Use of Implanted Radiotransmitters to Estimate Survival of Greater Sage-Grouse Chicks

ABSTRACT Reduced chick survival has been implicated in declines of greater sage-grouse (Centrocercus urophasianus) populations. Because monitoring survival of unmarked sage-grouse chicks is difficult, radiotelemetry may be an effective technique to estimate survival rates, identify causes of mortality, and collect ecological data. Previous studies have used subcutaneous implants to attach radiotransmitters to hatchlings of several species of birds with precocial young. Previous researchers who used subcutaneous implants in free-ranging populations removed chicks from the capture location and implanted transmitters at an alternate site. Because logistics precluded removing newly hatched greater sage-grouse chicks from the field, we evaluated a method for implanting transmitters at capture locations. We captured 288 chicks from 52 broods and monitored 286 radiomarked chicks daily for 28 days following capture during May and June 2001–2002. Two (>1%) chicks died during surgery and we did not radiomark them. At the end of the monitoring period, 26 chicks were alive and 212 were dead. Most (98%, 207/212) radiomarked chick mortality occurred < 21 days posthatch and predation (82%, 174/212) was the primary cause of death. Necropsies of 22 radiomarked chicks did not indicate inflammation or infection from implants, and they were not implicated in the death of any chicks. Fate of 48 chicks was unknown because of transmitter loss (n = 16), radio failure (n = 29), and brood mixing (n = 3). Overall, the 28-day chick survival rate was 0.220 (SE = 0.028). We found that mortalities related to the implant procedure and transmitter loss were similar to rates reported by previous researchers who removed chicks from capture sites and implanted transmitters at an alternate location. Subcutaneous implants may be a useful method for attaching transmitters to newly hatched sage-grouse chicks to estimate survival rates, identify causes of mortality, and collect ecological data.     

Yearling Greater Sage-Grouse Response to Energy Development in Wyoming

ABSTRACT Sagebrush (Artemisia spp.)-dominated habitats in the western United States have experienced extensive, rapid changes due to development of natural-gas fields, resulting in localized declines of greater sage-grouse (Centrocercus urophasianus) populations. It is unclear whether population declines in natural-gas fields are caused by avoidance or demographic impacts, or the age classes that are most affected. Land and wildlife management agencies need information on how energy developments affect sage-grouse populations to ensure informed land-use decisions are made, effective mitigation measures are identified, and appropriate monitoring programs are implemented (Sawyer et al. 2006). We used information from radio-equipped greater sage-grouse and lek counts to investigate natural-gas development influences on 1) the distribution of, and 2) the probability of recruiting yearling males and females into breeding populations in the Upper Green River Basin of southwestern Wyoming, USA. Yearling males avoided leks near the infrastructure of natural-gas fields when establishing breeding territories; yearling females avoided nesting within 950 m of the infrastructure of natural-gas fields. Additionally, both yearling males and yearling females reared in areas where infrastructure was present had lower annual survival, and yearling males established breeding territories less often, compared to yearlings reared in areas with no infrastructure. Our results supply mechanisms for population-level declines of sage-grouse documented in natural-gas fields, and suggest to land managers that current stipulations on development may not provide management solutions. Managing landscapes so that suitably sized and located regions remain undeveloped may be an effective strategy to sustain greater sage-grouse populations affected by energy developments.     

