Song and plumage effects on aggressive display by the European Robin Erithacus rubecula

The response was observed of territory-holding male European Robins ETithacus rubecula to red— and brown-breasted model Robins which were presented either simultaneously with tape-recorded Robin song, or with no song. Robins displayed and sang at the silent red-breasted model, but sang and displayed much more at the models that were presented with song. It is suggested that, while the red breast is one important feature of Robins eliciting aggressive behaviour, song is also important. The relationship of these findings to studies of stimulus features eliciting behaviour in other species is discussed.     

Sex-determination of Continental European Robins Erithacus r. rubecula

Existing morphological sex-determination criteria for Continental European Robins proved inadequate when applied to birds captured in western Denmark. Only 49% of captured birds met with the wing-length criteria resulting in the classification of 2% males and 47% females respectively. This article presents a primer for correct sexing of Robins by amplification of a female-specific DNA element by a polymerase chain reaction (PCR) on genomic DNA. Further, two new criteria for sex-determination are proposed based on logistic regression and discriminant function analysis using morphological measures from birds with DNA-verified sex. Success rates of 80–81% for the new criteria were a considerable improvement compared with a success rate of only 36% for the old criteria.     

Territoriality as a paternity guard in the European robin, Erithacus rubecula

To investigate the relative importance of paternity defences in the European robin we used behavioural observations, simulated intrusions and temporary male removal experiments. Given that paired males did not increase their mate attendance, copulation rate or territory size during the female's fertile period, the most frequently quoted paternity assurance strategies in birds were absent. However, males with fertile females sang and patrolled their territories more regularly, suggesting that territorial motivation and vigilance were elevated when the risk of cuckoldry was greatest. In addition, there was a significant effect of breeding period on response to simulated intrusions: residents approached and attacked freeze-dried mounts more readily in the fertile period. During 90-min removals of the pair male in the fertile period, neighbours trespassed more frequently relative to prefertile and fertile period controls and appeared to seek copulations with unattended females. When replaced on their territories, males immediately increased both song rate and patrolling rate in comparison with controls. We propose that male robins sing to signal their presence, and increase their territorial vigilance and aggression in the fertile period to protect paternity. Copyright 2000 The Association for the Study of Animal Behaviour.     

Song characteristics and response to playback of male and female Robins Erithacus rubecula

Male and female Robins, from a population occupying deciduous woodland habitat, were recorded during autumn, winter and spring. Song characteristics were compared between individuals with respect to sex and reproductive success. For the parameters measured, male song did not vary with reproductive success; however, male song was more complex and less redundant than female song. Furthermore, male song in response to playback was more complex in response to female than to male song. Both males and females could discriminate between male and female song. Males approached playback of male song more readily in the autumn than in the spring. The possible significance of these results is discussed.     

Anatomo-histologic characteristics of the epiphysis of the female silver fox outside the breeding season

The epiphysis of mature relatively wild silver foxes of various age living in the fur-farming farm have been studied during November--December. The organ mass (2.72 +/- 0.14 mg) has rather essential individual differences and its statistically significant changes with age are not revealed. There are not any connective septa deviding parenchyma of the organ into lobules. There is also not any definite division between the cortex and medulla. Under all histological methods of treatment the borders of the parenchymal cells are poorly defined. Nuclei of the pinealocytes can be divided into light and dark according to their coloration. The average diameter of the nuclei in the pinealocytes does not practically change with age, nor does the linear density of cellular elements arrangement. A definite individual variability is noted in amount of the cells including secretory granules. The phenomena mentioned demonstrate that the fox epiphysis maintains its constant level of physiological activity during the whole reproductive period of life.     

