Best squirrels trade a long life for an early reproduction

Age at primiparity plays a crucial role in population dynamics and life-history evolution. Long-term data on female North American red squirrels were analysed to study the fitness consequences of delaying first reproduction. Early breeders were born earlier, had a higher breeding success and achieved a higher lifetime reproductive success than females who delayed their first reproduction, which suggests a higher quality of early breeders. However, early breeders had similar mass when tagged, and similar number of food caches available at one year of age as late breeders. Nevertheless, we found evidence of survival costs of early primiparity. Early breeders had a lower survival between one and two years of age than late breeders and a lower lifespan. Our study points out that two reproductive tactics co-occurred in this population: a tactic based on early maturity at the cost of a lower survival versus a tactic based on delayed maturity and long lifespan. High quality individuals express the most profitable tactic by breeding early whereas low quality individuals do the best of a bad job by delaying their first reproduction.     

Sex-specific costs of reproduction on survival in a long-lived seabird

Costs of reproduction on survival have captured the attention of researchers since life history theory was formulated. Adults of long-lived species may increase survival by reducing their breeding effort or even skipping reproduction. In this study, we aimed to evaluate the costs of current reproduction on survival and whether skipping reproduction increases adult survival in a long-lived seabird. We used capture–mark–recapture data (1450 encounters) from two populations of Bulwer''s petrel (Bulweria bulwerii), breeding in the Azores and Canary Islands, North Atlantic Ocean. Using a multi-event model with two different breeding statuses (breeders versus non-breeders), we calculated probabilities of survival and of transitions between breeding statuses, evaluating potential differences between sexes. Females had lower survival probabilities than males, independent of their breeding status. When considering breeding status, breeding females had lower survival probabilities than non-breeding females, suggesting costs of reproduction on survival. Breeding males had higher survival probabilities than non-breeding males, suggesting that males do not incur costs of reproduction on survival and that only the highest quality males have access to breeding. The highest and the lowest probabilities of skipping reproduction were found in breeding males from the Azores and in breeding males from the Canary Islands, respectively. Intermediate values were observed in the females from both populations. This result is probably due to differences in the external factors affecting both populations, essentially predation pressure and competition. The existence of sex-specific costs of reproduction on survival in several populations of this long-lived species may have important implications for species population dynamics.     

Heterogeneity in individual quality overrides costs of reproduction in female reindeer

Reproductive allocation at one age is predicted to reduce the probability of surviving to the next year or to lead to a decrease in future reproduction. This prediction assumes that reproduction involves fitness costs. However, few empirical studies have assessed whether such costs may vary with the age at primiparity or might be overridden by heterogeneities in individual quality. We used data from 35 years’ monitoring of individually marked semi-domestic reindeer females to investigate fitness costs of reproduction. Using multi-state statistical models, we compared age-specific survival and reproduction among four reproductive states (never reproduced, experienced non-breeders, reproduced but did not wean offspring, and reproduced and weaned offspring) and among contrasted age at primiparity. We assessed whether reproductive costs occurred, resulting in a trade-off between current reproduction and future reproduction or survival, and whether early maturation was costly or rather reflected differences in individual quality of survival and reproduction capabilities. We did not find any evidence for fitness costs of reproduction in female reindeer. We found no cost of gestation and lactation in terms of future reproduction and survival. Conversely, successful breeders had higher survival and subsequent reproductive success than experienced non-breeders and unsuccessful breeders, independently of the age at primiparity. Moreover, it was beneficial to mature earlier, especially for females that successfully weaned their first offspring. Successful females at early primiparity remained successful throughout their life, clearly supporting the existence of marked among-female differences in quality. The weaning success peaked for multiparous females and was lower for first-time breeders, indicating a positive effect of experience on reproductive performance. Our findings emphasize an overwhelming importance of individual quality and experience to account for observed variation in survival and reproductive patterns of female reindeer that override trade-offs between current reproduction and future performance, at least in the absence of harsh winters.     

