Colony size, reproductive success, and colony choice in Cave Swallows Petrochelidon fulva

We investigated how annual reproductive success, as measured by the number of nestlings surviving to day 10 and the percentage of nests that were successful, varied with colony size of the Cave Swallow Petrochelidon fulva in south central Texas. We also studied whether Cave Swallows chose colonies, in part, on the basis of reproductive success at a site the previous year. Neither measure of reproductive success varied significantly with colony size for either first-wave or second-wave nestings. Mean clutch size per colony did not vary significantly with colony size. Mean nestling body mass, an index of parental foraging efficiency, was unrelated to colony size, except for broods of five, in which nestling mass declined significantly with colony size. Colony size was not significantly affected by reproductive success at the site the previous year, although sites with more successful nests during the first wave declined less in size during the second wave within the same season than did sites that had fewer successful first-wave nests. Unlike the closely related Cliff Swallow Petrochelidon pyrrhonota , Cave Swallows did not use breeding performance of conspecifics in choosing nest-sites, because they did not preferentially aggregate at sites that were the most successful the previous year. Coloniality in Cave Swallows did not appear to have a net negative effect on annual fitness, suggesting that colonial nesting was not solely a response to nest-site limitation, but the benefits of breeding colonially (if any) were unclear.     

Prospecting enhances breeding success of first-time breeders in the great cormorant, Phalacrocorax carbo sinensis

In many species of colonial seabirds, young birds visit colonies in the years before they start breeding. This prospecting behaviour may allow them to obtain information that could enhance their future breeding success. We examined the reproductive consequences of prospecting behaviour in the colonial great cormorant, and found support for this idea. New breeders that had been prospecting actively in the previous year obtained breeding sites of higher quality (i.e. closer to sites where conspecifics had fledged young in the previous year) and had higher breeding success than those that had been less active. Prospecting occurred mostly late in the breeding season, and coincided with the time when the majority of the eggs had hatched but before the chicks started fledging, that is, when breeding success in the colony reflected habitat suitability. These results are thus consistent with the use of conspecific reproductive performance as a cue for the quality of a breeding habitat as expected from the 'performance-based conspecific attraction hypothesis'. Copyright 1999 The Association for the Study of Animal Behaviour.     

Heritable choice of colony size in cliff swallows: does experience trump genetics in older birds?

The variation in breeding-colony size seen in populations of most colonial birds may reflect heritable choices made by individuals who are phenotypically specialized for particular social environments. Although a few studies have reported evidence for genetically based choice of group sizes in birds, we know relatively little about the extent to which animals potentially rely on experience versus innate preferences in deciding with how many conspecifics to settle at different times of their lives. We conducted a cross-fostering experiment in 1997-1998 on cliff swallows (Petrochelidon pyrrhonota) in southwestern Nebraska, USA, in which some individuals were reared in colonies different in size from those in which they were born. Breeding-colony sizes chosen by this cohort of birds were monitored by mark-recapture throughout their lives. A multistate mark-recapture analysis revealed that birds in their first breeding year chose colony sizes similar to those of their birth, regardless of their rearing environment, confirming a previous analysis. Beyond the first breeding year, however, cliff swallows' colony choice was less dependent on where they were born. Birds born in small colonies and reared in large colonies showed evidence of a delayed rearing effect, with these birds overwhelmingly choosing large colonies in later years. Heritabilities suggested strong genetic effects on first-year colony choice but not in later years. Cliff swallows' genetically based colony-size preferences their first year could be a way to ensure matching of their phenotype to an appropriate social environment as yearlings. In later years, familiarity with particular colony sites and available information on site quality may override innate group-size preferences when birds choose colonies.     

Costs and benefits of colony size vary during the breeding cycle in Black-headed Gulls Chroicocephalus ridibundus

We studied the effects of colony size on individual reproductive success in a multi-site population of Black-headed Gulls Chroicocephalus ridibundus where colony size ranged from 10 to 5,000 pairs. By focusing on family size, the number of chicks attended by individually marked parents, and accounting for between-individual variation, we detected a negative colony-size effect during the very first days of life of the chicks that was compensated by a subsequent increase in the proportion of surviving chicks with colony size. We suggest that this result originates in the interplay between overcrowding costs acting on hatching success, and benefits of colonial breeding, most probably more efficient food-searching (foraging enhancement), acting on chick survival. However, the frequency of complete colony failure increased with decreasing colony size. Taking this hazard risk into account yielded a corrected estimate of the effect of colony size on breeding success, and indicated that the largest colonies were the most productive. This pattern is congruent with the previous finding that larger colonies are more attractive to dispersing breeders.     

