Density-dependent host selection in ectoparasites: An application of isodar theory to fleas parasitizing rodents

Parasites should make the same decisions that every animal makes regarding fitness reward. They can maximize reproductive success by selection of those habitats that guarantee the greatest fitness output. We consider the host population as a habitat of a parasite population. Consequently, hosts (=habitats) that differ quantitatively or qualitatively will support different numbers of parasites. The nature of habitat selection can be detected by isodars, lines along which habitat selection yields equivalent fitness reward. We applied this approach to study host selection of five fleas, each infesting two desert rodents. Xenopsylla conformis, Xenopsylla ramesis, Nosopsyllus iranus theodori and Stenoponia tripectinata medialis parasitize Gerbillus dasyurus and Meriones crassus. Synosternus cleopatrae pyramidis parasitizes Gerbillus andersoni allenbyi and Gerbillus pyramidum. Three fleas ( X. conformis, X. ramesis and S. c. pyramidis) were able to perceive quantitative (amount of the resource; e.g. organic matter in the nest for flea larvae) and/or qualitative (pattern of resource acquisition; e.g. host defensiveness) differences between hosts. Two other fleas did not perceive between-host differences. X. conformis was a density-dependent host selector that showed sharp selectivity at low density. X. ramesis and S. c. pyramidis were density-independent host selectors with a direct correspondence of density with habitat quality. N. i. theodori and S. t. medialis were non-selectors with no relationship at all between density and host quality. The results of the application of the isodar theory suggest that ectoparasites, like other animals, behave as if they are able to make choices and decisions that favour environments in which their reproductive benefit is maximized.     

Geographical range size and host specificity in ectoparasites: a case study with Amphipsylla fleas and rodent hosts

Aim  To identify the factors that determine the geographical range sizes of ectoparasites with different degrees of host specificity.
Location  The study used data on the distributions of fleas of the genus Amphipsylla and their rodent hosts across the Holarctic.
Methods  All known points of occurrence of 32 flea species and 51 species of their rodent hosts were mapped. The shape and size of the geographical range of each species were estimated using a combination of the minimal convex polygon technique and modelling with the garp algorithm. Factors determining the geographical range sizes of the fleas were identified using stepwise multiple regression analysis.
Results  The geographical range size of fleas that are strongly host-specific across their entire ranges correlated positively with the geographical range size of the fleas' principal hosts, and negatively with the geographical range size of the fleas' potential competitors. The geographical range sizes of both (1) fleas that are locally host-specific but that shift their host preferences geographically, and (2) host-opportunistic fleas were positively correlated only with the area of the geographical ranges of their principal hosts. Strongly host-specific fleas occupied 0.2–80.0% of the geographical range of their principal hosts, whereas this figure was 0.9–83.7% in locally host-specific fleas and 16.6–63.7% in host-opportunistic fleas.
Main conclusions  The main determinant of the geographical range size of a flea species is the size of the geographical range of its hosts. The role of potential competitors in determining the geographical range size is stronger in host-specific than in host-opportunistic fleas. Cases in which the geographical range of a parasite is smaller than the geographical range(s) of its host(s) owing to narrower parasite environmental tolerances are much more frequent in host-opportunistic than in host-specific fleas.     

