Survival of Greater Sage‐Grouse broods: survey method affects disturbance and age‐specific detection probability

Investigators rely on brood surveys to estimate annual fecundity of game birds. However, investigators often do not account for factors that influence brood detection probability nor rarely document how much females and their broods are disturbed (flush rates) during surveys, which could lead to biased survival estimates. We used 45 radio‐tagged female Greater Sage‐Grouse (Centrocercus urophasianus) with broods to compare detection probabilities and document disturbance among four survey methods to allow future investigators to select the method that best meets their objectives. These methods included daytime flush, daytime visual, nocturnal spotlight, and fecal surveys at nocturnal roost sites, with the latter being a novel method. We used Cormack–Jolly–Seber (CJS) models to compare detection probability and daily survival estimates for visual and fecal surveys of broods 0–47 d post‐hatch and a double‐survey approach to compare detection probabilities among flush, fecal, and spotlight surveys ~42 d post‐hatch when investigators often determine brood fate. From CJS models, detection probability for visual surveys increased with brood age (0.618–0.881), whereas detection probability for fecal surveys did not (0.748). Daily survival probability estimates increased with brood age and differed annually based on fecal surveys (2016: 0.978–1.000 and 2017: 0.839–0.998). We detected age‐specific daily survival probability with visual surveys (0.956–0.997), but not annual differences. Based on the double‐survey approach, detection probability was high (0.857–1.000) for all methods. We flushed ~310–750% fewer females and broods during fecal and spotlight surveys than during both types of daytime surveys. Our results highlight the need to account for detection probabilities among methods and document disturbance to hens and broods that can help investigators design surveys to minimize impacts to birds. Furthermore, our result suggest that actions to improve brood survival during the first week post‐hatch may improve local recruitment.     

7. SHORT NOTES FROM MEMBERS

Cooper, J., Ross, G. J. B. &; Shaughnessy, P. D. 1978. Seasonal and spatial distribution of Rockhopper Penguins ashore in South Africa. Ostrich 49:40-44. There are 30 records of Rockhopper Penguins Eudyptes chrysocome ashore in South Africa. Both the southern subspecies E.c. chrysocome, and the northern subspecies E.c. moseleyi have been recorded. The northern subspecies has occurred more frequently. Most records are of moulting juveniles in January and February. Records of adult birds are more scattered throughout the year. Rockhopper Penguins in South Africa have been recorded only south of 30S. Birds of the northern subspecies probably originate from South Atlantic islands (Tristan da Cunha group and Gough). Birds of the southern subspecies are probably from the Prince Edward Islands.     

Increasing trend in the number of Southern Rockhopper Penguins (Eudyptes c. chrysocome) breeding at the Falkland Islands

The Falkland Islands currently supports one of the largest Southern Rockhopper Penguin (Eudyptes c. chrysocome) populations. Archipelago-wide censuses conducted in 2000 and 2005 revealed that the number of breeding pairs had declined by 30 % during this period. To establish whether the breeding population continued to decline, an archipelago-wide census was conducted in 2010. We report a conservative estimate of 319,163 ±SD 24,820 pairs breeding at the Falkland Islands in 2010. This represents a 51 % increase when compared with the number counted in 2005. A simple stochastic population model was developed to investigate the extent to which changes in demographic parameters between 2005 and 2010 could account for the increase in breeding pairs. The population model predicted a 38 % increase in the number of breeding pairs over a 5-year period (289,431 ±SD 24,615). The increase in the number of breeding pairs was therefore probably attributed to improved vital rates in the period between the 2005 and 2010 archipelago-wide censuses in combination with other factors such as a reduction in the proportion of adult birds that abstained from breeding. Based on the 2010 Falkland Islands estimate, the global population of the subspecies E. c. chrysocome is now closer to 870,000 breeding pairs of which the Falkland Islands accounts for approximately 36 %, the second largest proportion after Chile. We conclude that despite fluctuations, the number of Southern Rockhopper Penguins breeding at the Falkland Islands has increased over the last 15 years and suggest that the ‘Vulnerable’ conservation status of the species be re-assessed.     