Seasonal shifts in sex ratios of fledgling American kestrels (Falco sparverius paulus): The Early Bird Hypothesis

We document a seasonal shift in the sex ratios of broods produced by resident southeastern American kestrels (Falco sparverius paulus) breeding in nest boxes in Florida. Early in the breeding season, most biased broods were biased towards males, whereas later in the season, most biased broods were biased towards females. Computer-simulated broods subjected to sex-biased egg and/or nestling mortality demonstrate that it is possible that differential mortality produced the pattern of bias that we observed. However, these simulations do not exclude the possibility that female kestrels were manipulating the primary sex ratio of the broods. We present evidence that this sex ratio shift is adaptive: for males we detected breeding as yearlings, all had fledged early the previous season. No such relationship between season and the probability of breeding as a yearling was found for females. We propose the Early Bird Hypothesis as the ecological basis for the advantage of fledg ing early in males. We hypothesize that pre-emptive competition among post-fledging, dispersing males for breeding sites confers an advantage to males fledged early in the season. This hypothesis may explain why a non-migratory population of the Eurasian kestrel (F. tinnunculus) and non-migratory American kestrels breeding in Florida (F. s. paulus) exhibit this seasonal shift in sex ratios, whereas migratory American kestrels (F. s. sparverius) breeding in Saskatchewan, Canada, do not. We discuss the relevance of the Early Bird Hypothesis for other animal species.     

Evidence of prey depletion around lesser kestrel Falco naumanni colonies and its short term negative consequences

Predation may reduce prey numbers in such extent that prey may be depleted, which has negative effects on predator populations. Prey depletion would be more likely when the number of predators increase and/or concentrate their activity in a certain area, as is the case of colonial birds. As a matter of fact, food depletion due to intraspecific competition is considered a major cost of coloniality, and several studies have shown indirect evidence of this. However, no direct measures of food depletion have been provided along with its consequences for the fitness of the colony inhabitants. We carried out a field study with the lesser kestrel Falco naumanni, a raptor that breeds in colonies ranging from two to dozens of pairs. During the nestling period we sampled the main prey of the kestrels around different sized colonies at increasing distances. At the same time, we recorded hunting distances and prey delivery rates to the nest. In addition, we monitored the reproductive success in colonies of different sizes. Lesser kestrels feed their nestlings mainly with grasshoppers and these prey became depleted through the season in the surroundings of the large colonies. Prey depletion made kestrels fly longer distances to forage and prey delivery rates to the nest decreased. Lower feeding rates were not compensated by bringing larger prey, hence, the net amount of energy provided to the chicks decreased with the date in large colonies. By contrast, none of this occurred around small colonies, where both prey abundance and hunting distance remained constant throughout the season. As a consequence, the seasonal decline in the reproductive success (number of fledglings and fledgling body condition) was greater the larger the colony. Thus, these results evidence that food depletion and its fitness costs are related to colony size, as they are suffered by the kestrels breeding in large colonies but not by those settled at small ones. Finally, the consequences of prey depletion on the demographic dynamics and the regulation of colony size are discussed.     

Attendance off breeders and prospectors reflects the quality off colonies in the Kittiwake Rissa tridactyla

Attendance of breeders and prospectors (prebreeders, failed breeders and non-breeding adults) was studied from the time of hatching to the end of the breeding season in parts of two neighbouring Kittiwake Rissa tridactyla colonies in Brittany (France). Many clutches in one subcolony had been predated by a Carrion Crow Corvus corone , leading to a lower breeding success. Attendance of breeders and prospecting activity were greater in the more successful colony, and the number of breeding pairs increased in the following year in this colony only. Local reproductive success and social behaviour contributed to the attraction exerted by a colony. The demographic trend was explained by a performance-based conspecific attraction mechanism, acting through the effect of local reproductive success on recruitment, non-breeding and fidelity. Study of breeding success and prospecting activity allowed both evaluation of the quality of a colony and prediction of local demographic trends for the following year.     

