Eight new polymorphic microsatellite loci from the eastern phoebe (<Emphasis Type="Italic">Sayornis phoebe</Emphasis>)

We describe eight new polymorphic microsatellite loci (5 dinucleotide and three trinucleotide) for the eastern phoebe (Sayornis phoebe) to complement five previously published loci. The number of alleles per locus ranged from 3 to 15 and observed heterozygosities ranged from 0.33 to 0.91. Preliminary screening revealed that loci were polymorphic in other Tyrannidae: Empidonax virescens (n = 10), Tyrannus tyrannus (n = 10), Tyrannus vociferans (n = 5), and Tyrannus melancholicus (n = 28).     

Development of hypervariable microsatellite loci for use in eastern phoebes (Sayornis phoebe) and related Tyrannids

In order to complement on‐going research on reproductive strategies and dispersal in the eastern phoebe (Sayornis phoebe), we developed five polymorphic microsatellite loci which would permit us to assign parentage to offspring and to evaluate population genetic structure. The number of alleles per locus ranged from 3 to 15; probability of exclusion, with a known maternal genotype, was 0.995. Preliminary screening revealed that primers were polymorphic in other Tyrannidae.     

Correlation between two components of parental investment: nest defence intensity and nestling provisioning effort of willow tits

Nest defence intensity and nestling provisioning effort of female willow tits (Parus montanus) were significantly correlated at the end of nestling period: well-fed young were defended most intensely. Increased effort was rewarded, since broods with the highest female per-offspring provisioning rates were the most likely to produce local recruits. This suggests that the feeding ability is an important cue for parental investment decisions, at least in a species like the willow tit which has adopted the clutch adjustment strategy. Thus, the most valuable broods would not necessarily be the largest ones, but the ones in which the original number of young could be fed most adequately. However, no associations were found between the level of parental effort and offspring weight, size or condition, nor did the broods producing recruits differ from other broods in timing of breeding or number and size of offspring. The female behaviour may suggest that they invest the most time, energy and risk in the young whose chances of joining the winter flock are the best. The first well-fed young also gain an advantage of prior residency in joining the flock. The first to join normally obtain higher social status, and hence better winter survival, than latecomers. The corresponding patterns in male parental investment behaviour were weak or absent, which suggested that the male effort was affected by the female behaviour. Males seemed to invest in nestling provisioning in such a way as to supplement the female effort. During nest defence action males also seemed to invest in protection of females against predation.     

The effects of variable foraging conditions on common murre (Uria aalge) corticosterone concentrations and parental provisioning

This study investigated how total corticosterone concentrations, chick-feeding rates, and adult body mass changed with food availability from 1998 to 2000 in the same individually marked common murres (Uria aalge). Capelin, the main prey species, arrived inshore by the onset of murre chick hatching in 1998 and 1999 (prey match years); whereas in 2000, hatching began approximately 1 week before the capelin arrived inshore to spawn (prey mismatch year). Serum corticosterone concentrations were higher in the same individuals in the prey mismatch year than they were in either of the match years. Birds sampled before peak capelin spawning in the mismatch year had higher corticosterone levels than murres sampled after peak spawning. Murres with higher corticosterone levels had higher chick-feeding rates and less mass loss in the mismatch year (compared to the match year 1999) than birds with lower levels. Corticosterone levels did not differ between birds that had not foraged for at least 12 h (brooded chick overnight) and those that had, suggesting that short-term food deprivation did not affect corticosterone concentrations. Taken together, these findings suggest that the difference between years reflects a baseline shift in corticosterone levels, particularly in the high-quality birds that were able to increase both corticosterone concentrations and foraging effort.     

Manipulation of egg production reveals costs of reproduction in the tree lizard (Urosaurus ornatus)

Life-history theory predicts that the allocation of energy to current reproduction is associated with a decrement in future fecundity, future survival, or both. I treated this notion as the cost hypothesis, and tested the assumption that current reproduction exacts a cost in future survival and fecundity. Surgical manipulations of egg production were applied to natural populations of the tree lizard, Urosaurus ornatus, in south western New Mexico by yolkectomy surgery in two different years. I reduced the number of eggs produced in the first clutch during vitellogenesis by approximately 50% in yolkectomized females relative to controls. Subsequent survival, fecundity, and growth of females were followed for two or three years, depending on the cohort. Treated females in both cohorts showed significantly higher growth and survivorship throughout the experiment than in controls. After 2 years, yolkectomized females had grown an additional 2 mm (snout-vent length) compared to controls, enough for them to add on average an additional egg to their next clutch. This demonstrated a cost in terms of future fecundity through a reduction in growth and an increase in mortality in these lizards.     

