Female within-nest spawning-site preference in a paternal brooding blenny and its effect on the female mate choice

Female spawning-site preference within a nest and its effect on the female mate choice in a paternal brooding blenny Rhabdoblennius ellipes , were examined in rocky intertidal pools using artificial nests. The number of eggs deposited at the nest entrance site was lower than the number deposited at the center and at deep sites. Moreover, the within-nest difference in the egg developmental stage indicated that eggs at the nest entrance site were deposited after those at the center and deep sites. These results indicated that females prefer to spawn eggs in the center and deep sites rather than at the entrance site. Owing to the higher egg mortality rate at the entrance site, females may avoid spawning at such sites. An analysis of the nests in the study area suggested that the within-nest site preference affects female mate choice that is females avoided nests where only the entrance site was available for spawning and instead spawned in nests where the center and deep sites were available.     

Reproductive behavior and spawning success of female Amblyglyphidodon leucogaster (Pisces: Pomacentridae) from the Red Sea

The reproductive behavior of female whitebelly damselfish, Amblyglyphidodon leucogaster, was investigated in the Gulf of Aqaba, Red Sea over two breeding seasons. Females were promiscuous, mating with 7–10 different males throughout the season. Females lay eggs in distinct batches, defined as the total number of eggs laid in a day. Generally females deposit a batch of eggs with one male (87.2%) and are capable of laying a new batch every other day. Egg batch size averaged 4009 eggs and females laid from 2 to 22 egg batches per season. The variation in spawning success was not correlated to body size. Females preferred to deposit eggs in nests that already contained early stage eggs (0–2 days old). Within a nest, females chose to lay eggs contiguous to the youngest egg batch, regardless if the nest contained either a single batch or multiple batches of different ages. Female within-nest spawning patterns appear to be a consequence of between nest preferences for nests with young eggs. It is proposed that the strong within-nest preference is a consequence of mate selection where females may use new egg batches as a visual cue as part of a copying style. Such a style may reduce the risk of predation and increase feeding opportunities, because less time is expended in mate selection, which would provide additional resources for egg production and ultimately increase female spawning success over the breeding season. This revised version was published online in July 2006 with corrections to the Cover Date.     

Sequential settlement by nesting male and female Baya weaverbirds Ploceus philippinus: the role of monsoon winds

Theory predicts that females should track resources while making decisions about where to reproduce, and males should track females. However, when males create the resource that females require, female settlement follows that of males. In this situation, how do individuals make settling decisions and what are the fitness consequences of these decisions? Here, I test the hypothesis that environmental factors (wind direction and velocity) affect optimal nest placement and orientation, and thus male and female settlement, in Baya weaver Ploceus philippinus colonies. Nests experimentally placed in windward locations with entrance tubes oriented toward the wind were more likely to lose clutches of artificial eggs than were nests placed in other combinations of placement and orientation. Correspondingly, males built their nests strongly clustered on the leeward side of colony trees, with entrance tubes oriented away from the wind. However, females did not discriminate amongst natural nests on the basis of nest placement or orientation. This may be because the effect of wind on the success of natural nests was small and was overwhelmed by other causes of nest loss. I suggest that, in some situations, female preference may remain undetected because the options available to females are constrained by preceding settlement decisions by males. In these cases, it is necessary to manipulate male distribution outside the range of natural variation to gain a better understanding of selective pressures imposed by female preferences.     

Multiple maternity in black-headed gull Larus ridibundus clutches as revealed by protein fingerprinting

Social monogamy with biparental care is the norm in gulls Laridae , but egg colour variation suggests that some nests may contain mixed clutches laid by more than one female. Here we use protein fingerprinting of egg albumen to assess the occurrence of mixed maternity clutches in three colonies of black-headed gulls. Among 160 analysed clutches with >1 egg, 34% contained eggs from more than one female, and 15% of the eggs in clutches >1 came from other females than the major female (laying most eggs in nest). Among clutches with 2–3 eggs 28% were mixed, and among clutches with 4 or more eggs 89% contained eggs from two or more females. There were significantly fewer eggs from the major female in mixed nests (mean=2.06±0.63 SD) than in non-mixed nests (mean=2.82±0.43 SD). In nests without evidence of female conflict, hatching success of minority eggs was similar to that of eggs from the major female (12.5 and 8.4%, respectively). In 21% of mixed maternity nests, one or more minority eggs was buried or punctured, and 25% of eggs from major females were also found evicted, suggesting conflict between females and rejection of eggs. Intra-specific nest parasitism seems the most likely cause of mixed clutches, but there are also other possible causes.     

