Modelling age‐ and density‐structured reproductive biology and seasonal survival of Apanteles galleriae Wilkinson (Hym., Braconidae)

Abstract: The biology of Apanteles galleriae Wilkinson, an important biological control agent of wax moths, is well described in the literature. We developed models simulating the functional response of fecundity and female progeny proportion of adult females as a function of age, host and/or parasitoid density to integrate current knowledge. Daily pattern of age‐related fecundity and female progeny proportion of different parasitoids was also examined. We investigated the effect of sex, mating status and seasonal time on adult longevity. We derived survival data of females in two different seasonal periods. We found that 50% of adult life span is important for an efficient fecundity. The patterns of host and/or parasitoid density‐dependent fecundity and sex ratio varied considerably. The highest fecundity and female progeny proportion occurred with one parasitoid and one female equivalent host. Longevity of adults was affected by sex and month periods, but mating status did not affect their longevity. Female survival was greater between December and May relative to June and November. Our results indicated that age, host and parasitoid density, and the timing of rearing influenced the life processes of parasitoids.     

Shifting body weight-fecundity relationship in a capital breeder: maternal effects on egg numbers of the autumnal moth under field conditions

In the literature, various environmental factors are described as being capable of influencing the reproductive output of insect females irrespective of their body size. Still, female body size or weight is widely used as a proxy for fecundity. In the present study, a seven-year data set on the autumnal moth, Epirrita autumnata (Borkhausen) (Lepidoptera: Geometridae), was used to analyze whether the body weight-fecundity relationship in this capital breeding, cyclic forest defoliating lepidopteran is constant across years. Ambient temperature conditions and density of conspecifics during larval development, the length of the pupal period, as well as moth densities in the parent generation were examined as factors capable of modifying the body weight-fecundity relationship. While the regression slope of potential fecundity (total egg numbers per female) on pupal mass was constant across years, the mean total egg number per given body weight (the regression intercept) was significantly different between years. This residual variance in egg numbers after controlling for the effect of pupal mass was best explained by the pooled geometrid density (autumnal and winter moths) in the parent generation. The total egg number per given body weight decreased with increasing density of geometrid moths in the parent generation. Thus, maternal density effects on offspring fecundity were found in this system. Their rather weak nature suggests, however, that this maternal effect alone does not have the potential of causing cyclic population dynamics in the autumnal moth.     

Inbreeding avoidance in spiders: evidence for rescue effect in fecundity of female spiders with outbreeding opportunity

Selection by inbreeding depression should favour mating biases that reduce the risk of fertilization by related mates. However, equivocal evidence for inbreeding avoidance questions the strength of inbreeding depression as a selective force in the evolution of mating biases. Lack of inbreeding avoidance can be because of low risk of inbreeding, variation in tolerance to inbreeding or high costs of outbreeding. We examined the relationship between inbreeding depression and inbreeding avoidance adaptations under two levels of inbreeding in the spider Oedothorax apicatus, asking whether preference for unrelated sperm via pre- and/or post-copulatory mechanisms could restore female fitness when inbreeding depression increases. Using inbred isofemale lines we provided female spiders with one or two male spiders of different relatedness in five combinations: one male sib; one male nonsib; two male sibs; two male nonsibs; one male sib and one male nonsib. We assessed the effect of mating treatment on fecundity and hatching success of eggs after one and three generations of inbreeding. Inbreeding depression in F1 was not sufficient to detect inbreeding avoidance. In F3, inbreeding depression caused a major decline in fecundity and hatching rates of eggs. This effect was mitigated by complete recovery in fecundity in the sib-nonsib treatment, whereas no rescue effect was detected in the hatching success of eggs. The rescue effect is best explained by post-mating discrimination against kin via differential allocation of resources. The natural history of O. apicatus suggests that the costs of outbreeding may be low which combined with high costs of inbreeding should select for avoidance mechanisms. Direct benefits of post-mating inbreeding avoidance and possibly low costs of female multiple mating can favour polyandry as an inbreeding avoidance mechanism.     

