Bounded hybrid superiority in an avian hybrid zone: effects of mate, diet, and habitat choice

There has been considerable debate in the study of hybrid zones as to whether hybrids may be superior to parental types within the area of contact (bounded hybrid superiority). In birds, naturally occurring hybridization is relatively common, and hybridization within this group always involves mate choice. If hybrids are superior, females choosing heterospecific mates should be expected to show higher fitness under the conditions prevalent in the hybrid zone. Hybrid superiority under these circumstances would reduce reinforcement and thereby help to maintain the hybrid zone. To examine this issue, we studied reproductive performances of hybrids and parental species of gulls (Larus occidentalis and Larus glaucescens) at two colonies within a linear hybrid zone along the west coast of the United States. This hybrid zone contains predominantly gulls of intermediate phenotype. Previous studies indicated that hybrids were superior to one or both parental types, but provided no data on possible mechanisms that underlie this hybrid superiority. Using a hybrid index designed specifically for these species, we identified to phenotype more than 300 individuals associated with nests, including both individual males and females within 73 pairs in the central portion of the hybrid zone and 74 pairs in the northern portion of the hybrid zone. There was little evidence of assortative mating, and what little there was resulted solely because of pairings within intergrades. In the central hybrid zone, females paired with hybrid males produced larger clutches and hatched and fledged more chicks compared with females paired to western gull males. This was a result of heavy predation on eggs in sand habitat, where male western gulls established territories. In contrast, many hybrid males established territories in vegetated cover that was less vulnerable to predation. In the northern part of the hybrid zone, clutch size did not differ among pair categories, however, there were differences in hatching and fledging success, with females paired to hybrid males showing better success compared to females paired to glaucous-winged gull males. Hybrids showed better hatching and fledging success in the north because hybrids are more like western gulls than glaucous-winged gulls in foraging behavior, taking a higher percentage of fish in their diet, which enhances chick growth and survival. This is believed to be the first documentation of bounded hybrid superiority that delineates the mechanisms that underlie hybrid superiority.     

NARROW HYBRID ZONE BETWEEN TWO SUBSPECIES OF BIG SAGEBRUSH (ARTEMISIA TRIDENTATA: ASTERACEAE). IV. RECIPROCAL TRANSPLANT EXPERIMENTS

Does endogenous or exogenous selection stabilize the big sagebrush (Artemisia tridentata) hybrid zone? After two years of study, our reciprocal transplant experiments showed significant genotype by environment interactions for a number of fitness components, including germination, growth, and reproduction. Hybrids were the most fit within the hybrid garden. In the parental gardens, the native parental taxon was more fit than either the alien parental or hybrids. These results are consistent with the bounded hybrid superiority model, which assumes exogenous selection, but are clearly at odds with the dynamic equilibrium model, which assumes endogenous selection and universal hybrid unfitness.     

Narrow hybrid zone between two subspecies of big sagebrush (ARTEMISIA TRIDENTATA: Asteraceae). IX. Elemental uptake and niche separation

The concentrations of selected elements and their biological absorption coefficients were determined for leaves from plants in native stands and reciprocal transplant gardens to determine whether niche differentiation occurs among the parental taxa and their hybrids in the big sagebrush hybrid zone in Utah. The bounded hybrid superiority model predicts such niche differentiation, while the ecologically neutral dynamic equilibrium model predicts complete niche overlap, at least in the vicinity of the hybrid zone. The concentrations of elements in the leaves of site-indigenous sagebrush and the biological absorption coefficients differed significantly between the subspecies and between either parental taxon and hybrids. Within reciprocal transplant gardens, both the elemental concentrations and the biological absorption coefficients differed among the gardens and taxa. Significant genotype-by-environment interactions were observed for several essential elements. Niche differentiation was evident as correspondence analyses ordinated the parental taxa and hybrids into separate groups even when raised in the same garden. These findings support the ecologically based bounded hybrid superiority model and suggest that the big sagebrush parental taxa and their hybrids have adapted to their respective unique habitats.     

