Variation in reproductive output of marine turtles

Most marine turtle species are non-annual breeders and show variation in both the number of eggs laid per clutch and the number of clutches laid in a season. Large levels of inter-annual variation in the number of nesting females have been well documented in green turtle nesting populations and may be linked to environmental conditions. Other species of marine turtle exhibit less variation in nesting numbers. This inter-specific difference is thought to be linked to trophic status. To examine whether individual reproductive output is more variable in the herbivorous green turtle (Chelonia mydas Linneaeus 1758) than the carnivorous loggerhead (Caretta caretta Linneaeus 1758), we examined the nesting of both species in Cyprus over nine seasons. Green turtles showed slower annual growth rates (0.11 cm year⁻¹ curved carapace length (CCL) and 0.27 cm year⁻¹ curved carapace width (CCW)) than loggerhead turtles (0.36 cm year⁻¹ CCL, 0.51 cm year⁻¹ CCW). CCL was highly correlated to mean clutch size in both green (R²=0.51) and loggerhead turtles (R²=0.61) and maximal clutch size of green turtles (R²=0.58). Larger females did not lay a greater number of clutches or have a shorter remigration interval than smaller females of either species. On average, the size of green turtle clutches increased and that of loggerhead turtles decreased as the season progressed. Individual green turtles, however, produced more eggs per clutch through the season to a maximum in the third or fourth clutch. In loggerhead turtles, clutches 1-4 were very similar in size but the fifth clutch was 38% smaller than the first. No individuals of either species were recorded laying more than five clutches. Green turtles may not be able to achieve their maximum reproductive output with respect to clutch size throughout the season, whereas only loggerhead turtles laying five clutches (n=5) appear to become resource depleted. Green turtles nesting in years when large numbers of nests were recorded laid a greater number of clutches than females nesting in years with lower levels of nesting.     

On a Poorly Known Rookery of Green Turtles (<Emphasis Type="Italic">Chelonia mydas</Emphasis>) Nesting at the Chabahar Beach,Northeastern Gulf of Oman

There is a poorly known rookery of green turtles (Chelonia mydas) nesting in sandy beaches of Chabahar town, northeastern Gulf of Oman, Iran.This study has been carried out to evaluate nesting activity of this small rookery in 2014 nesting season (June to October). In this study, total clutches were collected and transferred to an artificial hatchery. The peak of nesting occurred from the third parts of August to the end of September. Mean CCL was 106.3 ± 6 cm and mean CCW was 94.5 ± 5 cm. Females laid on average of 99.42 ± 47.8 eggs per clutch. The mean inter-nesting interval was 18.5 days. The observed clutch frequency was 3.4. Mean hatching success was 36.63 ± 4.1%. The incubation period was 61.07 ± 5.4 days. Nest status evaluation represented that major causes for the failure was unhatched egg with no obvious embryo followed by unhatched eggs with obvious but undeveloped embryos > unhatched eggs with developed embryos > hatched eggs but dead embryos. The results achieved in this study are a valuable contribution to cognition of the reproductive ecology of the green turtle population globally and regionally.     

Not putting all their eggs in one basket: bet‐hedging despite extraordinary annual reproductive output of desert tortoises

Bet‐hedging theory makes the counter‐intuitive prediction that, if juvenile survival is low and unpredictable, organisms should consistently reduce short‐term reproductive output to minimize the risk of reproductive failure in the long‐term. We investigated the long‐term reproductive output of an Agassiz's desert tortoise (Gopherus agassizii) population and conformance to a bet‐hedging strategy of reproduction in an unpredictable but comparatively productive environment. Most females reproduced every year, even during periods of low precipitation and poor germination of food plants, and the mean percentage of reproducing females did not differ significantly on an annual basis. Although mean annual egg production (clutch size × clutch frequency) differed significantly among years, mean clutch size and mean clutch frequency remained relatively constant. During an El Niño year, mean annual egg production and mean annual clutch frequency were the highest ever reported for this species. Annual egg production was positively influenced by maternal body size but clutch size and clutch frequency were not. Our long‐term results confirm earlier conclusions based on short‐term research that desert tortoises have a bet‐hedging strategy of producing small clutches almost every year. The risk of long‐term reproductive failure is minimized in unpredictable environments, both through time by annually producing multiple small clutches over a long reproductive lifespan, even in years of low resource availability, and through space by depositing multiple annual clutches in different locations. The extraordinary annual reproductive output of this population appears to be the result of a typically high but unpredictable biomass of annual food plants at the site relative to tortoise habitat in dryer regions. Under the comparatively productive but unpredictable conditions, tortoises conform to predictions of a bet‐hedging strategy of reproduction with relatively small but consistent clutch sizes. Published 2015. This article is a U.S. Government work and is in the public domain in the USA, Biological Journal of the Linnean Society, 2015, 115 , 399–410.     