Winter segregation of migrant European robins Erithacus rubecula in relation to sex, age and size

Birds often show some form of social segregation during winter, both at large geographical scales (a consequence of differential migration) and at the regional or local level, when comparing different habitats or micro-habitats. However, our understanding of the mechanisms underlying such patterns is still poor. These issues have been rarely investigated in migratory Old-World passerines, particularly with respect to differences between the sexes. In this study, we show that female European robins Erithacus rubecula (sexed by molecular techniques) greatly outnumber males in southern Iberia, which confirms that this species is a differential migrant with a strong latitudinal segregation of the sexes. Furthermore, sex, age and body size influence the habitat distribution of robins in winter. Subordinate birds (females, juveniles and small individuals) were generally more common in habitats with a greater shrub development, and comparatively scarce in woodlands with relatively little undergrowth. Birds wintering in woodlands were in better condition (assessed by breast-muscle scoring) than birds wintering in shrubland. These results are consistent with the hypothesis that proposes that social dominance, mediated by differences in size and experience, is important in determining the habitat segregation of sex and age classes. The alternative hypothesis (habitat specialization), although not specifically supported by our findings, cannot be ruled out on the basis of the available evidence.     

The effect of yellow and blue light on magnetic compass orientation in European robins, Erithacus rubecula

To analyze the wavelength dependency of magnetic compass orientation, European robins were tested during spring migration under light of various wavelengths. Under 565-nm green light (control) the birds showed excellent orientation in their migratory direction; a 120° deflection of magnetic North resulted in a corresponding shift in the birds' directional tendencies, indicating the use of the magnetic compass. Under 443-nm blue light, the robins were likewise well oriented. Under 590-nm yellow, however, oriented behavior was no longer observed, although the activity was at the same level as under blue and green light. The spectral range where magnetic orientation is possible thus differs from the range of vision, the former showing parallels to that of rhodopsin absorption. The interpretation of the abrupt change in behavior observed between 565 green to 590 yellow is unclear. There is no simple relationship between magnetoreception and the known color receptors of birds. Accepted: 17 December 1998     

Frugivory of transient and wintering European robins Erithacus rubecula in a Mediterranean region and its relationship with ornithochory

Stomach contents of 209 dead and droppings of 57 mist-netted migrating or wintering European robins Erithacus rubecula in a Mediterranean area in southern France showed the following seasonal trends: (1) a dominance of invertebrates in early autumn, (2) a dominance of fruits in mid-autumn, (3) acorns being important in midwinter, and (4) a strong dominance of invertebrates at the end of winter and in early spring.
The fruits of 23 plant species had been consumed. The most widespread and abundant ones were also the most commonly taken ( Rubia peregrina at the regional level and Viburnum tinus at a local one). The size of these fruits ranged from 4 to 12 mm. mostly between 6 and 8 mm. Owing to the amount of fruits consumed and the numbers of birds involved, the European robin is one of the few main dispersers of the numerous fleshy-fruited plant species of the Mediterranean area. Nevertheless, a correct evaluation of its role is only possible within the dense network existing between these plant species and all their dispersers (mainly birds but also mammals).     

Territorial aggression and song of male European robins (Erithacus rubecula) in autumn and spring: effects of antiandrogen treatment

Male robins aggressively defending a feeding territory in winter have low levels of testosterone, while males defending a breeding territory in spring have elevated levels of testosterone. Song is an integrated part of territorial defense during both phases. We investigated whether testosterone is involved in the expression of these behaviors by treating free-living and captive male robins during both phases with the antiandrogen flutamide. Results suggest that, similar to species in which territoriality is restricted to the reproductive phase, aggressive defense of a breeding territory by male robins is facilitated by androgens. Territorial defense during the nonbreeding season, however, does not require androgenic activity. Singing frequency, on the other hand, was not significantly reduced during either phase by flutamide application. Since the quality of male song changes with season we suggest that the incorporation of sexual signals into male song during breeding depends on testosterone, but that the year-round production of territorial "keep out" signals is independent of testosterone.     