Nest, but Not Egg, Fidelity in a Territorial Salamander

Egg recognition and subsequent egg brooding are costly forms of parental investment in many species of vertebrates. Life history factors, such as coloniality or risk of brood parasitism, may constrain egg recognition in vertebrates. Female red-backed salamanders ( Plethodon cinereus ) from my study site are territorial and do not share nest sites with other females. They are terrestrial and neither they nor their eggs are likely to be displaced by environmental factors such as flooding. I experimentally tested, in the laboratory, the hypothesis that female red-backed salamanders can discriminate between their own eggs and the eggs of unfamiliar females. Each female was allowed to move about a test chamber containing two clutches of eggs, one clutch with which it was found in the forest and one that had been found with a distant female. Most females remained with one clutch of eggs, which they brooded during the entire observation period. However, they did not significantly prefer to brood their own eggs over the eggs of another female. In a corollary field experiment, I tested whether brooding females that were displaced 1 m from their nest sites would return to their territories and commence brooding behaviour within 3 d. All 10 displaced females returned to their own nest within this time period and were found brooding their eggs. Because female red-backed salamanders at my study site do not tend to share nest sites with other females and because their eggs remain in stationary nests, selection may not have favoured egg recognition. However, the results suggest that female salamanders indirectly recognize their own eggs by actively recognizing their territorial nest sites.     

Reproductive ecology of the jacky dragon (Amphibolurus muricatus): an agamid lizard with temperature‐dependent sex determination

Abstract The jacky dragon, Amphibolurus muricatus (White, ex Shaw 1790) is a medium sized agamid lizard from the southeast of Australia. Laboratory incubation trials show that this species possesses temperature‐dependent sex determination. Both high and low incubation temperatures produced all female offspring, while varying proportions of males hatched at intermediate temperatures. Females may lay several clutches containing from three to nine eggs during the spring and summer. We report the first field nest temperature recordings for a squamate reptile with temperature‐dependent sex determination. Hatchling sex is determined by nest temperatures that are due to the combination of daily and seasonal weather conditions, together with maternal nest site selection. Over the prolonged egg‐laying season, mean nest temperatures steadily increase. This suggests that hatchling sex is best predicted by the date of egg laying, and that sex ratios from field nests will vary over the course of the breeding season. Lizards hatching from eggs laid in the spring (October) experience a longer growing season and should reach a larger body size by the beginning of their first reproductive season, compared to lizards from eggs laid in late summer (February). Adult male A. muricatus attain a greater maximum body size and have relatively larger heads than females, possibly as a consequence of sexual selection due to male‐male competition for territories and mates. If reproductive success in males increases with larger body size, then early hatching males may obtain a greater fitness benefit as adults, compared to males that hatch in late summer. We hypothesize that early season nests should produce male‐biased sex ratios, and that this provides an adaptive explanation for temperature‐dependent sex determination in A. muricatus.     

Mate choice and cannibalism in a natural population of a stream goby,Rhinogobius sp.

Breeding ecology of the stream goby,Rhinogobius sp. LD (Large Dark), was investigated under natural conditions. Males selectively courted females of similar size to lead them to the nests, whereas females followed courting males preferentially when the relative male size was greater. Male-male competition for a female was relatively infrequent and not severe. Developmental stages of eggs and egg numbers in one nest indicated that males receive 1–3 clutches during one breeding cycle. Males guarding multiple clutches frequently ate some of the eggs, but those guarding single clutches rarely did so. Gravid females in the nest also frequently cannibalized eggs laid by a previous female, probably in order to extend the area available for egg deposition. Mate choice in this species is discussed in relation to paternal ability, limitation of available spawning area and the female-biased sex ratio.     

A molecular genetic analysis of the communal nesting of the ostrich (Struthio camelus)

The ostrich breeding system is complex and unique; communal clutches are laid by several females, although only one female, the major female, and the resident territorial male provide parental care. More eggs are laid in the nest than can be incubated and the major female ejects surplus eggs from the incubated central clutch. Microsatellite markers were used to analyse the parentage of communal nests in Nairobi National Park. This revealed that major females contributed a disproportionate number of fertile eggs to the central, incubated clutch and that multiple paternity and maternity within a nest were common; 68.9% of all incubated eggs on a nest were not parented by both the resident territorial male and the major female of that nest. All the males fertilized eggs on the clutches of neighbouring males. Unexpectedly, every major female with her own nest was also simultaneously a minor female with incubated eggs on neighbouring clutches. The relatedness between females laying in the same nest was not significantly different from the population average and significantly less than that between chicks hatched from the same nest.     