Laying date,body mass and tick infestation of nestling tropical Roseate Terns Sterna dougallii predict fledging success,first‐year survival and age at first return to the natal colony

Both intrinsic and extrinsic factors recorded at individual nests can predict offspring fitness and survival but few studies have examined these effects in the tropics. We recorded nestling survival, post‐fledging survival and age at first return of Roseate Terns breeding at Aride Island, Seychelles, over a 12‐year period (1998–2009). Nest data recorded at the egg, nestling and fledging stages were collected during six breeding seasons (1998, 2001–2005) and a capture‐mark‐recapture dataset of six cohorts of fledglings was obtained from 2001–2009. Logistic regression models were used to assess the predictive effect of reproductive variables on fledging success, while multistate capture‐mark‐recapture models were used to estimate post‐fledging survival and return–recruitment probabilities to the natal site. Nestling survival probability increased with earliness of laying and was negatively affected by tick infestation during the growth period (0–23 days). Fledging probability was also positively related to chick body condition, whereas other pre‐fledging reproductive parameters such as clutch size and egg size were not influential. A multistate modelling of age‐specific survival and return–recruitment (transition) rates found that first‐year survival differed between cohorts and was also negatively affected by tick infestation. Annual survival stabilized from age 2 onwards at 0.83 ± 0.02. Transition rates were positively related to body condition at fledging, with heavier individuals returning for the first time to the natal colony at a younger age compared with lighter individuals. These results highlight the importance of local conditions encountered by tropical seabirds during the breeding season in shaping demographic parameters.     

THE LAYING INTERVAL AND INCUBATION PERIOD OF ROCKHOPPER AND MACARONI PENGUINS

Williams, A. J. 1981. The laying interval and incubation period of Rockhopper and Macaroni Penguins. Ostrich 52: 226–229.

The laying interval and incubation period of Rockhopper Penguins Eudyptes chrysocome and Macaroni Penguins E. chrysolophus were studied at Marion Island in 1974–75 and 1976–77. On average, the laying interval was 4,4 and 4,5 days, the incubation period of second-laid eggs was 34,2 and 35,9 days and that of first-laid eggs was 39,1 and 38,0 days in Rock-hopper and Macaroni Penguins respectively. The laying interval in this genus is longer than that in other penguins. The incubation period is similar to that of most other penguins but the second-laid egg normally hatches before the first-laid egg. The long laying interval and the hatching sequence of the eggs both have important affects upon the mortality of eggs in the genus Eudyptes.     

Inter-annual variation in provisioning behaviour of Southern Rockhopper Penguins Eudyptes chrysocome chrysocome at Staten Island, Argentina

Understanding the relationship between reproductive performance and food availability requires knowledge about many different variables, including such factors as the length of incubation shifts, provisioning rates and patterns, as well as how variability in these factors affects reproductive output. To examine some of the most important aspects of parental investment, we studied the provisioning behaviour and patterns of adult Southern Rockhopper Penguins Eudyptes chrysocome chrysocome breeding at Staten Island, Argentina. We investigated foraging trip duration, provisioning rates and chick survival using adult foraging patterns. Our results show that Rockhopper Penguins had clear sex-specific differences in their provisioning behaviour. Females provision chicks throughout chick rearing. By contrast, males provision chicks only during the crèche stage and at a slightly lower rate than females during this period. Foraging trips increased in length as the breeding season progressed. Rockhopper Penguins from Staten Island performed longer trips throughout the breeding season than do other species of Eudyptes at several other locations. Our results also show differences in parental investment between years that were related to differences in chick survival. We suggest that this was most likely to be related to female rather than male foraging behaviour as only females showed inter-annual differences in their provisioning rates.     

Habitat use and spatial fidelity of male South American sea lions during the nonbreeding period

