Consequences of the size structure of fish populations for their effects on a generalist avian predator

Size-structured interspecific interactions can shift between predation and competition, depending on ontogenetic changes in size relationships. I examined the effects of common carp (Cyprinus carpio), an omnivorous fish, on the reproductive success of the red-necked grebe (Podiceps grisegena), an avian gape-limited predator, along a fish size gradient created by stocking distinct age-cohorts in seminatural ponds. Young-of-the-year (0+) carp were an essential food source for young grebes. Only adult birds were able to consume 1-year-old (1+) fish, while 2-year-old (2+) fish attained a size refuge from grebes. Amphibian larvae were the principal alternative prey to fish, followed by macroinvertebrates, but the abundance of both dramatically decreased along the carp size gradient. Fledging success was 2.8 times greater in ponds with 0+ versus 1+ carp; in ponds with 1+ carp, chicks received on average 2.6–3 times less prey biomass from their parents, and over 1/3 of broods suffered total failure. Breeding birds avoided settling on 2+ ponds. These results show that changes in prey fish size structure can account for shifts from positive trophic effects on the avian predator to a negative impact on the predator’s alternative resources. However, competition did not fully explain the decrease in grebe food resources in the presence of large fish, as carp and grebes overlapped little in diet. In experimental cages, 1+ carp totally eliminated young larvae of amphibians palatable to fish. In field conditions, breeding adults of palatable taxa avoided ponds with 1+ and older carp. Non-trophic interactions such as habitat selection by amphibians or macroinvertebrates to avoid large fish may provide an indirect mechanism strengthening the adverse bottom-up effects of fish on birds.     

Fish reduce habitat coupling by a waterbird: evidence from combined stable isotope and conventional dietary approaches

Aquatic consumers can function as habitat couplers by using allochthonous subsidies of prey that migrate across ecosystem boundaries. We examined the relative use of allochthonous (invertebrates—terrestrial or living on littoral vegetation; immigrating amphibians) versus autochthonous (aquatic invertebrates, fish) resources by the red-necked grebe Podiceps grisegena, a generalist predator, on fishless ponds versus ponds stocked with common carp Cyprinus carpio. We combined conventional methods of diet estimation with stable carbon and nitrogen analyses of egg yolks and putative prey of grebes. Prelaying grebes were observed to take mainly adult amphibians on fishless ponds and fish on stocked ponds. Alimentary tract analyses gave more weight to invertebrate prey, especially leaf beetles Donaciinae, apparently picked off water or emergent plants. Bayesian isotopic mixing models did not reveal predominance of a single food source but indicated that in the presence of fish grebes received relatively less energy for egg formation from amphibians and leaf beetles. Overall, our results show that grebes relied more on allochthonous resources (range of means 50–97 % of the biomass contribution estimated by different assessment methods) in the absence than in the presence of fish (8–23 %). We suggest that habitat coupling by waterbirds may be controlled by fish, which can suppress external prey subsidies, apart from being an attractive food for piscivorous birds.     

Effects of small-bodied fish on invertebrate prey and foraging patterns of waterbirds in Aspen Parkland wetlands

Competition between large-bodied fish and waterbirds for aquatic invertebrates is well documented in oligotrophic lakes. Recent evidence suggests that small-bodied fish that colonize eutrophic, hypoxia-prone wetlands such as prairie potholes can also reduce aquatic invertebrates, but the effects of these reductions on breeding waterbirds have so far not been directly documented. We added brook stickleback (Culaea inconstans) and fathead minnow (Pimephales promelas) to a fishless wetland in Aspen Parkland potholes in central Alberta, Canada. We monitored invertebrate biomasses and the foraging effort of blue-winged teal (Anas discors) and red-necked grebe (Podiceps grisegena) before and after the addition, relative to reference wetlands with and without fish. Fish reduced the biomass of gastropod prey of blue-winged teal, and teals increased foraging effort when fish were added. When the fish failed to overwinter due to hypoxic conditions, gastropod biomass increased, but teal foraging effort did not return to pre-treatment levels. Amphipods and chironomids increased following fish addition, possibly due to indirect positive effects of fish. Red-necked grebes did not exhibit any changes in foraging effort as a result of the fish addition or the subsequent fish extirpation. Grebes in Aspen Parkland appear to treat fish and invertebrates as equivalent prey. This study suggests that small-bodied fish in eutrophic systems can reduce some important invertebrate prey and change foraging behaviour of blue-winged teal and other waterbirds that rely on those invertebrates. Land-use practices that encourage survival of colonizing fish through drought years in Aspen Parkland wetlands, such as wetland consolidation, should not be encouraged.     