云南省玉龙鼠疫疫源地野外鼠形动物寄生蚤丰盛度影响因素分析

【目的】分析云南省玉龙鼠疫疫源地野外鼠形动物寄生蚤丰盛度的影响因素。【方法】选取云南省玉龙鼠疫疫源地3个海拔区域,按4个季节进行野外捕鼠,捕获的鼠形动物用梳检法收集体表寄生蚤并在显微镜下分类鉴定。通过实际测量和实地观察相结合的方式收集潜在影响鼠形动物寄生蚤丰盛度的因素包括鼠形动物特征变量指标(如种类、年龄、性别、体长、体重)、环境和气象因子(如海拔、季节)等数据。采用EpiData 3.02软件建立数据库,在R软件下使用跨栏负二项分布回归分析鼠形动物寄生蚤丰盛度的影响因素。【结果】从捕获的884只鼠形动物中检获寄生蚤9种484头,以特新蚤指名亚种、方叶栉眼蚤、无值大锥蚤、云南栉眼蚤为主(86.16%)。回归分析显示： 2 700-3 000 m和3 000 m以上海拔鼠形动物染蚤概率较2 400-2 700 m分别增加1.27和3.72倍;湿度高于70%时,鼠形动物染蚤概率较湿度≤70%时减少41%;与齐氏姬鼠的染蚤概率相比,中华姬鼠的染蚤概率降低50%,大绒鼠的染蚤概率增加79%;体长超过104 mm的鼠形动物染蚤概率较体长≤104 mm的鼠形动物染蚤概率增加76%;气温高于15℃时,鼠形动物染蚤数量较温度≤15℃时降低67%;成年鼠形动物的染蚤数量较未成年鼠形动物的染蚤数量增加2.25倍;与春季相比,夏季的染蚤数量增加2.00倍,秋季的染蚤概率减少48%,冬季的染蚤概率和染蚤数量分别增加1.44和1.06倍。【结论】玉龙鼠疫疫源地野外鼠形动物寄生蚤以特新蚤指名亚种、方叶栉眼蚤、无值大锥蚤、云南栉眼蚤为优势蚤种。鼠形动物寄生蚤丰盛度与海拔、季节、气温、湿度等环境气象因子及鼠形动物种类、体长、年龄等鼠形动物特征变量密切相关。     

Fitness effects of sex ratio response to host quality and size in the parasitoid wasp Spalangia cameroni

The parasitoid wasp Spalangia cameroni oviposited a greaterproportion of daughters in stable fly pupae than in house flypupae, even when I controlled for stable flies being smallerthan house flies. Sex ratio manipulation in response to hostquality has been modeled as being adaptive through an effectof host quality on the size and hence offspring production ofdaughters. 5. cameronis response to host species may insteadbe adaptive through an effect on larval survivorship, the developmenttime of daughters, and the size of sons. There was greater survivalof daughters than sons on stable flies. Controlling for hostsize, I found that development time of daughters was about 2%less on stable flies than on house flies. The decrease in developmenttime corresponds to a 2% increase in fitness as estimated byr, the intrinsic rate of increase, and is equivalent to abouta 9% increase in offspring production. Sons were about 2% largerfrom house flies than stable flies, which may increase offspringproduction by up to 3%. Host species had no consistent effecton size of daughters or development time of sons. In additionto the response to host species, mothers oviposited a greaterproportion of daughters in larger stable fly hosts. Whetherthis behavior is adaptive is unclear. Although offspring werelarger when they developed on larger stable flies, the rateof increase was less for daughters dian for sons. Effects ofstable fly size on offspring development time were negligible.     

Fitness consequences of pheromone production and host selection strategies in a tree-killing bark beetle (Coleoptera: Curculionidae: Scolytinae)

Timing of arrival at a resource often determines an individual’s reproductive success. Tree-killing bark beetles can reproduce in healthy trees by attacking in adequate numbers to overcome host defences that could otherwise be lethal. This process is mediated by aggregation and antiaggregation pheromones. Beetles that arrive early in such a “mass attack” must contend with undiminished tree defences, and produce enough pheromones to attract more beetles, but have a head start on gallery construction and egg-laying. Beetles that arrive late may be impeded by competition and diminishing availability of phloem, but should experience fewer costs associated with pheromone production and battling tree defences. We investigated relationships between timing of arrival, body size, pheromone production and fitness in the southern pine beetle, Dendroctonus frontalis. In field experiments, we captured beetles that arrived early (pioneers) and late on slash pine trees, Pinus elliottii, and measured pheromone amounts in their hindguts. We marked gallery entrances of beetles as they landed on a tree and measured their reproductive success after the attack terminated. We found no difference in body size or pheromone amounts between early and late arrivers. Most beetles arrived at the middle of the attack sequence, and excavated longer galleries per day than early arrivers. The number of offspring produced per day by beetles that established galleries midway through mass attack was higher than those that arrived early or very late in the sequence. Our results suggest that beetles do not exhibit adaptive phenotypic plasticity in pre-landing pheromone production, depending on the extent of previous colonisation of a host. Rather, it appears that stabilising selection favours beetles that attack in the middle of the sequence, and contributes to attack synchrony. Synchronous attack on trees is essential before population booms characteristic of tree-killing bark beetles can occur in nature.     

