Riparia Riparia Research Articles

The function and evolution of the tail streamer in hirundines

The morphology of a bird’s tail may result from compromises between aerodynamic efficiency, phylogenetic constraints and selection for non-aerodynamic characteristics, such as mate attraction. A good example of a trait shaped by trade-offs between aerodynamic efficiency and reproductive benefits mediated through female preference is the tail streamer of the barn swallow. Here we use a standardized task to measure the impact of manipulated tail streamer lengths on maneuvering flight in the barn swallow and in the sand martin, a closely related species that lacks a streamer. Our results show that the tail streamer of the barn swallow has a role in maneuvering flight. However, the outer tail feather is approximately 12 mm (9–20%) longer than the aerodynamic optimum for maneuvering flight. Furthermore, we show that the addition of artificial tail streamers to the sand martin, enhances maneuverability even at small streamer lengths, thereby implying that tail streamers may have evolved via natural selection for increased flight performance. Our results therefore suggest that initial tail streamer elongation in the barn swallow has a functional explanation in terms of increased aerodynamic performance. However, female choice has become associated with this trait, promoting the development of a costly handicap.

Sexy streamers? The role of natural and sexual selection in the evolution of hirundine tail streamers.

Matyjasiak et al. (2000) reported recently on experiments designed to mimic the early stages of tail streamer evolution in barn swallows (Hirundo rustic) by adding small streamers onto a closely related species that lacks them, the sand martin (Riparia riparia). This work was interpreted as providing evidence that there is a cost associated with the initial evolutionary stage of a tail streamer and that higher-quality individuals are better able to withstand this cost than lowerquality individuals. The authors suggest that these results give support to the hypothesis that tail streamers initially evolved as a handicap (Zahavi 1975), rather than as a result of Fisherian selection (Fisher 1958), to advertise male quality. We would like to raise a number of issues, which may offer an alternative interpretation.

Bird and butterfly communities associated with two types of urban riparian areas

We examined the distribution of birds and butterflies at two types of urban riparian areas in the metropolitan area of Denver, Colorado, USA. The sites consisted of relatively natural areas containing native woody vegetation, and those that had been highly modified with trees and shrubs removed and planted with lawn grass. Species richness of both butterflies and birds differed significantly between the two types of sites. Species richness, however, was not significantly correlated between birds and butterflies. Bird abundance was similar between site types but consisted of two assemblages. Brewer's blackbirds, common grackles, bank swallows, and mourning doves were abundant at wooded sites while American robins, European starlings, house sparrows, house finches, and mallards were more abundant at lawned riparian sites. Butterfly abundance was less at lawned sites. This contrast in bird and butterfly response to habitats suggests they respond to different habitats in distinct ways. Both taxon groups, however, were important discriminators between habitat types. Overall, butterfly and bird species richness was 44% less at highly modified riparian sites.

Interspecific variability of prevalence in blood parasites of adult passerine birds during the breeding season in Alaska.

Blood parasite prevalence based on microscopic examination of stained blood smears was determined in adults of 11 passerine bird species sampled during their breeding season (May and June 1997-98) in interior Alaska (USA). These species included primarily Nearctic migratory species such as the dark-eyed junco (Junco hyemalis) and neotropical migratory species such as the blackpoll warbler (Dendroica striata), alder flycatcher (Empidonax alnorum), Swainson's thrush (Catharus ustulatus), northern waterthrush (Seiurus noveboracensis), and bank swallow (Riparia riparia) as well as one long-distance palearctic migrant, the arctic warbler (Phylloscopus borealis). The more prevalent parasites were Leucocytozoon dubreuili (73% of the sampled turdinids), L. fringillinarum (42% of the sampled fringillids and parulids), and Trypanosoma avium (39% of the sampled hosts). Other parasites (H. fallisi: 18% of the sampled turdinids; Haemoproteus paruli: 14% of the sampled parulids; H. fringillae: 5% of the sampled fringillids; microfilariae: 4% of the sampled hosts) were observed less frequently. Plasmodium vaughani was found only in two yellow warblers (Dendroica petechia). Overall parasite prevalence varied between 0% in the alder flycatcher to >80% in Swainson's thrush, arctic warbler, and Townsend's warbler (Dendroica townsendi). Prevalence of various hematozoa also was bird species-dependent. No relationship was observed between prevalence and either foraging (aerial versus trees/shrubs) or nesting habits (ground versus arboreal) or general location of the wintering area of the different species examined. Prevalence also was unrelated to average dates of arrival on breeding grounds and, therefore, to potential duration of exposure to local insect vectors before capture. Differences in blood parasite prevalence among species breeding in a same region and in the same type of habitat may result from differences in host specificity such as immunological resistance to infection or blood meal preference by potential vectors and/or in behavioral adjustments/physiological traits that alter exposure to vectors.