Narrow hybrid zone between two subspecies of big sagebrush,Artemisia tridentata (Asteraceae). III. Developmental instability

Stable hybrid zones are believed to occur because gene dispersal is opposed by selection, but the nature of this selection remains unclear. The dynamic equilibrium model postulates that hybridization disrupts coadapted gene complexes, leading to alterations in development and subsequent hybrid unfitness. Alternatively, the bounded hybrid superiority model assumes that selection is due to exogenous factors. Here, we examine the developmental instability of sagebrush across a narrow hybrid zone. The results show no difference among the parental and hybrid taxa for the majority of comparisons examined. When significant differences in developmental instability were observed, one of the parental taxa (mountain big sagebrush) was typically the least developmentally stable. Hybrids were found to be the least developmentally stable for only two of the 28 comparisons studied. Interestingly, for some comparisons, hybrids were the most developmentally stable. These results contradict predictions of the dynamic equilibrium model. The sagebrush hybrid zone does not appear to be maintained by endogenous selection, as there is little indication that development is disrupted by hybridization.     

Predator–prey dynamics of bald eagles and glaucous‐winged gulls at Protection Island,Washington, USA

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Extrinsic and intrinsic factors influence fitness in an avian hybrid zone

The effects of hybridization on evolutionary processes are primarily determined by the differential between hybrid and parental species fitness. Assessing the impacts of hybridization can be challenging, however, as determining the relationship between individual fitness and the extent of introgression in wild populations is difficult. We evaluated the fitness consequences of hybridization for pure and hybrid females in a hybrid zone between two tidal marsh birds, the saltmarsh sparrow (Ammodramus caudacutus), a salt marsh obligate, and Nelson's sparrow (A. nelsoni), which has a broader ecological niche and a much younger evolutionary association with salt marshes. Biotic stressors associated with nesting in tidal environments suggest an important role for differential adaptation in shaping hybrid zone dynamics, with saltmarsh sparrows predicted to be better adapted to nesting in salt marshes. We collected DNA samples from adults (n = 394) and nestlings (n = 431) to determine the extent of introgression using 12 microsatellite loci and tested for the influence of extrinsic (nest placement) and intrinsic (genotype) factors on female reproductive success. We monitored nests (n = 228), collected data on reproductive output, and estimated daily nest survival rates using female genotype and nest characteristics as covariates. To test for reduced survival of hybrid females, we also used capture data to assess the distribution of admixed male and female individuals across age classes. Reproductive success of females varied by genotypic class, but hybrids did not have intermediate success as predicted. Instead, we found that pure Nelson's sparrows had, on average, 33% lower hatching success than any other genotype, whereas F1/F2 hybrids, backcrossed Nelson's sparrows, and backcrossed and pure saltmarsh sparrows all had similar hatching success. We found no effect of genotype or nest placement on daily nest survival probabilities. However, hybrid individuals with a higher proportion of saltmarsh sparrow alleles exhibit nesting behaviours better suited to nesting successfully in tidal marshes. Further, while the proportion of F1/F2 individuals was similar between nestling and adult males, we found that the proportion of F1/F2 individuals was 2.3 times greater in nestling females compared with adult females, indicating reduced survival of F1 females. We conclude that differences in reproductive success among pure and admixed individuals coupled with intrinsic mechanisms (reduced survival in F1 females) shape hybrid zone dynamics in this system.     