广聚萤叶甲成虫取食经历对后续 产卵寄主选择的影响

【背景】取食经历对植食性昆虫的寄主选择行为具有较大影响,影响天敌昆虫寄主专一性测定的设计和结果解释。【方法】采用选择性试验,观察了入侵豚草的重要天敌——广聚萤叶甲成虫羽化后取食不同植物对其后续产卵寄主选择的影响。【结果】与取食豚草的试虫相比,有取食三裂叶豚草、苍耳或菊芋经历的成虫选择苍耳产卵的频次增加,不再对豚草表现出明显的选择偏好性。对产卵识别期的Cox模型分析结果表明,成虫早期取食不同植物,对后续产卵选择有显著影响,成虫羽化后如果先取食豚草或三裂叶豚草,则选择苍耳产卵的倾向显著低于豚草;但如果先取食苍耳、菊芋和农家向日葵,则选择苍耳产卵的倾向与豚草无显著差异。【结论与意义】由此推测,广聚萤叶甲初羽化成虫取食的植物对其后续产卵选择具有较大影响,因而在寄主专一性测定中应关注测试前饲喂的植物种类。     

Riding across the selection landscape: fitness consequences of annual variation in reproductive characteristics

Evolutionary models estimating phenotypic selection in character size usually assume that the character is invariant across reproductive bouts. We show that variation in the size of reproductive traits may be large over multiple events and can influence fitness in organisms where these traits are produced anew each season. With data from populations of two orchid species, Caladenia valida and Tolumnia variegata, we used Bayesian statistics to investigate the effect on the distribution in fitness of individuals when the fitness landscape is not flat and when characters vary across reproductive bouts. Inconsistency in character size across reproductive periods within an individual increases the uncertainty of mean fitness and, consequently, the uncertainty in individual fitness. The trajectory of selection is likely to be muddled as a consequence of variation in morphology of individuals across reproductive bouts. The frequency and amplitude of such changes will certainly affect the dynamics between selection and genetic drift.     

New host,geographic records,and histopathologic studies of Angiostrongylus spp (Nematoda: Angiostrongylidae) in rodents from Argentina with updated summary of records from rodent hosts and host specificity assessment

To date, 21 species of the genus Angiostrongylus (Nematoda: Angiostrongylidae) have been reported around the world, 15 of which are parasites of rodents. In this study, new host, geographic records, and histopathologic studies of Angiostrongylus spp in sigmodontine rodents from Argentina, with an updated summary of records from rodent hosts and host specificity assessment, are provided. Records of Angiostrongylus costaricensis from Akodon montensis andAngiostrongylus morerai from six new hosts and geographical localities in Argentina are reported. The gross and histopathologic changes in the lungs of the host species due to angiostrongylosis are described. Published records of the genus Angiostrongylus from rodents and patterns of host specificity are presented. Individual Angiostrongylusspecies parasitise between one-19 different host species. The most frequent values of the specificity index (STD) were between 1-5.97. The elevated number of host species (n = 7) of A. morerai with a STD = 1.86 is a reflection of multiple systematic studies of parasites from sigmodontine rodents in the area of Cuenca del Plata, Argentina, showing that an increase in sampling effort can result in new findings. The combination of low host specificity and a wide geographic distribution of Angiostrongylus spp indicates a troubling epidemiological scenario although, as yet, no human cases have been reported.     

