Costs of large communal clutches for male and female Greater Rheas Rhea americana

The breeding system of the Greater Rhea Rhea americana is almost unique among birds as it combines harem polygyny and sequential polyandry, with communal egg-laying and uniparental male care. In this species, large communal clutches (more than 30 eggs) are rare and have a lower hatching success than smaller clutches. Here we analyse the proximate causes of hatching failures and the costs of large communal clutches (and therefore the costs of extensive polygyny) for males and females. We evaluated if length of the nesting period, egg viability, egg losses during incubation and male parental activity at the nest were affected by clutch size. We also evaluated if chicks hatched from large clutches have a lower survival during the first 2 months after hatching. Large clutches had longer nesting period and lower hatching success, mainly as a result of bacterial contamination of the eggs and increased hatching asynchrony. In addition, large clutches tended to lose more eggs as a result of accidental breakage or predation. Male activity at the nest and chick survival were not related to clutch size. Low hatching success, nest predation risk and energetic costs associated with large clutches penalize females that join large harems and males that accept additional eggs into the nest.     

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.     

Male body size and brooding success under natural conditions in the freshwater Japanese goby, Rhinogobius sp. OR (orange type)

Males of the Japanese goby Rhinogobius sp. OR (orange type) establish nests under stones in streams. They provide care to egg masses until hatching. We carried out field observations and a field experiment in a natural stream to examine the effect of male body size on brooding success. An experiment excluding guarding males indicated that their presence has a positive effect on nest defense. Field data showed that male body size was positively correlated with the size of the egg mass that was initially found at the nest. Furthermore, a sequential increase in egg mass size seen in the nests of larger males indicated that multiple females spawned during a single breeding cycle of larger males. To examine male brooding success, two dependent variables, egg mass persistence (days from initial egg mass to disappearance or hatching) and hatching success were analyzed. The results indicated that the variation found in both dependent variables was significantly explained by two independent variables, namely, initial egg-mass size and the number of matings. It is also suggested that male body size indirectly influenced brooding success through positive correlations with these independent variables.     

Artificial incubation of eggs of the green iguana (Iguana iguana)

Successful hatching of large numbers of artificially incubated eggs from the green iguana, Iguana iguana, are reported. Gravid females were captured at a nesting site in Summit Gardens, Soberania, Parque Nacional, Panama, and released into an enclosure with an artificial nesting area. Females dug their own tunnels and nest chambers or used artificial nest chambers for egg deposition. Eggs (n = 829) from 21 clutches were removed from the nests and artificially incubated. Average hatching success per clutch was 94.6% and the mean clutch incubation time was 92.1 days. A distinct odor was noticed in the incubation containers several days before the eggs began to hatch. At this time, the substrate layer was removed exposing the top surface of all eggs, and the eggs desiccated to some degree prior to hatching. High incubation temperature, hydric conditions, and egg arrangement are all implicated as contributing factors in low hatching success in previous studies using artificial incubation techniques.     

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.     

Induced responses of cherry trees to periodical cicada oviposition

Cherry trees (Prunus serotina) responded to oviposition by periodical cicadas (Magicicada spp.) by depositing gum at the egg nest. The proportion of cicada eggs that hatched successfully was significantly reduced at egg nests with visible gum compared to non-gummed egg nests. The number of egg nests with gum increased in proportion to the total number of egg nests on a tree. The probability of an egg nest having visible gum increased as the total number of egg nests increased. Mortality at hatching due to gum deposition increased as a direct density-dependent function of the number of cicada eggs laid in the tree. Although statistically significant, this relationship was weak and appeared to hold only at densities above 100 egg nests per tree. Gum deposition is discussed as an induced plant response to cicada attack. A cherry may reduce the number of cicada nymphs that will parasitize it up to the next oviposition period (17 or 13 years later) by reducing cicada hatching through gum deposition at the site of oviposition.     

Clutch size limitation in waders: experimental test in redshank Tringa totanus

Several hypotheses have been raised to explain the upper limit of clutch size at four eggs in waders (suborder Charadrii), which may play an important role in the evolution of the variety of mating and parental care systems in this group. Experimental tests of the hypotheses have produced conflicting results. It was recently suggested that the combined effects of several incubation costs of a larger clutch suffice to limit its size to four eggs in this group. Here we test the incubation-limitation hypothesis in a field experiment, in redshank Tringa totanus. We created five-egg clutches by adding one egg from another nest to a just completed four-egg clutch. Four-egg control clutches were created by replacing one of the eggs by an egg from another nest. All egg removals, additions and replacements were done before incubation started. Incubation time in five-egg clutches increased by 1 day to 24.3ǂ.23 days, compared to 23.3ǂ.32 days in four-egg clutches. Egg hatchability and nest predation rates did not differ significantly between treatments. On average five-egg clutches produced one extra chick at hatching (4.5ǂ.26 chicks) compared to four-egg clutches (3.5ǂ.27 chicks). Also when several additional costs from incubating enlarged clutches are added, redshanks by laying a fifth egg would on average increase their reproductive success at hatching by an estimated 22%. The incubation-limitation hypothesis therefore is clearly rejected in this species. Possible mechanisms behind the four-egg clutch limit in waders and ways of testing the alternatives are discussed.     

