Gliding flight in the American Kestrel (Falco sparverius): An electromyographic study

Electromyographic (EMG) activity was studied in American Kestrels (Falco sparverius) gliding in a windtunnel tilted to 8 degrees below the horizontal. Muscle activity was observed in Mm. biceps brachii, triceps humeralis, supracoracoideus, and pectoralis, and was absent in M. deltoideus major and M. thoracobrachialis (region of M. pectoralis). These active muscles are believed to function in holding the wing protracted and extended during gliding flight. Quantification of the EMG signals showed a lower level of activity during gliding than during flapping flight, supporting the idea that gliding is a metabolically less expensive form of locomotion than flapping flight. Comparison with the pectoralis musculature of specialized gliding and soaring birds suggests that the deep layer of the pectoralis is indeed used during gliding flight and that the slow tonic fibers found in soaring birds such as vultures represents a specialization for endurant gliding. It is hypothesized that these slow fibers should be present in the wing muscles that these birds use for wing protraction and extension, in addition to the deep layer of the pectoralis. © 1993 Wiley-Liss, Inc.     

Energetic limitation of avian parental effort: Field experiments in the kestrel (Falco tinnunculus)

We studied the limiting factors for brood size in the kestrel, Falco tinnunculus, by measuring parental effort in natural broods of different size and parental response to manipulation of food satiation of the brood. Parental effort was quantified as total daily time spent in flight, and total daily energy expenditure, from all-day observations. During nestling care males with different natural brood sizes (4 to 7 chicks), spent an average of 4.75 h · d^?1 in flight independent of brood size, and expended an average total daily energy of 382 kJ · d^?1. Due to a higher flight-hunting yield (mammal-prey caught per hour hunting), males with larger natural broods were able to provision their broods with the same amount of food (mainly Microtus arvalis) per chick (62.6 g · d^?1), with the same effort as males with smaller broods. This provisioning rate was close to the mean feeding rate of hand-raised chicks in the laboratory, that were fed ad libitum, (66.8 g · d^?1 · chick^?1). Our food deprivation experiments revealed that male kestrels strongly respond to food shortage in the nest. In the older nestling phase males on average increased their daily rate of food delivery to the nest as a response to experimental food deprivation by almost three times to 646.4 g · d^?1, by increasing their flight activity level from 4.46 to 8.41 h · d^?1. This increased energy expenditure was sustained, for as long as eleven days, by increasing the metabolizable energy intake up to what is presumed to be the maximum rate. Even under considerable experimental food stress (chicks not being satiated due to continuous removal of delivered food by the observers) about half of the available daylight time remained unused for foraging. We conclude 1) that the mean daily energy expenditure of males during nestling care — to which clutch size is apparently initially adjusted — is well below the maximum they are able to sustain and 2) that the energy expenditure they can sustain under extremely high nestling demand is not set by the available time for foraging or the available energy in the environment. Thus the birds normally operate well below their presumed maximum, and only during food shortage, e.g., as caused by our experiments, do they increase activity up to this maximum. Therefore we conclude that the kestrels have costs other than energy expenditure, such as parental survival, that are involved in the increased “cost” of parental effort. We discuss possible generalisations about existing energetic limitations during parental care in altricial birds. From published estimates of daily energy expenditure during parental care (DEE_par) in 30 different bird species we derived the equation: DEE_par = 14.26 kg^0.65 Watt. This relationship differs significantly in slope (T = 2.49; p > 0.02) from the allometric equation for the maximum rate of energy assimilation (DME_max) as provided by Kirkwood (1983): DME_max = 19.82 kg^0.72 Watt. In smaller species (ca. 25 g) DEE_par about equals DME_max, while in the larger species (ca. 10 kg) DEE_par represents only about 60% of the predicted DME_max. This suggests that limitations in parental effort are more frequently set by the maximum sustainable energy intake in the smaller species than in larger species. Our allometric equations for DEE_par suggests that the relation between BMR, estimated using the equations of Aschoff and Pohl (1970), and the observed parental energy expenditure, is such that on average bird parents work at a daily level somewhere between 3 and 4 times BMR.     