Effects of melengestrol acetate and P.G. 600 on fertility in Rambouillet ewes outside the natural breeding season

The effects of melengestrol acetate (MGA) and P.G. 600 on ewe fertility outside the natural breeding season were evaluated. Rambouillet ewes were assigned to one of four groups: (1) control (C; n=92); (2) PG600 (n=86); (3) MGA (n=99); and (4) MGA+PG600 (n=92). A pellet with or without MGA (0.3mg/ewe/d) was fed at 0.15kg/ewe/d for 7d. On the last day of pellet feeding, ewes were given either saline or 5mL of P.G. 600 i.m. (400IU equine chorionic gonadotropin (eCG) and 200IU human chorionic gonadotropin (hCG)). Ultrasonography was performed between Days 20 and 25 of gestation for ewes that were mated during the first 6 d of the breeding period from the MGA (n=15) and MGA+PG600 (n=8) groups, and the number of luteal structures and embryos were counted. During the first 6d of the breeding period, MGA increased (P<0.05) the percentage of ewes that mated and conceived when compared to C and PG600 (24.2% vs. 3.3% and 10.5%, respectively). Relative to MGA, the mean (+/-S.E.M.) number of luteal structures per ewe was enhanced (P<0.03) in MGA+PG600 (1.53+/-0.13 vs. 2.38+/-0.42, respectively), however as pregnancy progressed, the number of embryos (1.5+/-0.13 vs. 1.8+/-0.16, respectively) and lambs born (1.3+/-0.15 vs. 1.5+/-0.27, respectively) did not differ. Treatment with MGA reduced (P<0.01) the interval from ram introduction to lambing relative to groups that did not receive MGA (168+/-0.8d vs. 171+/-0.6d, respectively). In conclusion, treatment with MGA increased the percentage of ewes conceiving early in the breeding period. Although P.G. 600 increased the number of luteal structures present per ewe, it did not significantly enhance ewe prolificacy.     

Studies of pituitary-adrenal-testis interaction in sheep. II. The effects of repeated injections of adrenocorticotrophic hormone outside the breeding season

The administration of 0.5 mg of long-acting adrenocorticotrophic hormone (ACTH, Synacthen-Depot) twice daily for 5.5 d to four rams outside the breeding season caused marked rises in plasma cortisol without any evidence of adrenal depletion. This treatment also caused marked rises in basal plasma follicle stimulating hormone (FSH) concentrations which remained high even after cessation of treatment. Plasma FSH responses to 5 ug of gonadotrophin releasing hormone (GnRH) were consistently observed and ACTH treatment increased the FSH response to GnRH. In contrast, spontaneous fluctuations in the plasma luteinizing hormone (LH) and testosterone concentrations were abolished by ACTH treatment. The quantity of testosterone released after GnRH (estimated by the maximum values reached and by the area under the response curve) was also suppressed while that of LH was only slightly lower. A comparison of the results of this experiment with those obtained in rams during the breeding season showed that the effects of ACTH on LH and testosterone were more marked during the breeding season. In contrast, the effect of ACTH on FSH is to increase the latter during the nonbreeding season, whereas no effect was observed during the breeding season.     

Short term, repeated exposure to rams during the transition into the breeding season improves the synchrony of mating in the breeding season

The ram effect is widely used in Mediterranean breeds of sheep but its use in temperate genotypes is restricted by breed seasonality. However, ewes from these highly seasonal genotypes are sensitive to stimulation by rams close to the onset of the natural breeding season. In this study we developed a pre-mating protocol of repeated, short-term exposure to rams (fence-line contact or vasectomised rams) beginning during late anoestrus and continuing into the breeding season. We hypothesised that this pre-mating protocol would synchronise the distribution of mating of North of England Mule ewes during the breeding season above that observed in ewes isolated from rams prior to mating. Ram-exposed ewes were given contact with rams (Experiment 1: fence-line; FR, n = 94 and Experiment 2: vasectomised rams; VR; n = 103) for 24 h on Days 0 (10 September), 17 and 34 of the experiment. Control ewes (Experiment 1; FC, n = 98 and Experiment 2; VC; n = 106) remained isolated from rams prior to mating. In Experiment 2, a subset of VR (n = 35) and VC ewes (n = 35) were blood sampled twice weekly to monitor their pre-mating progesterone profiles. At mating, harnessed entire rams were introduced, 17 or 16 days after the last ram exposure (Experiments 1 and 2) and raddle marks were recorded daily. The median time from ram introduction to mating was reduced in ewes given both fence-line and vasectomised ram contact (P < 0.001), leading to a more compact distribution of mating and lambing (At least P < 0.01). In the blood sampled VR ewes, there was a progressive decline in the number of days from ram exposure to the onset of dioestrus (at least P < 0.05). This observation indicates that the cycles in VR ewes became increasingly synchronised over the pre-mating period, a pattern not evident in VC ewes. In conclusion, repeated, short-term exposure of ewes to rams during the transition into the breeding season is an effective method of synchronising the distribution of mating during the breeding season.     