Survival of Greater Sage-Grouse Chicks and Broods in the Northern Great Basin

ABSTRACT Reduced annual recruitment because of poor habitat quality has been implicated as one of the causative factors in the range-wide decline of sage-grouse (Centrocercus urophasianus) populations since the 1950s. Because chick and brood survival are directly linked to annual recruitment and may be the primary factors that limit sage-grouse population growth, we estimated 28-day survival rates of radiomarked chicks and broods from 2000 to 2003. We examined relationships between survival and several habitat variables measured at brood sites, including food availability (insects and forbs); horizontal cover of sagebrush, grasses, and forbs; and vertical cover of sagebrush and grass. We monitored 506 radiomarked chicks from 94 broods; chick survival was 0.392 (SE = 0.024). We found evidence that both food and cover variables were positively associated with chick survival, including Lepidoptera availability, slender phlox (Phlox gracilis) frequency, total forb cover, and grass cover. The effect of total grass cover on chick survival was dependent on the proportion of short grass. The hazard of an individual chick's death decreased 8.6% (95% CI = −1.0 to 18.3) for each percentage point increase in total grass cover when the proportion of short grass was >70%. Survival of 83 radiomarked broods was 0.673 (SE = 0.055). Lepidoptera availability and slender phlox frequency were the only habitat variables related to brood survival. Risk of total brood loss decreased by 11.8% (95% CI = 1.2–22.5) for each additional Lepidoptera individual and 2.7% (95% CI = −0.4 to 5.8) for each percentage point increase in the frequency of slender phlox found at brood sites. Model selection results revealed that temporal differences in brood survival were associated with variation in the availability of Lepidoptera and slender phlox. Years with high brood survival corresponded with years of high Lepidoptera availability and high slender phlox frequency. These foods likely provided high-quality nutrition for chicks during early growth and development and enhanced survival. Habitat management that promotes Lepidoptera and slender phlox abundance during May and June (i.e., early brood rearing) should have a positive effect on chick and brood survival in the short term and potentially increase annual recruitment.     

Mismatches between breeding phenology and resource abundance of resident alpine ptarmigan negatively affect chick survival

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Managing multiple vital rates to maximize greater sage-grouse population growth

Despite decades of field research on greater sage-grouse, range-wide demographic data have yet to be synthesized into a sensitivity analysis to guide management actions. We reviewed range-wide demographic rates for greater sage-grouse from 1938 to 2011 and used data from 50 studies to parameterize a 2-stage, female-based population matrix model. We conducted life-stage simulation analyses to determine the proportion of variation in population growth rate (λ) accounted for by each vital rate, and we calculated analytical sensitivity, elasticity, and variance-stabilized sensitivity to identify the contribution of each vital rate to λ. As expected for an upland game bird, greater sage-grouse showed marked annual and geographic variation in several vital rates. Three rates were demonstrably important for population growth: female survival, chick survival, and nest success. Female survival and chick survival, in that order, had the most influence on λ per unit change in vital rates. However, nest success explained more of the variation in λ than did the survival rates. In lieu of quantitative data on specific mortality factors driving local populations, we recommend that management efforts for greater sage-grouse first focus on increasing female survival by restoring large, intact sagebrush-steppe landscapes, reducing persistent sources of human-caused mortality, and eliminating anthropogenic habitat features that subsidize species that prey on juvenile, yearling, and adult females. Our analysis also supports efforts to increase chick survival and nest success by eliminating anthropogenic habitat features that subsidize chick and nest predators, and by managing shrub, forb, and grass cover, height, and composition to meet local brood-rearing and nesting habitat guidelines. We caution that habitat management to increase chick survival and nest success should not reduce the cover or height of sagebrush below that required for female survival in other seasons (e.g., fall, winter). The success or failure of management actions for sage-grouse should be assessed by measuring changes in vital rates over long time periods to avoid confounding with natural, annual variation. © 2011 The Wildlife Society.     

Landscape-Level Assessment of Brood Rearing Habitat for Greater Sage-Grouse in Nevada

Abstract: Loss of quality brood rearing habitat, resulting in reduced chick growth and poor recruitment, is one mechanism associated with decline of greater sage-grouse (Centrocercus urophasianus) populations. Low chick survival rates are typically attributed to poor-quality brood rearing habitat. Models that delineate suitability of sage-grouse nesting or brood rearing habitat at the landscape scale can provide key insights into the relationship between sage-grouse and the environment, allowing managers to identify and prioritize habitats for protection or restoration. We used Southwest Regional Gap landcover types to identify early and late greater sage-grouse brood rearing in east-central Nevada. We conducted an Ecological Niche Factor Analysis to 1) examine the effect these landcover types and other ecogeographical variables have on sage-grouse selection of brood rearing habitat, and 2) generate landscape-scale suitability maps. We also evaluated if incorporating a fitness component (brood survival) in landscape spatial analyses of habitat quality influenced our assessment of habitat suitability. Because 36% of our 6,500-km² study area was identified as early brood rearing habitat, we believe this habitat may not be limiting greater sage-grouse populations in east-central Nevada, USA, at least in wet years. We found strong selection for particular landcover types (e.g., higher elevation, moist sites with riparian shrubs or montane sagebrush) during late brood rearing. Late brood rearing habitat on which broods were successfully reared represented only 2.8% of the study area and had a restricted distribution, suggesting the potential that such habitat could limit sage-grouse populations in east-central Nevada.     

