Hatch or wait? A dilemma in reptilian incubation

Animals often form groups to reduce the risk of predation through the per capita dilution of their individual predation risk. The advantages of grouping also influence the timing of reproduction in many species. In particular, synchrony in the timing of births may have evolved as a predator-avoidance strategy as it dilutes the risk of predation upon vulnerable newborn and naive young. Eggs of an Australian freshwater turtle, Emydura macquarii , can hatch synchronously despite developmental asynchrony among eggs of a clutch and hatchlings have a reduced predation risk by emerging from the nest as a group. Developmental asynchrony within clutches was induced to reflect natural nests by dividing clutches and incubating them at either 25°C or 30°C. Some eggs were then reunited with their clutch-mates and hatching occurred synchronously in some of these groups. In groups where synchronous hatching did not occur, less advanced eggs still hatched earlier than the normal incubation period. Synchrony occurred because the less advanced eggs hatched up to five days earlier than the control embryos. We conclude that the less advanced embryos within a clutch either accelerate their development or hatch prematurely to ensure synchrony of hatching and hatchling group formation may facilitate emergence from the nest and dilute predation risk.     

Juvenile proportion as a predictor of freshwater turtle population change

The extreme longevity of turtles and tortoises can make it difficult to determine the conservation status of their populations because high annual adult survival may mask gradual attrition due to low levels of recruitment. When long-term demographic trends are unknown and available data are insufficient for population modelling, it may be assumed that a scarcity of juveniles indicates low recruitment that will result in population ageing and numerical decline. However, the reliability with which the proportion of juveniles foreshadows demographic change is uncertain. We tested the hypothesis that a low proportion of juveniles in a turtle population presages its ageing by analysing over 20 years of survey data for five discrete populations of the Australian western saw-shelled turtle (Myuchelys bellii: Chelidae), a listed threatened species. The analysis tested whether the initial proportion of juvenile turtles in each population was related to its temporal trend in average body size. The five populations had varied structure and trends, with the initial proportion of juvenile turtles ranging from 10% to 39% and average body size increasing over time in some populations and decreasing in others. Contrary to expectation, the initial proportion of juveniles was unrelated to the trend in average body size and, by inference, average age, indicating that effective trend forecasting requires more detailed demographic information than merely population structure.     

Sitting ducklings: Timing of hatch,nest departure,and predation risk for dabbling duck broods

For ground‐nesting waterfowl, the timing of egg hatch and duckling departure from the nest may be influenced by the risk of predation at the nest and en route to wetlands and constrained by the time required for ducklings to imprint on the hen and be physically able to leave the nest. We determined the timing of hatch, nest departure, and predation on dabbling duck broods using small video cameras placed at the nests of mallard (Anas platyrhynchos; n = 26), gadwall (Mareca strepera; n = 24), and cinnamon teal (Anas cyanoptera; n = 5). Mallard eggs began to hatch throughout the day and night, whereas gadwall eggs generally started to hatch during daylight hours (mean 7.5 hr after dawn). Among all species, duckling departure from the nest occurred during daylight (98%), and 53% of hens typically left the nest with their broods 1–4 hr after dawn. For mallard and gadwall, we identified three strategies for the timing of nest departure: (a) 9% of broods left the nest the same day that eggs began to hatch (6–12 hr later), (b) 81% of broods left the nest the day after eggs began to hatch, and (c) 10% of broods waited 2 days to depart the nest after eggs began to hatch, leaving the nest just after the second dawn (27–42 hr later). Overall, eggs were depredated at 10% of nests with cameras in the 2 days prior to hatch and ducklings were depredated at 15% of nests with cameras before leaving the nest. Our results suggest that broods prefer to depart the nest early in the morning, which may best balance developmental constraints with predation risk both at the nest and en route to wetlands.     