Multiple paternity assessed using microsatellite markers, in green turtles Chelonia mydas (Linnaeus, 1758) of Ascension Island, South Atlantic

Paternity was determined for three clutches and up to 20 offspring per clutch in the green turtle (Chelonia mydas) from Ascension Island, South Atlantic, using microsatellite markers. All three clutches were sired by at least two different males. The results were compared with those of previous studies of multiple paternity in turtles. No significant difference among studies was observed in the mean contribution of the males siring the largest proportion of progeny per clutch. The present study also provides evidence for segregation distortion (meiotic drive) in turtles.     

Strong male‐biased operational sex ratio in a breeding population of loggerhead turtles (Caretta caretta) inferred by paternal genotype reconstruction analysis

Characterization of a species mating systems is fundamental for understanding the natural history and evolution of that species. Polyandry can result in the multiple paternity of progeny arrays. The only previous study of the loggerhead turtle (Caretta caretta) in the USA showed that within the large peninsular Florida subpopulation, multiple paternity occurs in approximately 30% of clutches. Our study tested clutches from the smaller northern subpopulation for the presence of multiple paternal contributions. We examined mothers and up to 20 offspring from 19.5% of clutches laid across three nesting seasons (2008–2010) on the small nesting beach on Wassaw Island, Georgia, USA. We found that 75% of clutches sampled had multiple fathers with an average of 2.65 fathers per nest (1–7 fathers found). The average number of fathers per clutch varied among years and increased with female size. There was no relationship between number of fathers and hatching success. Finally, we found 195 individual paternal genotypes and determined that each male contributed to no more than a single clutch over the 3‐year sampling period. Together these results suggest that the operational sex ratio is male‐biased at this site.     

Metabolic heating and the prediction of sex ratios for green turtles (Chelonia mydas)

We compared incubation temperatures in nests (n=32) of the green turtle (Chelonia mydas) on Ascension Island in relation to sand temperatures of control sites at nest depth. Intrabeach thermal variation was low, whereas interbeach thermal variation was high in both control and nest sites. A marked rise in temperature was recorded in nests from 30% to 40% of the way through the incubation period and attributed to metabolic heating. Over the entire incubation period, metabolic heating accounted for a mean rise in temperature of between 0.07 degrees and 2.86 degrees C within nests. During the middle third of incubation, when sex is thought to be determined, this rise in temperature ranged between 0.07 degrees and 2.61 degrees C. Metabolic heating was related to both the number of eggs laid and the total number of hatchlings/embryos produced in a clutch. For 32 clutches in which temperature was recorded, we estimate that metabolic heating accounted for a rise of up to 30% in the proportion of females produced within different clutches. Previous studies have dismissed any effect of metabolic heating on the sex ratio of marine turtle hatchlings. Our results imply that metabolic heating needs to be considered when estimating green turtle hatchling sex ratios.     

Genetic structure and diversity of green sea turtle (Chelonia mydas) from South China Sea inferred by mtDNA control region sequence

We analyzed 88 control region sequences of green sea turtle (Chelonia mydas) from around Hainan Island in the South China Sea. These sequences had a length of 489 bp and revealed 8 mtDNA haplotypes of which four haplotypes (CMC1, CMC4, CMC7, and CMC8) had not been discovered before. Haplotype diversity (h) and nucleotide diversity (π) were 0.45 ± 0.054 and 0.0035 ± 0.0014, respectively. Neighbor-Joining tree based on control region sequences revealed that genetic relationship between green sea turtles from the South China Sea and from Japan Sea were very close. Clustering relationship based on control region sequences indicated that the South China Sea is an important breeding site and feeding habitat for green sea turtles, which connects with the Middle East Pacific, the Southwest Pacific and the Indian Ocean.     

