Time constraints in temperate‐breeding species: influence of growing season length on reproductive strategies

Organisms that reproduce in temperate regions have limited time to produce offspring successfully, and this constraint is expected to be more pronounced in areas with short growing seasons. Information concerning how reproductive ecology of endotherms might be influenced by growing season length (GSL) is rare, and species that breed over a broad geographic range provide an opportunity to study the effects of time constraints on reproductive strategies. We analyzed data from a temperate‐breeding bird, the lesser scaup Aythya affinis; hereafter scaup, collected at eight sites across a broad gradient of GSL to evaluate three hypotheses related to reproductive compensation in response to varying time constraints. Clutch initiation date in scaup was unaffected by GSL and was unrelated to latitude; spring thaw dates had a marginal impact on timing of breeding. Clutch size declined during the nesting season, as is reported frequently in bird species, but was also unaffected by GSL. Scaup do not appear to compensate for shorter growing seasons by more rapidly reducing clutch size. This study demonstrates that this species is remarkably consistent in terms of timing of breeding and clutch size, regardless of growing season characteristics. Such inflexibility could make this species particularly sensitive to environmental changes that affect resource availabilities.     

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.     

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.     

Male behavior through the breeding season in Saimiri boliviensis boliviensis

The Bolivian squirrel monkey (Saimiri boliviensis boliviensis) is a seasonal breeder. Male squirrel monkeys show distinct morphological and behavioral changes prior to and during the breeding season. A “fatting syndrome” includes increased body weight, increased levels of androgens, and in the Bolivian subspecies, an increasingly active role in the social organization of the group. In this study, the behavior of ten adult male Bolivian squirrel monkeys was analyzed over a 6-month period prior to, during, and after the breeding season. Each was housed as the only adult male in a breeding unit with six to ten adult females and one juvenile male. Employing a principle components method, 11 behavioral clusters were generated from 27 responses. Their activity clusters were identified as follows: sexual activity that showed a peak around the time of peak conceptions; excitatory activity that was initially high but decreased throughout the breeding season; and maintenance activity that did not change across the breeding season. The changing social behavior of the male squirrel monkey parallels physiological changes and is correlated with changing androgen levels.     

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.     

Intra- and interspecific agonistic behaviour in sympatric harriers during the breeding season

We analysed the temporal and sexual patterns of intra- and interspecific aggression in sympatric harriers during the breeding season, to determine the main resource defended (food, nest sites, mates) and how factors such as body size or breeding system (territorial versus colonial) influence aggressive behaviour. We predicted that if aggression is (at least partly) related to competition for food, the hen harrier, Circus cyaneus, because of its large size and territorial system, should be more aggressive (both intra- and interspecifically) than the smaller, colonial Montagu's harrier, Circus pygargus. The intraspecific aggression rate of both harriers peaked early in the season, was mainly intrasexual and increased with the number of neighbours. These patterns support the mate competition hypothesis to explain intraspecific aggression in both species. Montagu's harriers were more aggressive towards conspecifics than hen harriers. Their aggression rates towards heterospecifics were high at the start of the season then decreased, supporting the hypothesis that interspecific aggression serves primarily for nest site defence. Hen harriers showed lower interspecific aggression rates in the prelaying period and a gradual increase throughout the breeding cycle, particularly by females, who hunt around the nest for food for the nestlings. These patterns correspond to food competition, as food resources around the nest are probably more important and interspecific intrusion more costly for territorial hen harriers than for colonial Montagu's harriers. Copyright 2002 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.     

繁殖期白冠长尾雉占区雄鸟的活动区

20 0 0年 3～ 8月采用无线电遥测技术对董寨国家级自然保护区的白冠长尾雉 (Syrmaticusreevesii)繁殖期占区雄鸟的活动区进行了研究。对遥测时间超过 6 0天的 7只个体的研究表明 ,繁殖期白冠长尾雉占区雄鸟对活动区具有较强的依恋性 ;90 %调和平均转换法 ( 90 %HMT)计算的繁殖期活动区大小为 46 0± 13 7hm2 (n=7) ;繁殖初期、中期和末期的活动区大小分别为 2 4 5± 6 3、 19 6± 7 5和 17 9± 6 2hm2 (n =7) ;随着繁殖期的结束活动区与日活动区呈减小趋势。白冠长尾雉占区雄鸟对活动区的 3个亚区 (夜栖区、觅食休憩区和过渡区 )的利用时间存在明显差异 ,活动区所在地的植被结构的不同可能是造成这种差异的主要原因.     