Assessing the aerodynamic effects of tail elongations in the house martin ( Delichon urbica ): implications for the initial selection pressures in hirundines

Of the three species of hirundine that breed sympatrically across the U.K., one, the barn swallow, has outer tail feathers elongated into streamers, whereas the other two species, the house martin and the sand martin, do not. The tail streamer of the barn swallow is regarded as a classic example of a sexually selected trait. Recent evidence, however, has suggested that streamers may have evolved largely through natural selection for enhanced flight performance and increased maneuverability. We tested the hypotheses that small streamers (1) increase performance in turning flight, but (2) decrease performance in flight variables related to velocity. We manipulated the lengths of house martin outer tail feathers and measured changes in their free-flight performance, using stereo-video to reconstruct the birds" three-dimensional flight paths. Five flight variables were found to best describe individual variation in flight performance. Of these five, the three variables determining maneuverability predicted that flight performance would be optimized by a 6- to 10-mm increase in the length of the outer tail feathers. In contrast, for mean velocity and mean acceleration, extension of the outer tail feathers appears to have a detrimental effect on flight performance. We suggest that the initial selection pressure for streamers in ancestral short-tailed "barn swallows" was via natural selection for increased maneuverability. In addition, we propose that the benefits of increased maneuverability have differed between hirundines in the past, such that the cost of increasing the length of the outer tail feather has, to date, outweighed the benefits of doing so in streamerless hirundines.

Exposure to ectoparasites increases within-brood variability in size and body mass in the sand martin

Parasites often have detrimental effects on their hosts, and only host individuals able to cope with parasitism are likely to display induced or genetic resistance. Hosts may respond to parasitism by differential investment in offspring depending on their ability to cope with parasitism, because offspring that perform better than their siblings are themselves likely to have superior induced or genetic resistance. We tested whether nestlings of the highly colonial sand martin Riparia riparia were affected by the haematophagous tick Ixodes lividus by experimentally manipulating parasite loads of nests [nests sprayed with pyrethrum to remove parasites (sprayed), or nests sprayed with water (control)] at three stages of the breeding season. Prevalence and intensity of ticks were significantly affected by treatments. Breeding success was not significantly affected by treatment, although post-fledging survival was twice as high among nestlings from sprayed nests than from controls. Mean phenotypic traits of nestlings generally did not differ significantly among treatments, while within-brood variance in keel length (a skeletal character) and body mass were higher in control treatment broods than sprayed ones. Sedimentation rate, which reflects blood protein and immunoglobulin content, was significantly higher and less variable in sprayed than control broods. These findings are consistent with the suggestion that parasitism effects on host reproductive success act through an increase in the variance of offspring quality.

Examining the Cost of Experimentally Imitated Ornaments

Examining the Cost of Experimentally Imitated Ornaments

Imitating the initial evolutionary stage of a tail ornament.