Using Human-Imprinted Chicks to Evaluate the Importance of Forbs to Sage-Grouse

Abstract Although several studies have indicated the importance of forbs in brood habitats, no studies have quantified direct effects of the amount of forb cover on sage-grouse (Centrocercus urophasianus) chicks. In 2002 and 2003, we conducted field experiments in Middle Park and Moffat County, Colorado, USA, respectively. Our objective was to quantify effects of 3 levels of forb cover in brood habitat on mass gain and feather growth of human-imprinted sage-grouse chicks. The results indicate that increasing forb cover in brood areas with <20% forb cover may lead to increased chick survival and grouse productivity.     

Adverse effects of agricultural intensification and climate change on breeding habitat quality of Black‐tailed Godwits Limosa l. limosa in the Netherlands

Agricultural intensification is one of the main drivers of farmland bird declines, but effects on birds may be confounded with those of climate change. Here we examine the effects of intensification and climate change on a grassland breeding wader, the Black‐tailed Godwit Limosa l. limosa, in the Netherlands. Population decline has been linked to poor chick survival which, in turn, has been linked to available foraging habitat. Foraging habitat of the nidifugous chicks consists of uncut grasslands that provide cover and arthropod prey. Conservation measures such as agri‐environment schemes aim to increase the availability of chick foraging habitat but have not yet been successful in halting the decline. Field observations show that since the early 1980s, farmers advanced their first seasonal mowing or grazing date by 15 days, whereas Godwits did not advance their hatching date. Ringing data indicate that between 1945 and 1975 hatching dates advanced by about 2 weeks in parallel with the advancement of median mowing dates. Surprisingly, temperature sums at median mowing and hatching dates suggest that while the agricultural advancement before 1980 was largely due to agricultural intensification, after 1980 it was largely due to climate change. Examining arthropod abundance in a range of differently managed grasslands revealed that chick food abundance was little affected but that food accessibility in intensively used tall swards may be problematic for chicks. Our results suggest that, compared with 25 years ago, nowadays (1) a much higher proportion of clutches and chicks are exposed to agricultural activities, (2) there is little foraging habitat left when chicks hatch and (3) because of climate change, the vegetation in the remaining foraging habitat is taller and denser and therefore of lower quality. This indicates that for agri‐environment schemes to make a difference, they should not only be implemented in a larger percentage of the breeding area than the current maxima of 20–30% but they should also include measures that create more open, accessible swards.     

Brood Reduction in White Storks Mediated through Asymmetries in Plasma Testosterone Concentrations in Chicks

We hypothesized that increasing chick plasma testosterone concentrations, transmitted from the mothers via their eggs, enhances survival of their offspring and that the fitness of the young, depending on the maternal hormones, is influenced by parental quality. To test our hypotheses we distinguished the broods of white storks Ciconia ciconia L. where chicks died and those where all chicks survived. We analysed the plasma testosterone concentrations in the chicks, the ability of the chicks to be first to receive food and the mass of chicks before fledging in relation to their hatching order and recorded the body mass of parents and food mass delivered by them.
Female storks used the asymmetries in testosterone concentrations within a brood to control brood size and adjusted the number of young hatched to match the parental ability to rear offspring. Females of poor condition altered the testosterone concentrations to produce large differences between the chicks: The first-hatched chicks, which had high plasma testosterone levels, responded faster to the feeding parent and received more food than did their younger siblings. One or two later-hatched chicks, which had lower testosterone levels, died in these broods. Females in good condition produced small differences in testosterone concentrations between the chicks and all chicks survived in their brood. Chicks that were raised by the females of poor condition in reduced broods were heavier than chicks that were raised by females of good condition in broods where all chicks survived.
We suggest that the control of brood size by testosterone concentration, transmitted by the mother to the chicks, is a hormonal means of condition-dependent reproductive strategy in the white stork.     