Lower reproductive success in hybrid fur seal males indicates fitness costs to hybridization

Hybridization among organisms can potentially contribute to the processes of evolution, but this depends on the fitness of hybrids relative to parental species. A small, recently formed population of fur seals on subantarctic Macquarie Island contains a high proportion of hybrids (17-30%) derived from combinations of three parental species: Antarctic, subantarctic and New Zealand fur seals. Mitochondrial control-region data (restriction fragment length polymorphisms) and nine microsatellites were used to determine the species composition of breeding adults, and hybrid male fitness was measured by comparing reproductive success (number of genetically inferred paternities) of hybrid and pure-species territory males over 6 years. No correlations were found between male reproductive success and three genetic measures of outbreeding, but this may be due to a relatively small number of dominant males analysed. Territory males fathered 63% of pups, but hybrid males had lower reproductive success than pure-species males despite having the same ability to hold territories. A greater proportion of females in hybrid male territories conceived extra-territorially than those in territories of pure-species males, and most (70 of 82) mated with conspecifics. This suggests the presence of reproductive isolating mechanisms that promote positive assortative mating and reduce the production of hybrid offspring. Although we found no evidence for male sterility in the population, mechanisms that reduce lifetime reproductive success may act to decrease the frequency of hybrids. Our study has identified a disadvantage of hybridization - reduced reproductive success of hybrid sons - that may be contributing to the persistence of pure lineages at Macquarie Island and the temporal decline in hybridization observed there.     

Fertility in first-generation hybrids of roach, Rutilus rutilus (L.), and silver bream, Blicca bjoerkna (L.)

Fertility in first‐generation hybrids of roach, Rutilus rutilus, and silver bream, Blicca bjoerkna, was investigated. Sperm and egg production of hybrids at first sexual maturity were examined. Eggs from female hybrids were artificially fertilized with the sperm of a corresponding hybrid male; a hybrid male from the reciprocal crossbreeding; a parental species male R. rutilus; and a parental species male B. bjoerkna. The results revealed that gametogenesis was normal in female hybrids. However, in male hybrids, a low efficiency of gametogenesis was observed. The semen of male hybrids was extremely dilute, with spermatozoa concentration lower than that in parental species. Nevertheless, these F1 hybrids (males and females) from reciprocal crossbreeding were fertile. F2 and backcross generations were produced, but F2 crosses from the female hybrid and corresponding hybrid male displayed a drastically slower hatching rate. Also higher proportions of deformed embryos were hatched than in other post‐F1‐generation crosses.     

Tests of reproductive isolation among species in the <Emphasis Type="Italic">Fundulus notatus</Emphasis> (Cyprinodontiformes: Fundulidae) species complex

We assessed prezygotic (probability of spawning) and postzygotic (hatching success) reproductive isolation among the three ecologically and morphologically similar species in the Fundulus notatus species complex. We employed a multi-generation breeding experiment to test the hypotheses that karyotypic differences, body size differences, or geographic isolation among populations will increase pre or postzygotic reproductive barriers. Overall, prezygotic barriers were strong and postzygotic barriers weak in crosses of non-hybrid heterospecifics (F1 hybrid crosses) while prezygotic barriers were weaker and postzygotic barriers stronger in crosses involving hybrid individuals (F2 hybrid crosses and backcrosses). Prezygotic barriers among the two smaller species (Fundulus notatus and F. euryzonus) broke down rapidly; first generation hybrids spawned (F2 hybrid crosses and backcrosses) as frequently as parental forms in intraspecific crosses. There was no increase in postzygotic barriers among species with cytogenetic differences. There were increased prezygotic, but not postzygotic, barriers among geographically isolated populations of one species. While pure males and females were just as likely to spawn with hybrids, some types of hybrid females suffered from increased sterility, but not inviability, over hybrid males. Female sterility was only seen in hybrids with a Fundulus euryzonus parent, while other female hybrids produced viable eggs.     