Effect of clutch size on male egg-fanning behavior and hatching success in the goby, Eviota prasina (Klunzinger)

In fish that exhibit paternal care, the females often choose their mates on the basis of male traits that are indicative of the parental ability of the males. In a marine goby, Eviota prasina, males tend their eggs within their nests until hatching, and females prefer males that have longer dorsal fins and exhibit courtship behavior with a higher frequency as their mates. In order to clarify the relationship between these sexually selected traits and the parental ability of males of E. prasina, the factors affecting the hatching success of eggs within male nests and the male parental care behavior were examined in an aquarium experiment. Females spawned their eggs in male nests and the clutch size of females showed a high individual variation (range = 88-833 eggs). The hatching success of eggs within male nests showed a positive correlation with the time spent by males in fanning eggs and the clutch size. In contrast to the prediction, however, the hatching success did not show a significant correlation with the sexually selected traits, i.e., the male dorsal fin length and the frequency of courtship displays. Moreover, multiple regression analysis indicated that the time spent by the males in fanning was the most important factor affecting the survival rate of the eggs. The time spent by males in fanning behavior was influenced by the clutch size within their nests; the fanning behavior of males occurred with a higher frequency when they tended larger clutches. Males are required to invest a greater effort in egg-tending behavior to achieve a higher hatching success when they receive larger clutches, probably due to the greater reward for their parental behavior. Based on their mate choice, females may obtain other benefits such as high quality offspring.     

The effects of supplemental food on reproduction in parental male smallmouth bass

Synopsis We provided supplemental food for parental male smallmouth bass,Micropterus dolomieu, to determine if food supply limits the reproductive performance of nesting males as measured by care duration, reproductive success and survival of adult males. Although supplementing the diet of parental males had a positive effect on all three reproductive measures, the experiment generated contrasting results in different years. In the first year, supplemental feeding only improved the survival of fed males versus unfed males. In the second year, supplemental feeding increased care duration and reproductive success of fed males but reduced their survival relative to unfed males. Our supplemental feeding improved current or future measures of reproductive performance, but not both simultaneously. The results appear to demonstrate phenotypic plasticity in the extent to which energy from supplemental feeding can be allocated to present or future reproduction.     

Patterns of provisioning and iteroparity in a solitary halictine bee,Lasioglossum (Evylaeus) fratellum (Perez), with notes onL. (E.) calceatum (Scop.) andL. (E.) villosulum (K.)

Summary On Dartmoor, UK the temperate halictine beeLasioglossum (Evylaeus) fratellum is solitary and univoltine. A significant proportion of individually marked foundresses reproduce during two different seasons (iteroparity). This could represent a form of risk-spreading in environments where conditions are harsh and unpredictable. Perhaps associated with iteroparity, foundresses put in relatively low reproductive effort, often failing to provision their nests on suitable days, despite the small number of days available on Dartmoor. Foundresses stopped provisioning asynchronously, then remained in their nests until well after offspring emergence. There was a small proportion of digynous nests, in which nest-sharing appeared to be temporary.L. (E.) calceatum had a similar life-cycle on Dartmoor.L. villosulum was also solitary, and was able to provision its nests under less favourable weather conditions than the other two species. Information is also provided on nest productivity andSphecodes parasites.     

Weight loss,reproductive output,and the cost of reproduction in the common frog,Rana temporaria

Summary The consequences of reproduction for body weight, growth and survival were studied in a Swiss population of the explosive breeder, Rana temporaria. Males and females continuously loss weight in the range of 0.5% of total body weight per day from the breeding migration throughout May. Females also lost about 33% (1983) and 29% (1984) due to spawning. In addition to this significant year-to-year variation, there was also considerable individual variation in reproductive output. Skeletochronological techniques indicated that breeding male or female frogs experienced a growth reduction of several millimeters relative to non-breeding frogs of the same body size. There was no relationship between an individual female's reproductive output in consecutive years or with her subsequent growth or survival. It was concluded that weight loss is caused by a seasonally elevated metabolism in combination with a lack of feeding and represents a basic energetic cost of reproduction, resulting in lowered growth. Individual variation in relative reproductive output is mostly environmentally induced and is not an expression of different reproductive strategies. This may explain the lack of trade-offs that are predicted by the cost-of-reproduction-hypothesis.     

