive covariance of preference and performance). This suggestion stems from the assumption that natural selection has driven the system toward optimal use of resources.
2. Our study of the galling aphid Smynthurodes betae Westw. demonstrates that the distribution of galls on leaves is not due to preference, and can be altered by manipulating the aphid arrival time or the shoot growth rate.
3. We found no correlation between gall density and performance (aphid clone size) at different positions along the shoot.
4. Because leaves on the growing shoot are not equally responsive to aphid stimulation, the colonizers have no choice but to settle on leaves that are at the right stage when they arrive.
5. S.betae colonizers did not discriminate between shoots of their host and a congeneric non-host, on which their fitness is invariably zero.
6. Synchronization between galler and host plant phenologies seems to be the key to the observed distribution of galls on the tree. The data give no support to the preference-performance hypothesis.     

Specialist by preference,generalist by need: availability of quality hosts drives parasite choice in a natural multihost–parasite system

Encountering suitable hosts is key for parasite success. A general assumption for disease transmission is that the contact of a parasite with a potential host is driven by the density or relative frequency of hosts. That assumption ignores the potential role of differential host attractiveness for parasites that can drive the encounter of hosts. It has been posited that hosts may be chosen by parasites as a function of their suitability, but the existing literature addressing that hypothesis is still very scarce. In a natural system involving a parasitic Philornis botfly and its multiple bird hosts, there are profound differences in host quality. The Great Kiskadee tolerates and does not invest in resisting the infection, which makes it an optimal host. Alternative hosts are frequently used, but whilst some of them may be good options, others are bad alternatives. Here we examined the host selection processes that drive parasite dynamics in this system with 8 years of data from a longitudinal study under natural conditions. We found that the use of an alternative host was not driven by its density or relative frequency, but instead selection of these hosts was strongly dependent on availability of more suitable hosts. When optimal hosts are plentiful, the parasite tends to ignore alternative ones. As broods of optimal hosts become limited, good alternative hosts are targeted. The parasite chooses bad alternative hosts only when better alternatives are not sufficiently available. These results add evidence from a natural system that some parasites choose their hosts as a function of their profitability, and show that host selection by this parasite is plastic and context-dependent. Such findings could have important implications for the epidemiology of some parasitic and vector-borne infections which should be considered when modelling and managing those diseases. The facultative host selection observed here can be of high relevance for public health, animal husbandry, and biodiversity conservation, because reductions in the richness of hosts might cause humans, domestic animals, or endangered species to become increasingly targeted by parasites that can drive the encounter of hosts.