Energetics of reproduction: consequences of divergent selection on egg size, food limitation, and female age for egg composition and reproductive effort in a butterfly

Using lines artificially selected on egg size and being subjected to a restricted and an unrestricted feeding treatment, we examined the relationships between egg size, egg number, egg composition, and reproductive investment in the butterfly Bicyclus anynana . Despite a successful manipulation of egg size, correlated responses to selection in larval time, pupal mass, pupal time, longevity, fecundity, or the amount of energy allocated to reproduction were virtually absent. Thus, there was no indication for an evolutionary link between offspring size and reproductive investment. Egg composition, in contrast, was affected by selection, with larger eggs containing relatively more lipid and water, but less protein and energy compared to smaller eggs. Hence, females producing large eggs did not have to sacrifice fecundity due to adjustments in egg composition. Food limitation per se caused only minor changes in egg composition, and there was no general reduction in egg provisioning with female age. The latter was restricted to food-limited females, whereas egg quality remained remarkably similar throughout the females' life in control groups. We conclude that neglecting changes in biochemical egg composition, depending on genetic background, food availability, and female age, may introduce substantial error when estimating reproductive effort, and may ultimately lead to invalid conclusions.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 91 , 403–418.     

Within‐plant variation in reproductive investment: consequences for selection on flowering time

Variation among the leaves, flowers or fruit produced by a plant is often regarded as a nuisance to the experimenter and an impediment to selection. Here, we suggest that within‐plant variation can drive selection on other plant‐level traits. We examine within‐plant variation in floral sex allocation and in fruit set and predict that such variation generates variation in male success among plants, thereby driving selection on flowering time. We tested this prediction in a simulation model estimating selection on flowering time through male fitness when floral sex allocation and/or fruit set vary directionally among flowers on plants. We parameterized the model through a quantitative literature survey of within‐plant change in sex allocation. As predicted, within‐plant variation in floral sex allocation and in fruit set probability can generate selection on flowering time through male fitness. Declining fruit set from first to last flowers on plants, as occurs in many species, selected for early flowering onset through male fitness. This result was robust to self‐incompatibility and to varying returns on male versus female investment. Selection caused by declining fruit set was strong enough to reverse the selection for late flowering that can be caused by intrafloral protandry. Our model provides testable predictions regarding selection on flowering time through male fitness. The model also establishes the intriguing possibility that within‐plant variation may influence selection on other traits, regardless of whether that variation is under selection itself.     

Sex ratio manipulation by the parasitoid waspSpalangia cameroni in response to host age: A test of the host-size model

Summary A sex ratio response to host resources as measured by external host dimensions has been demonstrated in many parasitoid wasps, includingSpalangia cameroni. The responses generally are in the direction predicted by sex ratio theory, specifically the host-size models. Here I show that femaleS. cameroni also respond to differences in resource availability not associated with changes in external host dimensions, and this response is in the direction predicted by host-size models. When given old and young hosts simultaneously, femaleS. cameroni oviposit a greater proportion of sons in old than in young host pupae, at least for 0-day old versus 3-day old hosts. Old hosts weigh less than young hosts but are not significantly different in external width. Thus it appears that the offspring sex ratio response may result from mothers detecting physical or chemical changes within the host which are associated with host age. No evidence is found that the manipulation in response to host age has been selected for via an effect of host age on wasp size; there was no significant effect of host age on either male of female wasp size. A second prediction of the host-size models is also supported by this study: when each female is presented with only a single host age, rather than two host ages simultaneously, host age has no effect on offspring sex ratio.     