Condition dependence of nestling mouth colour and the effect of supplementing carotenoids on parental behaviour in the hihi (Notiomystis cincta)

Carotenoids are integument pigments that often reflect foraging efficiency, disease resistance and body condition. In contrast to the widespread attention this relationship has received in adult birds, the condition dependence of nestling colouration remains an understudied component of animal communication. Here we assess the condition dependence of carotenoid pigmentation in nestling hihi (Notiomystis cincta, an endangered New Zealand bird) and examine the influence of carotenoid supplementation on nestling quality and parental visitation rates. Our results show that carotenoids provided to breeding adult hihi were transferred to their offspring and resulted in an intensified orange-yellow flange colour. After accounting for carotenoid supplementation the parameter that most consistently explained variation in nestling flange colour was nestling tarsus length at 23 days, indicating condition dependence of this trait. We did not, however, detect direct effects of carotenoid supplementation on nestling mass or immune response (or any other fitness parameter measured). Carotenoid supplementation did, however, result in an increased paternal provisioning rate.     

Hormonal manipulation of offspring number: maternal effort and reproductive costs

We used exogenous gonadotropin hormones to physiologically enlarge litter size in the bank vole (Clethrionomys glareolus). This method allowed the study design to include possible production costs of reproduction and a trade-off between offspring number and body size at birth. Furthermore, progeny rearing and survival and postpartum survival of the females took place in outdoor enclosures to capture salient naturalistic effects that might be present during the fall and early winter. The aim of the study was to assess the effects of the manipulation on the growth and survival of the offspring and on the reproductive effort, survival, and future fecundity of the mothers. Mean offspring body size was smaller in enlarged litters compared to control litters at weaning, but the differences disappeared by the winter. Differences in litter sizes disappeared before weaning age due to higher mortality in enlarged litters. In addition to the effects of the litter size, offspring performance was probably also influenced by the ability of the mother to support the litter. Experimental females had higher reproductive effort at birth, and they also tended to have higher mortality during nursing. Combined effects of high reproductive effort at birth and high investment in nursing the litter entailed costs for the experimental females in terms of decreased probability of producing a second litter and a decreased body mass gain. Thus, enlarged litter size had both survival and fecundity costs for the mothers. Our results suggest that the evolution of litter size and reproductive effort is determined by reproductive costs for the mothers as well as by a trade-off between offspring number and quality.     

Determinants of reproductive success in voles: space use in relation to food and litter size manipulation

Spacing behaviour of female mammals is suggested to depend on the distribution and abundance of food. In addition, food limitation has been found to constrain the reproductive success of females. However, whether females maximize their reproductive success by adjusting space use in relation to current food availability and reproductive effort (e.g. litter size) has not been experimentally studied. We examined these questions by manipulating simultaneously food resources (control vs. food supplementation) and litter sizes (control vs. plus two pups) of territorial female bank voles (Clethrionomys glareolus) in large outdoor enclosures. Females with supplementary food had smaller home ranges (foraging area) and home range overlaps than control females, whereas litter size manipulation had no effect on space use. In contrast, the size of territory (exclusive area) was not affected by food supplementation or litter size manipulation. As we have previously shown elsewhere, extra food increases the reproductive success of bank vole females in terms of size and proportion of weaned offspring. According to the present data, greater overlap of female home ranges had a negative effect on reproductive success of females, particularly on survival of offspring. We conclude that higher food availability increases the reproductive success of bank vole females, and this effect may be mediated through lower vulnerability of offspring to direct killing and/or detrimental effects from other females in the population. Moreover, it seems that when density of conspecifics is controlled for, home range sizes of females, but not territoriality, is related to food resources in Clethrionomys voles.     