Mating system, multiple paternity and effective population size in the endemic flatback turtle (Natator depressus) in Australia

In recent years, genetic studies have been used to investigate mating systems of marine turtles, but to date no such research has been conducted on the flatback turtle (Natator depressus). This study investigates paternity of flatback turtle clutches at two rookeries in Queensland, Australia; Peak Island (Keppel Bay), and Mon Repos (Bundaberg). In the 2004–2005 nesting season, tissue samples were taken from either single or multiple clutches (n = 16) of nesting females (n = 8) representing a sampling effort ranging from 25% to 50% offspring per nest. Determination of the extent of multiple paternity was done using a comparative approach that included initial inferences based on observed alleles, Chi-square tests for deviations from Mendelian expectations, and three software programs (PARENTAGE1.0, GERUD2.0 and MER3.0). Results varied depending on the approach, but by calculating a consensus value of the output from these different methods, the null hypothesis of single paternity could be rejected in at least 11 of the 16 clutches (69%). Multiple paternity was thus observed in the clutches of six of nine females (67%), with two or three fathers being the most likely outcome. Analyses of successive clutches illustrated that paternal contribution to clutch fertilization can vary through time, as observed for two females. This first evidence regarding the mating system of flatback turtles indicates that multiple paternity is common in this species and that the observed frequency of multiple paternity is among the higher values reported in marine turtle species. Application of these results to estimates of effective population size (N _e) suggests that population size may have been relatively stable over long periods. Continued monitoring of population dynamics is recommended to ensure that future changes in the east coast can be detected.     

Factors influencing reproductive performance of northern bobwhite in South Florida

Reproductive success is a critical component of individual fitness, and also an important determinant of growth rates of populations characterized by early maturity and high fecundity. We used radiotelemetry data collected during 2003–2008 to estimate reproductive parameters in a declining northern bobwhite (Colinus virginianus) population in South Florida, and to test hypotheses regarding factors influencing these parameters. The overall clutch size was 12.10 ± 0.22, but females laid more eggs in their first clutch (12.43 ± 0.24) than in subsequent clutches (10.19 ± 0.53) within a nesting season. Daily nest survival was higher for first (0.966 ± 0.003) than subsequent nests (0.936 ± 0.011). Hatchability (proportion of laid eggs that hatched conditional upon nest survival to hatching) was 0.853 ± 0.008, but was higher for nests incubated by females (0.873 ± 0.009) than those incubated by males (0.798 ± 0.018). The proportion of individuals attempting a second nest was 0.112 ± 0.024 and 0.281 ± 0.040 when the first nest was successful and failed, respectively. Hatchability was lower when the nesting habitat was burned the previous winter. We found no evidence that food strip density (a management practice to provide supplemental food) influenced any of the reproductive parameters. Mean summer temperature affected hatchability, nest survival, and proportion of nests incubated by males. Overall, the reproductive output in our study population was lower than that reported for most other bobwhite populations, indicating that low reproductive performance may have contributed to bobwhite population declines in our study site. These results suggest that current management practices, particularly those related to habitat and harvest management, need careful evaluation.     

Impact of Clutch Relocation on Green Turtle Offspring

ABSTRACT For species with temperature-dependent sex determination, such as marine turtles, global climate change poses numerous threats. At the nesting beach, rising temperatures are predicted to further skew already female-biased sex ratios and increase embryonic mortality; sea-level rise and resultant coastal squeeze may leave few alternative breeding habitats in developed regions. As a result, clutch relocation, a commonly used management tool to reduce egg loss, may become necessary for safeguarding populations. Although studies have examined the impact of relocation on clutch success, few have examined the impact of this practice on the sex or phenotypic characteristics of hatchlings produced. We used a randomized block design experiment to examine effects of relocation on green turtle (Chelonia mydas) clutches. We compared hatching success, thermal conditions, and size (length and mass) of hatchlings from in situ control clutches with those subjected to 2 relocation methods, while controlling for maternal and other environmental effects. Relocated clutches did not vary significantly from control clutches in incubation temperature or inferred sex ratios during the critical middle third of incubation when sex is thought to be determined. Hatchling size was also unaffected by relocation. Both relocation methods, however, resulted in a 20% reduction in hatching success in comparison to in situ clutches. Clutch relocation is, however, likely to affect the population primary sex ratio, when clutches are relocated from sites in proximity to the sea where tidal inundation is a threat. Here, cooler conditions are likely to produce more males than are the warmer female-producing temperatures higher up the beach. For clutches at risk, relocation is a viable process and does not appear to affect hatchling size or predicted sex ratios if relocation sites are selected in areas utilized by other females. We urge caution, however, when moving clutches from potentially male-producing sites, particularly given predicted impacts of climate change on already female-biased sex ratios.     