Reindeer bull introduction affects the onset of the breeding season

Reindeer are seasonally polyestrus, short day breeders, with estrous cycles of approximately 20 days in length. The objective of this study was to investigate the effects of reindeer bull exposure on the onset of the reindeer cow breeding season and to investigate whether cows with calving experience responded differently than cows with no previous reproductive experience. During year 1, blood samples were collected weekly beginning 14 July and continuing until 15 September (n = 8) or 30 September (n = 8) in order to determine the onset of the breeding season, based on ovarian function, with no bull present. Plasma was stored frozen for later assay of progesterone (P(4)) following the conclusion of sample collection. Eight randomly selected cows were allowed to breed with a bull during year 1. The mean onset of ovarian activity in the first year was 15 September (range: 8-29 September). The bull was removed from cows more than 2 months prior to the start of the experimental period during year 2 and housed at a separate facility approximately 2 km distant. Blood samples were collected during year 2 from 14 cows 3x weekly beginning on 11 August (day 1) and continuing until 29 September (day 50) and plasma was stored frozen for later assay of P(4) following the conclusion of sample collection. On day 6, cows were divided into two groups such that group 1 (early bull exposure; EBE), consisted of four cows that had calved the previous spring and four cows that had no reproductive experience (n = 8). Group 2 (late bull exposure; LBE), consisted of three cows that had calved the previous spring and three cows that had no reproductive experience (n = 6). EBE experienced bull introduction on day 13, 23 days earlier than the average onset of ovarian activity during year 1. LBE experienced bull introduction on day 46, 10 days later than the average onset of ovarian activity during year 1. Progesterone concentrations were analyzed by ANOVA procedures for repeated measures. Bull presence was not requisite for the onset of ovarian activity during either year. Previous reproductive status had no effect on the onset of ovarian activity within EBE (P = 0.61) or within LBE (P = 0.92). Time of bull introduction had a significant effect on the onset of ovarian activity when EBE was compared to LBE (P<0.001). The first sustained increase in mean P(4) concentration above 1 ng/ml occurred on day 25 in EBE reindeer and day 41 in LBE reindeer. EBE reindeer initiated ovarian activity 12 days after bull introduction while LBE reindeer initiated ovarian activity 5 days before bull introduction. All cows penned with the bull conceived during both breeding seasons, demonstrated by production of live calves during the subsequent spring. Cows bred during year 1 all calved within a 9 day-period. During year 2, EBE displayed a more synchronous calving period compared to LBE (P<0.05). Results indicate that bull management affects the onset of the breeding season in reindeer cows, regardless of previous reproductive experience.     

An increase in parental investment during the breeding season

The intensity of nest-defence aggression by female redwinged black birds (Agelaius phoeniceus) with eggs decreased late in the breeding season, while aggression at this time increased amongst females defending broods. High predation pressure late in the breeding season decreased the probability of survival of the late nesting attempts, so that females with young had a much greater chance of success than females with eggs at this time. The observed changes in aggressive responses are predicted by the theory of parental investment.     