Between-lake differences in the diet and provisioning behaviour of Black-throated Divers Gavia arctica breeding in Scotland

Surveys at 23 Black-throated Diver breeding lakes in Scotland showed large between-lake differences in the species and size range of potential prey. The study lakes were classified into four types according to the main size-taxa prey classes present. Type 1 lakes lacked small fish, Type 4 lacked salmonids < 300 mm and Types 2 and 3 both had all four of the main prey classes. Diver diet for 30 families at the study lakes was quantified from 7943 prey items seen fed to chicks, and 153 items seen eaten by adults, during 662 h of observations. At all lake types adults mostly ate salmonids of 120–240 mm in length, especially Brown Trout Salmo trutta , and European Eels Anguilla anguilla up to c.  350 mm. The prey eaten by chicks ranged widely, from c.  0.03 g to 300 g, with important implications for feeding behaviour. Young chicks (days 1–8) rejected items greater than 70 mm long. The diet of young chicks consisted mostly of small fish, either Three-spined Sticklebacks Gasterosteus aculeatus (Type 2 lakes) or Minnow Phoxinus phoxinus (Type 3 and Type 4 lakes). At other lakes (Type 1) it consisted mostly of mayfly larvae (Ephemeroptera); invertebrates were the only abundant potential small prey in these lakes. At all lakes the chick diet became more similar to that of adults as chicks grew. Adults provisioning chicks used two foraging strategies. In 'excursion foraging', adults hunted away from the chicks and carried prey back for them; in 'attendance foraging', the chicks accompanied the foraging adult(s). Excursion foraging was used mostly before day 4 and was strongly and positively correlated with the abundance of fish < 80 mm in length. This behaviour is probably adaptive, but is probably tenable only where small fish are relatively abundant. Young chicks fed mainly on small fish had higher survival rates than those fed on invertebrates.     

Fish stocking creates an ecological trap for an avian predator via effects on prey availability

In anthropogenic landscapes animals may be lured into low‐quality habitats where they survive or reproduce poorly (‘ecological traps’). I investigated breeding habitat selection in relation to intra‐seasonal changes in food availability and reproductive output in red‐necked grebes Podiceps grisegena, a size‐limited predator, of common carp Cyprinus carpio ponds. Carp farms constitute highly heterogeneous habitat mosaics due to separate stocking of different age/size fish. Pond features significant for grebe settling decisions, i.e. hydroperiod and emergent vegetation cover, had no obvious effects on prey abundance for chicks and on fledging success. Breeding grebes avoided ponds containing fish too large for them to ingest but exhibited little preference between ponds with medium‐sized one‐year‐old carp that could be exploited by pre‐laying birds, and ponds designated for young‐of‐the‐year carp, where only invertebrates and amphibians were available as prey in early spring. Red‐necked grebes settling on ponds with medium‐sized fish failed to predict future shifts in interactions with carp stocks; carp exceeded the prey‐size threshold of chicks and adversely affected their non‐fish prey levels. The resulting food shortage led to severe egg‐to‐fledging mortality rates compared to fishless ponds or those containing young‐of‐the‐year fish. This study shows that waterbirds vulnerable to competition from fish can risk maladaptive habitat selection due to unrecognised spatial and temporal variation in food resources caused by fish stocking practices. Ecological traps created by perturbations to trophic interactions may be common but difficult to detect because altered dynamics of trophic resources can affect wildlife indirectly. As with other types of ecological traps, manipulation of habitat features identified as attractive cues for settling animals, but not related to critical food resources, may help to reduce perceptual pitfalls. For example, wetland management to mitigate trap effects driven by commercially stocked fish should preserve abundant emergent vegetation in habitats with weak fish impact and extend their hydroperiod.     