Airborne‐pollen pool and mating pattern in a hybrid zone between Pinus pumila and P. parviflora var. pentaphylla

The reproductive isolation barriers and the mating patterns among Pinus pumila, P. parviflora var. pentaphylla and their hybrids were examined by flowering phenology and genetic assays of three life stages: airborne‐pollen grains, adults and seeds, in a hybrid zone on Mount Apoi, Hokkaido, Japan. Chloroplast DNA composition of the airborne‐pollen was determined by single‐pollen polymerase chain reaction. Mating patterns were analysed by estimating the molecular hybrid index of the seed parent, their seed embryos and pollen parents. The observation of flowering phenology showed that the flowering of P. pumila precedes that of P. parviflora var. pentaphylla by about 6 to 10 days within the same altitudinal ranges. Although this prezygotic isolation barrier is effective, the genetic assay of airborne‐pollen showed that the two pine species, particularly P. pumila, still have chances to form F₁ hybrid seeds. Both parental species showed a strong assortative mating pattern; F₁ seeds were found in only 1.4% of seeds from P. pumila mother trees and not at all in P. parviflora var. pentaphylla. The assortative mating was concluded as the combined result of flowering time differentiation and cross‐incompatibility. In contrast to the parental species, hybrids were fertilized evenly by the two parental species and themselves. The breakdown of prezygotic barriers (intermediate flowering phenology) and cross‐incompatibility may account for the unselective mating. It is suggested that introgression is ongoing on Mount Apoi through backcrossing between hybrids and parental species, despite strong isolation barriers between the parental species.     

Azure‐winged Magpies Cyanopica cyanus trade off reproductive success and parental care by establishing a size hierarchy among nestlings

It is common in birds that the sizes of nestlings vary greatly when multiple young are produced in one nest. However, the methods used by parents to establish size hierarchy among nestlings and their effect on parental provisioning pattern may differ between species. In the Azure‐winged Magpie Cyanopica cyanus, we explored how and why parents controlled the sizes of nestlings. Asynchronous hatching was the main cause of size hierarchy within the brood, although the laying of larger eggs later in the laying sequence reduced this effect. Parents with asynchronous broods produced more eggs and fledged more nestlings than those with synchronous broods but their brood provisioning rates, food delivery per feeding bout and feeding efficiency did not differ. We performed a cross‐fostering experiment to synchronize some asynchronous broods. Provisioning rates of asynchronous broods were lower than those of synchronized broods, but the daily growth rates and fledging body mass of their nestlings were not different. Our findings indicate that parents of asynchronous broods can achieve higher reproductive success than those of synchronous broods based on the same parental care, and the same reproductive success as those of synchronized broods based on less parental care. It appears that parent birds can better trade off reproductive success and parental care by establishing a size hierarchy among nestlings.     

Nestling sex ratio of golden‐winged warblers Vermivora chrysoptera in an introgressed population

Sex ratio biases in avian species remain controversial, although several studies have documented apparent facultative adjustment of offspring sex ratios. While hybridizing pied and collared flycatchers have exhibited sex ratio skews that may be a response to sex‐based costs associated with hybridization, this appears not to be true of a hybridized population of blue‐winged Vermivora pinus and golden‐winged V. chrysoptera warblers. We examined the primary sex ratio of nestlings in a population of hybrid and introgressed golden‐winged warblers. The sex ratio of 298 nestlings from 81 nests in the population was approximately 50:50. We conducted paternity assignments and analyzed groups of nestlings with shared genetic parents (“genetic broods”) and found no difference from the expected binomial distribution, and no statistically significant relationship between parental species phenotype and nestling sex ratio. We saw no evidence of preferential production of male or female nestlings, and female hybrids were found to mate and breed in the population. This suggests that heterogametic (female) hybrids are both viable and fertile, and thus that Haldane's Rule does not apply to this system. While populations of hybridizing golden‐winged warblers should be monitored for evidence of costs of heterospecific pairings, it is unlikely that adjustment of sex ratios would be the form of compensation for sub‐optimal mating conditions. Our results provide support for the emerging hypothesis that hybrids suffer no disadvantage relative to golden‐winged and blue‐winged warblers.     

