Song-type sharing in song sparrows: implications for repertoire function and song learning

One hypothesis for the function of song repertoires is that males learn multiple song types so that they may share songs with neighbors, allowing them to match during territorial interactions. In at least one song sparrow population, in Washington, territorial males share a high proportion of song types with their neighbors and use these shared songs in matching. We recorded song sparrows in Pennsylvania and quantified sharing of whole songs and song segments. We found that song sharing is an order of magnitude less common in the Pennsylvania population. We found sharing of song segments to be significantly more common than the sharing of whole songs in three of the five fields we examined, while we found no significant differences between whole and partial song sharing in the remaining two fields. Finally, we found no evidence that sharing is greater between birds in the same field compared to birds in different fields. Taken with the data from Washington song sparrows, these results provide evidence for intraspecific geographic variation in the organization of song repertoires, and suggest that song sharing has not been a strong selective force in the evolution of song repertoires in song sparrows as a species. Furthermore, Washington and Pennsylvania song sparrows differ in how they learn song, in that Washington birds copy whole songs, while Pennsylvania birds appear to copy and recombine song segments, as has been found in laboratory studies of song learning. Thus both song learning and the function of song repertoires differ between populations of song sparrows. Such intraspecific geographic variation offers a unique opportunity to explore the ecological and historical factors which have influenced the evolution of song. Received: 30 June 1997 / Accepted after revision: 8 March 1998     

Temporal patterning of within-song type and between-song type variation in song repertoires

In many songbird species, males sing a repertoire of distinct song types. Song sparrows typically are described as having repertoires of about a dozen song types, but these song types are themselves quite variable and some songs are produced that appear intermediate between two types. In this study we quantify the similarity between successive songs produced by song sparrows in order to determine if differences between song types are emphasized or deemphasized in bouts of continuous singing. In spite of the high degree of variation within song types and similarity between song types observed in this species, we show that transitions from one song type to the next are distinct as compared to transitions within sequences of the same type (Figs. 4 and 5). Variation does not accumulate across sequences of the same song type, and the average variation observed within a continuous sequence of the same song type is significantly less than is predicted from the total variation recorded for that type across many different bouts (Fig. 6). These results support the view that song sparrows include two levels of variation in their singing: differences between song types as is commonly observed in species with song repertoires, and an independent level of variation observed for songs of the same type.     

An experimental study on the causal relationships between (ecto-)parasites, testosterone and sexual signalling

Bird song is considered to have evolved via sexual selection and should as such honestly signal aspects of the quality of its bearer. To ensure honesty, the immunocompetence handicap hypothesis proposed a dual role of testosterone, having positive effects on sexual signalling but suppressive effects on immune function. However, recent studies showed that it is rather an immune activation that suppresses the androgen production. This reversed chain of causation may significantly alter the pathways, which translate the effects of parasites and pathogens into changes in the expression of male sexual traits. We infested male canaries with Ixodes ricinus tick nymphs to investigate the causal relationships between (ecto-)parasites, testosterone and sexual signalling, here singing behaviour. We focused on flexible song traits, which may quickly reflect changes in the infestation status, and tested whether these effects relate to changes in the plasma testosterone levels or health state. The experimental tick infestation altered the males’ song performance by reducing song consistency, a trait that had previously been identified to reflect male quality. The tick infestation lowered the plasma testosterone levels and had a negative effect on the health status in terms of a reduced hematocrit. Our pathway analysis then revealed that it is the parasite-induced reduction of the plasma testosterone levels but not of the health state that caused the changes in song consistency. Thus, our study supports the view that it is the effect of parasites and immune activation on plasma testosterone levels that generates the trade-off between immunocompetence and sexual signalling.     

An experimental analysis of lunule function in the sand dollar Mellita quinquiesperforata

Coefficients of lift and drag for fifty lunulate sand dollars [Mellita quinquiesperforata (Leske 1778)], poised over a flat, sandy surface, were determined at seven different air stream velocities. Coefficients of static friction were obtained experimentally on level sand in standing water. Critical velocity, at which a sand dollar would be dislodged, is defined as that current velocity in which the force of drag is equal to the product of residual weight (i.e. weight-lift) and the coefficient of static friction. Small individuals have lower critical velocities than large ones: 26 cm s^?1 and 43 cm s^?1 for individuals 48 and 94 mm in diameter, respectively. Measurements were repeated for 20 of the controls, their lunules smoothly filled with modelling clay, and 20 from which lunule spines were removed. In every case, blocking the lunules decreased critical velocity, by an average of 15%; this was shown to be statistically significant (P>0.01). Removal of spines, which partially occlude the lunules, significantly (P>0.01) increased critical velocity by an average of 13%. Direct measurements in a water tunnel showed that calculated values of critical velocity were slightly too high, but otherwise confirmed the role of the lunules. Allometric analysis showed that as sand dollars increase in size, the distances of the lunules from the center of the disk increase less rapidly than the radii. A principal components' analysis of profile measurements of nine species indicated that lunulate and non-lunulate sand dollars are quite different in form. Lunulate species tend to be thinedged, flat-domed and larger in diameter. Some alternative ideas of lunule function are discussed and it is shown that there are no allometric or experimental data to support any of the suggested feeding hypotheses.     