Conditions experienced during the nonbreeding period have profound long‐term effects on individual fitness and survival. Therefore, knowledge of habitat use during the nonbreeding period can provide insights into processes that regulate populations. At the Falkland Islands, the habitat use of South American sea lions (Otaria flavescens) during the nonbreeding period is of particular interest because the population is yet to recover from a catastrophic decline between the mid‐1930s and 1965, and nonbreeding movements are poorly understood. Here, we assessed the habitat use of adult male (n = 13) and juvenile male (n = 6) South American sea lions at the Falkland Islands using satellite tags and stable isotope analysis of vibrissae. Male South American sea lions behaved like central place foragers. Foraging trips were restricted to the Patagonian Shelf and were typically short in distance and duration (127 ± 66 km and 4.1 ± 2.0 days, respectively). Individual male foraging trips were also typically characterized by a high degree of foraging site fidelity. However, the isotopic niche of adult males was smaller than juvenile males, which suggested that adult males were more consistent in their use of foraging habitats and prey over time. Our findings differ from male South American sea lions in Chile and Argentina, which undertake extended movements during the nonbreeding period. Hence, throughout their breeding range, male South American sea lions have diverse movement patterns during the nonbreeding period that intuitively reflects differences in the predictability or accessibility of preferred prey. Our findings challenge the long‐standing notion that South American sea lions undertake a winter migration away from the Falkland Islands. Therefore, impediments to South American sea lion population recovery likely originate locally and conservation measures at a national level are likely to be effective in addressing the decline and the failure of the population to recover.     

SUMMER DISTRIBUTION OF PELAGIC BIRDS OFF THE COAST OF ARGENTINA

Eighteen species of sea–birds were observed from CSS 'Hudson' in the South Atlantic between the Rio de la Plata and Tierra del Fuego in late January and late February 1970. Major feeding areas were noted off Mar del Plata, the Valdes Peninsula and Tierra del Fuego, Argentina, the first two in rich modified or unmodified water of the subantarctic Falkland Current, associated with upwelling. The southernmost area had most birds, the northernmost fewest. A temperate–latitude assemblage including Yellow–nosed Albatross and Cory's Shearwater occurred off Mar del Plata in warmed water of subantarctic (Falkland Current) origin. South of 39°S these two species were replaced by Black–browed Albatross and Great Shearwater in cold offshore water of the Falkland Current. Off the Valdes Peninsula Black–browed Albatross, Great and Sooty Shearwaters, White–chinned Petrels and family groups of Magellanic Penguins occurred near the edge of highly–productive water of the Falkland Current. The largest feeding area, east of Tierra del Fuego, had Magellanic and Rockhopper Penguins, Great Shearwaters, King Cormorants, South American Terns and jaegers, but no oceanographic information is available. At 50°S prions and diving petrels appeared for the first time, marking a possible oceanographic discontinuity or proximity to colonies on the Falkland Islands. Several common species of the South Atlantic were not seen on the transect including Cape Pigeon, Southern Fulmar, Soft–plumaged Petrel, Schlegel's Petrel and Pediunker. Kelp Gull, South American Tern, Manx Shearwater and jaegers were rare or absent far from land. New southern limits for the western South Atlantic Ocean were established for Cory's and Manx Shearwaters, and the Great Shearwater was observed for the first time in Chilean waters.     

Algal Parasite Herpodiscus durvillaeae (Phaeophyceae: Sphacelariales) Inferred to have Traversed the Pacific Ocean with its Buoyant Host

The parasitic phaeophycean endophyte Herpodiscus durvillaeae (Lindauer) G. R. South has previously only been recorded from New Zealand, in association with a single host species, Durvillaea antarctica (Chamisso) Hariot (southern bull‐kelp). Here we use DNA sequence data from plastid and nuclear markers (chloroplast rbcL, ribosomal LSU, and a nuclear pseudogene copy of COI) to test for the presence of H. durvillaeae beyond the New Zealand region, and on host species other than D. antarctica. Analyses of samples from the Falkland Islands confirm the first record of H. durvillaeae from the Atlantic Ocean. We report that Falkland Islands H. durvillaeae are genetically indistinguishable from samples of this species from New Zealand's sub‐Antarctic Campbell Island, suggesting recent dispersal of the parasite across the Pacific Ocean, presumably by rafting with its buoyant macroalgal host. We also here record H. durvillaeae from New Zealand endemics Durvillaea poha Fraser et al. and D. willana Lindauer.     