Behavioural isolation may facilitate homoploid hybrid speciation in cichlid fish

Hybrid speciation is constrained by the homogenizing effects of gene flow from the parental species. In the absence of post‐mating isolation due to structural changes in the genome, or temporal or spatial premating isolation, another form of reproductive isolation would be needed for homoploid hybrid speciation to occur. Here, we investigate the potential of behavioural mate choice to generate assortative mating among hybrids and parental species. We made three‐first‐generation hybrid crosses between different species of African cichlid fish. In three‐way mate‐choice experiments, we allowed hybrid and nonhybrid females to mate with either hybrid or nonhybrid males. We found that hybrids generally mated nonrandomly and that hybridization can lead to the expression of new combinations of traits and preferences that behaviourally isolate hybrids from both parental species. Specifically, we find that the phenotypic distinctiveness of hybrids predicts the symmetry and extent of their reproductive isolation. Our data suggest that behavioural mate choice among hybrids may facilitate the establishment of isolated hybrid populations, even in proximity to one or both parental species.     

The effects of heterospecifics and climatic conditions on incubation behavior within a mixed‐species colony

Parental incubation behavior largely influences nest survival, a critical demographic process in avian population dynamics, and behaviors vary across species with different life history breeding strategies. Although research has identified nest survival advantages of mixing colonies, behavioral mechanisms that might explain these effects is largely lacking. We examined parental incubation behavior using video‐monitoring techniques on Alcatraz Island, California, of black‐crowned night‐heron Nycticorax nycticorax (hereinafter, night‐heron) in a mixed‐species colony with California gulls Larus californicus and western gulls L. occidentalis. We first quantified general nesting behaviors (i.e. incubation constancy, and nest attendance), and a suite of specific nesting behaviors (i.e. inactivity, vigilance, preening, and nest maintenance) with respect to six different daily time periods. We employed linear mixed effects models to investigate environmental and temporal factors as sources of variation in incubation constancy and nest attendance using 211 nest days across three nesting seasons (2010–2012). We found incubation constancy (percent of time on the eggs) and nest attendance (percent of time at the nest) were lower for nests that were located < 3 m from one or more gull nest, which indirectly supports the predator protection hypothesis, whereby heterospecifics provide protection allowing more time for foraging and other self‐maintenance activities. To our knowledge, this is the first empirical evidence of the influence of one nesting species on the incubation behavior of another. We also identified distinct differences between incubation constancy and nest attentiveness, indicating that these biparental incubating species do not share similar energetic constraints as those that are observed for uniparental species. Additionally, we found that variation in incubation behavior was a function of temperature and precipitation, where the strength of these effects was dependent on the time of day. Overall, these findings strengthen our understanding of incubation behavior and nest ecology of a colonial‐nesting species.     

Status,trends, and patterns of covariation of breeding seabirds at St Lazaria Island,Southeast Alaska, 1994–2006

Aim We examined patterns of covariation among piscivorous and planktivorous seabirds breeding at St Lazaria Island in order to evaluate their responses to interannual changes in sea surface temperature, a variable that affects marine food webs. In addition, we evaluated seabird population trends for responses to decadal‐scale changes in the marine ecosystem. Location St Lazaria Island, Sitka Sound, Alaska. Methods Established seabird monitoring protocols for the Alaska Maritime National Wildlife Refuge were followed in estimating population trends, the timing of nesting events and the reproductive success of eight species of seabirds between 1994 and 2006. Results  Population increases were noted for storm‐petrels (Oceanodroma furcata and O. leucorhoa), rhinoceros auklets (Cerorhinca monocerata) and glaucous‐winged gulls (Larus glaucescens). We found no population trend for pelagic cormorants (Phalacrocorax pelagicus), but it appeared that populations of common (Uria aalge) and thick‐billed (U. lomvia) murres and of tufted puffins (Fratercula cirrhata) declined. We detected no linear trends in either breeding chronology or reproductive success over the study period for any seabird. All species of piscivorous seabirds apparently responded similarly to environmental cues as there was a positive covariation among species in the timing of nesting. Piscivores tended to nest earlier, and most species had higher rates of reproductive success in years with relatively warm spring sea temperatures. In contrast, planktivorous Leach’s storm‐petrels (O. leucorhoa) tended to nest earlier when spring and summer sea temperatures were relatively cool. Clearly, seabirds at St Lazaria were responding to interannual changes in sea temperatures near the breeding colony, probably as a result of effects on the food webs. Main conclusions Every seabird species we monitored at St Lazaria exhibited significant population trends between 1994 and 2006. For most species there appeared to be a relationship between both the timing of nesting and reproductive rates and spring or summer sea surface temperatures. Responses at both decadal (populations) and interannual (timing and reproductive success) scales make seabirds useful candidates for helping to monitor change in the marine environment.     