Measuring temporal variation in reproductive output reveals optimal resource allocation to reproduction in the northern grass lizard, Takydromus septentrionalis

We measured the reproductive output of Takydromus septentrionalis collected over 5 years between 1997 and 2005 to test the hypothesis that reproductive females should allocate an optimal fraction of accessible resources in a particular clutch and to individual eggs. Females laid 1–7 clutches per breeding season, with large females producing more, as well as larger clutches, than did small females. Clutch size, clutch mass, annual fecundity, and annual reproductive output were all positively related to female size (snout–vent length). Females switched from producing more, but smaller eggs in the first clutch to fewer, but larger eggs in the subsequent clutches. The mass-specific clutch mass was greater in the first clutch than in the subsequent clutches, but it did not differ among the subsequent clutches. Post-oviposition body mass, clutch size, and egg size showed differing degrees of annual variation, but clutch mass of either the first or the second clutch remained unchanged across the sampling years. The regression line describing the size–number trade-off was higher in the subsequent clutch than in the first clutch, but neither the line for first clutch, nor the line for the second clutch varied among years. Reproduction retarded growth more markedly in small females than in large ones. Our data show that: (1) trade-offs between size and number of eggs and between reproduction and growth (and thus, future reproduction) are evident in T. septentrionalis ; (2) females allocate an optimal fraction of accessible resources in current reproduction and to individual eggs; and (3) seasonal shifts in reproductive output and egg size are determined ultimately by natural selection.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 91 , 315–324.     

<Emphasis Type="Italic">Fopius arisanus</Emphasis> oviposition in four <Emphasis Type="Italic">Anastrepha</Emphasis> fruit fly species of economic importance in Mexico

We studied the oviposition performance of Fopius arisanus (Sonan) (Hymenoptera: Braconidae) attacking eggs of four fruit flies of the genus Anastrepha Schiner (Diptera: Tephritidae) under laboratory conditions. The complete process of oviposition on an individual egg of Anastrepha ludens (Loew) lasts in average 85.4 ± 2.9 s, including a tremor (25.8 ± 1.03 s) observed in the middle of this process related to the egg’s descent. The average parasitism of A. ludens egg was 60.9 ± 7.5%, with only 1.2% of superparasitized eggs. During individual acts of oviposition, we noted that F. arisanus possesses a highly flexible ovipositor that curves easily as it searches for additional suitable eggs, which may be of particular benefit when a female finds large clutches of eggs. The individual oviposition of F. arisanus in host fruits attacked by Anastrepha spp. varies with the egg clutch size of each fruit fly species: A. serpentina laid the biggest egg clutches (21.3 ± 1.4), followed by A. ludens (14.2 ± 0.9), and A. striata (1.0 ± 0.0) (=A. obliqua). The time spent by F. arisanus in individual ovipositions was parallel to these findings, reinforcing the idea that F. arisanus attacks several eggs in each individual insertion of its ovipositor. Neither formal oviposition acts, nor adult emergences of F. arisanus were registered in A. obliqua. We discuss the potential of F. arisanus as natural enemy of fruit flies of the genus Anastrepha, and explore the eventual developing of its mass rearing. Handling Editor: Torsten Meiners.     

Clutch size in the tropical scincid lizard Emoia sanfordi, a species endemic to the Vanuatu Archipelago

The majority of species in the scincid genus Emoia (Squamata: Scincidae) have a fixed clutch size of two eggs per clutch and produce between two and four clutches per year. One lineage within Emoia, the Emoia samoensis species group, consists of 13 species occurring in Melanesia and the islands of the southwestern Pacific Ocean, and exhibits variation in clutch size, with previously reported clutch sizes of two to five eggs. Little is known about reproduction in several members of this lineage including Emoia sanfordi, a large-bodied lizard endemic to the archipelago of Vanuatu in the South Pacific. We analyzed reproduction and clutch size in E. sanfordi females and discovered that there is a substantial amount of intraspecific variation, with clutch size ranging from two to seven eggs, with a modal clutch size of five eggs. Females were reproductively active throughout the study period of June through October and appear to be laying multiple clutches. The variation in clutch size seen in E. sanfordi is congruent with the variation previously reported within other closely related species.     