Costs of a sexual ornament in its early evolutionary form and the relationship between these costs and individual condition may be an important influence in the likelihood of possible evolutionary mechanisms involved in the evolution of this ornament. We reconstructed the tail shape in hypothetical ancestors of recent hirundines (Aves: Hirundinidae), from which the elongation of tail feathers under sexual selection might have begun. By elongating the tail in sand martins (Riparia riparia, Hirundinidae), we simulated the early evolution of a long forked tail--the typical ornament of male hirundines. Birds with initial ornament captured smaller insects than controls, which suggests that this ornament imposed a cost in terms of impaired foraging. Furthermore, birds with naturally longer tails were better able to cope with initial ornament than naturally short-tailed birds. If length of tail in sand martins indicates the quality of individuals, our results suggest higher costs of this initial ornament for poorer than for higher quality individuals. We discuss the potential role of the handicap principle and other mechanisms in early evolution of a tail ornament.

IMITATING THE INITIAL EVOLUTIONARY STAGE OF A TAIL ORNAMENT

Costs of a sexual ornament in its early evolutionary form and the relationship between these costs and individual condition may be an important influence in the likelihood of possible evolutionary mechanisms involved in the evolution of this ornament. We reconstructed the tail shape in hypothetical ancestors of recent hirundines (Aves: Hirundinidae), from which the elongation of tail feathers under sexual selection might have begun. By elongating the tail in sand martins (Riparia riparia, Hirundinidae), we simulated the early evolution of a long forked tail—the typical ornament of male hirundines. Birds with initial ornament captured smaller insects than controls, which suggests that this ornament imposed a cost in terms of impaired foraging. Furthermore, birds with naturally longer tails were better able to cope with initial ornament than naturally short-tailed birds. If length of tail in sand martins indicates the quality of individuals, our results suggest higher costs of this initial ornament for poorer than for higher quality individuals. We discuss the potential role of the handicap principle and other mechanisms in early evolution of a tail ornament.Corresponding Editor: E. Martins

Activity patterns of insectivorous bats and birds in northern Scandinavia (69° N), during continuous midsummer daylight

Previous studies suggest that many species of insectivorous bats are nocturnal, despite the relatively low availability of their insect prey at night, because of the risk of predation by diurnal predatory birds. We hypothesised that if this was the case bats living above the arctic circle would alter their feeding behaviour during midsummer because there would no longer be any benefit to restricting their activity to the period when their prey are least abundant. Alternatively, if bats were more influenced by competition from aerial insectivorous birds they would continue to feed at ‘night’ to avoid such competition. In northern Norway (69° N), during continuous midsummer daylight, insectivorous sand martins (Riparia riparia) concentrated their aerial feeding activity when aerial insects were most abundant. The birds stopped feeding between 23:00 and 07:00 when aerial insects were least abundant. In contrast, northern bats (Eptesicus nilssonii), fed mostly between 22:00 and 02:00, coinciding with the lowest aerial insect availability, and with the period when light levels were lowest (ca 1000 lux). Bat activity patterns were closest to those predicted by the avian competition hypothesis. The low densities of both sand martins and Northern bats in the study area, however, were less consistent with this hypothesis. Possibly populations of both species were higher historically and the observed patterns reflected historical competition. Bat activity was most closely correlated to ambient light levels. This raised two alternative explanations that we could not eliminate. Perhaps there was differential predation risk, between the brightest and darkest parts of the day, because the visual capacities of falcons are strongly dependent on luminance. Alternatively the bats may have been entrained to emerge at given light levels by their behaviour at other times of year.

Foraging Cost of a Long Tail Ornament: An Experiment with Sand Martin Females

The handicap hypothesis assumes that sexual ornaments impose a viability cost upon the bearers. There have been few empirical tests of this assumption. Previous studies show evidence for the cost of a tail ornament in male birds: a negative relationship between an experimentally increased tail ornament (long tail streamers) and efficiency at foraging for nestlings. However, it must be admitted, that the apparent impairing effect of an elongated tail could be a result of a decrease in male parental effort in response to an increase of female parental effort, which might have occurred in response to increased male attractiveness (differential allocation of female parental effort). In this study, the effect of differential parental expenditure was eliminated by lengthening the tail in female, rather than male, sand martins (Riparia riparia). Tail‐elongated females decreased the rate at which they fed nestlings, and captured more but smaller insects. There was no simultaneous increase of feeding rate in the males that could explain the decrease of feeding rate in the females. These results confirm the existence of a cost of a tail ornament in birds feeding in flight, as is expressed in terms of impaired flight and foraging capacity.

