Song as an honest signal of past developmental stress in the European starling (Sturnus vulgaris)

Bird song is a sexually selected male trait where females select males on the basis of song quality. It has recently been suggested that the quality of the adult male song may be determined by nutritional stress during early development. Here, we test the 'nutritional–stress hypothesis' using the complex song of the European starling. Fledgling starlings were kept under experimental treatment (unpredictable short–term food deprivations) or control conditions (ad libitum food supply), for three months immediately after independence. We measured their physiological and immune responses during the treatment and recorded song production during the following spring. Birds in the experimental group showed increased mass during the treatment and also a significantly suppressed humoral response compared with birds in the control group. There was no difference between the groups in the cell–mediated response. Next spring, males in the experimental group spent less time singing, sang fewer song bouts, took longer to start singing and also sang significantly shorter song bouts. These data support the hypothesis that both the quality and quantity of song produced by individual birds reflect past developmental stress. The results also suggest the 'nutritional–stress hypothesis' is best considered as a more general 'developmental–stress hypothesis'.

Developmental stress, social rank and song complexity in the European starling (Sturnus vulgaris)

Bird song is a sexually selected trait and females have been shown to prefer males that sing more complex songs. However, for repertoire size to be an honest signal of male quality it must be associated with some form of cost. This experiment investigates the effects of food restriction and social status during development on song complexity in the European starling (Sturnus vulgaris). Birds that experienced an unpredictable food supply early in life produced a significantly smaller repertoire of song phrases than those with a constant food supply. Social status during development was also significantly correlated with repertoire size, with dominant birds producing more phrase types. This study therefore provides novel evidence that social as well as nutritional history may be important in shaping the song signal in this species.

Rapid behavioural adjustments to unfavourable light conditions in European starlings (Sturnus vulgaris)

Although it is known that many birds possess ultraviolet (UV)-sensitive vision, most commercially housed poultry species, as well as species held in zoos, laboratories, or bred for show, are maintained under lighting that is deficient in UV wavelengths compared with normal daylight. UV-sensitive vision has been shown to be important in both foraging and mate-choice decisions. UV-poor conditions, in which information in this waveband is eliminated, could, therefore, present an important welfare issue. Eight European starlings were given a series of preference tests (eight hours per trial, for six days), in which they could choose to feed in one of four channels. Four experimental trials were carried out, during which the channels were covered in either UV-transmitting (UV+) or UV-blocking (UV-) filters, to determine whether birds had an initial preference for feeding in UV-rich environments and whether there was any change in their preference over time. There was an initial preference for the UV+ environment, but this preference declined very rapidly with familiarity, and was absent by the final trial. These results imply that starlings can rapidly adjust their feeding behaviour if faced with unfamiliar light environments and that any initial behavioural changes attributable to UV-deficient environments may be short-lived. However, further work is necessary to establish whether these adjustments occur across a range of species and contexts before any welfare concerns can be ruled out.

Invasive species can't cover their tracks : using microsatellites to assist management of starling (Sturnus vulgaris) populations in Western Australia

Invasive species are known to cause environmental and economic damage, requiring management by control agencies worldwide. These species often become well established in new environments long before their detection, resulting in a lack of knowledge regarding their history and dynamics. When new invasions are discovered, information regarding the source and pathway of the invasion, and the degree of connectivity with other populations can greatly benefit management strategies. Here we use invasive common starling (Sturnus vulgaris) populations from Australia to demonstrate that genetic techniques can provide this information to aid management, even when applied to highly vagile species over continental scales. Analysis of data from 11 microsatellites in 662 individuals sampled at 17 localities across their introduced range in Australia revealed four populations. One population consisted of all sampling sites from the expansion front in Western Australia, where control efforts are focused. Despite evidence of genetic exchange over both contemporary and historical timescales, gene flow is low between this population and all three more easterly populations. This suggests that localized control of starlings on the expansion front may be an achievable goal and the long-standing practice of targeting select proximal eastern source populations may be ineffective on its own. However, even with low levels of gene flow, successful control of starlings on the expansion front will require vigilance, and genetic monitoring of this population can provide essential information to managers. The techniques used here are broadly applicable to invasive populations worldwide.