Comparative survival and recovery of Ross's and lesser snow geese from Canada's central arctic

Large increases in several populations of North American arctic geese have resulted in ecosystem-level effects from associated herbivory. Consequently, some breeding populations have shown density dependence in recruitment through declines in food availability. Differences in population trajectories of lesser snow geese (Chen caerulescens caerulescens; hereafter snow geese) and Ross's geese (C. rossii) breeding in mixed-species colonies south of Queen Maud Gulf (QMG), in Canada's central arctic, suggest that density dependence may be limiting snow goose populations. Specifically, long-term declines in age ratios (immature:adult) of harvested snow geese may have resulted from declines in juvenile survival. Thus, we focused on juvenile (first-year) survival of snow and Ross's geese in relation to timing of reproduction (annual mean nest initiation date) and late summer weather. We banded Ross's and snow geese from 1991 to 2008 in the QMG Migratory Bird Sanctuary. We used age-structured mark-recapture models to estimate annual survival rates for adults and juveniles from recoveries of dead birds. Consistent with life history differences, juvenile snow geese survived at rates higher than juvenile Ross's geese. Juvenile survival of both species also was lower in late seasons, but was unrelated to arctic weather measured during a 17-day period after banding. We found no evidence of density dependence (i.e., a decline in juvenile survival over time) in either species. We also found no interspecific differences in age-specific hunting vulnerability, though juveniles were more vulnerable than adults in both species, as expected. Thus, interspecific differences in survival were unrelated to harvest. Lower survival of juvenile Ross's geese may result from natural migration mortality related to smaller body size (e.g., greater susceptibility to inclement weather or predation) compared to juvenile snow geese. Despite lower first-year survival, recruitment by Ross's geese may still be greater than that by snow geese because of earlier sexual maturity, greater breeding propensity, and higher nest success by Ross's geese. © 2012 The Wildlife Society.     

Function of egg punctures by Shiny Cowbirds in parasitized and nonparasitized Creamy‐bellied Thrush nests

ABSTRACT Avian brood parasites usually remove or puncture host eggs. Several hypotheses have been proposed to explain the function of these behaviors. Removing or puncturing host eggs may enhance the efficiency of incubation of cowbird eggs (incubation‐efficiency hypothesis) or reduce competition for food between cowbird and host chicks in parasitized nests (competition‐reduction hypothesis) and, in nonparasitized nests, may force hosts to renest and provide cowbirds with new opportunities for parasitism when nests are too advanced to be parasitized (nest‐predation hypothesis). Puncturing eggs may also allow cowbirds to assess the development of host eggs and use this information to decide whether to parasitize a nest (test‐incubation hypothesis). From 1999 to 2002, we tested these hypotheses using a population of Creamy‐bellied Thrushes (Turdus amaurochalinus) in Argentina that was heavily parasitized by Shiny Cowbirds (Molothrus bonariensis). We found that 56 of 94 Creamy‐bellied Thrush nests (60%) found during nest building or egg laying were parasitized by Shiny Cowbirds, and the mean number of cowbird eggs per parasitized nest was 1.6 ± 0.1 (N= 54 nests). At least one thrush egg was punctured in 71% (40/56) of parasitized nests, and 42% (16/38) of nonparasitized nests. We found that cowbird hatching success did not differ among nests where zero, one, or two thrush eggs were punctured and that the proportion of egg punctures associated with parasitism decreased as incubation progressed. Thus, our results do not support the incubation‐efficiency, nest‐predation, or test‐incubation hypotheses. However, the survival of cowbird chicks in our study was negatively associated with the number of thrush chicks. Thus, our results support the competition‐reduction hypothesis, with Shiny Cowbirds reducing competition between their young and host chicks by puncturing host eggs in parasitized nests.     

Spatial and temporal dynamics of turtle nest predation: edge effects

We determined the effect of distance from ecological edges, both wooded and water edges, on nest predation for 862 painted turtle, Chrysemys picta , nests from 1995 to 1999 at an ∼1.5 ha study site. In three of five years and overall, nests closer to the water edge had a higher probability of predation; and in one year nests closer to the wooded edge had a higher probability of predation. Although more turtles nested closer to the water edge as the nesting season progressed in some years and overall, this behavior does not explain the observed patterns of nest predation. We present a novel application of the cubic spline analysis to address the dynamics of predation across continuous distances from an edge and identify threshold values where the predation rate levels off. Threshold values of ∼25–40 m were detected in 1995, 1998, and with all five years combined. However, even though a significant edge effect was detected in 1997, a threshold was not clear. While an edge effect on predation was not detected in each year, this study provides evidence for a strong effect of distance from the water edge on nest predation over significant ecological time. Focusing on turtle nest predation and smaller spatial scales addresses previous taxonomic and spatial bias in edge effects research, and provides further support for the ecological importance of edge effects.     