Winter feeding of birds increases productivity in the subsequent breeding season

Supplementary food given to birds can have contemporary effects by reducing the risk of starvation, increasing survival and altering movements and reproductive performance. There is, however, a widely held perception that birds benefit from extra food over winter, but that it is better that they 'look after themselves' during breeding. Here we describe a landscape-scale experiment showing for the first time that the effects of increasing food availability only during the winter can be carried over to the subsequent breeding season. Even though food supplementation stopped six weeks prior to breeding, birds living on sites provisioned over winter had advanced laying dates and increased fledging success compared with birds living on unprovisioned sites. Thus, supplemental feeding of wild birds during winter, in a manner mimicking householders provisioning in gardens and backyards, has the potential to alter bird population dynamics by altering future reproductive performance. With levels of bird feeding by the public continuing to increase, the impacts of this additional food supply on wild bird populations may be considerable.     

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.     

Sexual maturity and breeding season of the badger, Meles meles in Sweden

The paper describes the seasonal sexual activity of badgers collected in central Sweden. Ovulation- and pregnancy rates were determined for females in different age classes and males were examined for testis weight and occurrence of spermatozoa.     

Exclusive frugivory and seed dispersal of Rhamnus alaternus in the bird breeding season

We studied avian frugivory and seed dispersal in a dioecious shrub, Rhamnus alaternus, focusing on the quantitative and qualitative components of effectiveness. The study took place at three locations in the northeast of the Iberian Peninsula, and examined bird behaviour, intensity of feeding, and the consequences for seedling emergence. The coincidence between the bird breeding season and fruit ripening of R. alaternus in the absence of other ripe fruit, generates a monospecific interaction. The extant frugivorous species were mainly legitimate seed dispersers and their abundance was low. Sylvia melanocephala and S. undata were the most important at one site whereas S. atricapilla, Erithacus rubecula and Turdus merula predominated at the other two sites. Fruit handling took place directly on the branches. Bird species used microhabitats differently as first post-feeding perch, which usually was a short distance away. The low density of frugivorous birds in all localities, among others factors, resulted in satiation of the disperser community and many mature fruits unconsumed. Both adults and juveniles feed upon the plants and their foraging patterns are similar. Adults of S. melanocephala were observed to feed fruit to nestlings and consequently a second phase of dispersal potentially arises from the transport of fecal sacs. Pulp removal and passage through the digestive tract increased the probability of seedling emergence. This plant-dispersal interaction has important consequences, both positive and negative for the plant. Positively, the fruiting of R. alaternus at a time when other ripe fruits are not available avoids interspecific competition for seed dispersers. In addition, a low density of seed rain may reduce intraspecific competition. Negatively, the low density and small size of the breeding frugivorous bird community limit fruit handling and removal away from the parent plants, while the territorial behaviour of birds at that time of the year reduces the potential distances of seed dispersal.     

Seed preference and energy intake of Goldfinches Carduelis carduelis in the breeding season

The food of a population of Europeans Goldfinches breeding in Orchards was studied. More than 5% of the total foodstuff consumed was met by the seeds of seven food plants. Water, lipid, carbohydrate and protein content was determined from 20 different plants which Goldfinches used during the breeding season in order to calculate energy content. The energy of fresh seeds varied from 1.88 kJ/g in Tussilago farfara to 12.14 kJ/g in Dactylis glomerata. The dependence of the rate of energy intake (profitability) on the pecking rate and the size (weight) of the seeds was calculated. The profitability of various seeds is in agreement with the relative number of birds using them. Calculation of the feeding time that a Goldfinch needs during daylight hours, if it feeds only on a particular plant, indicates that there are several foods that cannot fill the energy demands of the bird during the breeding season. These are eaten only if others with a high energy content are not available. Thus food selection in Goldfinches depends, apart from availability, on seed profitability. The oxygen consumption of a male Goldfinch was significantly higher when the animal was feeding on seed heads of Tussilago farfara and Taraxacum officinale than when it fed on loose dry seeds, and so, different feeding techniques have to be taken into account in calculating net energy gain.     

