Differences in the nestling begging calls of hosts and host-races of the common cuckoo, Cuculus canorus

We compared nestling begging calls of four hosts (reed warbler, Acrocephalus scirpaceus; great reed warbler, A. arundinaceus; dunnock, Prunella modularis; and meadow pipit, Anthus pratensis) and the respective host-races of the common cuckoo. Note structure varied between host species, but not between cuckoo host-races, so cuckoos did not vary their call note structure to match that of their hosts' chicks. Call rate increased with age, but there were marked differences between both host species and cuckoo host-races. Dunnock-cuckoos called more rapidly than reed warbler-cuckoos despite growing at the same rate. We suggest this difference reflects how cuckoos tune into the way these host species respond to begging signals from their own young, because dunnock chicks called much more rapidly than reed warbler chicks. Great reed warbler-cuckoos called at a lower rate than reed warbler-cuckoos when young, but at a greater rate when older than 8 days. This could also result from the cuckoo chicks tuning into differences in the way these hosts respond to begging signals. However, great reed warbler-cuckoos grew at a faster rate than the other cuckoo host-races, so they may also call faster to demand higher provisioning rates from this larger host. To test these hypotheses critically, data are needed on how the different host species integrate visual and vocal begging signals from their own broods. We discuss how differences in cuckoo begging might develop, given that cuckoo host-races are restricted to female cuckoo lineages. Copyright 2003 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.      

Cleptoparasites, social parasites and a common host: Chemical insignificance for visiting host nests, chemical mimicry for living in

Social insect colonies contain attractive resources for many organisms. Cleptoparasites sneak into their nests and steal food resources. Social parasites sneak into their social organisations and exploit them for reproduction. Both cleptoparasites and social parasites overcome the ability of social insects to detect intruders, which is mainly based on chemoreception. Here we compared the chemical strategies of social parasites and cleptoparasites that target the same host and analyse the implication of the results for the understanding of nestmate recognition mechanisms. The social parasitic wasp Polistes atrimandibularis (Hymenoptera: Vespidae), and the cleptoparasitic velvet ant Mutilla europaea (Hymenoptera: Mutillidae), both target the colonies of the paper wasp Polistes biglumis (Hymenoptera: Vespidae). There is no chemical mimicry with hosts in the cuticular chemical profiles of velvet ants and pre-invasion social parasites, but both have lower concentrations of recognition cues (chemical insignificance) and lower proportions of branched alkanes than their hosts. Additionally, they both have larger proportions of alkenes than their hosts. In contrast, post-invasion obligate social parasites have proportions of branched hydrocarbons as large as those of their hosts and their overall cuticular profiles resemble those of their hosts. These results suggest that the chemical strategies for evading host detection vary according to the lifestyles of the parasites. Cleptoparasites and pre-invasion social parasites that sneak into host colonies limit host overaggression by having few recognition cues, whereas post-invasion social parasites that sneak into their host social structure facilitate social integration by chemical mimicry with colony members.     

Host nest defense against a color-dimorphic brood parasite: great reed warblers (Acrocephalus arundinaceus) versus common cuckoos (Cuculus canorus)

We tested great reed warbler (Acrocephalus arundinaceus) discrimination against two common cuckoo (Cuculus canorus) female color morphs (gray and rufous) in two areas with different parasitism rates and proportions of the two morphs. Hosts recognized the two cuckoo morphs from a control, the feral pigeon (Columba livia), at Apaj, Hungary (where brood parasitism was heavy), whereas no significant differences among the models were recorded at Lužice, Czech Republic (where the parasitism rate was moderate). At Apaj, the hosts discriminated the rufous morph (which is slightly predominant there) better than the gray morph from the control. Between-site comparison (after controlling for background aggression) revealed that great reed warblers were more aggressive towards the rufous morph at Apaj than at Lužice, whereas their responses to the gray morph did not differ, corresponding with much higher between-site difference in the relative abundance of the rufous morph. Our results suggest that both local parasitism pressure and relative abundance of two female color morphs of a brood parasite may significantly influence host nest defenses.     