Individual heterogeneity in life-history trade-offs with age at first reproduction in capital breeding elephant seals

Recruitment age plays a key role in life-history evolution. Because individuals allocate limited resources among competing life-history functions, theory predicts trade-offs between current reproduction and future growth, survival and/or reproduction. Reproductive costs tend to vary with recruitment age, but may also be overridden by fixed individual differences leading to persistent demographic heterogeneity and positive covariation among demographic traits at the population level. We tested for evidence of intra- and inter-generational trade-offs and individual heterogeneity relating to age at first reproduction using three decades of detailed individual life-history data of 6,439 capital breeding female southern elephant seals. Contrary to the predictions from trade-off hypotheses, we found that recruitment at an early age was associated with higher population level survival and subsequent breeding probabilities. Nonetheless, a survival cost of first reproduction was evident at the population level, as first-time breeders always had lower survival probabilities than prebreeders and experienced breeders of the same age. However, models accounting for hidden persistent demographic heterogeneity revealed that the trade-off between first reproduction and survival was only expressed in “low quality” individuals, comprising 35% of the population. The short-term somatic costs associated with breeding at an early age had no effect on the ability of females to allocate resources to offspring in the next breeding season. Our results provide strong evidence for individual heterogeneity in the life-history trajectories of female elephant seals. By explicitly modeling hidden persistent demographic heterogeneity we show that individual heterogeneity governs the expression of trade-offs with first reproduction in elephant seals.     

Now you see him, now you don't: experience, not age, is related to reproduction in kittiwakes

In long-lived species, individuals can skip reproduction. The proportion of breeders affects population growth rate and viability, there is a need to investigate the factors influencing intermittent breeding. The theory predicts that if lack of experience is an important constraint, breeding probabilities should increase with experience for individuals of the same age, whereas under the so-called restraint hypothesis, breeding probabilities should increase with age regardless of experience. However, because the probability of detecting individuals in the wild is generally less than 1, it is difficult to know exactly the number of previous breeding episodes (breeding experience). To cope with this issue, we developed a hidden process model to incorporate experience as a latent state possibly influencing the probability of breeding. Using a 22-year mark-recapture dataset involving 9970 individuals, we analysed simultaneously experience and age effects on breeding probabilities in the kittiwake (Rissa tridactyla). We did not detect an influence of age on adult breeding probabilities. We found that inexperienced birds breed less frequently than experienced birds. Our approach enables us to highlight the key role of experience on adults breeding probabilities and can be used for a wide range of organisms for which detection is less than 1.     

To breed or not to breed: a seabird's response to extreme climatic events

Intermittent breeding is an important life-history strategy that has rarely been quantified in the wild and for which drivers remain unclear. It may be the result of a trade-off between survival and reproduction, with individuals skipping breeding when breeding conditions are below a certain threshold. Heterogeneity in individual quality can also lead to heterogeneity in intermittent breeding. We modelled survival, recruitment and breeding probability of the red-footed booby (Sula sula), using a 19 year mark–recapture dataset involving more than 11 000 birds. We showed that skipping breeding was more likely in El-Niño years, correlated with an increase in the local sea surface temperature, supporting the hypothesis that it may be partly an adaptive strategy of birds to face the trade-off between survival and reproduction owing to environmental constraints. We also showed that the age-specific probability of first breeding attempt was synchronized among different age-classes and higher in El-Niño years. This result suggested that pre-breeders may benefit from lowered competition with experienced breeders in years of high skipping probabilities.     

What shall I do now? State-dependent variations of life-history traits with aging in Wandering Albatrosses

Allocation decisions depend on an organism's condition which can change with age. Two opposite changes in life‐history traits are predicted in the presence of senescence: either an increase in breeding performance in late age associated with terminal investment or a decrease due to either life‐history trade‐offs between current breeding and future survival or decreased efficiency at old age. Age variation in several life‐history traits has been detected in a number of species, and demographic performances of individuals in a given year are influenced by their reproductive state the previous year. Few studies have, however, examined state‐dependent variation in life‐history traits with aging, and they focused mainly on a dichotomy of successful versus failed breeding and non‐breeding birds. Using a 50‐year dataset on the long‐lived quasi‐biennial breeding wandering albatross, we investigated variations in life‐history traits with aging according to a gradient of states corresponding to potential costs of reproduction the previous year (in ascending order): non‐breeding birds staying at sea or present at breeding grounds, breeding birds that failed early, late or were successful. We used multistate models to study survival and decompose reproduction into four components (probabilities of return, breeding, hatching, and fledging), while accounting for imperfect detection. Our results suggest the possible existence of two strategies in the population: strict biennial breeders that exhibited almost no reproductive senescence and quasi‐biennial breeders that showed an increased breeding frequency with a strong and moderate senescence on hatching and fledging probabilities, respectively. The patterns observed on survival were contrary to our predictions, suggesting an influence of individual quality rather than trade‐offs between reproduction and survival at late ages. This work represents a step further into understanding the evolutionary ecology of senescence and its relationship with costs of reproduction at the population level. It paves the way for individual‐based studies that could show the importance of intra‐population heterogeneity in those processes.     