Host selection and egg allocation behaviour by Aphytis melinus and A.lingnanensis: comparison of two facultatively gregarious parasitoids

Abstract. 1. The ovipositional and egg allocation behaviour of individual females of Aphytis melinus DeBach and A.lingnunensis Compere were compared.
2. Both Aphytis species exhibit the same behavioural sequence during oviposition.
3. Aphytis melinus laid most of its female eggs on the dorsum of a scale-insect beneath its cover, and most of its male eggs under the scale-insect's body. Aphytis lingnanensis also oviposited both dorsally and ventrally on scale-insect hosts, but female and male progeny arose with equal frequency from eggs laid in both locations.
4. Both A.melinus and A. lingnanensis are facultatively gregarious parasitoids. The degree of gregariousness depends on host size, i.e. the larger the host, the more the Iikelihood that several eggs will be deposited at each visit by the parasitoid.
5. When two eggs were laid during the same host visit, both A.melinus and A.lingnanensis laid one female and one male egg more often than would be expected under an assumption of random allocation of sexes.
6. Because A.melinus successfulIy utilize smaller hosts than A.lingnanensis to produce progeny, these parasitoids should not be considered ecological homo-logues, as suggested by DeBach & Sundby (1963).     

Interactions among intraspecific competition, emergence patterns, and host selection behaviour in Ips pini (Coleoptera: Scolytinae)

Abstract.  1. Variations in developmental conditions of bark beetles, particularly intraspecific competition, can induce morphological and physiological modifications in the offspring. It is hypothesised that intraspecific competition could also affect host selection behaviour. Such behavioural changes might be manifested in response to host (alpha-pinene) or beetle (verbenone) compounds.
2.  Ips pini were bred at different densities. The offspring were measured for size, weight, and lipid concentration, and then subjected to tunnelling bioassays in agar media amended with varying amounts of alpha-pinene or verbenone.
3. High parental colonisation densities reduced emergence time of parents and offspring. Increasing colonisation density and emergence time had a negative influence on offspring vigour, resulting in a reduction of the distance tunnelled by the beetles.
4. Both alpha-pinene and verbenone were repellent. Surprisingly, verbenone was also toxic at high concentrations, with its effect being greater on beetles arising from high densities. The repellent effect of these compounds did not vary according to colonisation density, but for both compounds, at a 1.5 mg g^–1 concentration, it increased with emergence time.
5. The consequences of varying vigour and behaviour in relation to developmental conditions and emergence patterns on population dynamics of I. pini are discussed.     

Effects of early adult experience on host selection in insects: Some experimental and theoretical results

A field experiment with Drosophila melanogasterrevealed that when flies encounter a particular food type soon after emergence, the probability of their subsequently being attracted to such a resource is increased. In this experiment, the length of time flies experienced their postemergence environments was under the control of the flies themselves. The experiment thus realistically mimicked one form of experiential effect that may be important in nature. A theoretical model is developed which shows that enhanced adult preferences for the types of resources fed on as larvae can substantially increase the degree of host-based genetic subdivision within a polyphagous population.     

Predictable modification of body size and competitive ability following a host shift by a seed beetle

Interfertile populations of the seed beetle Callosobruchus maculatus differ genetically in several behavioral, morphological, and life-history traits, including traits that affect the intensity of larval competition within seeds. Previous studies have suggested that this variation depends on differences in host size. I performed a selection experiment in which replicate beetle lines were either maintained on a small, ancestral host (mung bean) or switched to a larger, novel host (cowpea). After 40 generations, I estimated survival, development time, and adult mass on each host, both in the presence and absence of larval competition. The shift to cowpea substantially reduced body size; irrespective of rearing host, adults from the cowpea lines were more than 10% lighter than those from the mung bean lines. Switching to cowpea also improved survival and reduced development time on this host, but without decreasing performance on the ancestral host. The most striking effect of the shift to a larger host was a reduction in larval competitiveness. When two even-aged larvae co-existed within a seed, the probability that both survived to adult emergence was > or = 65% if larvae were from the cowpea lines but < or = 12% if they were from the mung bean lines. The adverse effects of competition on development time and adult mass were also less severe in the cowpea lines than in the mung bean lines. By rapidly evolving smaller size and reduced competitiveness, the cowpea lines converged toward populations chronically associated with cowpea. These results suggest that evolutionary trajectories can be predictable, and that host-specific selection can play a major role in the diversification of insect life histories. Because host shifts by small, endophagous insects are comparable to the colonization of new habitats, adaptive responses may often include traits (such as larval competitiveness) that are not directly related to host use.     

