Experimental evidence for the influence of food availability on incubation attendance and hatching asynchrony in the Australian reed warbler Acrocephalus australis

The amount of time a bird allocates to incubation is likely to be limited by energetic constraints. If food is abundant, energetic constraints may be reduced and the time spent incubating (incubation attendance) may increase. Moreover, the onset of incubation in relation to clutch completion may be advanced, resulting in a higher degree of hatching asynchrony. We measured the effect of experimentally increased food availability on incubation attendance and an estimate of hatching asynchrony in the Australian reed warbler Acrocephalus australis . Supplementary food was provided every other day, from a few days before the start of egg laying until just prior to hatching. Incubation attendance was measured with temperature loggers at nests receiving supplementary food and control nests. Hatching asynchrony was inferred from mass and size differences between siblings shortly after hatching. We found that 1) food supplementation resulted in an increase in incubation attendance, when comparing both nests receiving supplementary food to control nests as well as feeding to non-feeding days in nests receiving supplementary food, and 2) food supplementation resulted in a greater hatching asynchrony, without affecting clutch size, average egg volume or the likelihood of eggs hatching. This suggests that food availability acts in a proximate way to modify the extent of incubation attendance and hatching asynchrony. We discuss the adaptive significance of increased incubation attendance and a shift in the degree of hatching asynchrony in relation to food availability.     

Egg discrimination in the Australian reed warbler (Acrocephalus australis): rejection response toward model and conspecific eggs depending on timing and mode of artificial parasitism

In a coevolutionary arms race between an interspecific broodparasite and its host species, both are expected to evolveadaptations and counteradaptations. We studied egg discriminationin the Australian warbler (Acrocephalus australis). This speciesis currently not significantly parasitized by the seven speciesof cuckoo for which it is a suitable host. However, experimentalbrood parasitism in the warbler revealed a fine tuned egg discriminationresponse towards non-mimetic and conspecific eggs, the firstsuch evidence in an Australian passerine: (1) non-mimetic eggswere significantly more often rejected than conspecific eggs;(2) only non-mimetic dummy eggs were rejected selectively,whereas rejection of conspecific eggs entailed a rejectioncost; (3) replacement of a host's egg with a conspecific eggduring egg laying resulted in a significantly higher rejectionrate than after the day of clutch completion; (4) by contrast,rejection rate after addition of a conspecific egg was independentof nest stage; (5) conspecific eggs introduced into a clutchduring the egg laying period led to a significantly highernest desertion rate and a lower egg ejection rate than afterthe day of clutch completion; and (6) addition of a conspecificegg led to egg ejection while egg replacement with a conspecificegg led to nest desertion. The fact that this species respondsdifferentially toward different modes of artificial parasitismsuggests that its egg discrimination has evolved to minimizethe costs of rejection and parasitism. The ability to rejecthighly mimetic conspecific eggs may explain the current paucityof brood parasitism in this species. The significance of thisfor brood parasite-host coevolution is discussed.     

An analysis of hatching success in the great reed warbler Acrocephalus arundinaceus

Hatching success is a potentially important fitness component for avian species. Previous studies of hatching success in natural populations have primarily focused on effects of inbreeding but a general understanding of variation in hatching success is lacking. We analyse data on hatching success in a population of great reed warblers Acrocephalus arundinaceus in Lake Kvismaren in south central Sweden. The effects of a range of covariates, including three measures of inbreeding as well as effects of classifications in the data (such as identities of individuals), on hatching success are analysed simultaneously. This is done by means of fitting Bayesian binomial mixed models using Markov chain Monte Carlo methods. Using random effects for each individual parent we check for unexplained variation in hatching success among male and female individuals and compare it to effects of covariates such as degree of inbreeding. Model selection showed that there was a significant amount of unexplained variation in hatching probability between females. This was manifested by a few females laying eggs with a substantially lower hatching success than the majority of the females. The deviations were of the same order of magnitude as the significant effect of parent relatedness on hatching success. Whereas the negative effect of parent relatedness on hatchability is an expression of inbreeding, the female individual effect is not due to inbreeding and could reflect maternal effects, that females differ in fertilisation and/or incubation ability, or an over representation of genetic components from the female acting on the early developing embryo.     

Lost and found: the enigmatic large-billed reed warbler Acrocephalus orinus rediscovered after 139 years

We present compelling evidence of the continued existence of the large-billed reed warbler Acrocephalus orinus , hitherto known only from the unique type specimen collected in NW India 139 years ago. Morphological and genetic analyses of an unusual Acrocephalus warbler mist-netted south-west of Bangkok, Thailand, on 27 March 2006, confirmed its identity as A. orinus , and revealed that it was heterozygous at four out of eight microsatellite markers, indicating the continued existence of a viable population whose breeding and wintering areas are still unknown.     