Indirect effects of parasitism: costs of infection to other individuals can be greater than direct costs borne by the host

Parasitic infection has a direct physiological cost to hosts but may also alter how hosts interact with other individuals in their environment. Such indirect effects may alter both host fitness and the fitness of other individuals in the host''s social network, yet the relative impact of direct and indirect effects of infection are rarely quantified. During reproduction, a host''s social environment includes family members who may be in conflict over resource allocation. In such situations, infection may alter how resources are allocated, thereby redistributing the costs of parasitism between individuals. Here, we experimentally reduce parasite burdens of parent and/or nestling European shags (Phalacrocorax aristotelis) infected with Contracaecum nematodes in a factorial design, then simultaneously measure the impact of an individual''s infection on all family members. We found no direct effect of infection on parent or offspring traits but indirect effects were detected in all group members, with both immediate effects (mass change and survival) and longer-term effects (timing of parents’ subsequent breeding). Our results show that parasite infection can have a major impact on individuals other than the host, suggesting that the effect of parasites on population processes may be greater than previously thought.     

Age-specific survival and reproductive performances in fur seals: evidence of senescence and individual quality

Life history theory hypothesises that breeding events induce reproductive costs that may vary among individuals. However, the growing number of studies addressing this question are taxonomically biased, therefore impeding the generalisation of this hypothesis, especially with regard to marine top predators. This study investigated age‐related survival and breeding performances in subantarctic fur seal (Arctocephalus tropicalis) females from Amsterdam Island, southern Indian Ocean. Using multistate capture–recapture models on data obtained from known‐age tagged females over eight consecutive years, we tested for evidence of senescence, individual quality, and reproductive costs in terms of future survival and fecundity. Adult female yearly survival appeared high and constant throughout time. While a two age‐class model was preferred in non‐breeders, breeding females exhibited three age classes with a maximum survival for the prime‐age class (7–12 years). Survival and reproductive probabilities decreased from 13 years onward, suggesting senescence in this population. Survival was lower for non‐breeders than for breeders, among both prime‐aged (0.938 vs 0.982) and older (0.676 vs 0.855) females. Furthermore, non‐breeders exhibited higher probabilities of being non‐breeders the following year than did breeders (0.555 vs 0.414). Such results suggest consistency in female breeding performance over years, supporting the hypothesis that non‐breeding tend to occur among lower quality individuals rather than representing an alternative strategy to enhance residual reproductive value. However, the high proportion of females that did not breed during two consecutive years, and the lower probability of being a successful breeder after a greater reproductive effort confirmed the existence of reproductive costs, especially during the second half of the lactation. These results also suggest that younger age‐classes included a higher proportion of lower quality individuals, which are likely to face higher costs of reproduction. Such hypotheses lead to consider the first breeding event as a filter generating a within‐cohort selection process in females.     

Survival costs of reproduction vary with age in North American red squirrels

The costs of reproduction are expected to be higher under unfavourable conditions, so that breeding in years of low food supply should have important costs. In addition, the costs of reproduction may be contingent on the age of individuals, and young growing and old senescent individuals should suffer higher costs than the prime-age ones. We tested these predictions by investigating the costs of reproduction as a function of food availability and age in female North American red squirrels using the long-term data on survival and reproduction. We found that the costs of reproduction were independent of food supply, and we did not detect any trade-off between the current and future reproduction. We also did not detect any survival cost of reproduction for the prime-age females, but found evidence for survival costs in yearlings and old (6 years or above) females with successfully breeding individuals having a lower chance of survival compared with unsuccessful or non-breeding ones. These results supported our prediction that the costs of reproduction depended on the age of female red squirrels and were higher in young growing and old senescent individuals. Our study also indicated that, in contrast to large herbivores, heterogeneity in individual quality and viability selection in red squirrels do not affect the study of trade-offs and of the age variation in life-history traits.     