Maternal behavior and clutch manipulation in the ring-legged earwig (Dermaptera: Carcinophoridae)

The expression and maintenance of maternal behavior in the earwig,Euborellia annulipes, was examined through manipulation of clutch size, age, and species and through observations of interactions between brooding females. Females underwent discrete gonadotrophic cycles culminating in oviposition of first clutches that were highly variable in size. Neither the head capsule width nor the age of the mother was correlated with clutch size. Maternal care extended through embryogenesis and for the week following hatching. Clutch removal significantly shortened the interclutch interval, indicating that the presence of brood inhibited the onset of the second gonadotrophic cycle. Brooding females readily accepted replacement clutches of the same age. Thus, mothers did not appear to distinguish their own eggs from those of other females. Experimental doubling of clutch size did not significantly reduce the proportion hatching or fledging. In contrast, reducing clutch size diminished the percentage successfully fledging. Manipulation of clutch age resulted in reduced hatching/fledging success. Placing two females, each with newly laid clutches, in the same cage usually resulted in egg transfer from the nest of one female to that of the other within 12 h. Nests of females with larger forceps were significantly more likely to contain both clutches. When mothers with first clutches were paired with mothers with third clutches, eggs were more likely to be transferred to the nest of the older female.E. annulipes females with newly laid clutches appeared to accept as replacement clutches eggs of the earwigDoru taeniatum. Alien clutches were maintained for the typical duration of embryogenesis; however, noD. taeniatum hatchlings were observed.     

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.     

Does paternity or paternal investment determine the level of paternal care and does female choice explain egg stealing in the fifteen-spined stickleback?

An earlier field study on the fifteen-spined stickleback (Spinachia spinachia) showed that frequent male—male interactionsresult in high frequencies of sneaking and egg stealing. Moreover,sneaking behavior was performed not only by males adoptingalternative mating strategies, but also by males with theirown nests. The advantage of sneaking is easily understood, but it is more difficult to explain the evolutionary benefitof stealing eggs from other males. I investigated whether malessuffering from sneaking adjust their paternal effort in relationto their degree of paternity. I also examined whether femalesprefer males that have more eggs in their nests, as this couldexplain egg stealing. There was no relationship between thedegree of paternity and fanning activity, hatching success,or nest defense. However, the older the eggs become, the morethe males increase their attack rate toward potential egg predators(goldsinny wrasse and shore crabs). Thus, males adjusted theirlevel of defense to the amount of energy and time already investedin the clutch. Females did not prefer males with more eggs intheir nests. On the contrary, females preferred males withreduced clutches over males with enlarged clutches. Therefore,female choice is unlikely to be a driving force behind eggstealing in this species.     

THE BIRDS OF LANGEBAAN LAGOON

Data are presented on breeding success of Red Bishops (Euplectes orix) collected over four breeding seasons at a colony in the Addo Elephant National Park, Eastern Cape, South Africa. Overall hatching and fledging success were 53.8% and 26.0% of all eggs laid, respectively, and the overall mean number of fledglings per breeding attempt was 0.77. Hatching and fledging success varied significantly among seasons, with both clutch and brood losses due to predation being the main reason for the observed differences. Hatching success also differed significantly among clutch sizes, being highest for four-egg clutches (63.2%), intermediate for three-egg clutches (55.5%) and lowest for two-egg clutches and five-egg clutches (33.2% and 34.3%, respectively). However, fledging success was not significantly different among clutch sizes. The mean number of fledglings per breeding attempt was 0.44 for two-egg clutches, 0.80 for three-egg clutches, 1.10 for four-egg clutches, and 0.57 for five-egg clutches. The height of accepted nests (i.e.nests in which at least one egg was laid) was significantly lower than the height of nests not accepted. In addition, accepted nests in which eggs hatched and young fledged were significantly lower than accepted nests in which no eggs hatched and no young fledged. These overall effects of nest height on nest acceptance and hatching and fledging success were, however, due only to nests built above water, since no such effects were found when nests built above ground (i.e.on dry land) were analysed separately. I detected no effect of nest coverage on the probability of a nest being accepted, nor was there any effect of nest coverage on hatching or fledging success. Nests above water were significantly more likely to be accepted than nests above ground; however, hatching and fledging success of nests that were accepted did not differ significantly between nests built above water and those built above ground.     