When to use public information for breeding habitat selection? The role of environmental predictability and density dependence

To select their future breeding site, individuals usually have to assess local quality by using environmental cues. One optimal cue may be ‘public information’, the local reproductive success of conspecifics in a breeding patch (patch reproductive success) because it integrates the effect of all environmental factors on breeding success. However, the quality of information conveyed by patch reproductive success is likely to depend on (1) environmental predictability and (2) interactions between individuals. We investigated how these two factors, ignored by previous models, affect the performance of individuals using patch reproductive success for breeding habitat selection compared with other information. We built a two-patch, game-theoretical model to compare the success of a strategy of breeding habitat selection based on patch reproductive success relative to four other strategies: (1) random patch choice; (2) philopatry; (3) choice based on the presence of conspecifics the previous year (conspecific attraction); and (4) choice based on intrinsic patch quality the previous year. The results illustrate how the efficiency of strategies in tracking variations in patch quality depend on environmental predictability and costs linked to density dependence, themselves linked to the dynamics of spatial aggregation of individuals. In particular, strategies based on measures of patch reproductive success perform the best for intermediate and high temporal predictability of patch quality, whereas philopatry and conspecific attraction then perform poorly. The ‘conspecific attraction’ strategy always coexists with other strategies by efficiently parasitizing the information they use. We discuss the implications of a better understanding of breeding habitat selection behaviours for evolutionary and conservation biology. Copyright 2003 The Association for the Study of Animal Behaviour. Published by Elsevier Ltd. All rights reserved.      

Does foraging efficiency vary with colony size in the fairy martin Petrochelidon ariel?

Colonial breeding occurs in a wide range of taxa, however the advantages promoting its evolution and maintenance remain poorly understood. In many avian species, breeding colonies vary by several orders of magnitude and one approach to investigating the evolution of coloniality has been to examine how potential costs and benefits vary with colony size. Several hypotheses predict that foraging efficiency may improve with colony size, through benefits associated with social foraging and information exchange. However, it is argued that competition for limited food resources will also increase with colony size, potentially reducing foraging success. Here we use a number of measures (brood feeding rates, chick condition and survival, and adult condition) to estimate foraging efficiency in the fairy martin Petrochelidon ariel, across a range of colony sizes in a single season (17 colonies, size range 28–139 pairs). Brood provisioning rates were collected from multiple colonies simultaneously using an electronic monitoring system, controlling for temporal variation in environmental conditions. Provisioning rate was correlated with nestling condition, though we found no clear relationship between provisioning rate and colony size for either male or female parents. However, chicks were generally in worse condition and broods more likely to fail or experience partial loss in larger colonies. Moreover, the average condition of adults declined with colony size. Overall, these findings suggest that foraging efficiency declines with colony size in fairy martins, supporting the increased competition hypothesis. However, other factors, such as an increased ectoparasitise load in large colonies or change in the composition of phenotypes with colony size may have also contributed to these patterns.     

Reproductive success and nestling diet in the Lesser Spotted Woodpecker ( Picoides minor ): the early bird gets the caterpillar

Populations of Lesser Spotted Woodpecker (Picoides minor) are decreasing in size, necessitating management strategies. However, data on the reproductive biology of this species are scarce. The aim of this study was to investigate the influence of timing of breeding and nestling diet on the reproductive success of the Lesser Spotted Woodpecker and thereby contribute to an understanding of the determinants influencing its reproductive success. During 6 study years between 1996–2003, we investigated various variables of reproductive success in a Lesser Spotted Woodpecker population in the Taunus low mountain range, Germany. We observed nestling feeding at breeding holes to assess the composition of nestling diet and how it changes during the season. Clutch size, number of fledglings as well as body mass of the nestlings declined with the start of egg laying. Pairs composed of individuals that had breed together in previous years started egg laying earlier than newly established pairs. Nestling diet consisted mainly of aphids, caterpillars, craneflies and wood-living larvae. The composition changed considerably within the breeding season in both early and late broods. However, early broods were provided with caterpillars more frequently, whereas late broods received wood-dwelling larvae more often. Our results suggest that Lesser Spotted Woodpeckers time their breeding so as to coincide with the early nestling stage with the highest availability of caterpillars, since this is the time of highest energy demand. Moreover, food availability declines during the breeding season, and the decline in reproductive performance seems to be an effect of this development.     