Consequences of mite feeding injury to beans on the fecundity and survivorship of the two-spotted spider mite (Acari: Tetranychidae)

Fecundity and survival of the two-spotted spider mite,Tetranychus urticae Koch, were examined on bean (Phaseolus vulgaris L.) plants that had been subjected to mite feeding injury in the laboratory. Different numbers ofT. urticae were restricted on the first two leaves of young bean plants, and spider-mite fecundity and survivorship was assayed on the third leaf. Each plant received four recently enclosed females, one female from each of four mite lineages. Using changes in the ratio of root mass to shoot mass of bean plants as a continuous measure of plant stress from spider-mite feeding, fecundity was positively related to stress for three out of four experiments. In two out of four experiments, survival of females was also positively related to stress, but reached an asymptote at slight or moderate stress levels. No evidence for induced resistance in beans was found. Mite lineage and the interaction between lineage and stress affected female survival but not fecundity. The implications of these results for understanding spider-mite outbreaks are discussed.     

Breeding performance of Tomicus destruens at different densities: the effect of intraspecific competition

In many bark beetles (Coleoptera: Curculionidae: Scolytinae), intraspecific competition is a limiting factor governing their population dynamics. The aim of this study was to evaluate the effect of increasing infestation densities on the breeding performance of Tomicus destruens (Wollaston), one of the main pests of Mediterranean pine forests. Six densities (ranging from 0.25 to 1.50 females   dm⁻²) were tested under laboratory conditions on stone pine logs, assessing the mortality of eggs, larvae, and pupae, and recording the number of emerging adults. Density significantly affected the fertile tract of the maternal galleries, i.e., those containing eggs, which was shorter at densities higher than 0.75. Numbers of eggs, larvae, pupae, and adults per female decreased markedly when density increased from 0.25 to 1.5, although differences were statistically significant only at a density of 0.75 and higher. The lowest mortality was found in larvae reared at a density of 0.25 (17.7%) and the highest in pupae at 1.5 (84.8%). Increasing density affected the pre-emergence stages in different ways, with pupae and larvae suffering the highest increases in mortality (39.5 and 163.6%, respectively). The breeding performance of T. destruens (number of eggs laid reaching adulthood) progressively decreased from 17 to 4% with increasing densities, although at low infestation, the mean number of emerging adults per dm² of bark increased with density, showing a positive trade-off between female fecundity and female density. In conclusion, breeding densities higher than 0.75 female dm⁻² decreased female fecundity and elicited high mortality, mainly in larvae.     

The effect of mating frequency and mating pattern on female reproductive fitness in cabbage beetle,Colaphellus bowringi

It is generally thought that females can receive more of the material benefits from males by increasing mating frequency and polyandry can lead to greater reproductive success. The cabbage beetle, Colaphellus bowringi Baly (Coleoptera: Chrysomelidae), is a highly promiscuous species, in which females or males can readily mate repeatedly with a given partner or multiple partners at a very high frequency. In the present study, the effect of mating frequency (number of matings) and mating pattern (polyandry vs. monogamy) on female reproductive fitness was investigated by measuring fecundity, fertility, and female longevity. The results indicated that increased female mating frequency with the same male did not result in variation in lifetime fecundity, but significantly increased fertility and decreased female longevity. Moreover, five copulations were sufficient to acquire maximal reproductive potential. Female lifetime fecundity also did not differ between polyandrous and monogamous treatments. However, monogamous females exhibited a significant increase in fertility and significant prolongation of longevity compared with polyandrous females, further demonstrating that monogamy is superior to polyandry in this beetle.     

SEX RATIO VARIATION IN A PARASITIC WASP I. REACTION NORMS

To understand genetic and phenotypic constraints on the sex ratio in a parasitic wasp that attacks fly pupae, I carried out a laboratory study of sex ratio variability in five strains of Muscidifurax raptor (Hymenoptera: Pteromalidae). I manipulated the environment through combinations of temperature and day length, and the numbers of females that attack a group of hosts. The change of phenotype in each strain over the range of environmental conditions describes the norm of each reaction for that strain, and measures how a strain responds to environmental variation to create phenotypic variability. Sex ratio in parasitic wasps is a complex trait that has several components—the numbers of eggs laid by an ovipositing wasp and the fraction of eggs that are fertilized (female). Further, sex ratio may be influenced by a female's reaction to other females exploiting the same hosts (superparasitism). I found no strain-environment interactions in either sex ratio or fecundity when I varied environmental conditions. Although strains differed in sex ratio and fecundity, all strains produced a more female-biased sex ratio and had higher fecundity when temperature and day length increased. Sex ratio and fecundity were phenotypically correlated, and strains with greater fecundity also produced a more female-biased sex ratio. All strains facultatively shifted sex ratio toward a higher fraction of males with increasing female density, despite apparent differences in superparasitism among strains. Males and females survived equally during development, so that mortality differences among strains and across environments could not account for sex ratio variability. This study indicates that sex ratio variability among strains is constrained by the correlation between sex ratio and fecundity, and that strains display similar facultative shifts in sex ratio as female density increases because sex ratio shifts are insensitive to differing levels of superparasitism.     

