Fear in grasslands: the effect of Eurasian kestrels on skylark abundances

Predation has received considerable theoretical and empirical support in population regulation. The effect of predators, however, could be achieved in direct (killing) or indirect effects (such as displacement). In this paper, we explored the relationship between Eurasian kestrels Falco tinnunculus and skylarks Alauda arvensis in Mediterranean grasslands. First, we analysed the presence of skylarks in the kestrel diet over 9 years. We also compared a grassland area of experimentally increased kestrel density and a second grassland as control area to evaluate the direct or indirect effect on skylark abundance. We also considered two different habitats, grazed and ungrazed plots. If skylark abundance decreased as the kestrel breeding season progressed in high-density kestrel area compared with the control area, it would suggest a direct effect (predator hypothesis). If skylark abundance remains constant in both areas of contrasting kestrel density, it would suggest that skylarks avoid kestrels (avoidance hypothesis). We found that skylark abundance decreased in the kestrel area from the beginning of kestrel nest-box installation to recent years. The rate of skylark consumption decreased in a 9-year period as kestrel abundance increased, although the total amount skylark consumption did not show a decreasing trend. In addition, skylarks were more abundant in the kestrel-free area than in the kestrel area. Finally, we found that skylark abundance did not change through the kestrel breeding period in relation to grazing. We suggest that an increased breeding density of kestrels during their breeding period may force the skylarks to breed in other areas, which may explain the decline of their abundance.     

Male-biased predation of western green lizards by Eurasian kestrels

Selective predation can be an important force driving the evolution of organisms. In particular, sex-biased predation is expected to have implications for sexual selection, sex allocation and population dynamics. In this study, we analysed sex differences in the predation of the western green lizard (Lacerta bilineata) by the Eurasian kestrel (Falco tinnunculus) during the reproductive season. In addition, we investigated whether the rate of predation differed during the 8-year study period and among the three habitats studied. We collected lizard remains from nest boxes of kestrels. Freshly killed lizards were sexed by visual inspection, whilst the sex of head remains was assigned by analysing the cephalic scale morphology using geometric morphometrics. Our results show that the risk of being predated by a kestrel in our population was overall about 3.55 times higher for males than for females. To our knowledge this is the first study showing a male-biased predation in a lizard species. The selective predation of males was consistent between years over the 8-year study period (1999-2006) and also consistent between the three types of kestrel hunting habitat. Overall predation rates on lizards differed between habitats, depending on the year. We propose that the observed sex-biased predation is mainly due to sex differences in lizard behaviour.     

Carotenoid-based bill and eye ring coloration as honest signals of condition: an experimental test in the red-legged partridge (<Emphasis Type="Italic">Alectoris rufa</Emphasis>)

Carotenoid pigments cannot be synthesized by vertebrates but must be ingested through the diet. As they seem to be a limited resource, carotenoid-based ornaments are particularly interesting as possible honest signals of individual quality, in particular of foraging efficiency and nutritional status. Some studies have demonstrated the condition dependence of carotenoid-based plumage in birds. However, many other carotenoid-pigmented bare parts (i.e. skin, caruncles, bills, cere, and tarsi) are present in birds but, in comparison with plumage, little is known about these traits as indicators of individual quality. Here, we show that the eye ring pigmentation and bill redness of the red-legged partridge (Alectoris rufa) are positively associated to body condition and recent changes in body mass. Also, we found a negative relationship between these two traits and heterophil-to-lymphocyte ratio, an indicator of physiological stress (the relationship with bill redness being significant only for males). In an experiment, we found that after a period of reduction in food intake (with the consequent loss of body mass), food-restricted birds showed lower eye ring pigmentation than ad-libitum-fed birds. Therefore, different ornaments seem to reflect changes in body condition but at different speeds or intensities (eye ring, a fleshy ornament, appears to respond more rapidly to changes in the nutritional status than a keratinized structure as the bill). These results indicate that carotenoid-based ornaments are condition-dependent traits in the red-legged partridge, being therefore susceptible to be employed as honest signals of quality in sexual selection.     