Intraspecific nest parasitism in the European starling Sturnus vulgaris

Biochemical genetic markers were used along with conventional methods (abnormal laying sequence/clutch size, unusual egg shape/pigmentation) to identify intraspecific nest parasitism at two British nestbox colonies of the European starling. Between 11 and 37% of first clutches were parasitized during 1977–1979. Parasitic females probably comprised all of the following categories: (1) paired females contesting a nestbox occupied by another pair; (2) previously paired females who had laid a clutch but had been unsuccessful; (3) unpaired females who had copulated with males that already had a mate and nest site; and (4) ‘professional’ nest parasites who distributed at lest some of their eggs in one or more nests other than their own. Although parasitized nests had higher clutch sizes, parasitism led to fewer host young fledging per egg laid, mainly through the eviction of eggs and subsequent nest desertion. Number of parasitic young fledged per egg laid was highest when eggs were laid synchronously with the host, when host clutches were larger, or a smaller number of parasite eggs were added to a nest, thus favouring parasites that distribute their eggs amongst a number of nests. A greater pressure on nest sites may have accounted for the higher levels of parasitism at the Aberdeen colony and for the greater number of parasite eggs laid in a nest. Although most parasitic female starlings appeared to be much less successful than non-parasitic ones, nest parasitism in the starling might evolve directly when one or more of the following advantages are present. (1) There are no constraints on the number of eggs a female may lay but there are constraints on the number of young she may feed adequately. (2) Female survival is increased by having fewer or no eggs/young to care for. (3) Current feeding conditions favour the survival of more young than would be produced by the most common clutch size. Intraspecific nest parasitism is considered to be a first stage in the evolution of interspecific nest parasitism.     

Egg Coloration and Recognition of Conspecific Brood Parasitism in Eastern Bluebirds

Individual eastern bluebird (Sialia sialis) females produce clutches of eggs with unique coloration and older females and females in better body condition lay more pigmented blue‐green eggs. Conspecific brood parasitism in this species is not uncommon and bluebirds occasionally reject what appear to be normal eggs by moving them to the periphery of the nest. I used UV‐visual reflectance spectrometry to objectively measure coloration of eggs and nest material. To estimate the conspicuousness of the trait, I calculated the contrast between eggs and background nest material. I found high achromatic and chromatic contrast between the coloration of eggs and of the nests, suggesting that bluebird eggs are highly conspicuous. To test the hypothesis that expression of blue‐green coloration eggs facilitates recognition of eggs laid by conspecific brood parasites, I cross‐fostered individual eggs into host nests during egg laying and monitored the fate of those eggs. I found no support, however, for the hypothesis that egg coloration facilitates discrimination of parasitic eggs from host eggs.     

Double nesting of the Red-legged Partridge Alectoris rufa

Some Red-legged Partridge females lay two clutches in separate nests, one immediately after the other, which are incubated separately by the male and female of the pair. Pairs remained together during the laying of both clutches so that there were delays between the end of laving and incubation at the first nests. Incubation of the two nests began at approximately the same time but discrepancies of up to ten days occurred. Males incubated first clutches and females usually the second, but probably the first if the second was destroyed during laying. The proportion of surviving eggs which hatched was similar in first and second clutches but declined if the delay between laying and hatching was exceptionally long.
Yearling females began laying late and few seemed to attempt two clutches compared with older females of which 60–80% showed the double nesting habit. A model predicting reproductive success for both sexes in relation to the rate of nest predation during laying, suggested that attempting two clutches rather than one would be disadvantageous at high predation rates. Females would produce more young if their mates incubated their first clutch immediately it was complete rather than accompanying them during the laying of the second. However, males may benefit by this delay, even though it exposes the nest to predators for a longer time, because they are able to guard their mate and prevent other males from mating with her and fertilizing eggs of her second clutch.     

What limits clutch size? A test of the incubation‐capacity hypothesis in a high‐elevation population of Mountain Bluebirds

Most studies of factors that limit the number of eggs that birds lay have focused on the disadvantages of having too many young to feed. Less attention has been paid to the consequences of having a large number of eggs to incubate. The incubation‐capacity hypothesis proposes that females lay as many eggs as they can effectively incubate. We tested this hypothesis in 2018 in a montane population of Mountain Bluebirds (Sialia currucoides). Most females in this population lay five or six eggs; clutches of seven occur, but are rare. We added eggs to some nests, forcing females to incubate seven eggs, while leaving other nests as controls. Among females completing incubation, those with enlarged clutches hatched as many eggs as did control females, and did so in the same amount of time. This was despite an extended period of unusually cold and often wet weather that occurred when many females were incubating. Our results firmly reject the suggestion that females typically lay no more than six eggs because they cannot effectively heat seven eggs. One or more other factors must limit clutch size. One possible factor is suggested by the fact that during the period of inclement weather, more females with enlarged clutches than control females appeared to abandon nests before completing incubation. Because larger clutches require more energy to incubate, females with seven eggs during energetically stressful conditions could more quickly reach the point where they lack sufficient energy for both incubation and self‐maintenance. Such conditions may occur frequently enough in the montane environment that, on average, laying seven eggs results in reduced lifetime reproductive success.     