An exceptionally high diversity of hoverflies (Syrphidae) in the food of the reed warbler (Acrocephalus scirpaceus)

Despite being considered a classical example of protective Batesian mimicry hoverflies (Syrphidae) are known to be preyed upon by various passerines. The aim of the present study was to examine in detail food brought by reed warblers Acrocephalus scirpaceus to their nests to better understand the importance of hoverflies in the diet of small passerines. Using neck collars, 273 food samples containing 8,545 food items delivered to reed warbler and parasitic common cuckoo Cuculus canorus nestlings in warbler nests were recorded. The study was conducted during three breeding seasons in South Moravia, Czech Republic. An unusually high diversity of hoverflies was found — 27 species, including Mesembrius peregrinus (critically endangered species in the Czech Republic) and Mallota cimbiciformis (endangered species) — a new taxon to the Czech Republic. This indicates that nestling diet analyses may provide not only information on avian foraging behaviour but also important faunistic data. Thus, without the detailed identification to species level of material from foraging behaviour studies valuable scientific information may be lost. Overall dominance of Syrphidae was 3.7%, the most common species being Episyrphus balteatus (55.7%, n = 318). However, this number seriously underestimates the importance of hoverflies in the diet of reed warblers as hoverflies are one of the largest prey taken by warblers. Both larvae and pupae were rare, imagines strongly dominating (92.7%). Both specific wasp mimics (e.g., Chrysotoxum verrali) and bee mimics (e.g., Eristalis spp.) were not avoided by foraging reed warblers. The presence of a parasitic cuckoo chick did not affect host foraging behaviour with respect to overall dominance of hoverflies in the diet (warbler 3.3%, cuckoo 3.8%).     

Female choice in the sedge warbler Acrocephalus schoenobaenus: multiple cues from song and territory quality

Recent models of animal signalling emphasize the evolution of complex displays containing ''multiple messages''. A variety of potential cues used in female choice were investigated during a three-year field study of the sedge warbler, Acrocephalus schoenobaenus. Twelve possible cues were investigated, and three were found to have a significant influence upon pairing date. Two were different measures of song (repertoire size and song flighting) and one a measure of territory (territory size). Repertoire and territory size had a significant influence on pairing date in all three years, and song-flighting in two. The three cues were not intercorrelated and so had independent effects upon pairing date. We suggest that females select males upon multiple cues as these reflect different aspects of male and territory quality.     

Low frequency of extrapair paternity in the polygynous great reed warbler, Acrocephalus arundinaceus

We carried out DNA fingerprinting on 553 young (130 broods)great reed warblers (Acrocephalus arundnaceus) in 1987–1991.In the study population, where 40% of the males become polygynous,there was a low frequency of extrapair fertilizations (EPF).When data from all five years were pooled, 3.1% of the youngwere sired by extrapair males (EPF-males) and 5.4% of the broodscontained extrapair young. We found no cases of extrapair maternity;young with 6–17 mismatched DNA bands (n= 17) had highband sharing with their putative mothers (range = 0.52–0.72)but low band sharing with their putative fathers (range = 0.24–0.40).In broods exposed to EPF, on average 53% of the young were siredby EPF-males. We found the genetic father to each of the illegitimateyoung. In all cases the same EPF-male sired all extrapair youngin a brood. Broods containing EPF-young tended to be initiatedlate during the breeding season. Breeding attempts were ratherevenly distributed over two months, thus this breeding asynchronywould have facilitated EPFs. There was no difference in EPFfrequency between broods where the pair males had left theirfemales unguarded during parts of their fertile periods andbroods where males guarded throughout the fertile periods. Nestswith extrapair young had significantly shorter mean distanceto the closest male neighbor and more male neighbors within100 m than nests without extrapair young. We found no indicationthat females engaged in EPF to get parental care from the EPF-males,or because they were forced to copulate with extrapair males.The low frequency of EPF suggested that females did not seekgenetic diversity to their brood. We cannot rule out the possibilitythat females engaged in EPF to insure fertility. However, datasupporting this hypothesis were weak. Instead, our data supportthe conclusion that females engaged in EPF to increase the geneticquality of their offspring, and that females may have used malesong repertoire size as a cue when choosing EPF partners.     