Short-sighted evolution of virulence in parasitic honeybee workers (Apis mellifera capensis Esch.)

The short-sighted selection hypothesis for parasite virulence predicts that winners of within-host competition are poorer at transmission to new hosts. Social parasitism by self-replicating, female-producing workers occurs in the Cape honeybee Apis mellifera capensis, and colonies of other honeybee subspecies are susceptible hosts. We found high within-host virulence but low transmission rates in a clone of social parasitic A. m. capensis workers invading the neighbouring subspecies A. m. scutellata. In contrast, parasitic workers from the endemic range of A. m. capensis showed low within-host virulence but high transmission rates. This suggests a short-sighted selection scenario for the host-parasite co-evolution in the invasive range of the Cape honeybee, probably facilitated by beekeeping-assisted parasite transmission in apiaries.     

Anatomic and acoustic sexual dimorphism in the sound emission system of Phoenicoprocta capistrata (Lepidoptera: Arctiidae)

Both sexes of Phoenicoprocta capistrata have functional tymbals. The scanning electron microscopy revealed differences in the morphology of these organs in males and females. Male tymbals have a well-developed striated band, constituted by 21 ± 2 regularly arranged striae whereas female tymbals lack a striated band. This type of sexual dimorphism is rare in Arctiidae. The recording of the sound produced by moths held by the wings revealed that while males produced trains of pulses organized in modulation cycles, females produced clicks at low repetition rate following very irregular patterns. Statistically, there are differences between sexes in terms of the duration of pulses, which were 355 ± 24 μs in the case of males and 289 ± 29 μs for females. The spectral characteristics of the pulses also show sexual dimorphism. Male pulses are more tuned (Q ₁₀ = 5.2 ± 0.5) than female pulses (Q ₁₀ = 2.7 ± 0.5) and have a higher best frequency (42 ± 1 kHz vs. 29 ± 2 kHz). To our knowledge, this is the first report on an arctiid moth showing sexual dimorphism in tymbal’s anatomy that leads to a best frequency dimorphism. Males produce sound at mating attempts. The sounds recorded during mating are modulation cycles with the same spectral characteristics as those recorded when males are held by the wings. The morphological and acoustic features of female tymbals could indicate a process of degeneration and adaptation to conditions under which the emission of complex patterns is not necessary. Frank Coro no longer works at Universidad de La Habana.     

Dancing to different tunes: heterospecific deciphering of the honeybee waggle dance

Although the structure of the dance language is very similar among species of honeybees, communication of the distance component of the message varies both intraspecifically and interspecifically. However, it is not known whether different honeybee species would attend interspecific waggle dances and, if so, whether they can decipher such dances. Using mixed-species colonies of Apis cerana and Apis mellifera, we show that, despite internal differences in the structure of the waggle dances of foragers, both species attend, and act on the information encoded in each other’s waggle dances but with limited accuracy. These observations indicate that direction and distance communication pre-date speciation in honeybees.     

Costs of female odour in males of the parasitic wasp <Emphasis Type="Italic">Lariophagus distinguendus</Emphasis> (Hymenoptera: Pteromalidae)

The display of female traits by males is widespread in the animal kingdom. In several species, this phenomenon has been shown to function adaptively as a male mating strategy to deceive sexual rivals (female mimicry). Freshly emerged males of the parasitic wasp Lariophagus distinguendus (Hymenoptera: Pteromalidae) are perceived by other males as if they were females because of a very similar composition of cuticular hydrocarbons which function as a sex pheromone in this species inducing courtship behaviour in males. Within 32 h, however, males deactivate the pheromone and are no longer courted by other males. In this paper, behavioural experiments were performed to test hypotheses on potential costs and benefits associated with the female odour in young males. We did not find any benefits, but demonstrated that young males were significantly more often outrivaled in male-male contests when competing with two older males for a female. Also, young males were significantly more often mounted in homosexual courtship events during these contests. Thus, display of female traits by males is not necessarily beneficial, and in fact, can be disadvantageous. We suggest that these costs have favoured the evolution of the pheromone deactivation mechanism in L. distinguendus males. The function of cuticular hydrocarbons as a female courtship pheromone in L. distinguendus might have evolved secondarily from a primary function relevant for both genders, and the deactivation of the signal in males might have caused a shift of specificity of the chemical signal from the species level to the sex level.     