Immunological development in nestling American kestrels Falco sparverius: post-hatching ontogeny of the antibody response

Avian research involving examination of immune function or testing of immunocompetence in wild birds has been based upon information on Galliforms, (chicken and quail) even though they are precocial, whereas most wild species with which ecologists, biologists and toxicologists work are altricial; blind, naked and completely dependent at hatching. Here we begin to address this gap in knowledge, offering insight into the early, post-hatching, humoral immune response in an altricial bird, the American kestrel (Falco sparverius). Over two breeding seasons, nestling kestrels were immunized with a non-pathogenic antigen, dinitrophenol keyhole limpet hemocyanin (DNP-KLH), between 3 and 9 days post-hatching and boostered 6 days later. Background levels, primary and secondary immune responses were measured using an enzyme linked immunosorbent assay. The specificity of our laboratory produced rabbit, anti-kestrel antibody was determined using a double immunodiffusion assay. Results showed the rabbit antiserum to have specific anti-kestrel IgG activity. Birds as young as three days old could successfully mount an antibody response, the magnitude of which increased with age at first vaccination. Early immunization did not compromise growth rate, nor did it affect the maximum secondary response. Comparatively, adult kestrels immunized during the same season and following the same protocol, had antibody levels four times higher than those of the nestlings.     

The effects of temperature and artificial rain on the metabolism of American kestrels (Falco sparverius)

The effect of rainfall on the metabolism of birds is poorly understood. We measured the metabolism as rate of oxygen consumption (VO2) of four male and four female American kestrels (Falco sparverius) using open-circuit respirometry. We measured VO2 during the spring at ambient temperatures (Ta) of 5, 10, 15, and 25 degrees C in air without rainfall and with simulated rainfall of 2.5 (low rainfall) and 6.1 cm h(-1) (high rainfall). Kestrel metabolism was significantly higher when exposed to the two rainfall levels compared to no rainfall. However, kestrel metabolism was not significantly different at the two rainfall levels. Body temperature (Tb) was significantly lower under high rainfall compared to low rainfall. In addition, under both rainfall levels Tb decreased with decreasing Ta. Calculated thermal conductance was significantly higher in kestrels exposed to rain compared to no rainfall. Kestrels may use sleeking behavior at high rainfall levels to decrease water penetration of the plumage. Daily energy expenditure (DEE) of kestrels exposed to rain may increase markedly, and kestrel energetics may be further exacerbated by wind that often accompanies natural rainstorms.     

Preliminary report of Haemoproteus tinnunculi infection in a breeding population of American kestrels (Falco sparverius)

A population of American kestrels breeding in southeastern Pennsylvania was examined for hematozoa. Haemoproteus tinnunculi infected 17 of 23 (74%) of the adults. Parasitemia ranged from two to 252, with a median of 32 infected erythrocytes per 10,000. Parasitemia and body weight of female kestrels were negatively correlated. This parasite was not observed in the six juvenile or 38 nestling kestrels examined. Trypanosoma sp. was detected by culture in three of seven (43%) adults, but not in the six juveniles and eight nestlings examined.     

Seasonal shifts in sex ratios of fledgling American kestrels (Falco sparverius paulus): The Early Bird Hypothesis

We document a seasonal shift in the sex ratios of broods produced by resident southeastern American kestrels (Falco sparverius paulus) breeding in nest boxes in Florida. Early in the breeding season, most biased broods were biased towards males, whereas later in the season, most biased broods were biased towards females. Computer-simulated broods subjected to sex-biased egg and/or nestling mortality demonstrate that it is possible that differential mortality produced the pattern of bias that we observed. However, these simulations do not exclude the possibility that female kestrels were manipulating the primary sex ratio of the broods. We present evidence that this sex ratio shift is adaptive: for males we detected breeding as yearlings, all had fledged early the previous season. No such relationship between season and the probability of breeding as a yearling was found for females. We propose the Early Bird Hypothesis as the ecological basis for the advantage of fledg ing early in males. We hypothesize that pre-emptive competition among post-fledging, dispersing males for breeding sites confers an advantage to males fledged early in the season. This hypothesis may explain why a non-migratory population of the Eurasian kestrel (F. tinnunculus) and non-migratory American kestrels breeding in Florida (F. s. paulus) exhibit this seasonal shift in sex ratios, whereas migratory American kestrels (F. s. sparverius) breeding in Saskatchewan, Canada, do not. We discuss the relevance of the Early Bird Hypothesis for other animal species.     