THE EFFECT OF AGE ON THE BREEDING BIOLOGY OF THE KITTTWAKE RISSA TRIDACTYLA

1. From 1954 to 1956 inclusive, the biology of individual marked Kittiwakes was studied at North Shields, Northumberland.
2. It was concluded that older Kittiwakes reacted to the breeding stimulus earlier, more intensively and with greater success than younger breeding birds.
3. Birds with previous breeding experience returned to the colony before birds breeding for the first time and these before non-breeders.
4. Before breeding started, birds which had bred previously spent more of their time at the colony than those about to breed for the first time.
5. Birds breeding for at least the second time laid the first egg 7·5 days earlier than those breeding for the first time.
6. Breeding started one day later for every four days the return to the colony was postponed.
7. Older breeding birds showed greater nest-site tenacity, laid larger clutches and had greater breeding success than younger birds.
8. The chicks in broods of two (but not of one) increased in weight more rapidly where the parents had previous experience.
9. Breeding Kittiwakes showed strong colony tenacity, but 24% of the marked non-breeding birds were subsequently seen in other colonies.
10. Over half the birds retained the same mate as in the previous year.     

Nosy neighbours: large broods attract more visitors. A field experiment in the pied flycatcher, <Emphasis Type="Italic">Ficedula hypoleuca</Emphasis>

Life is uncertain. To reduce uncertainty and make adaptive decisions, individuals need to collect information. Individuals often visit the breeding sites of their conspecifics (i.e., “prospect”), likely to assess conspecifics’ reproductive success and to use such information to identify high-quality spots for future breeding. We investigated whether visitation rate by prospectors and success of visited sites are causally linked. We manipulated the reproductive success (enlarged, reduced, and control broods) in a nest-box population of migratory pied flycatchers, Ficedula hypoleuca, in Finland. We measured the visitation rates of prospectors at 87 nest-boxes continuously from manipulation (day 3 after hatching) to fledging. 302 adult pied flycatchers prospected 9194 times on these manipulated nests (at least 78% of detected prospectors were successful breeders). While the number of visitors and visits was not influenced by the relative change in brood size we induced, the resulting absolute brood size predicted the prospecting behaviour: the larger the brood size after manipulation, the more visitors and visits a nest had. The parental provisioning rate at a nest and brood size pre-manipulation did not predict the number of visitors or visits post-manipulation. More visitors, however, inspected early than late nests and broods in good condition. Our study suggests that individuals collect social information when visiting conspecific nests during breeding and provides evidence that large broods attract more visitors than small broods. We discuss the results in light of individual decision-making by animals in their natural environments.     

Expansion of the Mediterranean Gull Larus melanocephalus in Poland

Since 1981, when the first breeding pair of Mediterranean Gull Larus melanocephalus was recorded in Poland, the population of this gull has increased considerably. Its population size was stable until 1997, not exceeding ten pairs annually; thereafter, an increasing number of sightings were made, and during the last 5 years between 26 and 39 breeding pairs have been recorded in Poland. To date, breeding sites have been established in a total of 45 sites (maximum of 19 sites in a given year), with 27% of these (43% of all broods) found on islands located in the middle course of the Vistula River. Breeding sites have also included artificial reservoirs, such as dam reservoirs (20% of sites, 19% of broods), gravel pits (9% of sites, 13% of broods) and fishponds (24% of places, 8% of broods). Mediterranean Gulls were found to nest only within the colonies of other Laridae. Single pairs were recorded at 40% of the breeding sites, whereas a maximum of two to five pairs were recorded at 47% of the other sites. The great majority of breeding attempts were recorded in Black-headed Gull L. ridibundus colonies. In two cases, Mediterranean Gulls bred within mono-specific colonies of Common Gulls L. canus consisting of 60–150 pairs. The biggest concentrations of breeding sites were along the middle course of the Vistula river and in the southern part of Poland.     