Variance in prey abundance influences time budgets of breeding seabirds: evidence from pigeon guillemots Cepphus columba

We use data on pigeon guillemots Cepphus columba to test the hypothesis that discretionary time in breeding seabirds is correlated with variance in prey abundance. We measured the amount of time that guillemots spent at the colony before delivering fish to chicks ("resting time") in relation to fish abundance as measured by beach seines and bottom trawls. Radio telemetry showed that resting time was inversely correlated with time spent diving for fish during foraging trips (r=−0.95). Pigeon guillemots fed their chicks either Pacific sand lance Ammodytes hexapterus , a schooling midwater fish, which exhibited high interannual variance in abundance (CV=181%), or a variety of non-schooling demersal fishes, which were less variable in abundance (average CV=111%). Average resting times were 46% higher at colonies where schooling prey dominated the diet. Individuals at these colonies reduced resting times 32% during years of low food abundance, but did not reduce meal delivery rates. In contrast, individuals feeding on non-schooling fishes did not reduce resting times during low food years, but did reduce meal delivery rates by 27%. Interannual variance in resting times was greater for the schooling group than for the non-schooling group. We conclude from these differences that time allocation in pigeon guillemots is more flexible when variable schooling prey dominate diets. Resting times were also 27% lower for individuals feeding two-chick rather than one-chick broods. The combined effects of diet and brood size on adult time budgets may help to explain higher rates of brood reduction for pigeon guillemot chicks fed non-schooling fishes.     

Prey profitability for adult Grey Herons Ardea cinerea and the constraints on prey size when feeding young nestlings

Patterns of handling time and profitability are examined for adult Grey Herons feeding on carp, eels and catfish. Handling times generally increased with prey size but were influenced markedly by the morphology and behaviour of the prey. Profitability was highest for carp (max. 0.9 g/s for 15–20 cm fish), lowest for catfish (max. 0.05 g/s) and intermediate for eels (max. 0.1 g/s). Nestlings were unable to ingest the sizes of fish most profitable for the adults to consume until aged 20 days; by the age of 30 days, they could consume the full size-range of prey taken by the adults. In order to feed their young chicks, adults must therefore either select smaller prey, or break their large prey into smaller pieces. The diet of nestling Grey Herons in the Camargue is examined for evidence to support or refute the former hypothesis.
Young nestlings (≤20 days) regurgitated smaller carp than old nestlings (> 20 days). Comparison of prey types in the diet of the two groups showed that small prey species occurred significantly more often in the diet of young chicks, while the converse was true for larger prey species. The occurrence of particular prey types in the diet only of young chicks suggests that adults may forage in more marginal, shallower water (where small prey are probably more abundant) to meet the requirements of their brood during the early part of the nestling phase. The second hypothesis, that the adults break down large prey into smaller pieces, was not examined, although evidence from other studies suggests that this does occur; both mechanisms may therefore be important.     

Influence of fish on the breeding of the red-necked grebe Podiceps grisegena (Boddaert, 1783)

I determined whether selection of breeding sites by thered-neckedgrebe Podiceps grisegena (Boddaert, 1783) was affectedbycompetition for food with fish. During 1987–89, ten ponds inScania, southern Sweden were investigated, five with(breedingsites) and five without breeding red-necked grebe(non-breedingsites). I quantified the biomass of fish and macrozoans(here:larger invertebrates and small vertebrates) and measured thephysical and chemical conditions of the ponds.Breeding sites had significantly lower relative fish abundancethannon-breeding sites. They were either devoid of fish or hadonly nine-spined sticklebackPungitius pungitius. The biomass (dry weight) ofmacrozoans,many of which represent a food resource for grebes, was 16timesgreater at breeding than at non-breeding sites. The physicalandchemical conditions of the ponds did not differsignificantly.These results are consistent with the hypothesis thatcompetitionfor macrozoans occurs between red-necked grebe and fish andthat adense population of many fish species caused red-necked grebestoavoid otherwise acceptable breeding sites.     

Aspects of the foraging behaviour of the Antarctic Tern Sterna vittata gaini at Harmony Point, South Shetland Islands

During January and February of 2002 and 2003, we studied the diet of the Antarctic Tern Sterna vittata gaini at two colonies in Nelson Island, South Shetland Islands, by identifying the prey fed to chicks by breeders. The fish Notothenia coriiceps was the main prey in both seasons, followed by the myctophid Electrona antarctica, Antarctic krill Euphausia superba and gammarid amphipods. The contribution of fish to the diet increased as chicks grew older. Fish and amphipods were brought to chicks during the day, whereas adults brought Antarctic krill at sunrise and sunset. Both the duration of the feeding trips and the number of trips per foraging bout varied according to the type of prey caught. Preliminary information suggests that, among other causes, the foraging strategy is strongly influenced by the predation pressure of skuas on chicks. Results are compared with the only two previous study on the diet of the Antarctic Tern at the South Shetland Islands.     