Relationship between abundance of rodents and damage to agricultural crops

Developing a relationship between pest abundance and damage to crops is essential for the calculation of economic injury levels (EIL) leading to informed management decisions. The crop modelling framework, APSIM, was used to simulate the impact of mouse damage on yield loss on wheat where a long-term dataset on the density of mice was available (1983–2003). The model was calibrated using results from field trials where wheat plants were hand clipped to imitate mouse damage. The grazing effect of mice was estimated using the population density, daily intake per mouse and the proportion of wheat grain and plant tissue in the diet to determine yield loss. The mean yield loss caused by mice was 12.4% (±5.4S.E.; range −0.5 to 96%). There were 7/21 years when yield loss was >5%. A damage/abundance relationship was constructed and a sigmoidal curve explained 97% of variation when accounting for different trajectories of mouse densities from sowing to harvest. The majority of damage occurred around emergence of the crop when mouse densities were >100 mice ha⁻¹. This is the first time that field data on mouse density and a crop simulation model have been combined to estimate yield loss. The model examines the efficacy of baiting and how to estimate EILs. Because the broadscale application of zinc phosphide is cheap and effective, the EIL is very low (<1% yield loss). The APSIM model is highly flexible and could be used for other vertebrate pests in a range of crops or pastures to develop density/damage relationships and to assist with management.     

Contrasting responses of seed yield to elevated carbon dioxide under field conditions within Phaseolus vulgaris

The rising concentration of carbon dioxide [CO₂] in the atmosphere represents an increase in a growth-limiting resource for C₃ crop species. Identification of lines or characteristics of lines which have superior yield at elevated [CO₂] could aid in adaptation to this global change. While intraspecific variation in responses to elevated [CO₂] has been found in several species, intraspecific differences in crop yield responses to elevated [CO₂] under field conditions have seldom been documented. In this 4-year study, the responses of photosynthesis, growth, pod number, seed number and size, and seed yield to the elevation of [CO₂] to 180 μmol mol⁻¹ above the current ambient concentration were examined in four varieties of Phaseolus vulgaris in the field, using open-top chambers. There was a significant variety by [CO₂] interaction for seed yield, with seed yield at elevated [CO₂] ranging from 0.89 to 1.39 times that at ambient [CO₂] (mean 1.17×) in the different varieties, when averaged over 4 years. The highest yielding variety at elevated [CO₂] was not the highest yielding variety at ambient [CO₂]. The varieties with the largest and smallest yield responses both had an indeterminate growth habit. Down-regulation of photosynthesis at elevated [CO₂] only occurred in the two indeterminate varieties, and there was no significant correlation between the response of single leaf photosynthetic rate and the response of seed yield to elevated [CO₂] among varieties, nor between the responses of stem mass and seed yield. The change in the number of pods at elevated [CO₂] was the primary determinant of the response of seed yield. These results indicate that significant variation in the response of seed yield to elevated [CO₂] under field conditions does exist among varieties of P. vulgaris, and that variation in the response of pod and seed number may be more important than variation in photosynthetic response.     

The response of Trissolcus basalis to footprint contact kairomones from Nezara viridula females is mediated by leaf epicuticular waxes