Effects of size at metamorphosis on stonefly fecundity, longevity, and reproductive success

Many organisms with complex life cycles show considerable variation in size and timing at metamorphosis. Adult males of Megarcyssignata (Plecoptera: Perlodidae) are significantly smaller than females and emerge before females (protandry) from two western Colorado streams. During summer 1992 stoneflies from a trout stream emerged earlier in the season and at larger sizes than those from a colder fishless stream, and size at metamorphosis did not change over the emergence period in either stream. We performed two experiments to determine whether variation in size at metamorphosis affected the fecundity, reproductive success and longevity of individuals of this stonefly species and if total lifetime fecundity was affected by the number of matings. In the first experiment, total lifetime fecundity (eggs oviposited) was determined for adult females held in small plastic cages in the field. Males were removed after one copulation, or pairs were left together for life and allowed to multiply mate. Most copulations occurred in the first few days of the experiment. Females in treatments allowing multiple matings had significantly lower total lifetime fecundity and shorter adult longevity than females that only mated once. Multiple matings also reduced longevity of males. Fecundity increased significantly with female body mass at emergence, but only for females that mated once. While multiple matings eliminated the fecundity advantage of large female body size, number of matings did not affect the significant positive relationship between body mass at metamorphosis and longevity of males or females. In a second experiment designed to determine if body mass at emergence affected male mating success, we placed one large and one small male Megarcys in an observation arena containing one female and recorded which male obtained the first mating. The large and the small male had equal probabilities of copulating with the female. Copulations usually lasted all night, and the unmated male made frequent, but unsuccessful attempts to take over the copulating female. Our data suggest that selection pressures determining body size at metamorphosis may operate independently on males and females, resulting in evolution of sexual size dimorphism, protandry, and mating early in the adult stage. We emphasize the importance of interpreting the fitness consequences of larval growth and development on the timing of and size at metamorphosis in the context of the complete life cycle. Received: 1 July 1997 / Accepted: 12 November 1997     

Effect of food concentration on the production and viability of resting eggs of the rotifer Brachionus: implications for the timing of sexual reproduction

1. Life‐table experiments with Brachionus calyciflorus test several hypotheses related to the idea that sexual reproduction in monogonont rotifers should occur when food resources are favourable. 2. The food concentration necessary for a fertilised mictic female to produce one phenotypically normal resting egg was higher than that for an amictic female to produce one daughter. At the lowest concentration of Cryptomonas erosa (1.25 × 10³ cells mL^?1), the lifetime fecundity of these two types of females was 0.9 and 1.4, respectively. 3. The lifetime fecundity of both fertilised mictic females and amictic females increased with food concentration to 3.4 resting eggs and 15.2 daughters female^?1, respectively. The approach to maximal fecundity with increasing food concentration was more rapid for fertilised mictic females, such that their lifetime fecundity relative to that of amictic females gradually decreased from 0.64 (at 1.25 × 10³ C. erosa mL^?1) to 0.22 (at 2.5 × 10⁴ C. erosa mL^?1). 4. The probability of a fertilised mictic female producing one or more abnormal resting eggs during her lifetime was high (approximately 75%). The mean proportion of abnormal eggs produced per female varied among the different food‐concentration treatments (26–38%) but was not higher at the low food concentrations. 5. The proportion of normal resting eggs that hatched was high (51–71%); those produced at low food concentrations were no less likely to hatch than those produced at high food concentrations. No abnormal resting eggs hatched. 6. The probability of a fertilised mictic female producing an abnormal resting egg increased rapidly with her age at all food concentrations. The probability of a normal resting egg hatching declined with maternal age at the low food concentration in one of two experiments. 7. The results support the idea that induction of mictic females should occur when food resources are good. Coincidence of sexual reproduction with low food availability risks low production of resting eggs for several reasons. Population size may be small, with a low probability of encounters between young mictic females and males, and fertilised mictic females may be unable to mature and produce resting eggs.     

