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.     

Discovery of ten new specimens of large‐billed reed warbler Acrocephalus orinus,and new insights into its distributional range

We here report the finding of ten new specimens of the poorly known large‐billed reed warbler Acrocephalus orinus. Preliminary identifications were made on the basis of bill, tarsus and claw measurements, and their specific identity was then confirmed by comparison of partial sequences of the cytochrome b gene with a large data set containing nearly all other species in the genus Acrocephalus, including the type specimen of A. orinus. Five of the new specimens were collected in summer in Afghanistan and Kazakhstan, indicating that the species probably breeds in Central Asia, and the data and moult of the others suggest that the species migrates along the Himalayas to winter in N India and SE Asia. The population structure suggests a stable or shrinking population.     

MHC diversity in two Acrocephalus species: the outbred Great reed warbler and the inbred Seychelles warbler

The Great reed warbler (GRW) and the Seychelles warbler (SW) are congeners with markedly different demographic histories. The GRW is a normal outbred bird species while the SW population remains isolated and inbred after undergoing a severe population bottleneck. We examined variation at Major Histocompatibility Complex (MHC) class I exon 3 using restriction fragment length polymorphism, denaturing gradient gel electrophoresis and DNA sequencing. Although genetic variation was higher in the GRW, considerable variation has been maintained in the SW. The ten exon 3 sequences found in the SW were as diverged from each other as were a random sub-sample of the 67 sequences from the GRW. There was evidence for balancing selection in both species, and the phylogenetic analysis showing that the exon 3 sequences did not separate according to species, was consistent with transspecies evolution of the MHC.     

The identification and distribution of the enigmatic Large‐billed Reed Warbler Acrocephalus orinus

We re‐evaluate the characteristics of the little‐known Large‐billed Reed Warbler Acrocephalus orinus based on plumage and biometrical comparisons of 10 newly discovered specimens of A. orinus with a large sample of Blyth’s Reed Warbler Acrocephalus dumetorum, the species most closely resembling A. orinus. Using specimens whose identity was verified by DNA analyses, we found that the most important criteria for identifying orinus are its long and comparatively broad bill, with typically uniformly dark upper mandible, and on average longer tarsi and claws with a thin and pointed hind claw. In contrast, previously suggested differences in wing formula, plumage colour and shape of flight‐feathers are very minor, variable and seem of limited use for identification of orinus. Graphical analysis of hind claw length against bill to skull length should assist researchers in locating museum or live specimens of orinus that otherwise might be overlooked as dumetorum, and multivariate analysis unambiguously separates the two species based on our present dataset. A review of the scattered literature and specimen records suggests that the breeding distribution of orinus lies between southeastern Kazakhstan and northeastern Afghanistan, within the breeding range of dumetorum, but that orinus may well winter predominantly east of the main wintering range of dumetorum. Further evidence to support this assessment of the species' breeding range was obtained in June 2009, with the discovery of a population in the Pamir Mountains of Afghanistan.     

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.     

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.     

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.     

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.     

Singing as a handicap: the effects of food availability and weather on song output in the Australian reed warbler Acrocephalus australis

Bird song is generally regarded as a sexually selected trait, and may represent a reliable handicap signal under at least certain conditions. Females may use the degree of male song production as a reliable cue to male condition or territory quality. We investigated the effect of supplementary feeding on song output in the migratory Australian reed warbler Acrocephalus australis . We experimentally increased the food availability on alternate days, and recorded several weather variables. We measured song rate and song length independently. Supplementary fed birds sang more on feeding days than on non-feeding days, while control birds did not show this effect. Song output was not significantly associated with any of the weather variables examined. Our results indicate that singing has the potential to serve as a reliable handicap signal to territorial food availability irrespective of the prevailing weather conditions. We discuss the role of energetic constraints and behavioural flexibility on the signaling function of song.     

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%).     

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.     

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.     

Effect of nest and nest site characteristics on the risk of cuckoo Cuculus canorus parasitism in the great reed warbler Acrocephalus arundinaceus

The risk of cuckoo Cuculus canorus parasitism for great reed warbler Acrocephalus arundinaceus nests was evaluated in respect to nest and nest site structure in the reedbeds along canals in central Hungary, In total 90 nests were analysed, from which 31 remained unparasitised. 37 were single parasitised and 22 multiple parasitised. Multiple discriminant analysis, based on eight structural variables, separated well the unparasitised clutches from the cases of single and multiple parasitism, but the latter two categories also showed a weaker separation. Nests close to cuckoo vantage points are most vulnerable to parasitism. The variable "distance from nest to closest available cuckoo perch site" (tree or electric wire) played the most important role in the risk of parasitism. It showed highly significant differences among the groups: lowest distances were found in the case of multiple parasitism, and the highest distances were measured when nests remained unparasitised. Additionally, "nest visibility" also showed significant differences among the three groups: a more visible nest is more likely to be parasitised. Risk of parasitism affects on the host on two levels: 1) female cuckoos search for nest-building hosts from a perch site, but 2) when they are in the act of parasitism, they can find more visible nests in the reed. Besides the robust effects of the variables "distance to cuckoo perch site" and "nest visibility", the variable "distance from nest to open water" may reduce, but "nest volume" increases the risk of multiple parasitism. Differences between cases of single and multiple parasitism are weak, mainly affected by chance. We explain it by the high abundance of the cuckoo, the parasitism rate was 65.6% (59/90), Cuckoos parasitised almost all of the available nests in the close vicinity of potential perch sites. There was no evidence that great reed warblers nested closer to each other when risk of parasitism was high.     

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.     

Great reed warbler Acrocephalus arundinaceus and reed warbler Acrocephalus scirpaceus respond differently to cuckoo dummy at the nest

A cuckoo Cuculus canorus dummy was exposed at 24 nests of great reed warbler Acrocephalus arundinaceus (GRW) and 34 nests of reed warbler Acrocephalus scirpaceus (RW) during the egg-laying stage. The eight GRW pairs attacked the cuckoo directly, striking the dummy, but such a behaviour was not recorded in RWs. Also, other behavioural measures (closest distance from the model, duration of distress calls and number of excitement calls) indicated a lower level of defence by RWs compared to GRWs. In the study area, the parasitism rate was much lower in GRWs (1.7% of nests) than in RWs (11.3%). We suggest that one of the reasons for the lower level of cuckoo parasitism on GRWs is its stronger nest defence and hence higher risk of injury or even death for the cuckoo during egg dumping.     

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.     

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.