Ecological mismatches are moderated by local conditions for two populations of a long‐distance migratory bird

Ecological mismatches between reproductive events and seasonal resource peaks are frequently proposed to be a key driver of population dynamics resulting from global climate change. Many local populations are experiencing reduced reproductive success as a consequence of mismatches, but few mismatches have led to species‐level population declines. To better understand this apparent paradox, we investigated the breeding phenology and chick survival of two disjunct populations of Hudsonian godwits Limosa haemastica breeding at Churchill, Manitoba and Beluga River, Alaska. Only one population experienced a mismatch: godwits bred nearly one week after the onset of the invertebrate peak at Churchill because of asynchronous climatic change occurring throughout their annual cycle. However, chicks were not uniformly affected by the mismatch — growth rates and survival of young chicks were not correlated with invertebrate abundance, but older chicks tended to suffer lower survival rates on days of low invertebrate abundance. Ecological mismatches thus resulted in a complex array of consequences, but nonetheless contributed to reductions in chick survival. In contrast, godwits at Beluga River hatched their chicks just before the invertebrate peak, such that the period of highest energetic need coincided with the period of highest invertebrate abundance. As a result, growth rates and survival of godwit chicks were unaffected by invertebrate abundance. Godwits at Beluga River were able to properly time their reproduction because of predictable rates of climatic change and strong selection imposed by high predation on late‐hatched chicks. Taken together, our results suggest that population‐specific, local‐scale selection pressures play a critical role in determining the degree and severity of ecological mismatches. The potential for global climate change to induce species‐level population declines may therefore be mediated by the spatial variation in the selection pressures acting across a species’ range.     

Parental and chick responses of Laughing Gulls (Leucophaeus atricilla) to increase of flight costs and brood size

One main line of thought in life history theory is that investment in offspring must be balanced to minimize negative impacts on adult survival and future breeding. Seabirds have been regarded as fixed investors, although they exhibit a whole gradient of life history traits. We studied the consequences of brood size (one and three chicks) and of increased flight costs to one mate of a pair (3- and 5-cm trimming of the edge of the primary feathers) on parental response and on survival and body condition of chicks of Laughing Gulls (Leucophaeus atricilla). Parent gulls modified their nest attendance when their mate was handicapped, in a pattern dependent on the sex of the latter. Trimming of males affected chicks more severely than that of females. On its part, brood size affected amount of feeding sessions. Both chick body condition and survival were negatively affected by larger broods and by increased flight costs of one of their parents, especially when it was the male. Overall, parental inversion exhibited adjustments depending on the requirements of the brood and the fact that males compensated better the increased flying costs of their mates than vice versa. Despite a certain capacity by the males to compensate for the increased flight costs of the females, compensation was insufficient, and much less so in females, especially in larger broods, affecting chicks’ body condition and survival.     

Greater Sage-Grouse Response to the Physical Footprint of Energy Development

Energy infrastructure and associated habitat loss can lead to reduced reproductive rates for a variety of species including the greater sage-grouse (Centrocercus urophasianus). Our goal was to refine our understanding of how the physical footprint of energy development relates to sage-grouse nest and brood survival. Our survival analyses were conditional upon the amount of surface disturbance female sage-grouse were exposed to during reproductive stages. We quantified levels of exposure and compared them to the surface disturbance levels of the surrounding area. From 2008–2014, we collected data in 6 study areas in Wyoming, USA, containing 4 primary types of renewable and nonrenewable energy development. Our research focused on press disturbance (i.e., disturbance sustained after initial disturbance and associated with existing energy infrastructure and human activity). Our results suggest exposure to press disturbance during nesting and brood-rearing was related to lower nest and brood survival, which manifested at different spatial scales. Our analysis of nest survival suggested that the likelihood of a successful nest was negatively associated with the amount of press disturbance within an 8-km² area. Broods exposed to any press disturbance within a 1-km² area were less likely to survive compared to broods not exposed to press disturbance. Female sage-grouse consistently used habitat with lower disturbance levels during reproductive periods. Greater than 90% of nest and brood-rearing locations were in habitat with <3% press disturbance within a 2.7-km² area. Our research links surface disturbance associated with press disturbance to reproductive costs incurred by sage-grouse exposed to diverse energy development. Our results demonstrate a pattern of female avoidance of areas where press disturbance was high during nesting and brood-rearing and survival of nests and broods were highest in areas that had the least amount of disturbance. Our findings underscore the importance of minimizing disturbance to maintain viable sage-grouse populations. © 2020 The Wildlife Society.     