Gregarious foraging in barn swallows after the breeding season

Aerial vertebrate foragers, e.g. insectivorous bats, martins and swallows, often show gregarious behavior such as colonial breeding, communal roosting and aggregating behavior during foraging. Studies of gregariousness in aerial foragers have mostly focused on colonial breeding or communal roosting, and only a few intensive studies have dealt with gregariousness during foraging. Here, we report on large and stable aggregations of barn swallows, Hirundo rustica, observed during daytime after the breeding season in Japan. Relatively large aggregations of 20 or more birds were located around poultry, pig and cattle farms and mounds of manure. The aggregations were stable in size both within-days and between-days. Their activities consisted only of repeated cycles of foraging and resting around the farms where their prey, the large black soldier fly Hermetia illuceus (>10 mm) and other medium (5–10 mm) flies, was dense. Distributions of swallows around the farms overlapped with the distributions of prey, and the size of the aggregations significantly correlated with the abundance of prey.     

Physiological constraints and latitudinal breeding season in the Canidae

Physiological strategies that maximize reproductive success may be phylogenetically constrained or might have a plastic response to different environmental conditions. Among mammals, Canidae lend themselves to the study of these two influences on reproductive physiology because all the species studied to date have been characterized as monestrous (i.e., a single ovulatory event per breeding season), suggesting a phylogenetic effect. Greater flexibility could be associated with environments that are less seasonal, such as the tropics; however, little is known for many of the species from this region. To compensate for this lack of data, two regressions were done on the length of the reproductive season relative to the latitudinal distribution of a species: one with raw data and another with phylogenetically independent contrasts. There was a significant negative relationship, independent of phylogeny, with canids that have longer breeding seasons occurring at lower latitudes. In contrast, the pervasiveness of monestrus within Canidae appears to be phylogenetically constrained by their pairing/packing life and is most likely associated with monogamy. The persistence of the monestrous condition is supported by a captive study where a tropical canid, the fennec fox, Vulpes zerda, never exhibited polyestrous cycles despite a constant photoperiod (12L : 12D).     

Ovarian function in ewes at the onset of the breeding season

Transrectal ultrasonography of ovaries was performed each day, during the expected transition from anoestrus to the breeding season (mid-August to early October), in six Western white-faced cross-bred ewes, to record ovarian antral follicles > or = 3 mm in size and luteal structures. Jugular blood samples were collected daily for radioimmunoassay (RIA) of follicle-stimulating hormone (FSH), oestradiol and progesterone. The first ovulation of the breeding season was followed by the full-length oestrous cycle in all ewes studied. Prior to the ovulation, all ewes exhibited a distinct increase in circulating concentrations of progesterone, yet no corpora lutea (CL) were detected and luteinized unovulated follicles were detected in only three ewes. Secretion of FSH was not affected by the cessation of anoestrus and peaks of episodic FSH fluctuations were associated with the emergence of ovarian follicular waves (follicles growing from 3 to > or = 5 mm). During the 17 days prior to the first ovulation of the breeding season, there were no apparent changes in the pattern of emergence of follicular waves. Mean daily numbers of small antral follicles (not growing beyond 3 mm in diameter) declined (P < 0.05) after the first ovulation. The ovulation rate, maximal total and mean luteal volumes and maximal serum progesterone concentrations, but not mean diameters of ovulatory follicles, were ostensibly lower during the first oestrous cycle of the breeding season compared with the mid-breeding season of Western white-faced ewes. Oestradiol secretion by ovarian follicles appeared to be fully restored, compared with anoestrous ewes, but it was not synchronized with the growth of the largest antral follicles of waves until after the beginning of the first oestrous cycle. An increase in progesterone secretion preceding the first ovulation of the breeding season does not result, as previously suggested, from the ovulation of immature ovarian follicles and short-lived CL, but progesterone may be produced by luteinized unovulated follicles and/or interstitial tissue of unknown origin. This increase in serum concentrations of progesterone does not alter the pattern of follicular wave development, hence it seems to be important mainly for inducing oestrous behaviour, synchronizing it with the preovulatory surge of luteinizing hormone (LH), and preventing premature luteolysis during the ensuing luteal phase. Progesterone may also enhance ovarian follicular responsiveness to circulating gonadotropins through a local mechanism.