An ecomorphological framework for the coexistence of two cyprinid fish and their hybrids in a novel environment

Niche variation between hybrid taxa and their parental species has been deemed imperative to the persistence of hybrid populations in nature. However, the ecological factors promoting hybrid establishment remain poorly understood. Through the application of a multidisciplinary approach integrating genetics, morphometry, life‐history, and trophic ecology, we studied the hybrids of roach (Rutilus rutilus L.) and bream (Abramis brama L.), and their parental species inhabiting an Irish lake. The roach × bream hybrid exhibited a body shape intermediate of that of the parental species. Diet analyses depicted the hybrid as a generalist, feeding on all prey items consumed by either parental species. Stable isotope data confirm the trophic niche breadth of hybrids. A significant correlation between body shape and diet was detected, suggesting that the intermediate phenotype of hybrids might play a role in their feeding abilities, resulting in the utilization of a broader trophic spectrum than the parental species. Growth and age class structure analyses also yielded a scenario that is consistent with the ecological success of hybrids. Genetic analyses suggest that the majority of hybrids result from first‐generation crosses between the parental species; however, a potentially significant proportion of back‐crosses with bream were also detected. The recent introduction of roach and bream into Irish waters, as well as the climatic and ecological features of the colonized habitats, can explain the remarkable success of the roach × bream hybrid in Ireland. The adaptive significance of hybridization and its demographic consequences for the parental species are discussed. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 99 , 768–783.     

Narrow hybrid zone between two subspecies of big sagebrush (Artemisia tridentata: Asteraceae). II. Selection gradients and hybrid fitness

The dynamic equilibrium hypothesis proposes that hybrid zones are stabilized by a balance between dispersal and selection against hybrids. A key prediction of this hypothesis is that hybrids should have lower fitness than either parental taxon, regardless of habitat. Hybrid big sagebrush (Artemisia tridentata ssp. tridentata × ssp. vaseyana) in two narrow hybrid zones do show greatly decreased recruitment. Hybrids in one zone also show increased browsing by grasshoppers, while those in the other zone have increased numbers of aphids. Overall herbivore loads, however, are not greater on the hybrids than on the parental subspecies. Browsing by mule deer is greatest on ssp. vaseyana in both hybrid zones. Incidence of galls is also greatest on ssp. vaseyana in one hybrid zone. Moreover, browsing by Artemisia weevils is greatest on ssp. tridentata in one hybrid zone. Hybrids produce more flowers than either ssp. tridentata or ssp. vaseyana, while seed production rates of hybrids do not differ from those of the parental taxa. Finally, hybrid seeds germinate as well as those of ssp. tridentata and better than those of ssp. vaseyana. Thus, our data do not support the dynamic equilibrium hypothesis.     