Demographic parameters of black-browed albatrosses Thalassarche melanophris from the Falkland Islands

Black-browed albatrosses Thalassarche melanophris are currently classified as globally endangered. The most important populations of this species are believed to be declining due to, amongst other factors, unsustainable levels of incidental mortality in fishing gear. However, detailed demographic data are lacking for several critical populations, including the largest of all, nesting in the Falkland Islands. Here, we present data from the first Falkland Islands detailed demographic study (at New Island) and show that, from 2003 to 2009, the mean adult survival probability was 0.942 (95% CI: 0.930–0.952). Nesting frequency of adults is amongst the highest recorded for Thalassarche albatrosses and breeding success (0.564 chicks per egg) is within normal values. The nesting population in the intensively studied plots experienced an increase of 4% per year from 2004 to 2009. These results indicate that the Falklands population may not be as threatened as previously supposed, although studies from more sites and a longer time series are needed to confirm or refute this. The high survival rates may partly reflect recent efforts to mitigate bycatch made by the Falkland Islands and other fisheries in the region. The reinforcement of such initiatives may be critical to buffer the black-browed albatross population against ecosystem shifts and natural disasters (such as harmful algal blooms) that will likely become more frequent with ongoing global changes.     

Intronic variation at the CHD1‐Z gene in Black‐tailed Godwits Limosa limosa limosa: correlations with fitness components revisited

Recently, Schroeder et al. (2010, Ibis 152: 368–377) suggested that intronic variation in the CHD1‐Z gene of Black‐tailed Godwits breeding in southwest Friesland, The Netherlands, correlated with fitness components. Here we re‐examine this surprising result using an expanded dataset (2088 birds sampled from 2004 to 2010 vs. 284 birds from 2004 to 2007). We find that the presence of the Z* allele (9% of the birds) is not associated with breeding habitat type, egg size, adult survival, adult body mass or adult body condition. The results presented here, when used in synergy with the previously reported results by Schroeder et al., suggest that there might be a tendency towards female adults with the Z* allele laying earlier clutches than adult females without the Z* allele. The occurrence of the Z* allele was also associated with a higher chick body mass and return rate. Chicks with the Z* allele that had hatched early in the breeding season were heavier at birth than chicks without the Z* allele and chicks with the Z* allele that had hatched late. Collectively, the results suggest that variation in the CHD1‐Z gene may indeed have arisen as a byproduct of selection acting on females during the egg fase and on chicks during the rearing stages of the reproductive cycle.     

Effects of food availability on fledging condition and post-fledging survival in king penguin chicks

Effects of summer food shortage on king penguin Aptenodytes patagonicus chicks were studied at South Georgia. Two cohorts were compared, fledging in the austral summers of 1992 (n = 32) and 1994 (n = 33) when availability of food was judged good and poor, respectively. The former cohort had a higher pre-fledging mean mass (12.78 kg vs ≤ 10.03 kg), fledged earlier (median 5 January vs 21 January), and a higher proportion was re-sighted within 2 years of fledging (28% vs 0%). Within 4 years, 47% of the former cohort had been re-sighted (i.e. post-fledge survival); in addition, one was observed at the Falkland Islands, and 22% had bred (i.e. recruitment) in their colony of origin. The re-sighted chicks of the 1992 cohort fledged earlier than those not re-sighted (median 24 December vs 10 January), but it remain unclear if they were heavier at fledging. All chicks in this study (n = 65) were marked with both transponders (subcutaneously implanted) and flipper bands (on one flipper), and no losses of any markings were found (controlled up to 4 years afterwards). Therefore, data on chick post-fledging survival and recruitment were not adjusted for losses of markings, as has been done in other studies. Received: 21 October 1996 / Accepted: 2 February 1997     

Increased lifetime reproductive success of first‐hatched siblings in Common Kestrels Falco tinnunculus

Order of birth has profound consequences on offspring across taxa during development and can have effects on individuals later in life. In birds, differential maternal allocation and investment in their progeny lead to variance in the environmental conditions that offspring experience during growth within the brood. In particular, laying and hatching order have been proposed to influence individual quality during the growing period, but little is known about the fitness consequences that these two factors have for offspring from a lifetime perspective. We explored the effect of laying and hatching order on post‐fledgling survival (measured as recruitment probability) and lifetime reproductive success (LRS) in Common Kestrels Falco tinnunculus, using a long‐term and individual‐based dataset. First‐hatched chicks showed higher survival probability and LRS than their siblings. This effect was not due to body condition of the individuals at adulthood, the quality of their mates or the reproductive outcome compared with later‐hatched individuals. Instead, first‐hatched chicks had a higher recruitment probability. This could be explained by the higher body condition attained by first‐hatched chicks at the end of the nesting period, perhaps due to an enhanced competitive advantage for food over their siblings at the time of hatching. Laying order, in contrast to hatching order, appeared to have little or no effect on LRS. Our results suggest that hatching order within siblings predicts fitness, and that better early‐life conditions during growth experienced by first‐hatched chicks improve first survival and then recruitment, resulting in an enhanced LRS.     