Multiple generation effects of high temperature on the development and fecundity of Bemisia tabaci (Gennadius) (Hemiptera: Aleyrodidae) biotype B

Insects are ectotherms and their ability to resist temperature stress is limited. The immediate effects of sub‐lethal heat stress on insects are well documented, but longer‐term effects of such stresses are rarely reported. In this study, survival, development and reproduction of the whitefly, Bemisia tabaci (Gennadius) (Hemiptera: Aleyrodidae) biotype B, were compared over five consecutive generations at 27, 31 and 35 °C and for one generation at 37 °C. Both temperature and generation significantly affected the fitness of the whitefly. These impacts were more dramatic with increasing generations and temperatures. Among the experimental temperatures, the most favorable for development and reproduction were 27 °C and 31 °C. At 27 °C, survival, development and fecundity were all stable over these five generations. At 31 °C, immature survival rate was the highest in the fifth generation, but female fecundities decreased in the fourth and fifth generations. At 35 °C, egg hatching rate, immature survival rate and female fecundity decreased significantly in the fourth and fifth generations. At 37 °C, survival of B. tabaci was not adversely affected, but female fecundity at 37 °C was less than 10% of that at 27 °C or 31 °C. These results demonstrate that the lethal high temperature for B. tabaci is over 37 °C, and the whitefly population continued expanding in the five generations at 35 °C. The ability of B. tabaci biotype B to survive high temperature stress will play an important role in its population extension under global warming.     

What maintains noncytoplasmic incompatibility inducing Wolbachia in their hosts: a case study from a natural Drosophila yakuba population

Cytoplasmic incompatibility (CI) allows Wolbachia to invade hosts populations by specifically inducing sterility in crosses between infected males and uninfected females. In some species, non-CI inducing Wolbachia, that are thought to derive from CI-inducing ancestors, are common. In theory, the maintenance of such infections is not possible unless the bacterium is perfectly transmitted to offspring--and/or provides a fitness benefit to infected females. The present study aims to test this view by investigating a population of Drosophila yakuba from Gabon, West Africa. We did not find any evidence for CI using wild caught females. Infected females from the field transmitted the infection to 100% of their offspring. A positive effect on female fecundity was observed one generation after collecting, but this was not retrieved five generations later, using additional lines. Similarly, the presence of Wolbachia was found to affect mating behaviour, but the results of two experiments realized five generations apart were not consistent. Finally, Wolbachia was not found to affect sex ratio. Overall, our results would suggest that Wolbachia behaves like a neutral or nearly neutral trait in this species, and is maintained in the host by perfect maternal transmission.     

Intraspecific and Interspecific Competition Between Cotesia flavipes and Cotesia sesamiae (Hymenoptera: Braconidae), Gregarious Larval Endoparasitoids of Lepidopteran Stemborers

Super and multiple parasitism of Chilo partellus (Lepidoptera: Pyralidae) and Sesamia calamistis (Lepidoptera: Noctuidae) by Cotesia flavipes Cameron and Cotesia sesamiae (Cameron) (Hymenoptera: Braconidae) were investigated in the laboratory. Progeny production of Co. flavipes increased as a result of increasing the number of ovipositions, from one to three per one Ch. partellus host larva, then decreased as a result of four and five ovipositions per larva. Cocoon weight, sex ratio and emergence of the parasitoid progeny were not affected by superparasitism. Low progeny production of Co. sesamiae and poor survival of Ch. partellus host larvae were found as a result of superparasitism. When S. calamistis was the host, the duration of immature stages of Co. flavipes , parasitoid emergence, progeny production and sex ratio were not affected by superparasitism, but cocoon weight, adult longevity and the potential fecundity of adult females decreased. Superparasitism of S. calamistis by Co. sesamiae did not affect emergence, longevity or sex ratio of adult progeny of the parasitoid, but prolonged immature development, lowered cocoon weight and decreased potential fecundity of adult female progeny. Co. flavipes out-competed Co. sesamiae when Ch. partellus was parasitized by both species. The potential for local displacement of Co. sesamiae by Co. flavipes in areas dominated by Ch. partellus in East Africa is discussed.     

Genetics of Life History in DROSOPHILA MELANOGASTER. II. Exploratory Selection Experiments

Two types of small-scale selection experiments were performed. (1) Artificial selection experiments were performed on age-specific female fecundity. Selection for early fecundity over three generations produced a statistically detectable direct response. There was no detectable indirect response in other life-history characters. Selection for late fecundity over three generations did not produce any detectable direct response. Indirect responses were detected: early egg-laying decreased and longevity increased. (2) Natural selection for late-age fitness components increased late fecundity, female longevity, and the duration of female reproduction, while early fecundity and mean egg-laying rate decreased.     