Male-biased sex ratio increases female egg laying and fitness in the housefly, Musca domestica

A biased operational sex ratio (OSR) can have multiple, confounding effects on reproductive fitness. A biased OSR can increase harassment and mating activity directed towards potential mates but may also increase the ability of potential mates to choose a good partner if lower quality mates are screened out through competitive interactions. Additionally, a biased OSR may affect reproductive fitness through changes in male ejaculate content or in female reproductive response. We quantified how a male-biased OSR (1:1, 2:1, or 5:1 male to female) affected the size of a female??s first egg clutch and her offspring??s survivorship in the housefly, Musca domestica. A male-biased OSR increased female fitness: females laid more eggs in their first clutch, had increased offspring survivorship at a 2:1 versus 1:1 OSR, and had equivalent fitness with a 5:1 male to female OSR. Courtship activity increased when the OSR was male-biased but was not a significant predictor of female fitness. Trials where females chose their mates versus trials where a random male was chosen for them had equivalent first clutch sizes and offspring survivorship. These results suggest that there are cryptic effects from a male-biased OSR on female fitness that are most likely driven by pre-copulatory social environment.     

Metabolic thermogenesis and evaporative water loss in the Hwamei Garrulax canorus

Basal metabolic rate (BMR) is thought to be a major hub in the network of physiological mechanisms connecting life history traits. Evaporative water loss (EWL) is a physiological indicator that is widely used to measure water relations in inter- or intraspecific studies of birds in different environments. In this study, we examined the physiological responses of summer-acclimatized Hwamei Garrulax canorus to temperature by measuring their body temperature (T_b), metabolic rate (MR) and EWL at ambient temperatures (T_a) between 5 and 40 °C. Overall, we found that mean body temperature was 42.4 °C and average minimum thermal conductance (C) was 0.15 ml O₂ g⁻¹ h⁻¹ °C⁻¹ measured between 5 and 20 °C. The thermal neutral zone (TNZ) was 31.8–35.3 °C and BMR was 181.83 ml O₂ h⁻¹. Below the lower critical temperature, MR increased linearly with decreasing T_a according to the relationship: MR (ml O₂ h⁻¹)=266.59–2.66 T_a. At T_as above the upper critical temperature, MR increased with T_a according to the relationship: MR (ml O₂ h⁻¹)=−271.26+12.85 T_a. EWL increased with T_a according to the relationship: EWL (mg H₂O h⁻¹)=−19.16+12.64 T_a and exceeded metabolic water production at T_a>14.0 °C. The high T_b and thermal conductance, low BMR, narrow TNZ, and high evaporative water production/metabolic water production (EWP/MWP) ratio in the Hwamei are consistent with the idea that this species is adapted to warm, mesic climates, where metabolic thermogenesis and water conservation are not strong selective pressures.     

Maternal investment during egg laying and offspring sex: an experimental study of zebra finches

To study whether offspring sex is related to the amount of resources invested in eggs we performed an experiment on zebra finches, Taeniopygia guttata. By manipulating their food supply, we forced two groups of females either to increase or to decrease investment in subsequent eggs. Since zebra finches are sexually dimorphic and the reproductive value of the sexes may vary with maternal nutritional status, we predicted that females would adjust the sex of their offspring to egg quality. Females that received poor-quality food for 7 weeks before egg laying, then food supplemented with proteins after they laid the first egg, significantly increased the mass of subsequent eggs. An increase of egg mass with laying sequence was less pronounced in females that received high-quality food before laying and experienced food deterioration after starting to lay. The proportion of sons in subsequent eggs tended to increase in the latter group (although this was marginally significant) but was not related to laying sequence in the other group: these patterns differed significantly between the groups. Offspring sex was not related to egg mass, but newly hatched male chicks were heavier than female chicks. Furthermore, the hatching success of male eggs was lower than that of female eggs. We suggest that differential hatching success of the sexes and sex differences in mass at hatching may constitute important factors shaping brood sex ratios. Copyright 2002 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.     