Synchronization by low-amplitude light-dark cycles of 24-hour pineal and plasma melatonin rhythms of hatchling European starlings (Sturnus vulgaris)

In young European starlings, as in other avian species, high-amplitude 24-hr rhythms in plasma and pineal melatonin are already present around the time of hatching. In chickens this rhythmicity results at least partly from the light sensitivity of the melatonin-producing and -secreting system. In contrast to the chicken, the starling is a hole-nesting bird, and it seemed questionable whether the low light intensities in the nest are sufficient to synchronize perinatal melatonin rhythms. We therefore exposed starling eggs to light cycles roughly simulating those measured in nest-boxes, i.e., an 11-hr phase of complete darkness and a 13-hr phase consisting of 15 min of dim light (10 lux) alternating with 30 min of darkness. For one group the photophase lasted from 0600 to 1900 hr; for the other group the photophase lasted from 1800 to 0700 hr. In approximately 10-hr-old hatchlings of both groups, plasma and pineal melatonin concentrations were high during the dark phase and low during the light phase. We conclude that perinatal low-amplitude light intensity changes of the kind experienced by hatching starlings in the field are sufficient for synchronizing the melatonin-producing and -secreting system in the pineal and possibly other organs.

Sturnus vulgaris, mitochondrial selection, MATLAB code and microsatellite data

Here we provide MATLAB code used to simulate drift and selection between and within individuals, which has been used to investigate mitochondrial haplotype frequency shifts in Sturnus vulgaris. Also provided is a microsatellite data set used to assess whether empirical allele frequency shifts were likely to be caused by admixture. These files support and upcoming publication, which concludes that within-individuals selection on mitochondrial DNA best explains empirical data.

Pollutants increase song complexity and the volume of the brain area HVC in a songbird

Environmental pollutants which alter endocrine function are now known to decrease vertebrate reproductive success. There is considerable evidence for endocrine disruption from aquatic ecosystems, but knowledge is lacking with regard to the interface between terrestrial and aquatic ecosystems. Here, we show for the first time that birds foraging on invertebrates contaminated with environmental pollutants, show marked changes in both brain and behaviour. We found that male European starlings (Sturnus vulgaris) exposed to environmentally relevant levels of synthetic and natural estrogen mimics developed longer and more complex songs compared to control males, a sexually selected trait important in attracting females for reproduction. Moreover, females preferred the song of males which had higher pollutant exposure, despite the fact that experimentally dosed males showed reduced immune function. We also show that the key brain area controlling male song complexity (HVC) is significantly enlarged in the contaminated birds. This is the first evidence that environmental pollutants not only affect, but paradoxically enhance a signal of male quality such as song. Our data suggest that female starlings would bias their choice towards exposed males, with possible consequences at the population level. As the starling is a migratory species, our results suggest that transglobal effects of pollutants on terrestrial vertebrate physiology and reproduction could occur in birds.

The effect of flicker from fluorescent lights on mate choice in captive birds

The visual systems of birds are hypothesized to have higher temporal resolution than those of humans, suggesting that they may be able to perceive the flicker emitted from conventional low-frequency fluorescent lights (LF; 100 Hz in Europe, 120 Hz in the U.S.A.). These lights are commonly used in the housing of captive birds and this may affect both their welfare and performance in experiments. We carried out mate choice experiments on European starlings, Sturnus vulgaris, under both low- and high-frequency fluorescent lights (HF; > 30 kHz, at which flicker is imperceptible). Indicators of male condition and size, together with the reflectance spectra and length of the males' throat feathers, were also recorded to ascertain which variables correlated with female preference. Females ranked males consistently under HF, but not LF, lighting, and individual females chose different males under the two lighting types. Under HF lighting, females chose to spend more time with males that had longer throat feathers. The flicker rate of the light clearly affected the choices made by the females, possibly because of nonspecific stress effects or decreased discrimination ability. Our results imply that careful interpretation of mate choice experiments is needed, especially with regard to the lighting types used, to elucidate the real cause behind any variation shown. (c) 2006 The Association for the Study of Animal Behaviour. Published by Elsevier Ltd. All rights reserved.

Do European starlings prefer light environments containing UV?

Many captive birds are kept in artificial lighting that is typically deficient in ultraviolet (UV) wavelengths. Most birds can perceive the range of light that humans see but also have an additional retinal cone type that is tuned to UV wavelengths. Consequently, artificial lighting may be detrimental as it might limit the functionality of their vision. We examined the preferences of European starlings, Sturnus vulgaris, for various artificial light environments. In our first experiment, groups of starlings showed a preference for environments that contained UV (UV+) over those where UV wavelengths had been removed (UV -). This preference was not affected by the sex of the individuals within the group or, as shown in a later experiment, by whether the birds had been previously housed in UV+ or UV - conditions. In contrast, individual starlings showed no preference for UV+ over UV - environments, although the power of our test was low. In a subsequent experiment, starling groups preferred the higher of two light intensities that were presented; however, equalizing the overall quantal flux between UV+ and UV - extinguished any preference for UV+ over UV -. The group preference for UV+ conditions in the first experiment may therefore have resulted from a preference for brighter conditions rather than a specific preference for UV. However, equalizing the quantal flux may not equalize perceived brightness, because it is not known how birds' visual systems weight input from each cone type. We conclude that, for nonbreeding, group-housed captive starlings, there is no positive evidence of a preference for the presence of UV as a specific wavelength. (C) 2002 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.