Prevalence of west Nile virus antibodies in a breeding population of American Kestrels (Falco sparverius) in Pennsylvania

West Nile virus (WNV) has been identified in nearly 300 species of wild birds, including raptors, in North America since its introduction in New York City in 1999. American Kestrels (Falco sparverius) are susceptible to WNV infection, and the numbers of these birds have declined along the Atlantic coast in recent years. We examined the population biology and WNV exposure of kestrels breeding in the area surrounding Hawk Mountain Sanctuary in Kempton, Pennsylvania, USA. The reproductive biology of kestrels in this area was studied from 1992 until 2004. The number of kestrels breeding in nestboxes in 2004 was only 44% of the 6-yr mean observed prior to 1999. During the 2004 nesting season (study period: 8 June through 22 July 2004), adult kestrels were trapped near the site of their nestboxes. Blood samples were obtained, and serum antibodies specific to WNV were quantified using a plaque reduction neutralization test. Of 22 birds tested, 21 exhibited serum antibodies to WNV, suggesting that most (95%) of the adult kestrels in the population had been exposed to WNV.     

The retinal pigment epithelium of the eye regulates the development of scleral cartilage

The majority of vertebrate species have a layer of hyaline cartilage within the fibrous sclera giving an extra degree of support to the eyeball. In chicks, this is seen as a cuplike structure throughout the scleral layer. However, the mechanisms that control the development of scleral cartilage are largely unknown.Here we have studied the phases of scleral cartilage development and characterised expression profiles of genes activated during the cartilage differentiation programme. CART1 and SOX9, the earliest markers of pre-committed cartilage, are expressed in the mesenchyme surrounding the optic cup. Later AGGRECAN, a matrix protein expressed during chondrocyte differentiation, is also expressed. The expression of these genes is lost following early removal of the optic cup, suggesting a role for this tissue in inducing scleral cartilage. By grafting young retinal pigment epithelium (RPE) and retina into cranial mesenchyme in vivo, it was found that RPE alone has the ability to induce cartilage formation.There are some exceptions within the vertebrates where scleral cartilage is not present; one such example is the placental mammals. However, we found that the cartilage differentiation pathway is initiated in mice as seen by the expression of Cart1 and Sox9, but expression of the later cartilage marker Aggrecan is weak. Furthermore, cartilage forms in mouse peri-ocular mesenchyme micromass culture. This suggests that the process halts in vivo before full differentiation into cartilage, but that murine scleral mesenchyme has retained the potential to make cartilage in vitro.RA, Wnts and Bmps have been linked to the cartilage development process and are expressed within the developing RPE. We find that RA may have a role in early scleral cartilage development but is not likely to be the main factor involved.These data reveal the course of scleral cartilage formation and highlight the key role that the optic cup plays in this process. The driving element within the optic cup is almost certainly the retinal pigmented epithelium.     

The kestrel (Falco tinnunculus L.) in Berlin: investigation of breeding biology and feeding ecology

Approximately 200–250 pairs of kestrels (Falco tinnunculus) breed in Berlin, preferentially in nest boxes. From 2002 to 2004, ten monitoring sites (breeding sites) characterised by different housing structure, land utilisation, vegetation cover and degree of building density were studied in Berlin: four in the city centre, three in a mixed zone and three in the outskirts. All pairs bred in nest boxes, so the reproductive success could easily be determined. Pellets, and feathers of bird prey species, were collected during the breeding seasons, and the food spectrum was determined based on these remains. There was no significant difference in the reproductive success of the kestrels between the three zones. Data on the number of fledged young indicated a sufficient food supply. In total, 9 species of mice and shrews, 23 bird species and 31 beetle species were identified as prey of kestrels. Urban kestrels specialise in hunting birds if mice and shrews are not readily available, with the house sparrow (Passer domesticus) as the favoured prey bird. Of note are anthropogenic food items, such as cutlet bones, that were found only in the city centre. This shows that the kestrel can adapt itself to humans with regard to its diet. There was no urban gradient with regard to reproductive success, but there was with the composition of food, such as the domination of bird prey in the city centre. The number of individual items of bird prey decreased from the centre to the outskirts. In conclusion, the results show that the kestrel is an opportunistic species which survives well anywhere in the city of Berlin.