Evaluation of reproductive costs for Weddell seals in Erebus Bay, Antarctica

1. Organisms balance current reproduction against future survival and reproduction, which results in life-history trade-offs. These trade-offs are also known as reproductive costs and may represent significant factors shaping life-history strategy for many species. 2. Using multistate mark-resight models and 26 years of mark-resight data (1979-2004), we estimated the costs of reproduction to survival and reproductive probabilities for Weddell seals in Erebus Bay, Antarctica and evaluated whether this species either conformed to the 'prudent parent' reproductive strategy predicted by life-history theory for long-lived mammals or alternatively, incurred costs to survival in order to reproduce in a variable environment (flexible-strategy hypothesis). 3. Results strongly supported the presence of reproductive costs to survival (mean annual survival probability was 0.91 for breeders vs. 0.94 for nonbreeders), a notable difference for a long-lived mammal, demonstrating that investment in reproduction does result in a cost to survival for Weddell seals, contrary to the prudent parent hypothesis. 4. Reproductive costs to subsequent reproductive probabilities were also present for first-time breeders (mean probability of breeding the next year was 31.3% lower for first-time breeders than for experienced breeders), thus supporting our prediction of the influence of breeding experience. 5. We detected substantial annual variation in survival and breeding probabilities. Breeding probabilities were negatively influenced by summer sea-ice extent, whereas weak evidence suggested that survival probabilities were affected more by winter sea-ice extent, and the direction of this effect was negative. However, a model with annual variation unrelated to any of our climate or sea-ice covariates performed best, indicating that further study will be needed to determine the appropriate mechanism or combination of mechanisms underlying this annual variation.     

Determinants and consequences of age of primiparity in bighorn ewes

Because variation in age of first reproduction can have major effects on individual fitness and population dynamics, it is important to understand what maintains that variability. Although early primiparity is assumed to be costly, it is sometimes associated with high lifetime reproductive success. We used a long‐term study on bighorn sheep Ovis canadensis to determine what variables affect age at first reproduction, investigate the impact of primiparity on body resources and quantify the reproductive performance of primiparous ewes. We then examined the consequences of delayed primiparity on adult body mass, longevity and lifetime reproductive success. Environmental conditions during early development, body mass as a yearling, genotype and maternal effects affected age of primiparity. Primiparous ewes lost more mass in winter and gained less mass in summer than multiparous ewes. Small yearling ewes that postponed reproduction attained similar adult mass than heavy yearling ewes who reproduced at a younger age. Early primiparity did not reduce longevity and was positively associated with lifetime reproductive success. Starting to reproduce as soon as possible appears to maximize fitness of females. When early life conditions are unfavorable, however, delayed primiparity allows greater body growth and likely maximizes survival. The combination of a conservative reproductive strategy and maternal effects on age of primiparity may partly delay population recovery following density‐dependent declines.     

Nest fidelity is driven by multi-scale information in a long-lived seabird

Although the reproductive success of most organisms depends on factors acting at several spatial scales, little is known about how organisms are able to synthesize multi-scale information to optimize reproduction. Using longitudinal data from a long-lived seabird, Monteiro''s storm-petrel, we show that average breeding success is strongly related to oceanic conditions at the population level, and we postulate that (i) individuals use proximal information (their own reproduction outcome in year t) to assess the qualities of their mate and nest and to decide to retain them or not in year t + 1; (ii) the intensity of these responses depends on the quality of the oceanic environment in year t, which affects the predictability of reproduction outcome in year t + 1. Our results confirm that mate and nest fidelities are higher following successful reproduction and that the relationship between the success of a given pair and subsequent nest fidelity is stronger in years with unfavourable oceanic conditions, suggesting that individuals rely on distant information to modulate their use of proximal information and adjust their breeding strategy.     