同时而非先后一雌多雄交配增加不同温度下食蚜瓢虫的适合度（英文）

【目的】尽管一雌多雄在瓢虫科中常见,但各研究中获得的数据不足以解释雌虫多次交配和一雌多雄的一般适应性意义或适合度后果。本研究以温度为胁迫因子,旨在评价一雌多雄的某些益处（如增加的适合度）是否可传递给后代。【方法】本研究检测了黄斑盘瓢虫Coelophora saucia (Mulsant) 3种交配处理中的适合度：一雌一雄（与同一雄虫交配5次,1次/d）,先后一雌多雄（与5头不同的雄虫依次交配5次,即每天与新的雄虫交配1次）,以及同时一雌多雄（放进5头雄虫,任由雌虫选择雄虫,交配5次,1次/d）。观察了各交配处理不同温度下（25, 27和 30℃）繁殖力、卵的育性、后代发育和存活。【结果】结果表明,经历一雌多雄然后进行交配选择或竞争的雌性的繁殖能力最强,后代能在更广温度范围内最好地适应发育和存活。但先后一雌多雄交配的雌性与一雌一雄交配的雌性的繁殖能力相似。【结论】结果说明,在无交配选择或雄性竞争的条件下,一雌多雄的益处不明显。这可能是由于在依次射精的雄性间存在精子竞争,或由于雌性的隐性选择。据我们所知,本研究中观察发现的无交配选择时不表现一雌多雄的益处的现象,之前在昆虫中未观察到过。     

Consequences of acclimation on survival and reproductive capacities of cold-stored mummies of Aphidius rhopalosiphi (Hymenoptera: Aphidiinae)

The cold storage of parasitoid mummies is a crucial point during mass production of parasitoids for aphid control in wheat, Triticum spp. In this study, the effect of acclimation to cold before storage of mummies containing Aphidius rhopalosiphi DeStefani-Peres (Hymenoptera: Braconidae: Aphidiinae) was evaluated on survival, sterility, and fecundity of parasitoids. Groups of 1-d-old and 3-d-old mummies were stored at -5 degrees C for 10 d without acclimation or after one of five different acclimation treatments. One-day-old and 3-d-old mummies contain prepupa and postmetamorphosis but not yet sclerotinized adult parasitoids, respectively. The offspring and sex ratio of stored parasitoids were compared with a control that was left at rearing temperatures (20 degrees C). Without acclimation, two-thirds of the parasitoids died during storage, for both ages of mummies tested. For stored 1-d-old mummies, subsequent survival increased under progressive exposure to low temperatures and reached 67.3% after 480 min of acclimation. In the same way, percentage of male sterility decreased with acclimation duration, whereas female fecundity increased. For stored 3-d-old mummies, subsequent survival improved with the duration of the acclimation treatment, but not male sterility and female fecundity.     

The effect of crowding on adult populations ofDelia (=Hylemya) antiqua (Meigen)

Using 5 density levels, the effect of increasing density was studied on several population statistics of adult Delia (=Hylemya) antiqua. Amongst the statistics studied were adult lifespan; female mating frequency, fecundity and egg hatchability. It was shown that females lived significantly longer than males and that increasing density significantly reduced lifespan. Density had no effect on mating frequency. Total fecundity/female was significantly reduced with increasing density but oviposition rate was density independent. Using these statistics, mean generation time T, net reproduction rate R₀ and capacity for increase r_c were calculated at the different density levels. Values of R₀ showed a one-tailed response but there was no clear effect of density on r_c. Multiple comparisons between variables revealed several important relationships.

1. Adult lifespan was the most important factor affecting egg production.
2. Number of mated females was more important in affecting total fecundity/cage than adult density.

It could also be calculated that to produce the effect of one single mated female on total fecundity/cage the overall density would have to be reduced by 0.0023 individuals/cm³. These results are discussed in the context of efficient laboratory rearing of the onion fly.     