Why do grebes cover their nests? Laboratory and field tests of two alternative hypotheses

Egg predation is a common feature influencing the reproductive success of open nesting birds. Evolutionary pressure therefore favours building cryptic, inconspicuous nests. However, these antipredatory pressures may be in conflict with thermoregulatory constraints, which select for dry nest material maintaining optimum temperature inside a nest cup during the absence of incubating parents. Here we examined possible trade-offs between nest crypsis and thermoregulation in Little Grebes (Tachybaptus ruficollis), which lay their eggs in floating nests built from wet plant material. As this species regularly covers its eggs with nest material, we experimentally examined (1) the rates of egg predation on covered and uncovered artificial nests and (2) possible thermoregulatory costs from nest covering by comparing temperature and relative humidity changes inside the nest cup. Results revealed that covering clutches is beneficial in terms of deterring predators, because uncovered eggs were more vulnerable to predation. Moreover, covering clutches also had thermoregulatory benefits because the mean temperature and relative humidity inside nest cups covered by dry or wet materials were significantly higher for covered compared to uncovered treatments. Covering clutches in Little Grebes therefore does not pose thermoregulatory costs.     

Brood parasitism,female condition and clutch reduction in the common eider Somateria mollisima

Conspecific brood parasitism (CBP) is an alternative reproductive tactic found in many animals with parental care. Parasitizing females lay eggs in the nests of other females (hosts) of the same species, which incubate and raise both their own and the foreign offspring. The causes and consequences of CBP are debated. Using albumen fingerprinting of eggs for accurately detecting parasitism, we here analyse its relation to female condition and clutch size in High Arctic common eiders Somateria mollissima borealis. Among 166 clutches in a Svalbard colony, 31 (19%) contained eggs from more than one female, and 40 of 670 eggs (6%) were parasitic. In 6 cases an active nest with egg(s) was taken over by another female. Many suitable nest sites were unoccupied, indicating that CBP and nest takeover are reproductive tactics, not only consequences of nest site shortage. Similarity in body mass between female categories suggests that condition does not determine whether a nesting female becomes parasitised. There was no evidence of low condition in parasites: egg size was similar in hosts and parasites, and parasitism was equally frequent early and late in the laying season. Meta‐analysis of this and 3 other eider studies shows that there is a cost of being parasitised in this precocial species: host females laid on average 7% fewer eggs than other females.     

Costs and response to conspecific brood parasitism by colonial red-breasted mergansers

Costs of conspecific brood parasitism (CBP) are expected to be influenced by a species’ life history traits. Precocial birds lay large clutches, and clutches that have been enlarged by CBP can affect host fitness through a longer incubation period, displaced eggs, and lower hatching success. We examined costs and response to CBP by hosts in a population of colonial red-breasted mergansers (Mergus serrator; n?=?400 nests over 8 years) within which 29% of parasitized clutches were enlarged considerably (≥?15 eggs). Length of the incubation period did not increase with clutch size. The mean number of eggs displaced from a parasitized nest during incubation (2.8) was 2×?greater than at an unparasitized nest (1.4). Hatching success declined by 2% for each additional egg in the nest. Thus, for a nest with?≥?15 eggs, one or more fewer host eggs hatch relative to an unparasitized nest with the same number of host eggs, assuming equal probability of success for all eggs. Hosts were 40% more likely to desert nests receiving 2 or 6 experimental eggs relative to unparasitized control nests, although it is unknown whether hens deserting a nest renested elsewhere. Our study indicates that costs of CBP to hosts during nesting may be limited to those red-breasted mergansers incubating the largest clutches (≥?15 eggs), and it raises questions about the adaptive significance of deserting a parasitized clutch.     