Blood parasites in sage-grouse from Nevada and Oregon

Peripheral blood smears from 196 adult and yearling female greater sage-grouse (Centrocercus urophasianus) were examined for blood parasites (167 from the breeding and 29 from the brood-rearing season) to determine prevalence of blood parasites, to attempt to correlate infection with chick survival, and to establish base-line values of prevalence in sage-grouse from Nevada and Oregon (USA). Birds were captured and released on two study areas during 1999-2001; Sheldon National Wildlife Refuge (SNWR) in northwestern Nevada, and Hart Mountain National Antelope Refuge (HMNAR) in southeastern Oregon. Birds from a third study area, Beaty's Butte grazing allotment (BB) in southeastern Oregon, were sampled in 2000 and 2001. Overall, 19 birds (10%) were positive for Leucocytozoon lovati (= L. bonasae), 1 (0.5%) for Plasmodium pedioecetii, and 2 (1%) for microfilariae. Although prevalence of L. lovati on HMNAR was 39% during the breeding season in 1999 and 100% during the brood-rearing season in 2000, statistically, prevalence of L. lovati among study areas and years was not different. However, there were statistical differences between capture periods. Overall, 31% of the hens were positive for L. lovati during the brood-rearing season compared to 6% during the breeding season. There was no difference in packed cell volume between infected and non-infected birds and no difference between age-classes. However, mean sage-grouse productivity on HMNAR was higher (1.6 chicks/hen) for non-infected (n = 10) compared to infected hens (0.7 chicks/hen; n = 7), during 1999. Based on these limited observations on HMNAR in 1999, the possible effects that L. lovati may have on young sage-grouse could be detrimental to sage-grouse populations in Nevada and Oregon.     

Successful adult willow grouse are exposed to increased harvest risk

Age and sex ratios in bag records are frequently used as indices of population composition for harvested populations. However, vulnerability to harvest may differ by age and sex thereby producing bias in population estimates. We assessed whether age and sex affected vulnerability to harvest for willow grouse (Lagopus lagopus) where adult density and brood size were known in the harvested populations. We collected bag records during 2 days of controlled hunting in 4 areas in 2 years (2007 and 2008) in Jämtland county, Sweden. We found that vulnerability to harvest was different for chicks and adults, but not between male and female adults. Hunters encountered broods at a higher rate than single birds compared to personnel conducting pre-harvest counts along line transects. Furthermore, the probability of shooting a grouse was higher in encounters of broods than individual grouse. Proportionally, we calculated about a 50% probability of a hunter shooting either a chick or an adult independent of encountering a single bird or broods of 2–10 grouse. Increasing adult density also increased the vulnerability to harvest for adults relative to chicks, independent of the chick to adult ratio in the pre-harvest population. The different vulnerability of adults and chicks to harvest observed in this study will dampen variation in age classes in bag records compared to the population, and we caution against extrapolation of age ratios in bag records to harvested populations. © 2012 The Wildlife Society.     

The Impact of Increased Food Availability on Reproduction in a Long-Distance Migratory Songbird: Implications for Environmental Change?