Discovery of a breeding area of the enigmatic large‐billed reed warbler Acrocephalus orinus

The present study reports the discovery of the hitherto unknown breeding grounds of the large‐billed reed warbler Acrocephalus orinus in the Badakhshan region of Tajikistan. A total of eight adult individuals were caught at three riverine woodland sites and breeding was confirmed at one locality. The identity of the birds was confirmed on the basis of partial sequences of the cytochrome b gene. Even among individuals sampled at the same locality, genetic diversity was surprisingly high. This was interpreted as the result of secondary contact between previously isolated or fragmented populations. The breeding range of the large‐billed reed warbler is likely small and the species’ habitat is under intense pressure from the collection of firewood and fodder. Consequently, the large‐billed reed warbler remains of considerable conservation concern.     

Influences of wind flow on stopover decisions: the case of the reed warbler Acrocephalus scirpaceus in the Western Mediterranean

Wind directions measured at two different heights (850 hPa and 700 hPa) and at different hours of the night were analysed during the spring migration passage at a bird stopover site located in the western Mediterranean, in order to evaluate the importance of wind components for a stopover decision. From a huge ringing campaign of bird migration in north-east Spain, data from the reed warbler Acrocephalus scirpaceus have been used for the analysis. From a total of 2,478 reed warblers captured between 1993 and 1997 data recording significant days, with a high number of captures, and decrease days, with few captures, have been selected to develop an analysis of wind direction in relation to stopover and flight resumption. On days with a high capture the winds had mainly a fourth-quadrant flow (from the north, north-west and west), these being mainly head winds. Winds with westerly component (from the north-west, west and south-west), which enhance the flight, account for the majority of the days when there was a low capture of reed warblers. Wind direction therefore appears to be a determining factor for stopover decisions and resumption of flight for the reed warblers at an intermediate stage of their spring migration where topographical characteristics govern the winds.     

Dynamics of parasitemia of malaria parasites in a naturally and experimentally infected migratory songbird, the great reed warbler Acrocephalus arundinaceus

Little is known about the development of infection of malaria parasites of the genus Plasmodium in wild birds. We used qPCR, targeting specific mitochondrial lineages of Plasmodium ashfordi (GRW2) and Plasmodium relictum (GRW4), to monitor changes in intensities of parasitemia in captive great reed warblers Acrocephalus arundinaceus from summer to spring. The study involved both naturally infected adults and experimentally infected juveniles. The experiment demonstrated that P. ashfordi and P. relictum lineages differ substantially in several life-history traits (e.g. prepatent period and dynamics of parasitemia) and that individual hosts show substantial differences in responses to these infections. The intensity of parasitemia of lineages in mixed infections co-varied positively, suggesting a control mechanism by the host that is general across the parasite lineages. The intensity of parasitemia for individual hosts was highly repeatable suggesting variation between the host individuals in their genetic or acquired control of the infections. In future studies, care must be taken to avoid mixed infections in wild caught donors, and when possible use mosquitoes for the experiments as inoculation of infectious blood ignores important initial stages of the contact between the bird and the parasite.     

Within‐ and between‐season repeatability of eggshell colouration in the great reed warbler Acrocephalus arundinaceus

While avian eggshell colouration has attracted biologists for decades, little is known about its variation within individuals. The main goal of this study was to explore within‐ and between‐season repeatability of eggshell appearance in the great reed warbler Acrocephalus arundinaceus. To achieve this, we measured eggshell reflectance of first and replacement clutches of individual females within one breeding season, and the reflectance of their first clutches in two following breeding seasons. As environmental conditions may affect egg colouration, repeatability was estimated from linear mixed‐effects models where we initially included temperature and rainfall during egg laying, as well as year, clutch order and egg‐laying date as fixed effects, and clutch identity nested in females as random effects. Eggshell appearance within clutches showed moderate repeatability, while both within‐ and between‐season repeatability in clutch colouration for individual females was low. Our findings indicate that low intra‐clutch variation may have a function within the context of brood parasitism, i.e. in facilitating host recognition of alien eggs. With variable eggs between successive clutches, however, host females may need to relearn the appearance of their eggs with every clutch they lay. This could represent a significant constraint for host egg‐discrimination abilities. Yet, whether environmental or intrinsic physiological factors are responsible for the variation in eggshell colouration between successive clutches of the same females still remain to be discovered.     