The problem of optimal clutch size in a tritrophic system: the oviposition strategy of the thistle gallfly Urophora cardui (Diptera,Tephritidae)

The problem of optimal clutch sizes is a central theme in life history theory. Optimal allocation of eggs is especially complicated for insects in tritrophic systems. In this study we analyze some of the processes determining clutch sizes of the thistle gallfly Urophora cardui, a monophagous tephritid fly associated with Cirsium arvense. U. cardui forms multilocular shoot galls, which vary broadly in their size and number of their gall cells. We investigate various fitness consequences of gall size. An analysis of the number of cells per gall (which is correlated with gall diameter and gall weight) showed that in U. cardui there is mutual facilitation rather than larval competition. Increasing numbers of larvae per gall led to a decreasing mortality and increasing larval weight. Larval weight in turn was positively correlated with the probability of survival to adulthood and with adult weight and fecundity. Thus, all fitness parameters measured favoured large galls. Clutch sizes in oviposition experiments were distinctly larger than the number of gall cells of field populations and in cage experiments, suggesting high mortality of eggs and/or early larval instars. There was a significant relationship between the internal structure (i.e., the size of the growing point) of the bud and clutch size, suggesting that U. cardui females are able to measure bud quality and adapt clutch sizes accordingly. Clutch size was positively correlated with the female's age at first oviposition and negatively with the number of previous ovipositions and previously laid eggs. Since the potential egg capacity per female is higher than the average number of larvae it is likely to produce during its short adult lifespan, U. cardui females tend to be time-limited rather than egglimited, which might favour large clutches once an appropriate oviposition site has been located. As the development of the gall and hence the fate of a clutch depends on a number of unpredictable factors, exclusive concentration of eggs in a few large clusters would involve risks which could be avoided by increasing the number of clutches. Therefore we interpret the high variation of clutch sizes in U. cardui as a mixed strategy of bet hedging and gambling.     

Aspects of breeding ecology of the eastern olivaceous warbler (Hippolais pallida)

The breeding ecology of eastern olivaceous warblers Hippolais pallida is poorly known. In this study, we provide data on nest site selection and breeding parameters of the species in a population in northwestern Bulgaria, the only one known to be regularly and heavily parasitized by the common cuckoo Cuculus canorus (hereafter cuckoo). Eastern olivaceous warblers only bred within human settlements of the study area, avoiding seemingly suitable habitat outside them. Nests were built in a wide range of plant taxa but ailanthuses Ailanthus altissima and mulberries Morus spp. were most frequently used (21%). After taking into account the availability of vegetation, there was an apparent preference for several plant taxa but not for mulberries. Mean nest height was 1.65 ± 0.98 (0.53–7.60) m, n = 217, and it varied significantly among different types of nesting substrate. Laying date and clutch size of first breeding attempts averaged 10 June ± 0.98 days, n = 101 and 3.9 ± 0.07 (2–5) eggs, n = 72, respectively. Hatching success, fledging success, and breeding success were 42.5, 86.4, and 36.7%, respectively. The main sources of nest mortality were predation and cuckoo parasitism, with no significant difference in the proportion of nests lost to each. Cuckoo parasitism seemed responsible for the relatively low hatching success in this population.     

Growth and reproductive responses of Laminaria longicruris (Laminariales,Phaeophyta) to nutrient enrichment

A series of comparative culture experiments were conducted in order to determine responses of Laminaria longicruris male and female gametophytes and juvenile sporophytes to several temperatures (5, 10, 15, 20 °C), light levels (10, 35, 75 µmol m^–2 s^–1) and media nitrogen concentrations (0, 20, 100 µM ammonium-nitrogen). Responses were measured as numbers of male and female gametophytes producing gametangia and number of sporophytes produced following fertilization. Both male and female gametogenesis was reduced at 5 and 20 °C versus 10 and 15 °C. At 20 °C gametogenesis inhibition was greater with higher levels of ammonium-nitrogen concentration (100 µM). Sporophyte production was more sensitive to light, temperature and nitrogen concentration than gametogenesis. Production of sporophytes was inhibited completely at 20 °C. At lower temperatures, increasingly higher nutrient concentrations produced greater inhibition of production of sporophytes.     