Experimental Infection of a North American Raptor,American Kestrel (Falco sparverius), with Highly Pathogenic Avian Influenza Virus (H5N1)

Several species of wild raptors have been found in Eurasia infected with highly pathogenic avian influenza virus (HPAIV) subtype H5N1. Should HPAIV (H5N1) reach North America in migratory birds, species of raptors are at risk not only from environmental exposure, but also from consuming infected birds and carcasses. In this study we used American kestrels as a representative species of a North American raptor to examine the effects of HPAIV (H5N1) infection in terms of dose response, viral shedding, pathology, and survival. Our data showed that kestrels are highly susceptible to HPAIV (H5N1). All birds typically died or were euthanized due to severe neurologic disease within 4–5 days of inoculation and shed significant amounts of virus both orally and cloacally, regardless of dose administered. The most consistent microscopic lesions were necrosis in the brain and pancreas. This is the first experimental study of HPAIV infection in a North American raptor and highlights the potential risks to birds of prey if HPAIV (H5N1) is introduced into North America.     

The development of ommatidial patterning in metamorphosed eye imaginal discs implants ofDrosophila melanogaster

Summary The development of the rhabdomeric pattern in the compound eye ofDrosophila has been studied using combined transplantation and electron microscope techniques. In a first series of experiments eye imaginal discs of increasing age were implanted into larvae ready to pupate, thus losing variable amounts of the normal time for development. A sequence of differentiative abilities was found in the metamorphosed test pieces. As far as the photoreceptor cells are concerned, the most prominent steps of this sequence are: ability to form groups with other similar elements, anatomical polarization of microvilli, establishment of the rhabdomeric pattern and formation of an equator line. The stability of determination of the equator line was tested in a second experimental series. Fragment of different topographical origin within the mature eye anlage were brought to metamorphosis by implantation into larvae ready to pupate. It was found that an equator line differentiates only in those pieces which according to the published anlage maps contain the prospective equator region prior to metamorphosis. The mitotic abilities of implanted eye imaginal discs were investigated by means of in vitro³H-thymidine pulse-labelling and light microscope autoradiography of the differentiated test pieces. During the third larval stage the eye anlage is traversed by two consecutive mitotic waves, each one of them producing different categories of receptor cells. The first, anterior wave predominantly produces cells oriented toward the poles of the eye within the ommatidia, while the second, posterior wave gives rise to elements exclusively in an equatorial position. The dynamics of this proliferation are discussed in relation to the findings in the implantation experiments. Silver-grain counts support the possibility that at least two successive cell divisions occur in the eye anlage between labeling with tritiated thymidine and beginning of morphological differentiation. The relevance of this finding for the understanding of the concept of acquisition of competence is discussed.     

The accessory lateral eye of a diplopod, Cylindroiulus truncorum (Silvestri, 1896) (Diplopoda: Julidae)

Using electron microscopy we describe an accessory lateral eye for Cylindroiulus, a diplopod. The accessory eye is situated at the cell body rind of the optic lobes, deep inside the head, and is composed of six R-cells; a dioptric apparatus is absent. Comparison reveals that many arthropods possess accessory lateral eyes in addition to the compound eyes or lateral ocelli. Their homology and distribution among the arthropod main lineages is discussed along with characters that may be useful for reconstructing phylogeny.     

Extensive polymorphism and geographical variation at a positively selected MHC class II B gene of the lesser kestrel (Falco naumanni)

Understanding the selective forces that shape genetic variation in natural populations remains a high priority in evolutionary biology. Genes at the major histocompatibility complex (MHC) have become excellent models for the investigation of adaptive variation and natural selection because of their crucial role in fighting off pathogens. Here we present one of the first data sets examining patterns of MHC variation in wild populations of a bird of prey, the lesser kestrel, Falco naumanni . We report extensive polymorphism at the second exon of a putatively functional MHC class II gene, Fana- DAB*1. Overall, 103 alleles were isolated from 121 individuals sampled from Spain to Kazakhstan. Bayesian inference of diversifying selection suggests that several amino acid sites may have experienced strong positive selection (ω = 4.02 per codon). The analysis also suggests a prominent role of recombination in generating and maintaining MHC diversity (ρ = 4 Nc  = 0.389 per codon, θ = 0.017 per codon). Both the Fana -DAB*1 locus and a set of eight polymorphic microsatellite markers revealed an isolation-by-distance pattern across the Western Palaearctic ( r  = 0.67; P  = 0.01 and r  = 0.50; P  = 0.04, respectively). Nonetheless, geographical variation at the MHC contrasts with relatively uniform distributions in the frequencies of microsatellite alleles. In addition, we found lower fixation rates in the MHC than those predicted by genetic drift after controlling for neutral mitochondrial sequences. Our results therefore underscore the role of balancing selection as well as spatial variations in parasite-mediated selection regimes in shaping MHC diversity when gene flow is limited.