Song sharing in two populations of song sparrows (Melospiza melodia)

Sharing song types with immediate neighbors is widespread in birds with song repertoires, and sharing songs may confer a selective advantage in some cases. Levels of song sharing vary between different geographical populations of several bird species, and ecological differences often correlate with differences in singing behavior; in particular, males in migratory subspecies often share fewer songs than males in resident subspecies. The song sparrow (Melospiza melodia) appears to fit this pattern: resident song sparrows in western North America generally share 20–40% of their repertoire (of about eight songs) with each neighbor, while migratory subspecies from eastern North America often share 10% or less. We compared song sharing in two populations within a single subspecies of song sparrow (M. m. morphna) in Washington State. These populations, separated by only 120 km, nonetheless differ in migratory tendencies and several other ecological and life history variables. We recorded complete song repertoires from 11 male song sparrows in a high-elevation, migrating population at Gold Creek in west-central Washington, and compared them to two samples (n = 15 and n = 36) from a coastal, resident population at Discovery Park, Seattle, Washington. Despite major differences in habitat, population density, and migratory tendencies, song sharing among Gold Creek males was as high as that among Discovery Park males. In both populations, sharing was highest between immediate neighbors, and declined with distance. We conclude that at the within-subspecies level, neither migration nor population density affect song sharing in song sparrows, a song repertoire species. Received: 26 November 1998 / Received in revised form: 1 May 1999 / Accepted: 29 May 1999     

An experimental study of paternal behavior in red-winged blackbirds

Summary The effect of brood size and female nesting status on male parental behavior was investigated in red-winged blackbirds Agelaius phoeniceus using brood size manipulation experiments. Male redwings allocated parental effort on the basis of brood size and nestling age. Males began assisting females only at nests with at least three offspring older than three days. Female nesting status had no singificant influence on male parental care. When females were unable to meet a brood's demand for food, males assisted females with nestling feeding. Females did not reduce the amount of food delivered to nestlings when males assisted. The amount of food brought to nestlings by the male was additional to the amount of food provided by the female. Male assistance increased fledgling success. When female provisioning was sufficient to meet a brood's demand for food males did not assist. The value of male parental care varied inversely with the ability of the female to meet nestling food demands. The ability of unassisted females to provide sufficient food and to raise a brood of nestlings successfully appeared to be influenced by resource abundance.     

Recognition of individual neighbors by song in the song sparrow,a species with song repertoires

Summary Previous theory and research have suggested that bird species with song repertoires in general, and song sparrows (Melospiza melodia) in particular, cannot readily discriminate between the songs of neighbors and strangers. In a recent study (Stoddard et al. 1991) we showed that song sparrows can in fact discriminate neighbors from strangers on the basis of song. In this study we sought to demonstrate that song sparrows can make the finer discrimination between individual neighbors and that they can do so on the basis of a single song type. We compared the response of territorial males to song playback of neighbors and strangers at three locations: the neighbor's regular boundary, the opposite boundary, and the center of the territory. The birds showed strong neighbor-stranger discrimination at the regular boundary but not at the opposite boundary, nor in the center of the territory. The differences in song discrimination between different boundary locations indicate that song sparrows associate particular songs with particular territories, effectively discriminating between individual neighbors on the basis of song. Song repertoires themselves do not interfere with neighbor recognition to the extent originally postulated. As speakers are moved inside the territory from the border, however, the degree of discrimination diminishes. We believe that differences in speaker placement may have contributed to the variability in neighbor-stranger discrimination observed in previous studies of the song sparrow and perhaps other repertoire species as well. This interpretation is consistent with data from another song sparrow population showing that half the territory takeovers are by immediate neighbors.     