Aspects of the reproductive biology ofPollicipes pollicipes (Cirripedia; Lepadomorpha) from the southwest coast of Portugal

The pedunculated cirripedePollicipes pollicipes is of considerable economic importance in Portugal, particularly along the southwest coast. Current knowledge of its biology is poor, as evidenced by the paucity of available literature. This paper reports on the reproduction of twoP. pollicipes populations, Castelejo and Zavial, on the southwest coast of the Algarve. Differences in reproductive patterns between the two sites were minor and not statistically significant, despite differences in environmental conditions between the two sites. Adult animals carried fertilised eggs from April throughout September, and the percentage of individuals with egg masses correlated positively with air but not with sea water temperature. Brooding success was greater with average air temperature in excess of 14°C during months of strong upwelling and presumed food availability. Functional ovaries and testes are available in the population all year round but fertilisation does not appear to occur during the winter months. Reproduction ofP. pollicipes on the southwest coast of the Algarve appears closely linked to the upwelling phenomenon with the rate of deposition of gonadal tissue probably being determined by environmental temperature. The cues which determine the cessation of fertilisation remain to be investigated.     

A reproductive tradeoff in an herbivorous lady beetle: egg resorption and female survival

A reproductive tradeoff between current egg production and subsequent survival in a lady beetle, Epilachna niponica, a specialist herbivore on a thistle, Cirsium kagamontanum, was investigated at the two study sites, A and F. Survival of reproductive females decreased consistently from early May until mid-June, but apparently increased thereafter. In contrast, males showed a consistent decrease in survival throughout the reproductive season, without any sign of recovery. Dissection of ovaries of sampled females revealed that egg resorption increased late in the reproductive season, coincident with increased female survival. Reproductive females stopped oviposition immediately after a large flood in 1979 at site F. Two weeks after the habitat perturbation, females resumed oviposition in response to a flush of new leaves on damaged plants. Female survival sharply increased during the nonoviposition period, and declined when egg-laying resumed. Approximately 40% of long-lived reproductive females at site F survived up to the following reproductive season in the next year. Also, some of these long-lived females were observed ovipositing in the following reproductive season. The long-lived reproductive females which had previously invested in reproduction survived equally well as newly emerged females which had not reproduced in summer. These results suggest that there is a reproductive tradeoff between current egg production and subsequent survival. Egg resorption may be an adaptive ovipositional response to habitat perturbation such as flooding, which considerably reduces offspring fitness due to absolute shortage of food. Also, increased female survival accompanied by egg resorption enhances the likelihood of the future oviposition in the second reproductive season, thereby increasing a female's lifetime reproductive success.     

Effects of clutch size manipulations on reproductive behaviour and nesting success in the cooperatively breeding Bell Miner (Manorina melanophrys)

Summary An experimental manipulation of clutch size was carried out on a wild population of the cooperatively breeding Bell Miner (Manorina melanophrys, Meliphagidae) to assess which factor(s) limit clutch size in this species. Results provide some support for the trade-off hypothesis since there is a cost of reproduction for the breeding female in terms of loss of body mass. The breeding female performs most of the nestling care. Clutches of three eggs are also laid during the mid-breeding season which is the period most favourable for breeding (i.e. nestlings grow faster). This evidence also supports the intrabrood competition hypothesis. Clutches that have lost an egg were more likely to be deserted; this may be an antipredator strategy since partial clutch predation has been recorded in the field. Nest predation was high in this study (64.9%), suggesting that many small clutches may be a strategy to decrease the effect of nest predation on reproductive success over the whole breeding season (nest predation hypothesis). Both the trade-off hypothesis and the nest predation hypothesis may apply in this case since they are not mutually exclusive. The size of the attending group did not greatly affect reproductive success in the short term, although if both age structure and size of the group are taken into account, reproductive success can be better predicted.     

The reproductive biology and effective pollinators of the endangered beardtonguePenstemon penlandii (Scrophulariaceae)

Penland's beardtongue, a rare endemic plant of the Colorado Plateau, displays a mixed breeding system. Plants are partially self-compatible but set more fruits when cross-pollinated than when self-pollinated. Fruit production is significantly increased by pollinators. However, in two years of study there was no indication that fruit set was being limited by inadequate pollinator visitation. Pollinator effectiveness was judged by correlating bee behavior at the flowers with analysis of the pollen carried on bee bodies. The most important pollinators were native megachilid bees, particularly in the genusOsmia. The bees that pollinate Penland's beardtongue are essential to its reproduction and must be preserved along with this rare plant.