Cooperative breeding in South American hunter–gatherers

Evolutionary researchers have recently suggested that pre-modern human societies habitually practised cooperative breeding and that this feature helps explain human prosocial tendencies. Despite circumstantial evidence that post-reproductive females and extra-pair males both provide resources required for successful reproduction by mated pairs, no study has yet provided details about the flow of food resources by different age and sex categories to breeders and offspring, nor documented the ratio of helpers to breeders. Here, we show in two hunter–gatherer societies of South America that each breeding pair with dependent offspring on average obtained help from approximately 1.3 non-reproductive adults. Young married males and unmarried males of all ages were the main food providers, accounting for 93–100% of all excess food production available to breeding pairs and their offspring. Thus, each breeding pair with dependants was provisioned on average by 0.8 adult male helpers. The data provide no support for the hypothesis that post-reproductive females are the main provisioners of younger reproductive-aged kin in hunter–gatherer societies. Demographic and food acquisition data show that most breeding pairs can expect food deficits owing to foraging luck, health disabilities and accumulating dependency ratio of offspring in middle age, and that extra-pair provisioning may be essential to the evolved human life history.     

Effects of age,territoriality and breeding on survival of Bonelli’s Eagle Aquila fasciata

Survival typically contributes most to population trends in long‐lived birds and its accurate estimation is therefore vital for population management and conservation. We evaluated the effects of age, territoriality and reproduction on survival in Bonelli’s Eagle Aquila fasciata through multistate capture‐mark‐recapture analyses on a long‐term dataset. Monitoring was carried out in southeast France (1990–2008) and involved the surveying of territorial Eagles, the marking of fledged chicks, and the recording of resightings and recoveries of marked non‐territorial and territorial birds. Survival improved with age, but territoriality was not retained in the best model; yearly survival was estimated at 0.479 for fledglings (to 1 year of age), 0.570 for 1‐ and 2‐year‐olds, and 0.870 for 3‐year‐old and older individuals. The second best model supported a further increase in survival from 3‐year‐olds (0.821) to older individuals (0.880). In the third best supported model, territoriality enhanced survival, but only in 2‐year‐olds (0.632 vs. 0.562 for non‐territorial). We found no correlation between the previous breeding stage and future survival, consistent with the long lifespan of the study species. Nevertheless, 4‐year‐old and older successful breeders were more likely to breed the following year than failed adult breeders (0.869 vs. 0.582), suggesting that the cost of reproduction is small in comparison with the variation in quality among individuals or their territories.     

Intercolony variation in reproductive skipping in the African penguin

In long‐lived species, reproductive skipping is a common strategy whereby sexually mature animals skip a breeding season, potentially reducing population growth. This may be an adaptive decision to protect survival, or a non‐adaptive decision driven by individual‐specific constraints. Understanding the presence and drivers of reproductive skipping behavior can be important for effective population management, yet in many species such as the endangered African penguin (Spheniscus demersus), these factors remain unknown. This study uses multistate mark‐recapture methods to estimate African penguin survival and breeding probabilities at two colonies between 2013 and 2020. Overall, survival (mean ± SE) was higher at Stony Point (0.82 ± 0.01) than at Robben Island (0.77 ± 0.02). Inter‐colony differences were linked to food availability; under decreasing sardine (Sardinops sagax) abundance, survival decreased at Robben Island and increased at Stony Point. Additionally, reproductive skipping was evident across both colonies; at Robben Island the probability of a breeder becoming a nonbreeder was ~0.22, versus ~0.1 at Stony Point. Penguins skipping reproduction had a lower probability of future breeding than breeding individuals; this lack of adaptive benefit suggests reproductive skipping is driven by individual‐specific constraints. Lower survival and breeding propensity at Robben Island places this colony in greater need of conservation action. However, further research on the drivers of inter‐colony differences is needed.     

Effects of age,sex, and ENSO phase on foraging and flight performance in Nazca boobies

Age‐related changes in survival and reproduction are common in seabirds; however, the underlying causes remain elusive. A lack of experience for young individuals, and a decline in foraging performance for old birds, could underlie age‐related variation in reproduction because reproductive success is connected closely to provisioning offspring. For seabirds, flapping flight during foraging trips is physiologically costly; inexperience or senescent decline in performance of this demanding activity might cap delivery of food to the nest, providing a proximate explanation for poor breeding success in young and old age, respectively. We evaluated the hypothesis that young and old Nazca boobies (Sula granti), a Galápagos seabird, demonstrate deficits in foraging outcomes and flight performance. We tagged incubating male and female adults across the life span with both accelerometer and GPS loggers during the incubation periods of two breeding seasons (years), during the 2015 El Niño and the following weak La Niña. We tested the ability of age, sex, and environment to explain variation in foraging outcomes (e.g., mass gained) and flight variables (e.g., wingbeat frequency). Consistent with senescence, old birds gained less mass while foraging than middle‐aged individuals, a marginal effect, and achieved a slower airspeed late in a foraging trip. Contrary to expectations, young birds showed no deficit in foraging outcomes or flight performance, except for airspeed (contingent on environment). Young birds flew slower than middle‐aged birds in 2015, but faster than middle‐aged birds in 2016. Wingbeat frequency, flap–glide ratio, and body displacement (approximating wingbeat strength) failed to predict airspeed and were unaffected by age. Sex influenced nearly all aspects of performance. Environment affected flight performance and foraging outcomes. Boobies'' foraging outcomes were better during the extreme 2015 El Niño than during the 2016 weak La Niña, a surprising result given the negative effects tropical seabirds often experience during extreme El Niños.     