An experimental and theoretical analysis of the effect of added weight on the energetics and hydrostatic function of the swimbladder of European sea bass (Dicentrarchus labrax)

We conducted experiments to determine the effect of the increasing ultrasonic/radio transmitter weight on the routine metabolic rate of sea bass. We measured the oxygen consumption (MO₂) of fish tagged externally with a dummy transmitter made of a hollow pipe, the weight of which was adjusted with lead to represent in water 0, 1 and 4% (R_tf) of the animal weight. We then developed a theoretical model to estimate, for a given fish size, the range of added weight that fish can compensate for through swimbladder regulation. When R_tfБ%, MO₂ of untagged and tagged fish did not differ significantly. However, when R_tf reached 4%, fish that carried a tag incurred a significant elevation of oxygen consumption, which represented 28% of their total useable power (or metabolic scope). This result strongly supports the view that a high R_tf ratio contributes to a decrease in available metabolic energy by diverting energy from, e.g., growth or swimming performance. A comparison between the tagged fish and the theoretical model reinforced the hypothesis that, when R_tf attained 4%, the increase in metabolic rate reflected a supplementary and costly swimming effort necessary to maintain vertical position. In this condition, the swimbladder cannot regulate the buoyancy of tagged fish.     

Sound degradation as a distance cue in great tit (Parus major) song

Summary We report an experiment designed to test the ideas that: 1. male songbirds can use cues from the distortion of song by environmental factors (degradation) to estimate the distance of another singing male; 2. song degradation is assessed by reference to an internal standard. Great tits respond more strongly to undegraded than to degraded songs when both are played at the same amplitude and from the same position in the territory. This difference in response is shown only if the playback song is familiar to the test bird; familiar songs being those sung either by the test bird or neighbours of the test bird. We interpret these results as evidence that cues from song degradation can be used to estimate the distance of a singing conspecific and that degradation assessment is only possible if the bird has an internal representation of the song (because either it and/or a neighbour sings the song). We discuss the implications of these results for Morton's (1982) ranging hypothesis, and for the distinction between learning and performance in bird song. Our results partially support the ranging hypothesis, but question the nature of unrangeable songs sensu Morton. The finding that birds can assess the degree of degradation of songs that they do not sing, supports the idea that birds learn more songs than they sing.     

An experimental study of the relationship between below-ground structure and meiofaunal taxa

An experimental study was conducted in a South Carolina, USA salt marsh to examine relationships between meiofaunal-sized tube structure and densities of burrowing meiofauna. Abundance of the tube-building polychaete Manayunkia aestuarina Bourne was increased by erecting exciosure areas from June–August 1979. Non-manipulated areas of identical dimensions were staked out in adjacent locales. Densities of burrowing meiofauna and nematode trophic groups, and volumes of Spartina alterniflora root mat and tube-builders were monitored approximately every 2 wk from each experimental treatment. Volumes of tube-builders were significantly higher inside exclosure sites on all dates in July and August. Root-mat volume and total number of nematodes were not different between treatments at any time. The dominant copepod species, Stenhelia (D.) bifidia, and the nematode species, Spirinia sp., first increased in number inside manipulated sites and then decreased in abundance as tube volume increased. No general trend was apparent for nematode trophic groups when areas with increased volumes of tubebuilders were compared to unmanipulated sites. The fact that densities of S (D.) bifidia and Spirina sp. decreased only when tube-builders attained highest abundance suggests that negative interactions between tube-builders and these meiofaunal taxa may be effective only at elevated abundances.     

Song overproduction and selective attrition lead to song sharing in the field sparrow (Spizella pusilla)

Summary Male field sparrows settling on particular territories for the first time often sing two or more song types. Vocal experience with neighboring males then leads in most cases to the deletion from the male's repertoires of all but one song type. In 2 years, settling males retained the song type that most closely resembled that of a neighbor. Focal observations in 1 year revealed that the most actively singing neighbor was the one the new birds resembled. Songs were not selected on the basis of their absolute acoustic characteristics, but rather by their resemblance to neighbors' songs. After the attrition process is complete, males retain their song for the rest of their lives. Evidence supports the hypothesis that the actual acquisition of a song need not take place at the time and place as males occupy territories. Rather, males may acquire an abundance of song models earlier in life and then select songs from this repertoire that most closely resemble those of their neighbors. The development of mature repertoires and of local dialects is the outcome of one process in this species.     

Developmental stress affects the attractiveness of male song and female choice in the zebra finch (Taeniopygia guttata)

Developmental stress has recently been shown to have adverse effects upon adult male song structure in birds, which may well act as an honest signal of male quality to discriminating females. However, it still remains to be shown if females can discriminate between the songs of stressed and non-stressed males. Here we use a novel experimental design using an active choice paradigm to investigate preferences in captive female zebra finches (Taeniopygia guttata). Nine females were exposed to ten pairs of songs by previously stressed and non-stressed birds that had learned their song from the same tutor. Song pairs differed significantly in terms of song complexity, with songs of stressed males exhibiting lower numbers of syllables and fewer different syllables in a phrase. Song rate and peak frequency did not differ between stressed and non-stressed males. Females showed a significant preference for non-stressed songs in terms of directed perching activity and time spent on perches. Our results therefore indicate that developmental stress affects not only the structure of male song, but that such structural differences are biologically relevant to female mate choice decisions.     