Cross-shoal variability in the feeding habits of migrating Atlantic cod (Gadus morhua)

Prey intake and selection were related to within-shoal position for Atlantic cod (Gadus morhua) engaged in annual migration across the Newfoundland shelf in the northwest Atlantic. Comparisons made among fish occupying five regions from the front to rear of a large (>10 km across) migrating shoal indicated that leading fish, or scouts, were larger, ate more food by weight, and had a more varied diet than did fish at other positions. Also, scouts consumed more preferred prey types (fish and pelagic invertebrates) than did fish at other positions. In contrast, trailing fish consumed few fish prey but a larger proportion of benthic invertebrates. Our results are the first to document systematic heterogeneous feeding success among members of a free-ranging and migrating fish shoal in the open ocean.     

Juvenile helping in the moorhen,Gallinula chloropus

In a 3-year study of the moorhen, Gallinula chloropus, some chicks from first broods stayed on their natal territory once they had reached independence, and helped to rear their younger, second brood, siblings. Juvenile dispersal was constrained by habitat saturation, and first brood young were forced to stay on their natal territory. Juveniles that hatched early in the year were forced to stay longer, and helped more than those that hatched late. The total feeding rates to broods with and without juvenile helpers were the same, but parents with helpers reduced their feeding rates relative to parents without helpers. Pairs with helpers (=pairs attempting second broods) reared more chicks per nesting, attempt than pairs rearing chicks at the same time of year without helpers (=pairs attempting first clutch renests). This was true both for all clutches, and for hatched clutches only, even when controlling for parental quality, territory size and scasonal effects.     

Brood reduction in the Red-necked Grebe Podiceps grisegena

Brood reduction in Red-necked Grebes Podiceps grisegena breeding on fish ponds in south-eastern Poland occurred either through the desertion of the last-laid eggs after partial hatching of the clutch and/or the selective starvation of the smallest chicks. Abandonment of unhatched eggs was not influenced by the number of young already hatched or by the breeding date, but it was more likely in larger clutches and in families suffering chick starvation. Chicks from the largest broods had a higher probability of survival until fledging than those from single-chick broods. Larger chicks obtained food more successfully through better positioning during food delivery. In families that did not suffer brood reduction, chicks were better provisioned with food than in reduced broods. Although allocation of food among chicks in reduced broods was more skewed to the disadvantage of the younger siblings, dominant chicks obtained less food prior to brood reduction than dominant siblings in unreduced broods. Sibling aggression did not differ between unreduced and reduced broods before death of the weakest chicks. Post-laying adjustment of the number of offspring to prevailing feeding conditions occurred at two stages: by parental manipulation of the number of hatched eggs at the time when parents and chicks leave the nest and by competition between chicks. It is suggested that late egg desertion may be an adaptive mechanism of brood-size adjustment, when elimination of the weakest chicks through sibling competition is not very efficient.     

Feeding habits of invasive American mink (Neovison vison) in northern Germany—potential implications for fishery and waterfowl

In order to collect ecological data of invasive American mink (Neovison vison) at a fishpond area in northeastern Germany, we conducted a telemetry study in which 14 mink were radio-tracked. During this project, 2,502 scats from radio-tracked individuals were collected in the period from October 2003 to October 2005. Investigated mink principally prey on fish, small mammals and birds (eggs inclusive), whereas amphibians, reptiles and invertebrates were caught infrequently. Analysing mink scats of different seasons, we found significant seasonal variations of diet composition. In spring, fish, mammals and birds were hunted in similar amounts. During summer, birds made up the main part of the diet followed by mammals. In autumn, the proportion of birds in the mink diet decreased, whereas fish gained in importance. This trend continued during the winter period, when mink preyed almost exclusively on fish. Amphibians, crustaceans, insects, molluscs and reptiles were found only occasionally in scat samples. Among birds, the mink preyed mainly on the Eurasian Coot (Fulica atra) followed by the Mallard (Anas platyrhynchos). Mammalian prey was clearly dominated by the water vole (Arvicola terrestris) and among fish, mink hunted especially perch (Perca fluviatilis), roach (Rutilus rutilus) and carp (Cyprinus carpio). Results clearly demonstrate that mink is an opportunistic predator, which hunts its prey according to availability and vulnerability, respectively. Despite the high portions of fish in their autumn and winter diet, the economic damage caused by mink seems to be negligible. However, high predation rates on birds during the breeding season indicate a potential negative impact of mink on waterfowl.     