First fossil gravid turtle provides insight into the evolution of reproductive traits in turtles

Here we report on the first discovery of shelled eggs inside the body cavity of a fossil turtle and on an isolated egg clutch, both referable to the Cretaceous turtle Adocus. These discoveries provide a unique opportunity to gain insight into the reproductive traits of an extinct turtle and to understand the evolution of such traits among living turtles. The gravid adult and egg clutch indicate that Adocus laid large clutches of rigid-shelled spherical eggs and established their nests near rivers, traits that are shared by its closest living relatives, the soft-shelled turtles. Adocus eggshell, however, was probably more rigid than that of living turtles, based on its great thickness and structure, features that may represent unique adaptations to intense predation or to arid nest environments. In light of the reproductive traits observed in Adocus, the distribution of reproductive traits among turtles reveals that large clutches of rigid-shelled eggs are primitive for hidden-necked turtles (cryptodirans) and that spherical eggs may have evolved independently within this group.     

The evolution of egg clustering in butterflies: A test of the egg desiccation hypothesis

Females of many insect species cluster their eggs. Egg clustering by lepidopteran species usually results in aggregation of larvae that are more often conspicuously coloured and apparently distasteful or unpalatable than larvae of solitary species. While the costs and benefits of aggregation in terms of larval survival and growth are well documented, the evolutionary ecology of egg clustering has been long debated and is still unresolved. We tested the egg desiccation hypothesis, first proposed by Stamp (1980), which to our knowledge has never been examined experimentally. The egg desiccation hypothesis proposes that egg clustering is adaptive per se (i.e. increases fitness of females) by reducing egg mortality via desiccation.We tested this hypothesis for the Nymphalid butterfly, Chlosyne lacinia, an egg-clustering species on its sunflower host plant, Helianthus annuus. We first documented natural variation in batch size for this butterfly. We then tested experimentally hatch success of varying batch sizes and egg-layering arrangements under controlled humidity levels. Hatch success was positively related to relative humidity. Eggs in larger groups with greater number of layers had greater hatch success than smaller, monolayered egg batches, especially when relative humidity was low. Our results indicate that, not only number of eggs, but also the arrangement (i.e. layering and density), increase batch survival by protecting eggs from desiccation. However, despite increased hatch success in dense, multilayered clusters, we found wide variation in layering and density in natural populations of C. lacinia. This variation is probably maintained by trade-offs in egg survival, such as increased cannibalism of eggs by siblings, in dense clusters. Nevertheless, protection from egg desiccation provides an alternative explanation for the origin and maintenance of egg clustering in lepidopterans and possibly other insects. The pattern of egg deposition in the Nymphalidae supports this hypothesis, since most North American species cluster their eggs tightly, whereas most species in tropical regions lay eggs singly or in loose monolayers.     

A retrospective analysis of sea turtle nest depredation patterns

Nest predation can significantly reduce hatchling recruitment in sea turtle populations. We examined 20 yr of data from Canaveral National Seashore, Florida, which has pristine and altered beaches. We used chi-squared test to determine if secondary predation events were related to the nests' primary predation events, and proportional hazard regression analysis to determine the relative risk of individual nest predation. To determine if human beach use and nest predation risk were spatially or temporally linked, we ranked human beach use and examined predation frequency across all screened and marked nests. We found that once a nest has experienced predation it has an increased likelihood of experiencing a subsequent predation event when compared to other nests on the beach. Primary and multiple predation events occurred with greater frequency in limited beach use areas and with lowest frequency in moderate use areas. Predation risk decreased by an average of 29.5% from 2000 to 2008, relative to the initial year of study. Nests deposited mid-season were 9.8% more likely to be predated than early or late season nests. We conclude that where anti-nest predator management efforts are required but restricted (e.g., by funds, available personnel, or multiple management goals) resources should be concentrated to protect mid-season nests and those in limited access areas, if enhancing hatchling numbers is a management goal. © 2012 The Wildlife Society.     