Hybrids between chum Oncorhynchus keta and pink Oncorhynchus gorbuscha salmon: age,growth and morphology and effects on salmon production

Mature hybrids between chum salmon Oncorhynchus keta and pink salmon Oncorhynchus gorbuscha, which were identified by an intermediate colour pattern, were caught at the Kurilsky Hatchery, Iturup Island, Russia. Most of them were female and 3 years old (a partial freshwater year and 2 marine years), which is intermediate between the ages of maturity of the parental species. The hybrids exceed both parental species in the rate of growth, are large in size and robust and might successfully compete for mating in the wild or be chosen for artificial reproduction. The ratio of the scale length over width, R, is oblate (R < 1), whereas scales of the parental species are prolate (R > 1). From scale analyses, the c.v. in body size of hybrid females at the second marine year is twice that of O. keta, which suggests developmental instability in the hybrid. A dynamic model predicted that continuing hybridization at a low rate does not produce a substantial hybrid load due to selection against advanced‐generation hybrids and backcrosses. A high hybridization rate, however, may be an additional risk for genetic management and should be taken into account in programmes of artificial reproduction of Pacific salmon Oncorhynchus spp., although such hybrids might have commercial use in confined production systems.     

Testing common assumptions in studies of songbird nest success

We studied Ovenbird Seiurus aurocapilla and Golden‐winged Warbler Vermivora chrysoptera populations in northern Minnesota, USA, to test two common assumptions in studies of songbird nest success: (1) that the condition of an empty nest on or near its expected fledge date is an indicator of nest fate; and (2) that the presence of a fledgling or family group within a territory confirms a successful nest in that territory. We monitored the condition of nests and used radiotelemetry to monitor juveniles through the expected fledging date and early post‐fledging period. Of nests that contained nestlings 1–2 days before the expected fledge date, fates were misidentified using nest condition alone for 9.5% of Ovenbird nests, but those misidentifications were made in both directions (succeeded or failed), yielding only a small bias in estimated nest success. However, 20% of Golden‐winged Warbler nests were misidentified as successful using nest condition during the final visit interval, biasing the nest success estimate upward by 21–28% depending on the treatment of uncertain nest fates. Fledgling Ovenbirds from 58% of nests travelled beyond their natal territory within 24 h, rising to 98% after 5 days, and those fledglings travelled up to 390 m from nests within 10 days of fledging. Fledgling Golden‐winged Warblers from 13% of nests travelled beyond their natal territory within 24 h, rising to 85% after 5 days, and those fledglings travelled up to 510 m from nests within 10 days of fledging. We conclude that nest condition and fledgling presence can be misleading indicators of nest fate, probably commonly biasing nest success estimates upward, and we recommend that these assumptions should be tested in additional species.     

Comparative reproductive biology of sympatric species: nest and chick survival of American avocets and black‐necked stilts

Identifying differences in reproductive success rates of closely related and sympatrically breeding species can be useful for understanding limitations to population growth. We simultaneously examined the reproductive ecology of American avocets Recurvirostra americana and black‐necked stilts Himantopus mexicanus using 1274 monitored nests and 240 radio‐marked chicks in San Francisco Bay, California. Although there were 1.8 times more avocet nests than stilt nests, stilts nonetheless fledged 3.3 times more chicks. Greater production by stilts than avocets was the result of greater chick survival from hatching to fledging (avocet: 6%; stilt: 40%), and not because of differences in clutch size (avocet: 3.84; stilt: 3.77), nest survival (avocet: 44%; stilt: 35%), or egg hatching success (avocet: 90%; stilt: 92%). We reviewed the literature and confirmed that nest survival and hatching success are generally similar when avocets and stilts breed sympatrically. In addition to species, chick survival was strongly influenced by age, site, and year. In particular, daily survival rates increased rapidly with chick age, with 70% of mortalities occurring ≤ 1 week after hatch. California gulls Larus californicus caused 55% of avocet, but only 15% of stilt, chick deaths. Differential use of micro‐habitats likely reduced stilt chick's vulnerability to gull predation, particularly during the first week after hatch, because stilts nested in vegetation 2.7 times more often than avocets and vegetation height was 65% taller at stilt nests compared with avocet nests. Our results demonstrate that two co‐occurring and closely related species with similar life history strategies can differ markedly in reproductive success, and simultaneous studies of such species can identify differences that limit productivity.