Cost of parasitism and host immune defence in the sand martin Riparia riparia: a role for parent-offspring conflict?

Parent-offspring conflict may arise because the lifetime reproductive success of the parent is more influenced by its life span than by reproductive success during a particular reproductive event, while the fitness of an offspring depends firstly on its own survival as a juvenile and only subsequently on its own reproductive success. The naive immune system of young animals may allow offspring to be much more affected by parasites than their parents, and thus cause an initial asymmetry in a potential parent-offspring conflict. We investigated this type of conflict by assessing the health status and the immune response of parent and offspring sand martins Riparia riparia infested with manipulated loads of ticks Ixodes lividus (nests either treated with pyrethrin, water, or just visited). The prevalence and the intensity of tick infestations differed among treatments, with low tick loads in nests with the pyrethrin treatment. Ticks reduced the reproductive success of the host and increased offspring wing length. Broods with ticks had higher leukocyte concentrations and concentrations of immunoglobulins. The concentration of immunoglobulins in nestlings was negatively related to brood size and nestling tarsus length. Nestlings receiving the control treatments had a positive association between wing length and the concentration of immunoglobulins and a negative association between tarsus length and immunoglobulins. In contrast, adult sand martins did not respond to the parasite treatment in terms of immune response. Hence, the naive immune system of nestlings may be the crucial factor causing the parent-offspring conflict over costs of parasitism to be resolved to the advantage of parents that may sacrifice nestlings in heavily parasitized nests.

The Influence of Repeated Releases on the Homing Behaviour of Sand Martins (Riparia riparia)

Data on the effect of repeated releases on the homing behaviour of sand martins released four times from the same site (beyond 50 km from the nesting colony) are presented. The results, obtained by two series of releases, show a progressive improvement of both the initial orientation and the homing performances. When the birds released four times from the same site were displaced in a new site, located roughly at the same distance, but in the opposite direction to the former, they do not change directional preferences with respect to the last experimental release; the homing speeds were also not different to those recorded for birds without any release experience. While the learning of orienting cues can not be excluded, the data point out that the habituation to the handling stress and a form of directional training seem to play a major role in determining the observed behaviour. The familiarity with a site does not seem to improve the homing behaviour of sand martins when released from a different one.

A LONG TERM STUDY ON INITIAL ORIENTATION IN SAND MARTINS (RIPARIA RIPARIA)

Abstract Over a period of 18 years sand martins (Riparia riparia) were displaced from their breeding colonies and released to investigate the relationship between distance of displacement and initial orientation. The results show significant homeward orientation at the shortest distances of displacement (7.1-18.2 km, series A), but not over longer distances (49-180 km, series B and C). Birds displaced over 49 km from their breeding colonies only seldom show homeward directedness, while pseudo-pooling and second order analysis of their directional choices seem to indicate the tendency to fly in a given compass direction. Possible causal factors that can induce this behaviour in sand martins are discussed, taking into account not only local biasing factors but also the existence of a so called preferred compass direction. These data show that sand martins are able to determine the correct homing direction inside a probably familiar area. The response over longer distances indicates difficulties in determining the appropriate homing course at takeoff.

Brood parasitism in the sand martin,Riparia riparia: evidence for two parasitic strategies in a colonial passerine

Sand martin parentage was studied at a large breeding colony in central Scotland. Multilocus DNA fingerprinting was used during the 3-year study to exclude some social parents as true parents, and thereby determine the frequency of monogamy, quasiparasitism (QP), intraspecific brood parasitism (ISBP) and extrapair fertilization (EPF) amongst 45 broods and 167 nestlings. Monogamous parentage characterized the majority of broods (60%), so most chicks were the offspring of their social parents (81%). QP (involving a male’s extrapair mate laying in his nest) was found in 9% of broods and 2.4% of chicks and ISBP (or ‘egg dumping’) in 4% of broods and 1.8% of chicks. A substantial proportion of offspring arose from EPFs, affecting 36% of broods and 14% of chicks. On present evidence, the relatively high frequency of QP found in sand martins is unusual. We propose that its observed frequency is unlikely to be due to chance events and may represent a female-driven strategy.