An example of character release in host selection and egg colour of cuckoos Cuculus spp. in Japan

We have studied in northern Japan the hitherto unidentified eggs of a species of cuckoo in the nests of the Bush Warbler Cettia diphone . The cuckoo in question appeared to be the Himalayan Cuckoo Cuculus saturatus which parasitizes mainly the Willow Warbler Phylloscopus occipitalis in southern Japan. The egg colour in this northern Cuckoo was chocolate-brown or orange-brown, similar to that of the Bush Warbler but unlike that of the southern Himalayan Cuckoo. Egg size was significantly larger than that of the southern Himalayan Cuckoo and instead similar to that of the Little Cuckoo C. poliocephalus which uses the same host species in southern Japan, to which the Little Cuckoo is confined. The shift in host species and egg colour in the northern Himalayan Cuckoo seems to be a case of character release in the absence of the Little Cuckoo.     

Dark nests and egg colour in birds: a possible functional role of ultraviolet reflectance in egg detectability

Owing to the conspicuousness of ultraviolet (UV) colour in dark environments, natural selection might have selected UV egg coloration because it would enhance egg detectability by parents in murky nests. Here, we tested this hypothesis by using comparative and experimental approaches. First, we studied variation in egg coloration of 98 species of European passerines measured using UV-visible reflectance spectrometry (300-700nm) in relation to nesting habits. Analyses based on raw data and controlling for phylogenetic distances both at the species and the family levels revealed that hole-nester species produced eggs with higher UV reflectance than those nesting in open habitats. The experimental approach consisted of the manipulation of UV reflectance of the experimental eggs introduced outside the nest-cup of the hole-nester spotless starling Sturnus unicolor and the study of the retrieval of these eggs. Ultraviolet-reflecting eggs (controls) were more frequently retrieved to the nest-cup than non-reflecting (-UV) eggs. These results were not due to '-UV' eggs being recognized by starlings as parasitic because when a parasitic egg is detected, starlings removed it from the nest-box. Therefore, these results are consistent with the hypothesis that UV egg colours are designed to provide highly detectable targets for parent birds in dark nest environments.     

Effects of Enterococcus faecalis on egg production,egg quality and caecal microbiota of hens during the late laying period

This study was conducted to determine the effects of diet supplementation of laying hens with Enterococcus faecalis (EF) on egg production, egg quality and caecal microbiota. A total of 360 Hy-Line Brown laying hens (72 weeks old) were divided into three groups with four replicates of 30 birds each. The laying hens were fed with the basal diet (Control), the basal diet + 3.75 · 10⁸ cfu EF/kg (Group I) or the basal diet + 7.5 · 10⁸ cfu EF/kg (Group II). The experiment lasted for 45 d. Eggs and caecal samples were collected at the end of the experiment. Results showed that dietary supplementation with EF did not affect the average daily egg weight, cracked egg rate, mortality and egg quality. However, EF supplementation caused a significantly increased laying rate and decreased feed/egg ratio (p < 0.05). The differences in caecal microbiota between Group II and the Control were significant. The relative abundance of Verrucomicrobia and Cyanobacteria at the phylum level, Rikenellaceae, Christensenellaceae and Veillonellaceae at the family level, and the Faecalibacterium, Christensenellaceae R-7 group and Eubacterium coprostanoligenes group at the genus level changed significantly in Group II compared with the Control (p < 0.05). In conclusion, the tested dietary supplementations with EF improved product performance and affected the caecal microbial community structure of laying hens during the late laying period.     

Omnivory in the calanoid copepod Temora longicornis: feeding, egg production and egg hatching rates