Male-only care and classical polyandry in birds: phylogeny, ecology and sex differences in remating opportunities

It has been argued recently that the combination of male-only parental care and classical polyandry in birds is the most interesting and yet the least understood of all avian breeding systems. Despite a huge number of hypotheses, careful comparative analyses have repeatedly failed to identify consistent ecological differences between species showing male-only care and closely related species showing other patterns of care. This has led to the suggestion that such analyses fail because the crucial differences are between ancient lineages rather than between closely related species. Here, therefore, I use comparisons between families to test three well-known hypotheses: that male-only care is associated with: (i) a low rate of fecundity; (ii) large egg size relative to female size; or (iii) female-biased opportunities for remating. Families showing male-only care do not differ from families showing female-only care with respect to rate of fecundity or relative egg size. There is, however, a significant difference between these two groups of families with respect to an index of remating opportunities, nesting density. Families showing female-only care nest at high density, while those showing male-only care nest at very low density. This is one of the first times a consistent ecological correlate has been identified for male-only care in birds. It suggests that female-only care arises (or persists) in families where remating opportunities are abundant for both sexes, whereas male-only care arises (or persists) in families where remating opportunities are rare for both sexes and particularly scarce for males. This in turn suggests that sex differences in remating opportunities are the key ecological factor in determining male-only care and classical polyandry in birds.     

Kin-biased dispersal behaviour in the mango shield scale, Milviscutulus mangiferae

When fitness decreases with increasing density in a habitat, dispersal behaviour is expected to evolve. To avoid competition between kin, dispersal behaviour based on kin recognition should be more likely to occur when the individuals in a habitat are closely related. I tested this prediction with first-instar larvae (crawlers) of the mango shield scale, Milviscutulus mangiferae. The body size of adult females, a measure of fecundity, was larger when only one female was present on a leaf than when two were present. When I placed two crawlers on a leaf, they emigrated more frequently when they were siblings than when they were not related. I discuss the implication of the results for kin recognition in thelytokous parthenogenetic animals. Copyright 2000 The Association for the Study of Animal Behaviour.     

Access to multiple mates increases fecundity but does not affect per‐offspring maternal investment in a marine gastropod

Females of many organisms mate more than once and with more than one male, suggesting that polyandry confers some advantage to the female or her offspring. However, variation in maternal investment in response to mate choice and mate number can confound efforts to determine if there are benefits of polyandry. Access to multiple mates could increase maternal investment in offspring via a number of different mechanisms. Few studies have determined if investment is influenced by mate choice and number, and data are particularly lacking for marine invertebrates. This study was designed to determine if maternal investment and offspring size increase with access to increasing numbers of mates in the protandrous intertidal slipper snail Crepidula cf. marginalis. Virgin female slipper limpets were exposed to one, three, or five potential mates and their fecundity, egg size, and hatchling size were measured for multiple clutches. Treatment had a significant effect on fecundity, with fecundity increasing with the number of potential mates. Treatment did not have an effect on the size of eggs or hatchlings, on the variation in egg size or hatchling size within broods, or on the frequency of oviposition. Treatment did alter the variation in average offspring size among females, but not in the way predicted by theory. The main result, that access to multiple mates does not have an effect on per offspring maternal investment, makes C. cf. marginalis an ideal candidate to study the effects of polyandry on offspring fitness without having to take into account confounding effects of variation in maternal investment.     

Density-dependent effects on the reproductive fitness of the New Zealand beech scale insect (Ultracoelostoma assimile) across multiple spatial scales

Abstract.  1. Intraspecific competition can be an important factor influencing the individual fitness of organisms. This study reports density-dependent effects on the fecundity of the beech scale insect ( Ultracoelostoma assimile , Hemiptera: Margarodidae) on naturally occurring, canopy-dominant red beech ( Nothofagus fusca , Fagaceae) trees in New Zealand. For the first time an increasing intensity of intraspecific competition at increasing spatial scales within individual host trees is demonstrated.
2. Beech scale insect 'tests' containing adult female scale insects and eggs were collected from the trunks of 10 red beech trees [17.7–48.5 cm in diameter at breast height (1.4 m), d.b.h.] with varying densities of scale insect infestation. The relationship between individual female fecundity and scale insect density at three spatial scales: local (within 5 cm), lower trunk (below 2 m above ground level), and whole tree, was tested.
3. Beech scale insect fecundity was density dependent, with total female egg load decreasing with increasing scale insect density. The strength of the density-dependent effect increased with increasing spatial scale, suggesting that scale insects are creating a tree-wide drain on the quality of phloem sap, rather than depleting nutrients from localised, high-density areas within trees. These results indicate that at high densities, the New Zealand beech scale insect can have a negative effect on the nutritional quality of the phloem of red beech, thus negatively affecting conspecifics elsewhere on individual host trees. The increasing intensity of the effect with increasing spatial scale within individual trees emphasises the importance of measuring density-dependent effects at the appropriate spatial scale.