4. A FIELD NATURALIST'S PEREGRINATIONS. BIRD NOTES FROM A NORTHERN SECTOR OF SOUTHERN RHODESIA—Part III

Boyer, H. J. 1988. Breeding biology of the Dune Lark. Ostrich 59:30-37.

The peak of the breeding season of the Dune Lark Mirafra erythrochlamys occurred in January and February and was not dependent on rainfall. Most nests were domed, although one undomed nest was recorded. Ninety-one percent of clutches were of two eggs (mean = 1,9; range 1–2; n = 11). The eggs are described and measurements given. Incubation, by the female only, began with the laying of the second egg, and hatching occurred after 13–14 days. Growth and development of nestlings are described. The young left the nest after 12–14 days, and post-nestling parental care lasted for approximately one month. Sixty-one percent of eggs hatched. and 28% produced young which successfully left the nest. Most losses of eggs and young were the result of predation.     

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.     

Postural Shifts During Egg‐Brooding and Their Impact on Egg Water Balance in Children’s Pythons (Antaresia childreni)

Parental care typically consists of distinct behavioral components that are balanced to address the multiple needs of offspring. Female pythons exhibit post‐oviposition parental care in which they coil around their parchment‐shelled eggs throughout incubation (40–80 d). Subtle postural shifts during egg‐brooding facilitate embryonic gas exchange but may entail hydric costs to the clutch. This study used a simple behavioral model to (1) further quantify the costs and benefits of specific parental behaviors to developing offspring and (2) determine the influence that developmental stage and relative clutch mass have on parental behavior. Although previous research has demonstrated that egg‐brooding as a whole reduces clutch water loss, we hypothesized that egg‐brooding female pythons specifically adopt a tightly coiled posture to conserve embryonic water, but must make postural adjustments to enhance gas exchange between the clutch and nest environments at the cost of increased clutch water loss. We measured rates of water loss in brooding Children’s pythons (Antaresia childreni) and their respective clutches (i.e., brooding units) and monitored changes in brooding posture. We conducted serial trials to elucidate the effect of developmental stage on postural adjustments and water loss. Results demonstrated that the proportion of time females spent in a tightly coiled posture was inversely related to mean water loss from the brooding unit. Analyses indicated that slight adjustments in posture led to bursts in brooding unit water loss. Indeed, brooding unit water loss during postural adjustments was significantly higher than during tight coiling. These findings imply that python egg‐brooding provides an adjustable diffusive barrier that leads to discontinuous gas exchange, which minimizes clutch water loss. Because females with larger relative clutch masses spent more time tightly coiled, egg‐brooding female pythons may use a ‘water first’ strategy in which they intentionally conserve clutch water at the cost of reduced embryonic respiratory gas exchange.     

Countertactics of the japanese aucha perchsiniperca kawamebari against brood parasitism by the japanese minnowpungtungia herzi

Japanese aucha perch males taking care of eggs and fry are often brood-parasitized by a Japanese minnow. When the minnows are ovipositing in the perch nests, some of the perch eggs in the nests are often eaten by other fishes. The minnows’ eggs are subsequently guarded by the perch males until hatching. Countertactics of the perch against such brood parasitism were examined in a river of western Japan. Perch males did not employ any countertactics such as ejection of parasites’ eggs or nest desertion. On the other hand, perch females avoided spawning in nests with greater numbers of perch eggs and produced smaller clutches during the parasitic period than in the non-parasitic period. These behavior can be regarded as anti-parasitism tactics because minnows prefer to spawn in nests with higher numbers of perch eggs.     

Egg attendance and brooding by males of the giant water bugLethocerus medius (Guerin) in the field (Heteroptera: Belostomatidae)

Males of the giant water bug Lethocerus medius(Guerin) typify their monobasic subfamily, the Lethocerinae, in that they do not brood eggs attached to their backs as do males of all members of the subfamily Belostomatinae. Exclusive male parental investment as expressed in the Belostomatinae is extremely rare behavior among animals, and evolution of the trait is obscure. Lethocerus mediusmales apparently remain with their mates through oviposition and are consistently found in attendance of eggs after the female has departed. This behavior may enhance paternity assurance at no cost in opportunity for polygyny. Two double clutches of eggs were found, from which we infer the potential for polygynous matings and shared parental investment. Male L. mediusbrood attended egg clutches above the surface of the water, where they may moisten them, shade them, and defend them against predation. Egg attendance/brooding by L. mediusand other Lethocerusspecies may represent a plesiomorphic state from which paternal back- brooding evolved in the Belostomatinae.     