An osteological study of the dugong Dugong dugon (Sirenia) from India

An osteological study of the dugong (sea cow) Dugong dugon (Müller) from India, based on complete skeletons of an adult male and a juvenile female indicated certain morphological variations with age. It has also revealed a close osteological similarity between the dugongs from India and the Red Sea. Based on a statistical analysis, regression equations for certain typical skull measurements have been calculated; these equations, characteristic of the dugong from India, could be used for comparing dugongs of different regions.     

An experimental study on recolonization and succession of marine macrobenthos in defaunated sediment

Hypoxia/anoxia in coastal waters is a world wide problem which often results in mass mortality and defaunation of benthos. In this study, field experiments were carried out to examine recolonization and succession of macrobenthic infauna in defaunated sediments, and the time required for recovery from complete defaunation to a stable community. Trays (33 cm length × 25.5 cm width × 11 cm depth) of defaunated sediment were exposed at the subtidal of a pristine site in subtropical Hong Kong. Temporal changes of macrobenthic communities in defaunated sediment were analyzed by univariate and multivariate statistics, and compared with those in undisturbed natural sediment at the same site. Initial colonization of macrobenthos occurred rapidly. A total of 42 species was found, with an average of 258 animals per tray and 24 species per tray recorded in the first month. Abundance showed a small peak (496 animals per tray) after 3 months, reached a sharp peak (1154 animals per tray) after 6 months, and declined thereafter. Species number increased gradually, reached a maximum (68 species per tray) after 9 months, and then decreased. Recolonization was predominantly contributed by larval settlement rather than adult migration. Temporal changes in abundance, species number and diversity of the macrobenthic community in defaunated sediment resemble the spatial changes along a decreasing pollution gradient previously defined by other authors. Results of this experiment suggest that newly available sediment may allow more species to colonize (or coexist) than sediment pre-occupied by an established community. This is probably due to less interspecific competition in the former habitat. No significant difference in abundance or species richness was observed between defaunated and natural sediments after 15 months, suggesting that a stable community had been achieved, although minor variations in species composition were still discernible between defaunated and natural sediments. Received: 24 May 1999 / Accepted: 20 October 1999     

Effects of copper on the faunas of marine soft-sediments: An experimental field study

Although laboratory experimental studies have shown that copper is toxic to marine organisms at concentrations found in contaminated sediments, there is little unequivocal evidence of undesirable ecological effects in the field, other than at extreme concentrations. We describe a study in Botany Bay, New South Wales, Australia, in which the concentrations of copper in marine sediments were experimentally enhanced. Changes in the abundance and taxonomic composition of the fauna of copper-treated sediments relative to those of two control treatments were monitored over a period of six months. Univariate (ANOVA) and multivariate (non-metric multidimensional scaling, MDS) analyses of the changes in the fauna showed that increased concentrations of copper (140 to 1200 g g^-1 compared with background concentrations of 29 to 40 g g^-1) had an impact on the fauna. The nature of the response varied among taxa. For example, in some taxa, numbers of individuals decreased through time relative to controls, whereas the abundance of another taxon remained fairly constant through time in the copper treatment while numbers of control individuals increased. Differences in the changes of the faunas through time among the control and copper treatments were not always consistent among replicate experimental units 5 m apart, nor were they consistent between replicate experimental sites 100 m apart. The magnitudes of the changes in the faunas caused by the copper treatment are considered in the context of the magnitude of previously measured natural temporal variation.     

Ranging of songs with the song type on use of different cues in Carolina wrens: effects of familiarity

Male territorial song birds are usually spaced far apart and most often hear conspecific song after it has been degraded by propagation through the environment. Their ability to use the degradation of songs to assess the distance of a singing rival without approaching (called ranging) presumably increases the efficiency of defending a territory. In order to assess degradation in a song the receiver needs to compare the characteristics of the received song to its characteristics at the source or at different distances. Earlier experiments on ranging in species with song repertoires have suggested that prior familiarity with the particular song type is necessary for ranging. Here I show that male Carolina wrens (Thryothorus ludovicianus) can use either temporal or spectral characteristics for ranging song types which they were unlikely to have heard previously. Playbacks consisting of only one song prevented subjects' close-range experience with the loudspeaker, and flights beyond the loudspeaker provided direct evidence for over-assessment of distance when songs were degraded. Because ranging of songs was not affected by the degree of familiarity with the song type, this experiment provides no evidence that song repertoires hinder ranging in Carolina wrens, as suggested by Morton's ranging hypothesis. Instead, at least approximate ranging of songs is evidently possible by assessment of degradation in general features of a species' songs. Received: 9 August 1996 / Accepted after revision: 15 February 1997     