Evidence of selection for egg crypsis in conspicuous nests

ABSTRACT The value of egg coloration as crypsis, once accepted as a general principle, has recently been questioned because most experiments have failed to show that egg coloration deters predation. The nest‐crypsis hypothesis postulates that, among species that build conspicuous nests, selection for egg crypsis is relaxed or absent because visually searching predators detect nests prior to eggs. I tested the nest‐crypsis hypothesis using the large, relatively conspicuous nests of American Robins (Turdus migratorius), and eggs that differed markedly in color that were collected from the nests of Red‐winged Blackbirds (Agelaius phoeniceus), Brewer's Blackbirds (Euphagus cyanocephalus), and Yellow‐headed Blackbirds (Xanthocephalus xanthocephalus). Each nest (N= 22) received a clutch of each species during three sequential predation trials that were 16 d in duration. The order of clutch presentation was randomized for each nest. Survival trends for Brewer's and Yellow‐headed Blackbirds were similar, and higher than those for clutches of Red‐winged Blackbirds. By the end of trials, overall survival of the three clutch types was roughly equivalent. However, clutches of Red‐winged Blackbird eggs, the most conspicuous egg type to the human eye, were discovered sooner by predators. Because the experimental design controlled for effects of nest crypsis, nest location, and nest size, this difference in egg survival can be attributed to differences in egg pigmentation. Thus, my results support a role for egg coloration as camouflage in conspicuous nests.     

Nest predation increases with parental activity: separating nest site and parental activity effects.

Alexander Skutch hypothesized that increased parental activity can increase the risk of nest predation. We tested this hypothesis using ten open-nesting bird species in Arizona, USA. Parental activity was greater during the nestling than incubation stage because parents visited the nest frequently to feed their young during the nestling stage. However, nest predation did not generally increase with parental activity between nesting stages across the ten study species. Previous investigators have found similar results. We tested whether nest site effects might yield higher predation during incubation because the most obvious sites are depredated most rapidly. We conducted experiments using nest sites from the previous year to remove parental activity. Our results showed that nest sites have highly repeatable effects on nest predation risk; poor nest sites incurred rapid predation and caused predation rates to be greater during the incubation than nestling stage. This pattern also was exhibited in a bird species with similar (i.e. controlled) parental activity between nesting stages. Once nest site effects are taken into account, nest predation shows a strong proximate increase with parental activity during the nestling stage within and across species. Parental activity and nest sites exert antagonistic influences on current estimates of nest predation between nesting stages and both must be considered in order to understand current patterns of nest predation, which is an important source of natural selection.     

A nondestructive method for estimating the age of Clapper Rail eggs

ABSTRACT.   Estimates of nest survival rate are often based on the assumption that the rate is constant during the nest cycle. Because violating this assumption can generate biased estimates, determining the age of a nest is important. For precocial marsh birds, including Clapper Rails ( Rallus longirostris ), current estimates of nest survival do not take this source of bias into account because a standardized, nondestructive method for aging eggs in not available. We developed a nondestructive flotation technique using known-age nests that can be used to estimate the projected hatch date of Clapper Rail eggs within 3 d of the actual hatch date. This precision is limited in part due to variance in the length of incubation between nests and the common occurrence of asynchronous hatching. However, for investigators assessing the fate of Clapper Rail nests, our technique provides an effective and nondestructive method for projecting the hatch date of Clapper Rail eggs, and will facilitate unbiased estimates of nest survival rates.     