Invertebrate prey availability limits reproductive success but not breeding population size in suburban House Sparrows Passer domesticus

Factors affecting avian demography and abundance in urban landscapes are poorly understood and this hinders attempts to manage urban bird communities. Several recent studies indicate that lack of invertebrate prey in urban landscapes may constrain avian productivity and fitness relative to that in other habitats. House Sparrow Passer domesticus populations have undergone large declines in many European urban centres and inadequate reproductive success linked to invertebrate availability has been postulated as a potential cause of these declines. We conducted a replicated supplementary feeding experiment to test whether the availability of invertebrate prey limits the breeding success and adult abundance (colony size) of House Sparrows in suburban London, where House Sparrow populations declined by 60% during the decade preceding our study. Daily mealworm provision over two successive breeding seasons, sufficient to provide 82% of chick energy requirements of House Sparrow pairs nesting within 50 m of feeders, had a large positive impact on the abundance of recently fledged birds (+62%), but only a small positive impact, limited mainly to small colonies, on the overall abundance of territorial males. Colony growth was only weakly related to fledgling abundance in the previous year and did not appear to be constrained by nest‐site availability. Conservation interventions that enhance invertebrate availability for suburban House Sparrows may increase reproductive success but are unlikely, on their own, to lead to population growth or recovery.     

Testing habitat copying in breeding habitat selection in a species adapted to variable environments

The habitat copying hypothesis states that animals use the reproductive performance of conspecifics to evaluate habitat quality and choose their future breeding site. We used data from Audouin's Gull Larus audouinii (1992–2003), a species adapted to unpredictable environments, to analyse subcolony (as patch) choice within a colony (small spatial scale). We also assessed the suitability of alternative hypotheses to the habitat copying hypothesis. The probability of subcolonies being reoccupied annually increased with their size (as number of nests), which suggests the existence of group adherence effects. Subcolony growth rate was related to its average reproductive success (or patch reproductive success) in the previous year: the higher the reproductive success in a colony, the higher the probability of growth the following year. However, this last result was obtained without considering the effect of colony size on the response variable because colony size is related to it. Therefore, results suggest at the population level that in this system habitat copying might either be one of the strategies used by the species in selecting its breeding habitat, or one of the possible strategies operating alone. The other strategies are group adherence mechanisms, and also the effect of conspecific attraction. At the individual level we failed to find evidence of habitat copying and only the previous success of an individual affected its fidelity to a subcolony. The importance of the lack of environmental predictability in the system is discussed, as predictability is a prerequisite of habitat copying.     

The effect of parental age,experience and historical reproductive success on wandering albatross (Diomedea exulans) chick growth and survival

Growth and survival of altricial young are influenced by their parents’ abilities to invest in a breeding attempt. As a result, chick growth and survival in one breeding season may be indicative of their parents’ long-term reproductive potential. To determine whether variation in long-term reproductive success is driven by differential breeding investment, parental care and chick growth in wandering albatrosses (Diomedea exulans) were correlated with parental historical reproductive success. Effects of age and breeding experience (determined from past breeding attempts) and pre-laying body condition (mass–size indices) on chick growth and survival also were tested. Longer brooding of chicks increased their survival, but length of chick brooding did not differ between historically unproductive and successful breeders. Past reproductive success also was not correlated with chick growth rates or fledging mass or size. Chick brooding period, chick growth rates, final mass and size were independent of parental body condition. Older and more experienced parents brooded chicks for longer and their chicks grew faster, supporting previous findings that breeding competence is a learnt skill. Chick care and growth characteristics differed more between than within pairs, suggesting that differences in these characteristics are driven by variation among pairs.     

Spatial differences in breeding success in the pied avocet Recurvirostra avosetta: effects of habitat on hatching success and chick survival

I studied the breeding biology of pied avocets Recurvirostra avosetta in natural habitats (alkaline lakes), and in semi-natural sites (dry fishpond, reconstructed wetlands) in Hungary to relate habitat selection patterns to spatial and temporal variation in breeding success. Colonies were initiated earlier in semi-natural sites than in natural habitats, and earlier on islands than on the mainland. Hatching success was higher on islands than on the mainland, in semi-natural sites than in natural habitats, in colonies of at least 15 pairs than in smaller colonies, and for nests initiated earlier than later within a colony. Fledging success was higher in the wet years (1999–2000) than in the dry year (1998), decreased strongly by season in both habitats and increased with average daily temperature in the first week post-hatch in 1999–2000. Most pairs hatching young in semi-natural sites attempted to lead their chicks to feeding areas in natural habitats, whereas no such movement occurred in the opposite direction. Chick mortality due to predation was high during brood movements and only 23% of the pairs moving their young produced fledglings, compared to 43% for pairs remaining in semi-natural sites and 68% for pairs hatching and rearing young in natural habitats (total n=192 broods). These results suggest that semi-natural sites were more suitable for nesting, whereas natural habitats were more suitable for chick-rearing. The opposing trends in habitat-related breeding success between nesting and chick-rearing suggest sub-optimal habitat selection by Pied Avocets due to an incorrect assessment of the potential for successful reproduction of semi-natural sites, which may thus function as ecological traps.     