Parent–offspring recognition in tree swallows,Tachycineta bicolor

Parent–offspring recognition appears to be highly developed in species in which the risk of misdirecting care is high (e.g. colonial species). Some of the best evidence for this relationship comes from comparative work on swallows of the family Hirundinidae. Using methods followed in earlier studies, we determined whether parent–offspring recognition occurs in the tree swallow,Tachycineta bicolora non-colonial species closely related to the highly colonial bank swallow,Riparia ripariaand the solitary rough-winged swallow,Stelgidopteryx ruficollisParents did not discriminate between playbacks of the calls of their own versus non-related nestlings. However, older nestlings called more in response to playback of parental calls than non-parental calls, suggesting that they recognized their own parents. Despite significant individual variation in parental and nestling calls, variation in tree swallow nestling calls was lower than analogous calls in the bank swallow. Our results provide further support for a positive relationship between recognition, individual variation in call structure and coloniality.

A Variant of the Hypergeometric Distribution for Large or Territorial Organisms

-The familiar Poisson distribution for the counts of cells occupied by 0, 1, 2 ... organisms over the N cells of a grid, is related to its binomial equivalent (q + p)'l in which p = 1/N, q = (N 1)/N, and the power n is the count of organisms summed across all N cells. This format of the binomial lends itself to modification as a hypergeometric variant where each cell has a maximum capacity M, such that p = M/MN and q = M(N 1)/MN initially, before successive reductions as conditional probabilities, and the power of the expansion remains n. Uses of this random generator are illustrated with a tree seedling problem, in which weak contagion is detected where both the Poisson and the binomial indicated acceptable randomness, and with a distribution of bank swallow tunnel entrances, in which significant uniformity is confirmed even after an allowance is made for the size of the tunnels and their supporting walls. Use of the hypergeometric variant in the bank swallow problem avoided apparent confirmation of uniformity as a statistical artifact. Chain calculation methods are presented which reduce the work of all three generators, especially that of the otherwise ponderously time-consuming hypergeometric.

Morphological Aspects of Avian Tail Movements: A Functional Approach in Hirundines

It has been proposed that the aerodynamically optimal tail shape in birds should be triangular when spread and forked when closed. According to this aerodynamic model, any area distal to the point of maximum width contributes drag, which is proportional to total tail area, but not lift. One assumption of this model seems to be uniformity in structures responsible for tail movements (i.e. tail muscles) among species. We tested for different muscular arrangements in species with different tail shapes. By dissecting tail muscles in three hirundine species (Barn Swallow [Hirundo rustica], Common House-Martin [Delichon urbica], and Bank Swallow [Riparia riparia]), we demonstrate anatomical differences associated with deeply forked tails and speculate that such differences compensate for the increased cost of longer feathers. Therefore, we propose that reliable assessment of the aerodynamic cost of tails should be based on tail shape as well as internal anatomical structures such as muscles.

Survival rates of Hungarian sand martins and their relationship with Sahel rainfall

Since 1986, I have carried out an intensive field survey of 10 000-30 000 pairs of a sand martin ( Riparia riparia ) population. Direct survey of the size and distribution of the breeding population and estimation of adult survival rates by SURGE, based on extensive ringing data sets, allow us to analyze the effects of different environmental factors with high precision. The model selection showed that the S s+t , P t model, in which the survival rates differ by sex for adults and vary in parallel by year and the capture rate varies by year, fits the data. The adult females had a lower survival rate compared to the males. The capture rate could be modelled as a quotient of the number of captured birds and the number of breeding birds along the upper part of the river Tisza. The survival rates of adults were related to the rainfall of the southern Sahel, which has an important role in the extension of the winter foraging habitat in the Sahel. Although the severe decrease in the population size, which may r...

An Experimental Test of Offspring Recognition in Western Bluebirds

An Experimental Test of Offspring Recognition in Western Bluebirds