We measured in laboratory experiments the ingestion, egg production and egg hatching rates of female Temora longicornis as a function of diet. The diets consisted of a diatom (Thalassiosira weissflogii), an autotrophic dinoflagellate (Heterocapsa triquetra), and a bacterivorous ciliate (Uronema sp.) given as sole foods, or combinations of these single-food items: diatom+dinoflagellate, diatom+ciliate, dinoflagellate+ciliate, and diatom+ciliate+dinoflagellate. For the three single-item diets, the functional response was similar; i.e., ingestion rate increased linearly with food concentration (food range: ∼25 to ∼600 μg C l⁻¹). When all diets were considered, maximum daily carbon ration (∼70% of body weight) was independent of food type. However, the maximum daily egg production rate (12% of body carbon) was obtained with the diatom diet. For all diets, both ingestion and egg production rates increased with food concentration. Ingestion and egg production rates were affected differently by the interaction of food concentration and food type: at low food concentrations, ingestion rates were highest on diets containing the diatom. At high food concentrations, egg production rates were highest on the two phytoplankter diets and their combination. The presence of the ciliate in the diet did not enhance ingestion rate or egg production. Mixed-food diets did not enhance egg production relative to single-food diets. Hence, dietary diversity did not appear to be particularly advantageous for reproduction. Carbon-specific egg production efficiency (EPE; egg production/ingestion) was independent of food concentration and type, and equaled 9%. Egg hatching success was low (mean<30%) and independent of food concentration and type, and egg production rates. Our results are consistent with previous observations that egg production in T. longicornis is enhanced during diatom blooms. However, the relatively low EPE and egg hatching success suggest that reproduction and recruitment in this study were severely constrained by the biochemical composition of the diet, or the physiological condition of the females towards the end of their season of growth in Long Island Sound.     

Development of an artificial host egg for in vitro egg laying of Trichogramma galloi and T. pretiosum using plastic membranes

An artificial host egg was developed as a first attempt to improve the techniques for in vitro rearing of Trichogramma galloi Zucchi and T. pretiosum Riley (Hymenoptera, Trichogrammatidae). Plastic membranes (polyethylene, polypropylene and polystyrene) of different thickness were tested for their usefulness in manufacturing artificial eggs, using a thermal system to produce semi-spherical cupules on the plastic surface. These artificial eggs were filled with larval hemolymph of Helicoverpa zea (Boddie) (Lepidoptera, Noctuidae), and the optimal size of the artificial egg and density of females/artificial egg for the highest host acceptance (percentage of artificial eggs with at least one parasitoid egg) as well as the ideal level of parasitism (eggs/artificial egg) were determined. High-density polyethylene 7–8 and 9–10 m thick resulted in the best acceptance and level of parasitization for both parasitoid species. Six females per artificial egg resulted in the best rate of parasitization for these parasitoids, although these data were not different when using 4 or 5 females of T. galloi. The size, shape and surface texture of the artificial eggs were found to provide enough cues to elicit parasitization behavior in Trichogramma females, since no chemical stimulus was provided.     

Changes in the muscle condition of female Zebra Finches Poephila guttata during egg laying and the role of protein storage in bird skeletal muscle

Samples of pectoral muscle tissue from Zebra Finches Poephila guttata were taken from birds at various stages of egg laying. As egg laying progressed, there was a significant loss of both water-soluble and alkali-soluble proteins. Gel filtration of the water-soluble proteins showed three peaks, only one of which changed significantly during the laving cycle. This was a high-molecular-weight protein that declined in concentration during the period of egg laying. This protein may function as a reserve, enabling female birds to provide amino acids for egg protein production from their muscle tissue without impairing normal muscle function.     

Individual changes underlie age-specific pattern of laying date and egg-size in female common terns Sterna hirundo

It has been suggested that breeding performance differs between young and old birds due to the appearance and disappearance of phenotypes through differential survival (selection hypothesis) or differential recruitment (delayed breeding hypothesis) of high-quality individuals, but each bird may show constant breeding performance over its life. We tested constant egg-volume and laying date by modelling their variability on the basis of the 109 known-age females of common tern Sterna hirundo with data available from 1 to 9 years. Longitudinal analyses showed a significant advancement of laying date, as well as a steady increase in egg-volume, in young age classes from 2 to 5–7 years old, indicating individual intrinsic changes in performance with age. In our model, female effect accounted for 74% and 8% of variance in egg-volume and laying date, respectively, suggesting that if correlation between breeding performance and survival or recruitment exists, population patterns of age-specific performance may emerge. However, we found no evidence that birds that did not return to breed during young age classes laid later or smaller eggs than returned breeders. Likewise, we found no evidence that recruiting birds laid earlier or larger eggs than same aged birds recruited in preceding years. Thus, this study shows that age-specific patterns in timing of breeding and egg-size in common terns result from individuals intrinsic changes, and we reject the selection and the delayed breeding hypotheses as a major factor shaping age-specific patterns at population level.Communicated by F. Bairlein     