Macrogeographical variation in the song of a widely distributed African warbler

The songs of oscine passerine birds vary on many spatial scales, reflecting the actions of diverse evolutionary pressures. Here we examine the songs of Cisticola erythrops, which effectively signal species identity across a geographical area spanning 6500 km in sub-Saharan Africa. Selection for species identification should promote stability in song traits, while sexual selection and geographical segregation should promote diversity. Cisticola erythrops share syllable types across the entire range of species and structure songs similarly, but individuals sing highly variable songs through improvisational recombination of syllables. Patterns of syllable use change gradually across the range of the species and do not show distinct breaks at subspecies boundaries. The acoustic properties of the most common syllable type also change gradually with distance. The results illustrate how songs can be simultaneously species-specific and highly variable at an individual level. At a larger level, patterns of variation indicate that cultural drift has generated song diversity through an isolation by distance mechanism.     

Factors influencing the appearance of spider (Araneae) and beetle (Coleoptera) assemblages in nests of great reed warbler Acrocephalus arundinaceus

The appearance of spider (Araneae) and beetle (Coleoptera) assemblages found in nests of great reed warbler Acrocephalus arundinaceus was studied, firstly to investigate breeding success and the amount of precipitation as potential factors which might affect the abundance and species richness of both groups. In addition, we compared the diversity of spider and beetle assemblages between nests found in different reed habitats, and considered the position of nests (above water or dry ground). In this study we selected five different randomly chosen reed habitats: two mining ponds, two small canals and one large canal. Great Reed Warbler nests were collected either shortly after fledging, or after the clutch had failed. Altogether, 12 species of spider and 19 species of beetle were collected. In both groups there was no significant difference in abundance between successful, lost and cuckoo-parasitized nests; however, there was a significant difference in species richness between the three nest categories in spider assemblages, which was not the case in beetle assemblages. The amount of precipitation did not affect beetle or spider abundance; only the species richness of spiders showed significant growth. Furthermore, we found no significant relationship between vegetation cover and the species richness and abundance of spiders and beetles. The diversity of both groups differed significantly according to reed habitat: beetle assemblages were most diverse by the large canal and spiders at the mining ponds.     

Transgenerational effects of food availability on age at maturity and reproductive output in an asexual collembolan species

Transgenerational effects of environmental conditions can have several important ecological and evolutionary implications. We conducted a fully factorial experiment manipulating food availability across three generations in the collembolan Folsomia candida, a springtail species that inhabits soil and leaf litter environments which vary in resource availability. Maternal and grandmaternal food availability influenced age at maturity and reproductive output. These effects appear to be cumulative rather than adaptive transgenerational life-history adjustments. Such cumulative effects can profoundly influence eco-evolutionary dynamics in both stable and fluctuating environments.     

A comparative study of territory in the Reed warbler (Acrocephalus scirpaceus) and Sedge warbler (A. schoenobaenus)

Various aspects of territoriality were studied comparatively in a marked population of Reed and Sedge warblers at the Attenborough Nature Reserve, Nottinghamshire. Differences in arrival patterns, establishment of territory and particularly territory size were detected. In view of the dynamic nature of territory boundaries, a method is described for obtaining measures of mean territory size for comparative purposes. The sympatric species were largely separated by different breeding and feeding habitats, but there was also a degree of interspecific competition in some areas and interspecific territorialism was found to occur. Adults of both these migratory species were found to exhibit strong site fidelity, and there was some evidence to suggest that early post-fledging experience may influence initial selection of a particular area and territory for first breeding.
The adaptive significance of territory is discussed in relation to the findings presented and current hypotheses. There was some evidence which suggests that reduction of predation pressure and intraspecific interference through spacing out, as well as provision of a nest-site may be important functions of territory in these species, but no evidence that food supply or population regulation are involved. The significance of interspecific territorialism is considered in relation to interspecific competition. The various factors effecting territory size and the patterns of nesting dispersion are also considered in an attempt to account for the striking interspecific differences found between these closely related species. It is concluded that although a multiplicity of factors may well be interacting in complex fashion, the more important of these stem from contrasts in the physical properties of the two habitats.     

Establishment and maintenance of the breeding territory in the polygynous great reed warbler

Settlement of male great reed warblers in the breeding ground was highly asynchronous at Lake Biwa in Japan. It took over 1 month from the appearance of the first male to a saturation in number of males. In resource-defense-polygyny, males are expected to try to defend as large an area as possible in the optimal habitat. In fact, a small number of the earliest settling males divided up the breeding ground almost completely as territories among themselves and these were later reduced in size with the addition of later settlers. The reduction was not due to a seasonal decline of aggressiveness on the part of the owners but to a higher level of intrusion pressure by later arriving males. The neighbor-neighbor relationship, once established, was rather stable. Home range overlap was small and territorial contacts were few between neighbors. Territorial boundaries seem not to shift despite the addition of new males as long as the neighbors were the same. The stable relationships between neighbors are expected to help the earlier resident to maintain his large and good territories throughout the breeding season and hence polygyny is favored in this species.     