Does the flicker frequency of fluorescent lighting affect the welfare of captive European starlings?

It is common practice for captive birds to be kept under fluorescent lighting, which typically flickers at either 100Hz (UK) or 120 Hz (USA). Such lighting was developed for human vision and it is thought that birds may be able to detect higher frequencies of flicker than humans. For humans, 100Hz fluorescent lighting has been linked to eyestrain, headaches and migraine, even though this rate of flicker is above the human perceptual flicker-fusion frequency of around 60 Hz. Keeping birds under 100 Hz lighting is therefore potentially detrimental to their welfare. We studied the preferences of wild-caught European starlings, Sturnus vulgaris, for high-frequency (HF, >30 kHz) fluorescent lighting versus conventional low-frequency (LF, 100 Hz) fluorescent lighting. The flicker frequency of the HF lighting would be undetectable to the nervous system of any animal. We also exposed starlings to either HF or LF light for 2 weeks, and investigated the degree of stress caused by each environment by monitoring their behaviour and plasma corticosterone levels. Groups of starlings showed a preference for HF lighting over LF lighting (P < 0.001), which indicates that they can detect a difference between the two lighting conditions and find the HF lighting preferable. However, there were no measurable differences in behaviour or plasma corticosterone levels when the birds were housed under either HF or LF for 2 weeks, thus providing initial evidence that housing starlings under 100Hz lighting may not be detrimental to the welfare of starlings during early captivity. (C) 2003 Elsevier B.V. All rights reserved.

'Anting' as food preparation : formic acid is worse on an empty stomach

Anting is a behavior common among passerine birds, yet its function is unknown. The behavior consists of a highly stereotyped set of movements which start when a bird picks up an ant, usually one which sprays formic acid as a defense, and sweeps it with frenzied motions through its feathers. The bird will often also eat the ant. As formic acid is toxic, we have tested the food-preparation hypothesis, that is, that the birds are anting to remove a distasteful or toxic substance from the ant before eating it. In a pair of experiments on starlings, Sturnus vulgaris, we have found evidence in support of this hypothesis.

Context-dependent visual preferences in starlings and blue tits: mate choice and light environment

There is considerable interest in the role that ultraviolet (UV) cues play in the foraging and mate choice decisions of birds. However, with the exception of the zebra finch, Taeniopygia guttata, it is not yet clear whether ultraviolet preferences are context specific, or whether birds show a general preference for full-avian-spectrum environments 320-700 nm) irrespective of the activity in which they are engaged. We investigated whether European starlings, Sturnus vulgaris, and blue tits, Parus caeruleus, show general (nonresource based) or context-specific preferences for full-spectrum environments. We found that neither species showed a general preference for UV-present (UV+) over UV-deficient (UV-) environments, when those environments contained no resources (experiment 1). Furthermore, neither species showed a UV+ preference when cages contained food, water and perches (starlings; experiment 2) or food, perches and heterospecifics (blue tits; Hunt et al. 1999. Animal Behaviour, 58, 809-815). However, both species did show highly significant preferences for UV+ conditions when viewing potential mates. Such experiments are necessary before one can conclude that particular wavebands have specific relevance to mate choice. In fact, our results suggest that the importance of particular wavelength compositions do indeed vary with behavioural context. (C) 2002 The Association for the Study of Animal Behaviour.

Mitochondrial DNA offers unique insights into invasion history of the common starling

Mitochondrial DNA (mtDNA) can be a powerful genetic marker for tracing origins and history of invasive populations. Here, we use mtDNA to address questions relevant to the understanding of invasion pathways of common starlings (Sturnus vulgaris) into Western Australia (WA) and discuss the utility of this marker to provide information useful to invasive species management. Mitochondrial sequence data indicate two geographically restricted genetic groups within Australia. Evidence of dispersal from genetically distinct sources outside the sampled range of starlings in Australia suggests increased vigilance by management agencies may be required to prevent further incursions from widely separated localities. Overall, genetic diversity in Australia was lower than in samples from the native range. Within Australia, genetic diversity was lowest in the most recently colonized area in the west, indicating that demographic bottlenecks have occurred in this area. Evidence of restricted dispersal between localities on the edge of the range expansion (ERE) in WA and other Australian sampling localities suggests that localized control within the ERE may be effective in preventing further range expansion. Signatures of spatial and demographic expansion are present in mismatch analyses from sampling localities located at the ERE, but neutrality indices did not support this finding, suggesting that the former may be more sensitive to recent expansion. Additionally, mismatch analyses support the presence of admixture, which is likely to have occurred pre-introduction. We compare our findings with those from a microsatellite study of the same samples and discuss how the mtDNA analyses used here offer valuable and unique insights into the invasion history of introduced species.