The use of ultrasonography to assess reproductive investment and output in pythons

Reproductive investment and output are integral fitness components, often incorporated into life‐history trade‐off models and important to population dynamics. The trade‐offs associated with reproduction can be dramatic in species such as snakes that make especially large investments into reproduction. Unfortunately, traditional methods used to determine reproductive investment and output are effective in many (but not all) situations. Thus, we used portable ultrasonography to serially estimate reproductive investment and reproductive output in three python species that exhibit significant variation in phylogeny, geographic range, body size, egg size, and clutch size: ball pythons (Python regius), Children's pythons (Antaresia childreni), and water pythons (Liasis fuscus). At each time point of measurement (range: 1–49 days pre‐oviposition), ultrasound estimates of viable clutch size were highly accurate in all three species. However, ultrasound estimates of mean viable egg mass, and thus viable clutch mass, significantly differed from the actual values (range: 23–73% error). Interestingly, this error was considerably smaller as females approached oviposition, suggesting that female pythons transfer a significant amount of water into their eggs during the week before oviposition. Thus, water balance during late‐stage egg development may be an integral part of reproductive success. The results obtained in the present study form the foundation for future assessments of reproductive investment, and also provide insight into the use of ultrasound technology to assist such efforts. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 103 , 772–778.     

Nesting Phenology and Clutch Characteristics of Captive Siamese Crocodiles (Crocodylus siamensis) in Cambodia

The Siamese crocodile (Crocodylus siamensis) is considered one of the least studied and most critically endangered crocodilians in the world. Although few wild populations remain, more than 700,000 C. siamensis are held on commercial crocodile farms in Southeast Asia. Despite conservation concerns, many aspects of C. siamensis life history remain poorly known, particularly with regards to its reproductive biology. We studied nesting phenology, clutch characteristics, and other aspects of C. siamensis reproductive biology on crocodile farms in Cambodia during 2000 and 2001. Oviposition among captive crocodiles began in February and continued into early June. The mean (±1 SD) oviposition date based on pooled data from 2000 and 2001 was 5 April ± 24 days. Mean oviposition date differed significantly between 2000 and 2001, possibly as a result of annual variability among nesting cues. The mean incubation period was 72 ± 3 days and eggs hatched from 5 May to 18 August. Mean clutch size (25.0 ± 8.8 eggs; n = 183) differed significantly between years, possibly resulting from the >2.5‐fold increase in sample size during 2001. There was no correlation between clutch size and oviposition date during either 2000 or 2001. A single female produced two clutches during 2001, complimenting previous reports of double‐clutching among C. siamensis. The mean length and width of 515 eggs were 78.2 ± 4.9 and 48.1 ± 2.5 mm, respectively; mean egg mass was 90.8 ± 16.5 g (n = 471). One unpipped egg contained a set of twins. Zoo Biol 31:534‐545, 2012. © 2011 Wiley Periodicals, Inc.     

Factors influencing double brooding in Eurasian Hoopoes Upupa epops

Double brooding may be a good strategy for short‐lived species to maximize annual and lifetime reproductive success (ARS and LRS, respectively). Nevertheless, there is typically individual variation in the probability of producing a second clutch. Here we evaluate factors that influence the decision to double brood in the Eurasian Hoopoe Upupa epops. Analyses of an 11‐year dataset showed that 36% of the females and 21% of the males produced a second clutch after successfully raising a first clutch. Double‐brooded females had higher ARS (9.1 ± 1.9 fledglings; mean ± se) and LRS (0.93 ± 0.08 recruits) than single‐brooded females (ARS: 4.5 ± 2.1 fledglings; LRS: 0.36 ± 0.03 recruits). This suggests that double brooding is adaptive in Hoopoes, and raises the question of why most individuals only produce one clutch per season. The probability of double brooding varied only slightly between years, suggesting that it is influenced by individual characteristics rather than by external, population‐level environmental factors. In both sexes, the probability of double brooding increased with earlier timing of the first clutch, and the timing of reproduction was the most important factor influencing reproductive success. The latter is likely to be mediated by changes in resources during the season. The probability of double brooding also increased slightly with female age, due to differences in intrinsic quality among females rather than to a gain in experience. In contrast to many other studies, the probability of double brooding increased with an increasing number of fledglings from the first clutch, suggesting that it is a strategy of individuals of high quality. Taken together, we show that the individual quality of the breeder and the timing of their first clutches are key factors influencing the decision to double brood, and thereby that they are important determinants of reproductive performance in Eurasian Hoopoes.     