Age‐specific survival and movement among major African Penguin Spheniscus demersus colonies

Reliable estimates of survival and dispersal are crucial to understanding population dynamics, but for seabirds, in which some individuals spend years away from land, mortality and emigration are often confounded. Multistate mark–recapture methods reduce bias by incorporating movement into the process of estimating survival. We used a multistate model to provide unbiased age‐specific survival and movement probabilities for the Endangered African Penguin Spheniscus demersus based on 5281 nestlings and 31 049 adults flipper‐banded and resighted in the Western Cape, South Africa, between 1994 and 2012. Adult survival was initially high (≥ 0.74) but declined after 2003–2004 coincident with a reduction in the availability of Sardine Sardinops sagax and Anchovy Engraulis encrasicolus on the west coast of South Africa. Juvenile survival was poorly estimated, but was lower and more variable than adult survival. Fidelity to the locality of origin varied over time, but was high in adults at Robben and Dassen islands (≥ 0.88) and above 0.55 for juvenile and immature Penguins at all localities. Movement occurred predominantly during 1994–2003 and was indicative of immigration to Robben and Dassen islands. Our results confirm that a prolonged period of adult mortality contributed to the observed decline in the African Penguin population and suggest a need for approaches operating over large spatial scales to ensure food security for marine top predators.     

Climate as a driver of population variability in breeding Gentoo Penguins Pygoscelis papua at the Falkland Islands

Detecting and predicting how populations respond to environmental variability are eminent challenges in conservation research and management. This is particularly true for wildlife populations at high latitudes, many of which demonstrate changes in population dynamics associated with global warming. The Falkland Islands (Southwest Atlantic) hold one of the largest Gentoo Penguin Pygoscelis papua populations in the world, representing c. 34% of the global population. The numbers of breeding Gentoo Penguins at the Falkland Islands have shown a high degree of inter‐annual variability since monitoring commenced in 1990. However, proximate causes of annual variability in breeding numbers have not been explored. Here we examine 21 years of Gentoo Penguin breeding surveys from the Falkland Islands and assess whether inter‐annual variability in the number of breeding pairs were correlated with proxies of environmental variability. There was a positive correlation between the number of breeding pairs and a broad‐scale climatic variation index, the Southern Oscillation Index (SOI). In turn, the SOI was significantly correlated with spring sea surface temperature anomalies, indicating a more immediate atmospherically forced response to El Niño Southern Oscillation variability in the Southwest Atlantic than previously reported. However, we also describe a non‐linear response to environmental variability that may highlight foraging plasticity and/or the complexity of regional ecosystem interactions that operate across a range of different scales.     

PREY OF NORTHERN ROCKHOPPER PENGUINS AT GOUGH ISLAND,SOUTH ATLANTIC OCEAN

Klages, N. T., Brooke, M. de L. & Watkins, B. P. 1988. Prey of Northern Rockhopper Penguins at Gough Island, south Atlantic Ocean. Ostrich 59:162-165.

The diet of Northern Rockhopper Penguins Eudyptes chrysocome moseleyi breeding on Gough Island, south Atlantic Ocean was studied, during November 1984, 1985 and 1986 by stomach content analysis. Rockhopper Penguins fed chiefly on the euphausiids Thysanoessa gregaria, Euphausia lucens and E. similis. Fish and squid were of minor importance by mass but constituted the largest individual prey items.     

Estimating age‐dependent survival when juveniles resemble females: Invasive ring‐necked parakeets as an example