ON THE LIFE-EXPECTANCY OF THE MATOPOS BLACK EAGLES

Oatley, T. B. 1982. The Starred Robin in Natal, Part 3: Breeding, populations and plumages. Ostrich 53: 206–221 The female Starred Robin Pogonocichla stellata constructs a domed nest of moss and dead leaves usually on sloping ground and well concealed in the herb layer. The normal clutch is three eggs laid on consecutive days. Incubation usually starts with the laying of the third egg. The mean size of 138 eggs was 22 x 16 mm. The female incubates the eggs for 16 to 18 days and intermittently broods the young for the fist five of the average 14-day nestling period. Both sexes feed the young from the time of hatching and parental care lasts for some 42 days after leaving the nest. Eggs are laid from October to December with 63% of clutches started in November. Data on sex ratios indicate a surplus of adult males in the population and annual survival rates are estimated at 0,84 for males and 0,76 for females. 51% of eggs laid in 60 nests give rise to fledged young. About 21.3% of eggs laid produce adults. The level of brood parasitism by cuckoos is relatively low. Most adult mortality occurs outside of the breeding seasons. Chilling and overnight starvation from January to March when incidence of late afternoon thunderstorms is highMaycause significant mortality. The subadult plumage appears to confer crypsis and enable the immature bird to reside in adult territories without harassment. AdultsMaybenefit through an effective reduction in competitive stress.     

Parental quality and egg size affect chick survival in a precocial bird, the lapwing Vanellus vanellus

Effects of egg size and parental quality on lapwing Vanellus vanellus chick survival were studied in southwestern Sweden over 6 years. Chicks from large eggs were heavier at hatching and survived significantly better than those from small eggs. To control for the confounding effect of parental quality on egg size and chick survival, we performed a cross-fostering experiment during 2 years, exchanging clutches between nests with large and small eggs. In control clutches, chicks from large eggs survived better than those from small eggs, but we found no significant difference in chick survival between exchanged clutches. Thus, egg size did not affect chick survival independently of parental quality. Fledging success increased with parental age and/or experience, and with female body mass. Hence, both egg size and parental quality affect chick survival in the lapwing. Received: 22 February 1996 / Accepted: 30 September 1996     

The lack of influence of exogenous testosterone on male parental behavior in a neotropical frog (Eleutherodactylus): a field experiment

Males of the Puerto Rican frog Eleutherodactylus coqui exhibit prolonged parental care (nest attendance and defense) of direct-developing eggs in nonaquatic nest sites. Earlier work showed that plasma androgen levels are significantly lower in males caring for eggs than in noncaring, calling males. In this study, we investigated the effects of elevated plasma androgens on parental care behavior by implanting either testosterone (T) or placebo (P) pellets in parental males brooding eggs in the field. Plasma androgen levels, assessed 4 days after pellet implantation, were six times higher in T males than in P males. Despite this increase, T males exhibited no changes in nest fidelity or brooding behavior. The results indicate that lower androgen levels, at least those corresponding to a decline in T during parental care, are not necessary to maintain parental behavior in this tropical frog.     

Why egg-caring males of Isaza (<Emphasis Type="Italic">Gymnogobius isaza</Emphasis>, Gobiidae) refuse additional females: preliminary field observations

Like many other gobies, males of the Isaza goby, Gymnogobius isaza endemic to Lake Biwa, Japan, conduct parental care of eggs at nests, and females are likely to choose mates while visiting nests. The reproductive strategy should induce polygyny, but Isaza males never accept additional females in one breeding cycle. Sampling data of broods indicated that the egg mass areas were much smaller than the nest sizes, suggesting that nest size is not the limiting factor in obtaining further eggs from additional females. The brood size greatly decreased as the duration of care progressed. Few individuals including caring males ate eggs, and heterospecific egg predators were rarely observed. Sixty percent of egg masses at the middle stages of egg development were infected with aquatic fungi, some being covered with a fungus mat that drastically reduced survivorship. Infected egg masses contained more eggs than non-infected ones at the same stage, indicating that large egg masses are prone to be frequently destroyed by fungi. It is likely that the activity of parental males is lowered during the long care periods at low water temperature in early spring. Such lowered activity of caring males might be responsible for infections in large broods that would have needed more care. We propose the hypothesis that male rejection of additional females may be related to optimal brood size, which will be less susceptible to fungus infection and produce more hatching young than otherwise. This hypothesis will explain not only male avoidance of additional females but also some unique reproductive behaviors of this fish such as some females spawning of a portion of of the mature eggs in one nest.