The food of the Nile perch, Lates niloticus (L.), after the disappearance of the haplochromine cichlids in the Nyanza Gulf of Lake Victoria (Kenya)

After the disappearance of the haplochromine species in the Nyanza Gulf of Lake Victoria as a result of predation by Lates niloticus , the latter has turned its attention to aquatic invertebrates and other fish. Changes in the diet of the Nile perch with increase in its size have been observed: young L. niloticus preyed mostly on invertebrates, including crustaceans and various small aquatic insects; large, immature L. niloticus supplemented the invertebrate diet with both young and small fish; adults above 80 cm total length were mainly piscivorous. L. niloticus feeds on fish prey of about one third its own length.
The tendency of L. niloticus to switch from one prey item to another, depending on availability, is reported; e.g., in the Nyanza Gulf, the prey diet has shifted from the haplochromine to Caridina nilotica and L. niloticus juveniles.     

Shared parental care is costly for nestlings of common cuckoos and their great reed warbler hosts

Obligate avian brood parasitism typically involves one of 2strategies: parasite chicks are either 1) virulent and evictall other eggs and nest mates to be raised alone or 2) moretolerant and share foster parental care with host chicks forsome or the entirety of the nestling period. We studied theconsequences of experimentally forced mixed broods of age-matchedone common cuckoo (Cuculus canorus) and 2 great reed warbler(Acrocephalus arundinaceus) chicks. In these broods, both cuckooand host chicks grew slower than did either individual cuckoosor great reed warblers in broods of 1 parasite or 3 host chicks,respectively. Video records showed that in mixed broods, cuckoochicks received feedings less frequently than the 33% predictedby chance at 4 days of age but parental food allocations increasedto chance levels at 8 days of age. The consistent patterns oflower growth rates arose even though chicks in broods of 1 parasiteand 2 hosts received the largest prey items per feeding. Inaddition, several other measures of parental provisioning alsodid not predict species and brood-specific differences in nestlinggrowth rates across the different treatments. However, variationin begging displays and its specific costs on host and parasitechicks in the different nest treatments were not quantifiedin this study. We conclude that young of nest mate–evictorcommon cuckoos benefit from the sole occupancy of host nestsin part owing to an initial competitive disadvantage for parentalcare in broods with age-matched great reed warbler chicks.     

Diet of the western carp gudgeon (Hypseleotris klunzingeri Ogilby) in an Australian floodplain lake: the role of water level stability

To examine temporal variation in diet, two size classes of western carp gudgeon Hypseleotris klunzingeri were collected from macrophyte habitats in Dugays 2 billabong, a floodplain lake on the River Murray, Victoria, Australia, for 1 year. Fish were collected bimonthly during the day and at night between October 1995 and September 1996. Western carp gudgeon consumed a range of prey, including caddisflies, odonates, shrimps and detritus. The bulk of their diet volume, however, consisted of chironomids and zooplankton. The diets of small and large western carp gudgeon were similar and did not change from day to night. There was, however, high variability across sampling trips in fish diets, both at the community level and among selected prey groups, particularly chironomids, zooplankton and detritus. Community analysis revealed that diets of fish collected from the billabong following stable antecedent water levels were similar and showed little variation, while those collected following fluctuating antecedent water levels showed much more variation. These patterns were mirrored in the volumes of individual diet components.     