Many populations of migratory songbirds are declining or shifting in distribution. This is likely due to environmental changes that alter factors such as food availability that may have an impact on survival and/or breeding success. We tested the impact of experimentally supplemented food on the breeding success over three years of northern wheatears (Oenanthe oenanthe), a species in decline over much of Europe. The number of offspring fledged over the season was higher for food-supplemented birds than for control birds. The mechanisms for this effect were that food supplementation advanced breeding date, which, together with increased resources, allowed further breeding attempts. While food supplementation did not increase the clutch size, hatching success or number of chicks fledged per breeding attempt, it did increase chick size in one year of the study. The increased breeding success was greater for males than females; males could attempt to rear simultaneous broods with multiple females as well as attempting second broods, whereas females could only increase their breeding effort via second broods. Multiple brooding is rare in the study population, but this study demonstrates the potential for changes in food availability to affect wheatear breeding productivity, primarily via phenotypic flexibility in the number of breeding attempts. Our results have implications for our understanding of how wheatears may respond to natural changes in food availability due to climate changes or changes in habitat management.     

Survival of resident and translocated greater sage-grouse in Strawberry Valley,Utah: A 13-year study

Survival of greater sage-grouse (Centrocercus urophasianus) has been well described in large populations across the species range. Very little published information exists, however, on survival rates of translocated sage-grouse or grouse from a long-term (>10 yr) study. Our objectives were to estimate seasonal and annual survival rates; assess differences in survival between resident and translocated, adult and yearling, and male and female sage-grouse; identify environmental and behavioral factors associated with survival; and assess the influence of mammalian predator control on survival rates of radio-marked sage-grouse in Strawberry Valley, Utah from 1998 to 2010. We used a 2-stage model selection approach using Akaike's Information Criterion corrected for sample size (AIC_c) with known-fate models in Program MARK to evaluate the influences of seasonal, annual, demographic, and behavioral effects on survival rates of sage-grouse. We captured and fitted 535 individual sage-grouse (male and female, resident and translocated) with radio transmitters over a 13-year period and monitored them weekly. The top model of survival, which accounted for 22% of the AIC_c weight, included 3 seasons that varied by year where rates were influenced by residency, sex, and whether a female initiated a nest. A group-level covariate for the number of canids killed each year received some support as this variable improved model fit compared to identical models without it, although confidence intervals around β estimates overlapped zero slightly. All other demographic or environmental variables showed little or no support. Annual estimates of survival for females ranged between 28% and 84% depending on year and translocation source. Survival was consistently highest during the fall–winter months with a mean monthly survival rate of 0.97 (95% CI = 0.96–0.98). The lack of a control site and other potential confounding factors limit the extent of our inference with respect to predator control. Nonetheless, we suggest managers consider enhancing nesting habitat, translocating sage-grouse, and possibly controlling predators to improve survival rates of sage-grouse. © The Wildlife Society, 2013     

Field experiments on the effect of ticks on breeding success and chick health of cattle egrets

Abstract Two experiments compared broods that were naturally tick-infested with an equal number that were rendered tick-free by application of an acaricide to their nests and of barriers against further infestation. In the first experiment conducted in 1991–92 nestlings in tick-infested broods had up to 159 larval ticks at once and a mean infestation of 23.6 larval ticks per chick per day. The chick's mean tick load was inversely correlated with its longevity. Chicks that survived to at least 18 days post-hatching had significantly lower larval tick loads than those that died by 18 days, excluding the third-hatched chick in each brood whose survival rate was low irrespective of tick-infestation. At 7 days post-hatching, tick-infested chicks had a lower haematocrit and higher polychromasia than tick-free chicks. I infer that blood loss anaemia caused the deaths of tick-infested chicks in their first week. None died in their second week and the demise of those in their third week may have been due to the paralysis manifested prior to their death. I conclude that heavy tick infestation of their chicks reduced the breeding success of the parent egrets below that of the egrets whose chicks were kept free of ticks. In the second experiment in the 1992–93 season the mean level of tick infestation was much lower (5.1 per chick) and these chicks survived equally with tick-free chicks through fledging.     