Juvenile exposure to predator cues induces a larger egg size in fish

When females anticipate a hazardous environment for their offspring, they can increase offspring survival by producing larger young. Early environmental experience determines egg size in different animal taxa. We predicted that a higher perceived predation risk by juveniles would cause an increase in the sizes of eggs that they produce as adults. To test this, we exposed juveniles of the mouthbrooding cichlid Eretmodus cyanostictus in a split-brood experiment either to cues of a natural predator or to a control situation. After maturation, females that had been confronted with predators produced heavier eggs, whereas clutch size itself was not affected by the treatment. This effect cannot be explained by a differential female body size because the predator treatment did not influence growth trajectories. The observed increase of egg mass is likely to be adaptive, as heavier eggs gave rise to larger young and in fish, juvenile predation risk drops sharply with increasing body size. This study provides the first evidence that predator cues perceived by females early in life positively affect egg mass, suggesting that these cues allow her to predict the predation risk for her offspring.     

Incidental nest predation in freshwater turtles: inter- and intraspecific differences in vulnerability are explained by relative crypsis

There has long been interest in the influence of predators on prey populations, although most predator–prey studies have focused on prey species that are targets of directed predator searching. Conversely, few have addressed depredation that occurs after incidental encounters with predators. We tested two predictions stemming from the hypothesis that nest predation on two sympatric freshwater turtle species whose nests are differentially prone to opportunistic detection—painted turtles (Chrysemys picta) and snapping turtles (Chelydra serpentina)—is incidental: (1) predation rates should be density independent, and (2) individual predators should not alter their foraging behavior after encountering nests. After monitoring nest survival and predator behavior following nest depredation over 2 years, we confirmed that predation by raccoons (Procyon lotor), the primary nest predators in our study area, matched both predictions. Furthermore, cryptic C. picta nests were victimized with lower frequency than more detectable C. serpentina nests, and nests of both species were more vulnerable in human-modified areas where opportunistic nest discovery is facilitated. Despite apparently being incidental, predation on nests of both species was intensive (57% for painted turtles, 84% for snapping turtles), and most depredations occurred within 1 day of nest establishment. By implication, predation need not be directed to affect prey demography, and factors influencing prey crypsis are drivers of the impact of incidental predation on prey. Our results also imply that efforts to conserve imperiled turtle populations in human-modified landscapes should include restoration of undisturbed conditions that are less likely to expose nests to incidental predators.     

Dynamics of Winter Survival of Eggs and Juveniles of Meloidogyne incognita and M. arenaria

Winter survival dynamics of Meloidogyne incognita and M. arenaria were studied at nine sites in Texas for 2 years. Population survival from October until April was variable among sites, ranging from 0.1% to 33%. A negative correlation (r = -0.86, P = 0.01) was observed between initial population, densities in October and survival percentage until the following April. Total population (eggs + J2) and population of eggs declined continuously during the survival period. Populations of juveniles (J2) increased initially, then declined. The total populations were 82% eggs in October; hatch of these eggs was believed responsible for the observed increase in the population density of J2. Viable eggs were recovered from the soil until March. Eggs are as important as J2 in winter survival of M. incognita and M. arenaria in Texas. Survival data were analyzed by a simple mathematical model.     

Fragmentation within and between wetland reserves: the importance of spatial scales for nest predation in reed buntings

Wetlands in many parts of the world are affected by fragmentation at multiple spatial scales. In Switzerland, most wetlands have been destroyed over the past two centuries and management of the remaining wetland reserves has intensified in the recent years leading to increased fragmentation of reed areas within reserves. Using four years of data on the reproductive performance of color-banded reed bunting Emberiza schoeniclus populations, we explored fragmentation effects on nest predation rates at four spatial scales ranging from the nest to the landscape scale. In the egg stage, predation rate was negatively related to vegetation cover, vegetation height and nest height, but positively linked to water cover and depth next to the nest (nest scale). Probability of predation declined with increasing size of reed patches containing the nests and distances of nests to the water and land sided reed edge, as well as with decreasing edge to area ratio (edge scale). There was a weak positive association between degree of fragmentation of reed patches within sites and nest predation rates (site scale). Finally, nest predation probability increased with distance to the nearest wetland (landscape scale). Jointly analyzing variables from different spatial scales revealed that a model combining variables from the nest, edge and landscape scale best explained predation probability in the egg stage. In the nestling stage, the single most important factor influencing nest predation probability was the distance to the nearest wetland (landscape scale), with nest predation decreasing with distance between sites. Our results show that the probability of nest predation in reed buntings is affected by fragmentation within and between wetland reserves and that the effects differ between breeding stages. Future management of wetland reserves should aim at sparing reed patches large and dense enough to provide safe nest sites for birds.     