An ultrastructural study of Crassostrea angulata (Mollusca,Bivalvia) spermatogenesis

The cytoplasm of Crassostrea angulata (collected from the Atlantic coast of Northern Spain, Galiza, between 1992 and 1993) spermatogonia contains a few proacrosomal dense vesicles and intermitochondrial round granulo-fibrillar dense bodies. In primary spermatocytes, large cytoplasmic spaces appear occupied by large round and more scattered granulo-fibrillar dense bodies which are intermingled with cord-like dense structures and are in close association with mitochondria and the rough endoplasmic reticulum. As a result of Golgian activity, the number of dense proacrosomal vesicles increases and a flagellum arises from the centrioles. During the second meiotic division, several round dense bodies surrounding cord-like dense structures as well as dense proacrosomal vesicles, were observed between chromosomes. During spermiogenesis, no Golgian activity was noticed, and only the rough endoplasmic reticulum was observed in close association with proacrosomal vesicles. Similar results were observed for Ostrea edulis (collected from the Atlantic coast of southern Portugal, Algarve, between 1992 and 1993). Spermatozoa of both species are of the aquasperm type, and although their ultrastructure is quite similar, silver and phosphotungstic acid-chromic acid stainings enabled us to clearly distinguish both species at the ultrastructural level. The different morphological aspects of spermatogenesis in the Pteriomorphia and the use of silver and PTA stainings in the Mollusca are reviewed and discussed.     

Effects of different levels of song overlapping on singing behaviour in male territorial nightingales (Luscinia megarhynchos)

In signalling interactions, animals can directly address information to a specific individual. Vocal overlapping is such a signalling strategy used in songbirds, anurans, and insects. In songbirds, numerous studies using high rates of song overlap to simulate an escalating situation have shown that song overlapping is perceived as a threatening signal by interacting and by listening (eavesdropping) individuals, indicating a high social relevance of song overlapping. Here we present a playback experiment on nocturnally singing male territorial nightingales (Luscinia megarhynchos). Using three different rates of song overlap (1, 25, or 50%), we tested whether or not lower levels of song overlapping act as a signal of aggressive intent and if birds would increase the intensity of their response with increasing level of song overlapping. Subjects did not vary song duration in response to the different playback treatments but increasingly interrupted their singing with increasing overlap by the three playback treatments. The effects persisted even after the playback ceased to overlap and switched to an alternating singing mode. These results expand on previous studies by showing that song overlapping is interpreted as an aggressive signal even when it is used at low or moderate levels. They suggest that, within the range tested here, increasing levels of song overlapping are perceived to be increasingly aggressive.     

Short- and long-term consequences of prenatal testosterone for immune function: an experimental study in the zebra finch

Hormone-mediated maternal effects play an important role in the formation of a differentiated phenotype. They have been shown to influence a wide array of offspring traits, both early in life and in adulthood. One important offspring trait that is under the influence of maternal androgens is the immune system. In birds, a growing number of studies show that yolk androgens modulate immune function during the chick stage. However, there is a lack of knowledge regarding long-term effects of prenatal androgens on offspring immunity. In this study, we therefore investigated the influence of prenatal testosterone (T) on several measures of immunity in fledgling and adult zebra finches (Taeniopygia guttata). Cell-mediated immune response (towards phytohaemagglutinin, PHA) of fledglings hatching from control eggs was negatively related to brood size, whereas there was no such association for fledglings hatching from eggs with experimentally elevated T levels (T fledglings). Male control fledglings showed reduced mass gain compared to female control fledglings within 24 h after the PHA injection. This pattern was reversed in T fledglings. Total antibody levels in fledglings were not affected by egg treatment. Neither cell-mediated immunity nor total antibody levels in sexually mature zebra finches were influenced by egg treatment. However, there was an immuno-enhancing effect of elevated egg T on both primary and secondary humoral immune responses toward diphtheria and tetanus antigens in ca 5 and 7 month old zebra finches. In addition, the covariation between different immune components differed between T and control offspring, suggesting that egg treatment may have altered the potential trade-offs between different parts of the immune system. Our results suggest that prenatal androgens could be an important factor contributing to individual variation in immune function even in adulthood.