The heterospecific habitat copying hypothesis: can competitors indicate habitat quality?

According to the "habitat copying" hypothesis, animals use thereproductive performance of conspecifics to assess habitat suitabilityand choose their future breeding site. This is because conspecificsshare ecological needs and thus indicate habitat suitability.Here, we propose the "heterospecific habitat copying" hypothesis,which states that animals should use public information (i.e.,information derived from the performance of others) from con-and heterospecifics sharing ecological needs. In a correlationalapproach we test some assumptions and predictions of this hypothesiswith a data set from two sympatric bird populations, rollers(Coracias garrulus) and kestrels (Falco tinnunculus), usingthe same nest-boxes and exploiting similar food resources. Sincekestrels are residents and breed earlier, we assumed that theyare dominant over rollers for nest-box acquisition. The environmentappears to be patchy for both species and temporally predictablefor kestrels only. Two results suggest that the use of heterospecificpublic information in breeding habitat selection may be at work:(1) an increase in the reoccupancy probability by kestrels ofprevious roller nests with increasing nest success, and (2)an increase in roller breeding population with increasing localkestrel success. Most of the other observed patterns could beexplained by alternative mechanisms such as natal philopatry,breeding fidelity, conspecific attraction, intraspecific habitatcopying, and the effect of interspecific competition.     

Increasing breeding behaviors in a captive colony of Northern Bald Ibis through conspecific acoustic enrichment

The goal of this study was to increase reproductive behaviors in a captive colony of Northern Bald Ibis (Geronticus eremita) at the Bronx Zoo. The Northern Bald Ibis is classified as critically endangered by the IUCN, with only about 100 known breeding pairs in the wild. Our analysis of zoo breeding and colony size data confirmed earlier observations that small captive colonies of this species do not breed as well as larger colonies, possibly because of insufficient social cues. Using principles from social facilitation theory, we attempted to provide such social cues by presenting acoustic enrichment in the form of conspecific breeding vocalization playbacks. In May 2009, we tracked multiple breeding behaviors during playbacks and control periods. Although breeding behaviors increased in response to playbacks, the colony produced no eggs. We repeated this experiment in April 2010 using playbacks of higher quality recordings acquired from a semi‐wild breeding colony of Northern Bald Ibis in Austria. Breeding behaviors again increased during playbacks. In addition, five pairs in this colony of 15 birds produced 13 eggs, and six chicks successfully fledged—the first reproductive success in this colony since 2007. The acoustic enrichment techniques we employed may encourage increased breeding activity in captive populations of colonial species as well as wild colonies with small populations. Zoo Biol 31:71;–81, 2012. © 2011 Wiley Periodicals, Inc.     

Effects of electromagnetic fields on the reproductive success of American kestrels

Reduced reproductive success of birds nesting near power lines has been documented but never directly attributed to electromagnetic fields (EMFs). Laboratory studies have identified EMF effects on embryonic development, but reproductive success of wild birds is dependent on additional factors, including fertility, egg size, hatching, and fledging success. We tested whether EMFs affect reproductive success of birds. Captive American kestrels (Falco sparverius) were bred for one season per year for 2 yr under either controlled or EMF conditions. EMF exposure was equivalent to that experienced by wild reproducing kestrels and was weakly associated with reduced egg laying in 1 yr only. In both years fertility was higher, but hatching success was lower in EMF pairs than control pairs. Fledging success was higher in EMF pairs than control pairs in 1995 only. Egg composition and embryonic development were examined in 1 yr only, but hatchlings were measured in both years. EMF eggs were larger, with more yolk, albumen, and water, but had thinner egg shells than control eggs. Late-term EMF embryos were larger and longer than control embryos, although hatchlings were similar in body mass and size. EMF exposure affected reproductive success of kestrels, increasing fertility, egg size, embryonic development, and fledging success but reducing hatching success.