Chemical footprints left behind by true bugs are perceived as contact kairomones by scelionid egg parasitoids. Female wasps encountering a contaminated artificial substrate display a characteristic arrestment posture, holding the body motionless and antennating the surface. In the system Nezara viridula (L.) and its egg parasitoid Trissolcus basalis (Wollaston), previous studies have shown that the kairomone mediating such behavior is part of N. viridula’s cuticular hydrocarbons (CHC) and furthermore that the wasp’s ability to discriminate host male and female footprints is mainly based on the presence/absence of nonadecane (nC₁₉). In this study, the effect of epicuticular waxes of leaves of broad bean, Vicia faba, on wasp responses to footprints of N. viridula females were investigated. Approximately 20% of T. basalis females displayed an arrestment posture when released on the adaxial leaf surfaces of broad bean plants with intact wax layer and without host chemical contamination; whereas ∼70% of wasps displayed the arrestment posture when intact leaves were contaminated by host female footprints. Adaxial leaf surfaces of broad bean plants dewaxed with an aqueous solution of gum arabic and afterwards contaminated by N. viridula females induced arrestment responses in about 10% of female wasps; the same percentage of arrestment (10%) was observed when the wasps were released on leaves contaminated by host females and subsequently dewaxed. The side of the polymer film that was appressed to the leaf surface, peeled from the contaminated leaves, induced an arrestment posture in about 95% of observed wasps. Scanning electron microscopy (SEM) revealed that the epicuticular waxes occurred as a film densely crystallized as irregularly shaped platelets with spherical granules randomly distributed. These findings demonstrated that epicuticular waxes of broad bean leaves can mediate the foraging behavior of T. basalis females by absorbing contact kairomones of the host.     

Settlement decisions in blue tits: difference in the use of social information according to age and individual success

Dispersers are expected to assess breeding habitat quality before settlement. Although cues reflecting habitat quality are well studied, social cues have not been as well evaluated. In this paper, we studied breeding habitat selection during 3 years in a natural population of blue tits, Cyanistes caeruleus, breeding in nest-boxes. Our aim was to investigate if this species used conspecific density and/or reproductive success of con- and heterospecifics (i.e., social cues) in settlement decisions. The patterns observed were consistent with the idea that juveniles, when dispersing from their natal patches, did not react to any of the cues that we tested. In contrast, breeders that dispersed seemed to respond to both conspecific mean patch reproductive success (PRS) and breeding density of the settlement patch in the year of dispersal, their response differing according to their own reproductive success. Indeed, failed breeders moved to areas with high PRS and low density relative to source patches, while successful breeders behaved the opposite. The comparison between juveniles and adults might be modulated by the limited time available to juveniles to gather information on PRS and density at the end of the dispersing year. Adults lacking these time constraints, however, seemed to rely on these conspecific cues although limited by their own quality. Additionally, breeders were more likely to be immigrants in patches with relatively low breeding success and density the previous year, suggesting that settlement is influenced by multiple cues, which may reveal information on different aspects of habitat and be available at different moments. Collectively, our results support the importance of social cues for blue tits’ settlement.     

Elevated CO2 reduces the response of Sitobion avenae (Homoptera: Aphididae) to alarm pheromone

The aphid alarm pheromone (E)-β-farnesene (EβF) is an efficient signal that warns aphids of attack by natural enemies. In this field study, eight open-top chambers (OTCs) located in Beijing, China (40°11′N, 116°24′E) with spring wheat Triticum aestivum were used to examine the response of the grain aphid Sitobion avenae to CO₂ (ambient vs. double ambient) and EβF (applied zero, two, or five times each day). We experimentally tested the hypotheses that, depending on frequency of EβF release, elevated CO₂ reduces the response (in terms of population density) of S. avenae to EβF, and that lower activity of acetylcholinesterase (AChE) in S. avenae may be involved in its reduced sensitivity to EβF under elevated CO₂. Numbers of S. avenae declined with increased frequency of EβF application under ambient CO₂ but were unaffected by EβF application under elevated CO₂. Additionally, the mean relative growth rate (MRGR) and the dry material and amino acid content of S. avenae increased with elevated CO₂ but declined when with EβF application. Activities of superoxide dismutase and catalase were higher in S. avenae under elevated vs. ambient CO₂. Under elevated CO₂, however, AChE activity remained low when S. avenae was exposed to the lower EβF frequency, while the highest AChE activity occurred in aphids exposed to the higher EβF frequency. These results indicate that aphids become insensitive to EβF under elevated CO₂, perhaps because of decreased AChE activity.