Territory structure of the Robin Erithacus rubecula outside the breeding season

On the basis of radio-tracking data from 51 individuals, the space use of Robins Erithacus rubecula outside the breeding season was non-uniform and multinuclear. On average, territorial Robins spent 78% of their time in one or more resource patches represented by bushes, which accounted for only 1% of a convex polygon encompassing each range. A suite of range polygons was developed to quantify range structure. One of these polygons showed similar size to, and high overlap with, defended-area polygons and so was a good estimate of territory. In addition, the area of bushes in each polygon suggested Robins occupied constant resource territories. There was undefended neutral ground between territories, which, although seldom visited, was crossed to achieve intrusions. Over the short term (days), Robins showed high fidelity to the resource patches they defended. Over the longer term (months), most individuals showed some degree of range shift. It is concluded that if the information concerning territories is to be maximized, empirically derived sampling protocols should be used when measuring territory parameters.     

Biochemical components of stallion seminal plasma before and after the breeding season

Seasonal and breed differences were determined for glyceryl-phosphoryl-choline (GPC), ergothioneine (EGT), fructose and total protein in the seminal plasma of 24 stallions from three breed groups.Significant differences were found between the GPC, EGT, fructose and total protein levels before and after the breeding season. During the post-breeding period, the concentrations of EGT, fructose and total protein increased more than 100% while that for GPC decreased by about 50%. This phenomenon was found in each of the breed groups investigated.Breed differences in the GPC level during the pre- and post-seasonal periods were not significant. Differences were found only during the pre-seasonal period in EGT, fructose and total protein levels.     

Ovarian function in ewes during the transition from breeding season to anoestrus

Transrectal ovarian ultrasonography was conducted in six Western white-faced ewes for 35 days from the last oestrus of the breeding season, to record the number and size of all ovarian follicles > or = 3 mm in diameter and luteal structures. Blood samples were collected once a day for estimation of serum concentrations of follicle-stimulating hormone (FSH), oestradiol and progesterone. Each ewe had five follicular waves (follicles growing from 3 to > or = 5 mm in diameter) over the scanning period. The duration of the growth phase of the largest ovarian follicles did not differ (P > 0.05) between waves, but follicular static and regressing phases decreased significantly (P < 0.05) after the decline in serum progesterone concentrations at the end of the last luteal phase of the breeding season. The intervals between the five follicular waves were: 9.2+/-0.4, 5.2+/-0.7, 8.3+/-0.8 and 5.8+/-0.7 days; the two shorter intervals differed (P < 0.05) from the two longer intervals. Using the cycle-detection program, rhythmic increases in serum FSH concentrations were detected in all ewes; the amplitude, duration and periodicity of FSH fluctuations did not vary (P > 0.05) throughout the period of study. The number of identified FSH peaks (7.8+/-0.5 peaks per ewe, per scanning period) was greater (P < 0.05) than the number of emerging follicular waves. Serum concentrations of oestradiol remained low (< or = 1 pg/ml) on most days, in five out of the six ewes studied, and sporadic elevations in oestradiol secretion above the non-detectable level were not associated with the emergence of follicular waves. The ovulation rate was lower than that seen during the middle portion of the breeding season (November-December) in white-faced ewes but the transitional ewes had larger corpora lutea (CL). Maximal serum concentrations of progesterone appeared to be lower and the plateau phase of progesterone secretion appeared to be shorter during the last luteal phase of the ovulatory season in comparison to the mid-breeding season of Western white-faced ewes. During the transition into anoestrus in ewes, the endogenous rhythm of FSH release is remarkably robust but the pattern of emergence of sequential follicular waves is dissociated from FSH and oestradiol secretion. Luteal progesterone secretion is suppressed because of fewer ovulations and diminished total luteal volume, but it may also result from diminished gonadotropic support. These season-related alterations in the normal pattern of ovine ovarian cycles appear to be due to reduction in ovarian responsiveness to gonadotropins and/or attenuation in secretion of luteinizing hormone (LH) occurring at the onset of the anovulatory season in ewes.