Multiple paternity and female-biased mutation at a microsatellite locus in the olive ridley sea turtle (Lepidochelys olivacea)

Multiple paternity in the olive ridley sea turtle (Lepidochelys olivacea) population nesting in Suriname was demonstrated using two microsatellite loci, viz., Ei8 and Cm84. The large number of offspring sampled per clutch (70 on average, ranging from 15 to 103) and the number of alleles found at the two loci (18 and eight alleles, respectively) enabled unambiguous assessment of the occurrence of multiple paternity. In two out of 10 clutches analysed, the offspring had been sired by at least two males, which was confirmed at both loci. In both clutches, unequal paternity occurred: 73% and 92% of the offspring had been sired by the primary male. The probability of detecting multiple paternity was 0.903, and therefore there is a small chance that multiple paternity occurred but remained undetected in some of the eight clutches that appeared to be singly sired. Analysis of 703 offspring revealed a high mutation rate for locus Ei8 (micro = 2.3 x 10(-2)) with all 33 mutations occurring in maternal alleles. In particular, one allele of 274 bp mutated at a high frequency in a clutch to which the mother contributed the allele, but in another clutch where the father contributed the same allele, no such mutations were observed. Inferred allele-specific mutation rates for Ei8 and expected numbers of mutations per clutch confirmed that maternal alleles for Ei8 are more likely to mutate in the olive ridley sea turtle than paternal alleles. Possible explanations are discussed.     

Multiple paternity in egg clutches of hawksbill turtles (Eretmochelys imbricata)

We present the first data collected on the genetic mating system of the hawksbill turtle Eretmochelys imbricata, the only marine turtle not studied to date. We examined paternity within 12 egg clutches from ten female hawksbill turtles from Sabah Turtle Islands, Malaysia. A total of 375 hatchlings were analysed using five microsatellite markers. Results demonstrated that clutches from two out of ten females were sired by multiple males (maximum of two). Although at a low frequency, observation of multiple paternity indicates that hawksbill turtles exhibit the same genetic mating system (polyandry) as observed for other species of marine turtles. Consistent paternity across multiple clutches laid by individual females in one breeding season supports the hypothesis that sperm are stored from mating prior to nesting and are then used to fertilize all subsequent clutches of eggs that season.     

Embryonic death is linked to maternal identity in the leatherback turtle (Dermochelys coriacea)

Leatherback turtles have an average global hatching success rate of ~50%, lower than other marine turtle species. Embryonic death has been linked to environmental factors such as precipitation and temperature, although, there is still a lot of variability that remains to be explained. We examined how nesting season, the time of nesting each season, the relative position of each clutch laid by each female each season, maternal identity and associated factors such as reproductive experience of the female (new nester versus remigrant) and period of egg retention between clutches (interclutch interval) affected hatching success and stage of embryonic death in failed eggs of leatherback turtles nesting at Playa Grande, Costa Rica. Data were collected during five nesting seasons from 2004/05 to 2008/09. Mean hatching success was 50.4%. Nesting season significantly influenced hatching success in addition to early and late stage embryonic death. Neither clutch position nor nesting time during the season had a significant affect on hatching success or the stage of embryonic death. Some leatherback females consistently produced nests with higher hatching success rates than others. Remigrant females arrived earlier to nest, produced more clutches and had higher rates of hatching success than new nesters. Reproductive experience did not affect stage of death or the duration of the interclutch interval. The length of interclutch interval had a significant affect on the proportion of eggs that failed in each clutch and the developmental stage they died at. Intrinsic factors such as maternal identity are playing a role in affecting embryonic death in the leatherback turtle.     

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.