Variation in the diet of the viperine snake Natvix mama in relation to prey availability

The diet of the viperine snake was compared with food availability in the Ebro Delta, a wetland largely occupied by rice fields, in 1990 and 1991. Snake selection of prey type and size was studied seasonally and by snake group: males, females and immature snakes. Overall, feeding activity (percentage of individuals with prey and number of prey per stomach) increased with food availability. Diet analysis showed that viperine snakes mainly foraged on the green frog Rana perezi (adults and tadpoles) and the carp Cyprinus earpio. Conversely, viperine snakes rejected the mosquito fish Gambusia holbroki which is the most abundant species in autumn, when Natrix maura has a low feeding activity. Statistical comparisons between viperine snake diet and prey availability showed that males selected small carp, immature snakes selected tadpoles and, in spring, females selected frogs. The selection of small carp by males may reflect a sexual divergence of trophic niche related to sexual size dimorphism, as females are larger than males. As tadpoles are presumably easier to catch than fish, tadpole selection by immature individuals may reflect variance in capture abilities. In spring, the selection of frogs by females overlapped with vitellogenesis, suggesting that females compensate for the cost of reproduction by selecting green frogs, which have a greater biomass and higher energy content than fish. Carps eaten in spring were smaller than in summer. Moreover, in summer viperine snakes selected smaller carp than the available mean size. This divergent tendency between carp size selection and carp size availability reveals how seasonal diet shifts in prey size selection may be a response to an increase in prey size.     

Translocation of threatened New Zealand falcons to vineyards increases nest attendance, brooding and feeding rates

Anthropogenic landscapes can be rich in resources, and may in some cases provide potential habitat for species whose natural habitat has declined. We used remote videography to assess whether reintroducing individuals of the threatened New Zealand falcon Falco novaeseelandiae into a highly modified agricultural habitat affected the feeding rates of breeding falcons or related breeding behavior such as nest attendance and brooding rates. Over 2,800 recording hours of footage were used to compare the behavior of falcons living in six natural nests (in unmanaged, hilly terrain between 4 km and 20 km from the nearest vineyard), with that of four breeding falcon pairs that had been transported into vineyards and nested within 500 m of the nearest vineyard. Falcons in vineyard nests had higher feeding rates, higher nest attendance, and higher brooding rates. As chick age increased, parents in vineyard nests fed chicks a greater amount of total prey and larger prey items on average than did parents in hill nests. Parents with larger broods brought in larger prey items and a greater total sum of prey biomass. Nevertheless, chicks in nests containing siblings received less daily biomass per individual than single chicks. Some of these results can be attributed to the supplementary feeding of falcons in vineyards. However, even after removing supplementary food from our analysis, falcons in vineyards still fed larger prey items to chicks than did parents in hill nests, suggesting that the anthropogenic habitat may be a viable source of quality food. Although agricultural regions globally are rarely associated with raptor conservation, these results suggest that translocating New Zealand falcons into vineyards has potential for the conservation of this species.     

Chick parasitism by blowflies affects feeding rates in a Mediterranean population of blue tits

Offspring fitness depends on interactions between parental care and environmental constraints. It has been suggested that in altricial birds parents are able to compensate for the detrimental effects of ectoparasites by improving food provisioning. We tested this prediction in a population of blue tits highly parasitized by blowfly larvae. The frequency of parental feeding visits was significantly higher in parasitized broods than in broods experimentally deparasitized. Despite a strong increase in parental care, chicks of parasitized broods were lighter, smaller, and more anaemic than chicks in deparasitized broods. Parents invest more in feeding parasitized young but cannot fully compensate for the negative effects of parasites, hence young are in poor condition at fledging.     

A novel strategy to escape a poor habitat: red-necked grebes transfer flightless young to other ponds

Animals confronted with the threat of the death of their offspring may exhibit unusual and risk-prone behaviours. Grebes (Podicipediformes) are water birds which cannot effectively walk, thus unfledged young are assumed to be unable to depart from their natal ponds by land. We provide evidence that red-necked grebes Podiceps grisegena, breeding on ponds with scarce food resources, transferred their flightless young (2–4 weeks old) to other, unconnected ponds by land or air. Although a large proportion of breeding grebes in the study area nested on food-poor fish ponds acting as ecological traps, where they suffered significant brood losses, brood movements to new ponds accounted for only 3.3% of such breeding attempts. The infrequency of this strategy may be explained by the lack of suitable territories in close proximity and the high risk of predation or fatal injury. The means of chick transfer remains unclear; the chicks may have followed or been carried by parents shuffling across the pond levees; alternatively, parents may have carried the young on their backs in flight. Our findings indicate that red-necked grebes assess the current level of resources available for chicks and may adopt novel and risky strategies to escape total brood failure.