Extra-ovariolar egg resorption in a dung beetle, Euoniticellus intermedius

Most oösorption in the dung beetle Euoniticellus intermedius takes place in the haemocoele, oöcytes being extruded from the ovariole before the deposition of the chorion. Oösorption can be induced in the laboratory both by prevention of oviposition and by starvation. For up to two days after the onset of starvation the terminal oöcyte appears normal. After three days the prechorionic oöcyte may move through the ovariole wall; the yolk spheres are then disrupted. On the fourth day little yolk remains in the extruded oöcyte, and most of the extruded cells are degenerating. We suggest that extra-ovariolar egg resorption may be a mechanism for ensuring that the single ovariole is not occluded when conditions are suitable for oviposition.     

Effects of fruit fly host, fruit species, and host egg to female parasitoid ratio on the laboratory rearing of Biosteres arisanus

The host suitability of the oriental fruit fly, Bactrocera dorsalis (Hendel), for development of Biosteres arisanus (Sonan), a braconid parasitoid, was compared with three other fruit fly species, namely, Mediterranean fruit fly, Ceratitis capitata Weidemann, melon fly, Bactrocera cucurbitae Coquilett, and Malaysian fruit fly, Bactrocera latifrons (Hendel). In addition, effects of five different fruit species, namely, Carica papaya L. (solo papaya), Musa sapientum (L.) O. Ktze. (apple banana), Mangifera indica (L.) (Haden mango), Terminalia catappa (L.) (false kamani), and Citrus aurantiifolia (Christman) Swingle (common lime), on the parasitization rate of B. dorsalis and sex ratio of parasitoid progenies were evaluated. Effects of host egg to female B. arisanus ratios on parasitoid progeny yields were likewise determined. The host suitability of fruit flies for development of B. arisanus was ranked as: B. dorsalis>C. capitata=B. latifrons=B. cucurbitae. Based on percent parasitization of B. dorsalis, preference of B. arisanus females for host eggs varied with fruit species, however, preferential oviposition displayed by female parasitoids did not influence sex ratios of subsequent parasitoid progenies. Increases in host egg to female parasitoid ratios of 5:1, 10:1, 20:1, 25:1, and 30:1 corresponded with increases in parasitoid progeny yield reaching a plateau at 20:1.     

Relationships between parasitoid host range and host defence: a comparative study of egg encapsulation in two related parasitoid species

Abstract. Parasitoid host range may proceed from traits affecting host suitability, traits affecting parasitoid foraging behaviour, or both. We tested the hypothesis that encapsulation can be used as a reliable indicator of parasitoid host range in two closely related larval endoparasitoids of Lepidoptera. Cotesia glomerata (L.) (Hymenoptera: Braconidae) is gregarious and a generalist on several species of Pieridae, whereas C. rubecula (Marshall) is solitary and specific to Pieris rapae (L.). We determined the effects of host species ( Pieris brassicae (L.), P. napi (L.) and P. rapae ) (Lepidoptera: Pieridae) and host developmental stage (early first, second and third instar) on encapsulation of parasitoid eggs. Host species and parasitoid species, as well as the resulting interaction between these two factors had significant effects on encapsulation of Cotesia eggs. Encapsulation in Pieris hosts was much lower for C. glomerata (<34%, except for second and third instar of P. rapae ) than for C. rubecula (>32%), even when the latter was parasitizing P. rapae. Encapsulation increased with the age of the larvae, although the only significant difference was for C. glomerata. Overall, P. rapae showed a stronger encapsulation reaction than P. brassicae and P. napi. Encapsulation levels of C. glomerata corresponded well to patterns of female host species and host age preference for oviposition and parasitoid larval performance. In contrast, percentages of encapsulation of C. rubecula were not consistent with host preference and host suitability. We argue that encapsulation alone is unlikely to provide a sufficient explanation for C. glomerata and C. rubecula host range.