FILIAL CANNIBALISM IN THE CORTEZ DAMSELFISH STEGASTES RECTIFRAENUM

Male cortez damselfish, Stegastes rectifraenum, in a central Gulf of California population, ate a large percentage (28.3%) of the clutches they received. This high rate of filial cannibalism permitted the testing of several predictions concerning the types of clutches that should be preferentially eaten and the mating tactics females should employ to reduce cannibalism rates. Males ate clutches that were smaller than average and that were at early stages of development. Experimentally reduced clutches were consumed at higher rates than controls. When multiple clutches were present, males preferentially ate the younger, smaller clutches. Females were more likely to deposit eggs with males who were caring for other early-stage eggs and to avoid males with late-stage eggs. This pattern was accentuated in females that deposited smaller clutches. These results provide evidence that filial cannibalism represents an adaptive response to clutches that do not provide adequate benefits to warrant the costs of parental care.     

Brood Parasitism and Nest Takeover in Common Eiders

Conspecific brood parasitism (CBP) is an alternative breeding tactic that occurs in many brood‐tending animals and can have important fitness effects for both host and parasite. We use protein fingerprinting of egg albumen to distinguish the eggs from different females and to estimate the frequency, pattern and tactics of CBP and other forms of mixed maternity in a Hudson Bay population of common eiders (Somateria mollissima sedentaria). Mixed clutches, containing eggs from more than one female, occurred in 31% of the 86 nests studied that progressed to clutch completion. Other females than the host laid 8% of the eggs. In 11 (41%) of the mixed clutches another female laid before the host started laying, corroborating the hypothesis that takeover of nests started by other females accounts for many of the mixed clutches in this population. Our results also indicate that traditional non‐molecular methods of identifying foreign eggs may considerably underestimate the frequency of mixed clutches.     

The influence of oxygen stress on female choice for male nest structure in the common goby

This study examines the role of dissolved oxygen in modifying female mate choice in the common goby, Pomatoschistus microps. Experimental manipulations of nests showed that under normal (saturated) oxygen conditions, females preferred to spawn in nests with the most elaborate construction, that is, those with the most sand on top and the smallest entrance. Such manipulated nests proved less vulnerable to detection by an egg predator, the shore crab, Carcinus maenas, but these small entrances may reduce oxygenation of eggs. Indeed, in a low oxygen environment females with a choice between nests of high build and nests of low build did not significantly prefer either. This may have been due to weakened preference or perhaps detection of a mismatch between the male's parental quality and the increased ventilation needs of eggs when nest entrance sizes are small and ambient oxygen levels are low. When nests were not manipulated, those males that built small entrances fanned their eggs more often, such that hatching success in the absence of a predator was not related to the initial size of the nest entrance area. Thus, under normal oxygen conditions males may initially build nests with the smallest entrance they are capable of ventilating successfully, and females choosing such males gain from nest camouflage. Under low oxygen, the risk that manipulated males may be unable to compensate fully may outweigh such benefits, and females may use other criteria that signal willingness to provide parental care. Copyright 1999 The Association for the Study of Animal Behaviour.     

Laying sequence,diploid males,and nest usurpation in the leafcutter bee,Megachile rotundata (Hymenoptera: Megachilidae)

Variations from the normal female-male sequence of eggs in nests of the leafcutter bee,Megachile rotundata, were examined. Three alternatives were considered: Out-of-sequence males (i) were diploids, (ii) were the result of supersedure of nests or intraspecific brood parasitism, or (iii) were the result of females occasionally laying a male-female sequence. Electrophoretic data provided definitive evidence of diploid males and of multiple females laying in 7 of 18 nests. In the others, the remaining explanation is that females occasionally lay male eggs before female eggs in a nest.     

Maternal Care in the Burrower Bug <Emphasis Type="Italic">Adomerus triguttulus</Emphasis>: Defensive Behavior

Some burrower bugs (Heteroptera: Cydnidae) show complex patterns of maternal care, including defense against predators and the provisioning of food to nymphs. Recently, the subsocial cydnid bugs have attracted the interest of researchers as model systems to study the behavioral ecology of parental investment. However, there have been few attempts to quantify the fitness benefits of maternal behavior other than provisioning. Here, we examined the maternal behavior of Adomerus triguttulus and its adaptive significance in terms of offspring survival in the field. A. triguttulus young depend on fallen nutlets of myrmecophorous mints, Lamium spp. Under field conditions, females attend offspring, from eggs to second instar nymphs, in nests on the ground under the litter. When disturbed, the females showed aggressive responses against the source of disturbance. The females often carried spherical clutches of eggs away from the nest when heavily disturbed. Female-removal experiments in the field demonstrated a defensive function of the female behavior; predators, such as ants, attacked egg clutches without females and the clutches often disappeared during the experiment. Egg clutches without females sometimes also suffered from fungal infection. Selective factors on maternal defensive behavior in A. triguttulus are discussed in terms of habitat properties possibly emerging from insect–plant associations.