Spatio-temporal variation in territory quality and oxidative status: a natural experiment in the Seychelles warbler (Acrocephalus sechellensis)

1. Fluctuations in the quality of the habitat in which an animal lives can have major consequences for its behaviour and physiological state. In poor-quality habitat with low food availability, metabolically intensive foraging activity is likely to result in increased generation of reactive oxygen species, while scarcity of food can lead to a weakening of exogenously derived antioxidant defences. The consequent oxidant/antioxidant imbalance may lead to elevated oxidative stress. 2. Although the link between food availability and oxidative stress has been studied in the laboratory, very little is known about this relationship in the wild. Here, we investigate the association between territory quality (measured through food availability) and oxidative stress in the Seychelles warbler (Acrocephalus sechellensis). 3. Seychelles warblers are insectivorous birds that inhabit a fixed feeding territory year round. Individuals experience profound and rapid local fluctuations in territory quality within these territories, owing to changing patterns of vegetation defoliation resulting from seasonal changes in prevailing wind direction and wind-borne salt spray. 4. As expected, oxidant generation (measured as reactive oxygen metabolites; ROMs) was higher when territory quality was low, but there was no correlation between territory quality and antioxidant capacity (OXY). The negative correlation between territory quality and ROMs was significant between individuals and approached significance within individuals, indicating that the pattern resulted from individual responses to environmental variation. 5. ROMs and OXY levels within individuals were positively correlated, but the relationship between territory quality and ROMs persisted after including OXY as a covariate, implying that oxidative stress occurs in low territory quality conditions. 6. Our results indicate that the oxidative stress balance of an individual is sensitive to relatively short-term changes in territory quality, which may have consequences for the birds' fitness.     

Responses of great reed warblers Acrocephalus arundinaceus to experimental brood parasitism: the effects of a cuckoo Cuculus canorus dummy and egg mimicry

Egg rejection behaviour towards parasitic eggs was studied in a great reed warbler Acrocephalus arundinaceus population in central Hungary, which was heavily (about 65%) parasitised by the common cuckoo Cuculus canorus . Clutches were experimentally parasitised during the egg-laying period with artificial, moderately mimetic cuckoo eggs or with conspecific eggs that were good mimics of the hosts' eggs. Great reed warblers rejected 76.2% of the artificial cuckoo eggs, mainly by ejection, but accepted most of the conspecific eggs (87.5%). Cuckoo eggs in naturally parasitised clutches were rejected at a lower rate (32%). When, in addition to the egg mimicry experiments, a stuffed cuckoo was placed near the nest, accompanied by the recording of a female cuckoo call, hosts' rejection rate of the artificial cuckoo egg increased from 76% to 96%. The sight of the cuckoo, on the other hand, did not influence host's rejection behaviour when a conspecific egg was used in the experiment. A stuffed collared dove Streptopelia decaocto , accompanied by its call, was used as a control, and did not cause any increased rejection. Great reed warblers were more aggressive towards the cuckoo than to the dove dummy. When the cuckoo eggs in naturally parasitised clutches were exchanged with artificial cuckoo eggs, we observed no increase in the rejection rate. We conclude that great reed warblers in our heavily parasitised population are capable of detecting brood parasitism in their clutch by identifying the parasitic egg. The efficiency of this identification depends mainly on the mimicry of the foreign egg. The sight of the cuckoo at the nest may increase rejection rate by stimulus summation, and this conditional effect is mainly affected by the degree of mimicry of the parasitic egg.     

Nestling growth and song repertoire size in great reed warblers: evidence for song learning as an indicator mechanism in mate choice

Females of many songbird species show a preference for mating with males that have larger song repertoires, but the advantages associated with this preference are uncertain. We tested the hypothesis that song complexity can serve as an indicator of male quality because the development of the brain regions underlying song learning and production occurs when young birds typically face nutritional and other stresses, so that song reflects how well a male fared during post-hatch development. A key prediction of this hypothesis is that variation in nestling condition should correspond to variation in the adult song repertoires of individuals. We used data from a long-term study of the great reed warbler (Acrocephalus arundinaceus) to test this prediction, correlating two measures of nestling development with subsequent repertoire size of males. We found that the length of the innermost primary feather, a standard measure of development, significantly predicted first-year repertoire size. The relationship between repertoire size and body mass was nearly significant, in spite of the large variance inherent in this measure. These data support the idea that song may provide females with information about a male's response to developmental stress, which in turn is expected to correlate with indirect or direct benefits she might receive.