Hatching date influences age at first reproduction in the black-headed gull

In long-lived colonial birds, age at recruitment is an important life-history character. Variation in this parameter may reflect differences in several factors, including competitive ability and breeding strategies. Further, these differences may be due to timing of hatching (for instance through differences in competitive ability). We investigated the age of first-time breeders in relation to hatching date in a black-headed gull Larus ridibundus colony situated in central France, from 1979 to 1993. Age at first breeding was estimated for four groups of individuals (total n=550) according to their hatching date, using a recent capture-recapture methodology which allowed us to estimate recruitment rate without the limiting assumptions of methods relying on simple return rates. The age at first breeding was negatively correlated with the hatching date of individuals: individuals hatched earlier in the season started breeding at a younger age than individuals born later. Proportionally more 2-year-old late-hatched individuals were seen breeding on small peripheral colonies than young early-hatched individuals. This difference disappeared after age 3 years. These results strongly suggest that individuals hatched late in the season start to breed on peripheral colonies before recruiting to their natal colony. A difference of few weeks in hatching date has consequences which can last for several years.     

Assessment of costs of reproduction in a pelagic seabird using multistate mark-recapture models

We used a long‐term population band‐resight survey database, a parallel reproduction database, and multistate mark–recapture analysis to assess the costs of reproduction, a keystone concept of life‐history evolution, in Nazca boobies (Sula granti) from Punta Cevallos, Isla Española, Galápagos, Ecuador. We used eight years of resight and breeding data to compare models that included sex‐ and state‐specific survival probabilities and probabilities of transition between reproductive states using multistate mark–recapture models. Models that included state‐specific effects were compared with models lacking such effects to evaluate costs of reproduction. The top model, optimizing the trade‐off of model simplicity and fit to the data using the Akaike Information Criterion (AIC), showed evidence of a temporally varying survival cost of reproduction: nonbreeders showed higher annual survival than breeders did in some years. Because increasing investment among breeders showed no negative association with survival and subsequent breeding success, this evidence indicates a cost to both males and females of initiating, but not of continuing, a reproductive attempt. In some cases, breeders reaching the highest reproductive state (fledging an offspring) showed higher survival or subsequent breeding success than did failed breeders, consistent with differences in overall quality that promote both survival and reproduction. Although a male‐biased adult sex ratio was observed in this population of Nazca boobies, models of state‐ and sex‐specific survival and transition probabilities were not supported, indicating that males and females do not incur different costs of reproduction, and that the observed sex ratio bias is not due to sex‐specific adult mortality.     

Cooperative breeding behaviors in the Hawaiian Stilt (Himantopus mexicanus knudseni)

Cooperative breeding, which is commonly characterized by nonbreeding individuals that assist others with reproduction, is common in avian species. However, few accounts have been reported in Charadriiformes, particularly island‐nesting species. We present incidental observations of cooperative breeding behaviors in the Hawaiian Stilt (Himantopus mexicanus knudseni), an endangered subspecies of the Black‐necked Stilt (Himantopus mexicanus), during the 2012–2020 nesting seasons on the Hawaiian islands of O‘ahu and Moloka‘i. We describe two different behaviors that are indicative of cooperative breeding: (a) egg incubation by multiple adults; (b) helpers‐at‐the‐nest, whereby juveniles delay dispersal and reproduction to assist parents and siblings with reproduction. These observations are the first published accounts of cooperative breeding in this subspecies and merit further investigation, as cooperative breeding may improve population viability of the endangered, endemic Hawaiian Stilt.