Brood reduction in the Blue-eyed Shag Phalacrocorax atriceps

Brood reduction is common in a population of Blue-eyed Shags on Signy Island, South Orkney Islands. This paper describes possible adaptations which may reduce the brood. In clutches of three, the last egg was smaller, and hatched 2.4 days later than its siblings. Whilst 78–84% of first and second ('A' & 'B') chicks fledged, only 11 % of 'C' chicks did. In a sample of artificially synchronized broods chick survival was as high as in normal asynchronously hatching broods, but there were more cases of total brood loss. The age at which the C chick died was related inversely to the length of the A-C hatching interval. Relative differences in sibling weights were highest during the first 12 days, when most of the C chick deaths occurred. At this age the daily food requirements of each brood of three was one-tenth that of each brood of two just prior to fledging. It is suggested that C chicks were unable to compete effectively for a food supply which was limited by the parents, rather than by the environment. The asymptotic weight attained by A chicks was inversely related to brood size, and was greater than that of B or C chicks. Normal asynchronous broods produced at least one heavy (A) chick and one medium weight (B) chick, whilst in synchronized broods the asymptotic weight attained was similar to that of B chicks in normal broods.     

Offspring desertion and parental care in the Whiskered Tern Chlidonias hybrida

In species with biparental care, a conflict of interest can arise if one mate tries to maximize its own reproductive success at the expense of the other's. One of the mates can desert the brood to accrue a number of benefits to enhance its own fitness, leaving parental care to the remaining parent. This study is the first to describe the desertion pattern in a tern species (Sternidae). We investigated offspring desertion in the Whiskered Tern Chlidonias hybrida, a species with semi‐precocial chicks. Offspring desertion was recorded in 52% of nests prior to fledging (n = 131 nests). Females also deserted during the post‐fledging period. Of the deserters, 97% were females. Desertions started when chicks were 5 days old and no longer required intense brooding. Desertions before fledging did not affect fledging success. Provisioning rates between pair members differed, and females supplied much less food than males. Female provisioning rate affected the chances of nest desertion significantly: daily desertion rates were lower when females supplied more food. After females had deserted, males increased their provisioning rates but compensated for the loss of female care only partly in two‐ and three‐chick broods. Only in small (one‐chick) broods was compensation full. We conclude that male and female Whiskered Terns adopt different reproductive strategies in the population studied here. Females invest much less in parental care than males, providing less food and deserting more frequently. Given the ready availability of food and low predation pressure, benefits appear to accrue to females that desert; selection forces may therefore not be acting against female desertion.     

Maternal effects as drivers of sibling competition in a parent–offspring conflict context? An experimental test

Maternal effects occur when the mother's phenotype influences her offspring's phenotype. In birds, differential allocation in egg yolk components can allow mothers to compensate for the competitive disadvantage of junior chicks. We hypothesize that the parent–older chick conflict peaks at intermediate conditions: parents benefit from the younger chick(s) survival, but its death benefits the older chick in terms of growth and survival. We thus expect maternal compensation to follow a bell‐shaped pattern in relation to environmental conditions. We studied a black‐legged kittiwake (Rissa tridactyla) population where previous results revealed increased allocation of yolk testosterone in younger as compared to older chicks in intermediate conditions, in line with our theoretical framework. We therefore predicted a maternally induced increase in aggressiveness, growth, and survival for younger chicks born in intermediate environmental conditions. Controlling for parental effects and chick sex, we manipulated food availability before egg laying to create a situation with intermediate (Unfed group) and good (Fed group) environmental conditions. Within each feeding treatment, we further created experimental broods where the natural hatching order was reversed to maximize our chances to observe an effect of feeding treatment on the younger chicks' aggressiveness. As predicted, we found that chick aggressiveness was higher in younger chicks born from the Unfed group (i.e., in intermediate environmental conditions), but only when they were put in a senior position, in reversed broods. Predictions on growth and survival were not confirmed. Mothers thus seem to favor the competitiveness of their younger chick in intermediate conditions via egg yolk components, but our study also suggests that hatching asynchrony need to be small for maternal compensation to be efficient. We emphasize the need for further studies investigating other chick behaviors (e.g., begging) and focusing on the relative role of different yolk components in shaping parent–offspring conflict over sibling competition.