Using artificial nests to predict nest survival at reintroduction sites

Artificial nests are frequently used to assess factors affecting survival of natural bird nests. We tested the potential for artificial nests to be used in a novel application, the prediction of nest predation rates at potential reintroduction sites where exotic predators are being controlled. We collected artificial nest data from nine sites with different predator control regimes around the North Island of New Zealand, and compared the nest survival rates with those of North Island robin (Petroica longipes) nests at the same sites. Most of the robin populations had been reintroduced in the last 10 years, and were known to vary in nest survival and status (increasing/stable or declining). We derived estimates of robin nest survival for each site based on Stanley estimates of daily survival probabilities and the known incubation and brooding periods of robins. Estimates of artificial nest survival for each site were derived using the known fate model in MARK. We identified the imprints on the clay eggs in the artificial nests, and obtained different estimates of artificial nest survival based on imprints made by different potential predators. We then compared the value of these estimates for predicting natural nest survival, assuming a relationship of the form s = αpβ, where s is natural nest survival and p is artificial nest survival. Artificial nest survival estimates based on imprints made by rats (Rattus spp.) and brushtail possums (Trichosurus vulpecula) were clearly the best predictors (based on AICc), and explained 64% of the variation in robin nest survival among sites. Inclusion of bird imprints in the artificial nest survival estimates substantially reduced their predictive value. We suggest that artificial nests may provide a useful tool for predicting the suitability of potential reintroduction sites for New Zealand forest birds as long as imprints on clay eggs are correctly identified.     

Demography of an alpine population of Savannah Sparrows

ABSTRACT.   Although Savannah Sparrows ( Passerculus sandwichensis ) have been well studied across their North American range, few data are available for populations that breed in high-elevation habitats. We collected data over six years on the demography of a population of Savannah Sparrows ( P. s. anthinus ) breeding in alpine tundra and sub-alpine meadows in northern British Columbia, Canada. The mean duration of the breeding season at our site was 45.5 d, and pairs produced a maximum of one brood per season. Clutch sizes varied annually (mean = 4.37, range = 3.90 – 4.71 eggs). Nest fate also varied among years (range = 33 – 92%) due to variation in abiotic (weather) and biotic (predators) conditions. Uncorrected return rates of banded birds were 68% for adults and 17% for juveniles ( N = 22 and 102, respectively). However, when resighting probability was taken into account, apparent annual survival was 75% for adults and 34% for juveniles. Compared to populations at lower elevations, Savannah Sparrows in our study had shorter breeding seasons, fewer broods per season, larger clutches, and higher adult and juvenile return rates. Our results suggest that Savannah Sparrows that breed in high-elevation habitats have adopted a low fecundity, high survival life history strategy that enables their persistence in these challenging environments.     

Conspecific nest parasitism is associated with inequality in nest predation risk in the common goldeneye (Bucephala clangula)

Previous studies of the role of nest predation in conspecificnest parasitism have not taken into account the possibilitythat predation risk may not be randomly distributed among nestsites and that breeding individuals may use different cues toassess the risk and adjust their reproductive tactic betweenyears accordingly. Especially in cavity-nesting species, therole of nest predation in conspecific nest parasitism has beendownplayed, while the role of nest site limitation has beenhighlighted. Using both observational and experimental data,I show that in the common goldeneye (Bucephala clangula), acavity-nesting species in which conspecific nest parasitism iscommon, predation risk varies considerably between nest sitesand does not follow a random expectation. The inequality inpredation risk between nest sites also showed up in the occurrenceof parasitized nests in an experimental setup. Nests parasitizedin year t were more frequent in those nest sites that were notdepredated during the previous nesting attempt in year t - nthan in nest sites that were depredated and in control nestsites that had not been used for nesting before. A nest siteaddition experiment revealed that conspecific nest parasitismwas not associated with nest site limitation. My findings givesupport for the hypothesis that nest predation is an importantecological factor explaining conspecific nest parasitism ingoldeneyes.