Fitness consequences of potential assortative mating inside and outside a hybrid zone in Chorthippus parallelus (Orthoptera: Acrididae): implications for reinforcement and sexual selection theory

We have examined the fitness consequences of random and potentially non-random matings within two populations taken from inside, and two from outside a hybrid zone in Chorthippus parallelus. When given the opportunity to mate non-randomly, females from all populations laid egg pods more quickly than females obliged to mate at random. A range of fitness parameters measured on the offspring did not show increased fitness following potential non-random mating for any population. However, in non-hybrid populations, the sons of non-randomly mated females had about twice the mating success of the sons of those females forced to mate at random, suggesting the existence of heritable variation for male reproductive success. Hybrid dysfunction did not occur amongst the offspring of randomly mated hybrid females, demonstrating that the lack of dysfunction within these populations is not due to the evolution of assortative mating within them.     

Rapid adaptation to a novel host in a seed beetle (Callosobruchus maculatus): the role of sexual selection

Rapid diversification is common among herbivorous insects and is often the result of host shifts, leading to the exploitation of novel food sources. This, in turn, is associated with adaptive evolution of female oviposition behavior and larval feeding biology. Although natural selection is the typical driver of such adaptation, the role of sexual selection is less clear. In theory, sexual selection can either accelerate or impede adaptation. To assess the independent effects of natural and sexual selection on the rate of adaptation, we performed a laboratory natural selection experiment in a herbivorous bruchid beetle (Callosobruchus maculatus). We established replicated selection lines where we varied natural (food type) and sexual (mating system) selection in a 2 x 2 orthogonal design, and propagated our lines for 35 generations. In half of the lines, we induced a host shift whereas the other half was kept on the ancestral host. We experimentally enforced monogamy in half of the lines, whereas the other half remained polygamous. The beetles rapidly adapted to the novel host, which primarily involved increased host acceptance by females and an accelerated rate of larval development. We also found that our mating system treatment affected the rate of adaptation, but that this effect was contingent upon food type. As beetles adapted to the novel host, sexual selection reinforced natural selection whereas populations residing close to their adaptive peak (i.e., those using their ancestral host) exhibited higher fitness in the absence of sexual selection. We discuss our findings in light of current sexual selection theory and suggest that the net evolutionary effect of reproductive competition may critically depend on natural selection. Sexual selection may commonly accelerate adaptation under directional natural selection whereas sexual selection, and the associated load brought by sexual conflict, may tend to depress population fitness under stabilizing natural selection.     

Occurrence of different development time patterns induced by photoperiod in Anagrus atomus (Hymenoptera: Mymaridae), an egg parasitoid of Empoasca vitis (Homoptera: Cicadellidae)

The wasp Anagrus atomus L. parasitizes eggs of the cicadellid Empoasca vitis (Göthe) on grapevines during the vegetative season. In vineyards, abundant parasitoid emergence occurs in the autumn as a result of a developmental delay in part of the A. atomus population. In this context, physiological times required by the parasitoid to develop in E. vitis eggs are recorded during the summer. In early summer, all individuals develop fully, with physiological times in agreement with those reported in the literature. From early August onwards, an increasing proportion of parasitoids show a delay in development involving pre‐imaginal stages. Under transmission microscopy, larvae of A. atomus inside the eggs of E. vitis can be clearly differentiated as ‘light’ and ‘dark’ types, which are associated with normal and delayed development, respectively. The decreasing photoperiod as autumn approaches appears to promote the retardation of growth. This phenomenon can be considered a typical case of risk spreading because, when the availability of E. vitis eggs in vineyards is high (i.e. in early summer), A. atomus shows (i) only a normal developmental pattern that allows a faster population increase, whereas (ii) when E. vitis egg availability begins to be scarce (i.e. from mid summer), an increasing proportion of individuals having delayed development provides better fitness because it allows the parasitoid to synchronize its life history with leafhopper species that lay overwintering eggs on plants other than grapevines.