Many species only show sexual dimorphism at the age of maturity, such that juveniles typically resemble females. Under these circumstances, estimating accurate age‐specific demographic parameters is challenging. Here, we propose a multievent model parameterization able to estimate age‐dependent survival using capture–recapture data with uncertainty in age and sex assignment of individuals. We illustrate this modeling approach with capture–recapture data from the ring‐necked parakeet Psittacula krameri. We analyzed capture, recapture, and resighting data (439 recaptures/resightings) of 156 ring‐necked parakeets tagged with neck collars in Barcelona city from 2003 to 2016 to estimate the juvenile and adult survival rate. Our models successfully estimated the survival probabilities of the different age classes considered. Survival probability was similar between adults (0.83, 95% CI = 0.77–0.87) and juveniles during their second (0.79, 95% CI = 0.58–0.87) and third winter (0.83, 95% CI = 0.65–0.88). The youngest juveniles (1st winter) showed a slightly lower survival (0.57, 95% CI = 0.37–0.79). Among adults, females showed a slightly higher survival than males (0.87, 95% CI = 0.78–0.93; and 0.80, 95% CI = 0.73–0.86, respectively). These high survival figures predict high population persistence in this species and urge management policies. The analysis also stresses the usefulness of multievent models to estimate juvenile survival when age cannot be fully ascertained.     

A recipe for postfledging survival in great tits Parus major: be large and be early (but not too much)

Survival of juveniles during the postfledging period can be markedly low, which may have major consequences on avian population dynamics. Knowing which factors operating during the nesting phase affect postfledging survival is crucial to understand avian breeding strategies. We aimed to obtain a robust set of predictors of postfledging local survival using the great tit (Parus major) as a model species. We used mark–recapture models to analyze the effect of hatching date, temperatures experienced during the nestling period, fledging size and body mass on first‐year postfledging survival probability of great tit juveniles. We used data from 5192 nestlings of first clutches ringed between 1993 and 2010. Mean first‐year postfledging survival probability was 15.2%, and it was lower for smaller individuals, as well as for those born in either very early or late broods. Our results stress the importance of choosing an optimum hatching period, and raising large chicks to increase first‐year local survival probability in the studied population.     

Equal nonbreeding period survival in adults and juveniles of a long‐distant migrant bird

In migrant birds, survival estimates for the different life‐history stages between fledging and first breeding are scarce. First‐year survival is shown to be strongly reduced compared with annual survival of adult birds. However, it remains unclear whether the main bottleneck in juvenile long‐distant migrants occurs in the postfledging period within the breeding ranges or en route. Quantifying survival rates during different life‐history stages and during different periods of the migration cycle is crucial to understand forces driving the evolution of optimal life histories in migrant birds. Here, we estimate survival rates of adult and juvenile barn swallows (Hirundo rustica L.) in the breeding and nonbreeding areas using a population model integrating survival estimates in the breeding ranges based on a large radio‐telemetry data set and published estimates of demographic parameters from large‐scale population‐monitoring projects across Switzerland. Input parameters included the country‐wide population trend, annual productivity estimates of the double‐brooded species, and year‐to‐year survival corrected for breeding dispersal. Juvenile survival in the 3‐week postfledging period was low (S = 0.32; SE = 0.05), whereas in the rest of the annual cycle survival estimates of adults and juveniles were similarly high (S > 0.957). Thus, the postfledging period was the main survival bottleneck, revealing the striking result that nonbreeding period mortality (including migration) is not higher for juveniles than for adult birds. Therefore, focusing future research on sources of variation in postfledging mortality can provide new insights into determinants of population dynamics and life‐history evolution of migrant birds.     

Restoration of indigenous dominance in exotic grassland by the establishment of juvenile Microlaena stipoides plants

Two experiments were conducted to determine the establishment success of reintroducing Microlaena stipoides (pātītī, weeping rice grass) into existing high‐fertility grassland on the volcanic cones of the Auckland Isthmus. The first experiment monitored the survival and development of juveniles planted in a factorial design including two planting densities, two slope classes and two aspects across three cones. Plant survival during establishment was consistently over 90%. Maximum M. stipoides cover after 2 years (>80%) was achieved on north‐facing steep slopes (>25°) at the greater planting density (40 plants/m²). However, results were particularly idiosyncratic to specific cone/topographical combinations. The second experiment, on a flat site on one cone, monitored the survival and development of juveniles planted across four post‐planting treatments designed to suppress resident vegetation recovery. Plant survival after 6 months was relatively low (50%), and none of the treatments achieved greater than 5% cover of M. stipoides after 1 year. The chosen post‐planting treatments were unable to suppress vigorous recovery of competitive exotic grasses on a moist fertile site. Overall, juvenile planting was shown to be a potentially successful method of Microlaena establishment and could restore indigenous dominance to exotic grassland in this environment, but individual site factors and the high cost of establishment must be considered.