Red fox trap success is correlated with piping plover chick survival

Predation management is an important component of managing species of conservation concern. The piping plover (Charadrius melodus; plover), a disturbance-dependent and conservation-reliant shorebird that nests on sandy beaches and barrier islands on the Atlantic Coast, was listed under the United States Endangered Species Act in 1986, with habitat loss and predation stated as key causes of its decline. We evaluated the relationship between a suite of predators and plover chick survival from 2015–2018. We used a camera grid to establish indices of the abundance of 3 known chick predators: red fox (Vulpes vulpes), raccoon (Procyon lotor), and domestic cat. We used camera detections in a survival model to assess potential relationships between predator species detection and plover chick survival. Plover chick survival was negatively related with red fox detection, but not with detection of the other 2 predators. In addition to the correlation with red fox detections, chick survival was negatively related to high plover nesting density. Our results suggest that red foxes were predators of piping plover chicks during our study, likely augmented by other density-dependent sources of mortality. Targeted predator management could aid in conservation of piping plovers in this system as a short-term solution, but long-term recovery plans must also address habitat limitation.     

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.     

Snowy plover nest site selection,spatial patterning,and temperatures in the Southern High Plains of Texas

Snowy plover (Charadrius nivosus) populations have declined throughout their range, in part because of habitat degradation and poor nest success, making information regarding regionally specific nest site selection and spatial patterns important when considering habitat conservation and management guidelines. We determined nest site selection characteristics (n = 180) and examined spatial patterns (n = 215) of snowy plover nests in saline lakes in the Southern High Plains (SHP) of Texas. At 104 nests, we examined the influence of substrate type on nest temperatures and heat mitigation. Snowy plover nests were more likely to be found near an object, on pebble substrate, and with fewer plants than random sites. High use areas were generally located in areas with pebble substrate and on human-made or natural islands, berms, and peninsulas. Overall, nests placed on pebble substrate had lower temperatures during the day than nests placed on sand substrates. Nest placement on pebble substrate may be valuable to nesting snowy plovers, providing thermal advantages to incubating adults and depressing potentially high nest predation rates. Management guidelines for this region should emphasize the importance of addressing key elements of snowy plover nesting habitat including the presence of pebble substrate and reducing vegetation encroachment. © 2012 The Wildlife Society.     

Piping plover habitat selection and nest success on natural,managed, and engineered sandbars

Loss of breeding habitat and nest predation have contributed to the decline of many shorebird species. The United States Army Corps of Engineers (USACE) initiated a piping plover (Charadrius melodus) habitat creation and augmentation program on the Missouri River in the summer of 2004. The USACE increased unvegetated sandbar habitat by depositing dredged material (engineered sandbars) and by clearing vegetation from existing sandbars (managed sandbars). We evaluated the effects of this increase in nesting and foraging habitat on habitat selection and nest daily survival rate (DSR) of piping plovers on Lewis and Clark Lake and the Gavins Point Reach of the Missouri River from 2005 to 2007 (n = 623 nests). Piping plovers selected engineered sandbars more often than expected based on area and selected natural and managed habitats less than expected based on area. Daily survival rate on engineered sandbars was significantly higher than on natural or managed sandbars (log odds: 2.50, 95% CI: 1.05–5.94). Thus, plovers' habitat selection may have increased their nesting success. Our results suggest that habitat augmentation may stave off declines in piping plover populations limited by insufficient habitat and low nesting success. © 2011 The Wildlife Society.     

Habitat use by Mountain Plovers in prairie dog colonies in northeastern New Mexico

ABSTRACT Mountain Plovers (Charadrius montanus) are grassland birds that often breed in close association with colonies of black‐tailed prairie dogs (Cynomys ludovicianus). However, not all colonies provide plover nesting habitat or habitat of equal quality, and the characteristics of colonies important for plovers remain poorly understood. Over two years, I used plover distribution surveys, territory mapping, and habitat sampling to study habitat use by plovers in prairie dog colonies in shortgrass prairie in northeastern New Mexico. My objective was to document important components of plover breeding habitat in colonies by comparing characteristics of used and unused habitats at three spatial scales: colony, territory, and nest‐site. I found evidence of plover breeding in 14 of 44 colonies in 2009 and 13 of 43 colonies in 2010. Based on logistic regression, the probability of a colony being occupied by plovers was positively associated with colony size, but negatively associated with mean vegetation height. Preference for larger colonies could relate to minimum habitat requirements, or a potential tendency of this species to nest in social clusters. Shorter vegetation height was strongly correlated with greater bare ground and lower forb/subshrub cover, all characteristics that may be related to plover predator avoidance and foraging microhabitat. At both the territory and nest‐site scale, areas used by plovers had shorter vegetation, more bare ground, and less forb/subshrub cover than unused areas. Nest sites were also more sloped, perhaps to reduce risk of flooding, and located further away from the nearest prairie dog burrow, perhaps to reduce risk of disturbance. Overall, my results show that plover use of prairie dog colonies was influenced by landscape and habitat features of colonies, and suggest that large colonies are particularly valuable because they are most likely to contain adequate areas with preferred habitat characteristics.     

Disturbance regimes and mountain plover habitat in shortgrass steppe: Large herbivore grazing does not substitute for prairie dog grazing or fire

Restoring historical disturbance regimes to enhance habitat for grassland birds can conflict with livestock production goals and has been controversial because of uncertainty in the frequency and pattern of different disturbances prior to European settlement. We studied nesting habitat for the mountain plover (Charadrius montanus) in relation to prescribed fire, grazing by large herbivores (cattle), and grazing by black-tailed prairie dogs (Cynomys ludovicianus) in the shortgrass steppe of northeastern Colorado. Breeding mountain plovers primarily occurred on black-tailed prairie dog colonies or areas burned during the previous dormant season. Vegetation surrounding mountain plover nests and foraging locations was characterized by a fine-scale mosaic of prostrate (<4 cm tall) vegetated patches interspersed with >35% bare soil in a given square meter, with this fine-scale pattern distributed over a broad (>100-m radius) area. Mountain plovers rarely occupied grassland lacking prairie dogs or recent fire, but those that did selected sites with similar vegetation height and bare soil exposure as sites on burns and prairie dog colonies. Vegetation structure at mountain plover-occupied sites was also similar to random sites on burns and prairie dog colonies, but differed substantially from sites managed only with cattle. Intensive cattle grazing at twice the recommended stocking rate during spring (Mar–May) or summer (May–Oct) for 6 years produced significantly less bare soil than burns and prairie dog colonies, particularly following years with average or above-average precipitation. Thus, intensive cattle grazing did not substitute for prairie dog grazing or fire in terms of effects on vegetation structure and mountain plover habitat. Both prescribed burning and increased size and distribution of black-tailed prairie dog colonies appear to be effective and complementary means to manage for mountain plover breeding habitat in shortgrass steppe. Provision of mountain plover habitat has tradeoffs with traditional management for livestock production. Thus, managers need to clearly define desired outcomes for management to provide multiple ecosystem goods and services. © 2012 The Wildlife Society.     

Mid-contract management of Conservation Reserve Program grasslands provides benefits for ring-necked pheasant nest and brood survival

Conservation Reserve Program (CRP) fields may provide good habitat for nesting and brood-rearing ring-necked pheasants (Phasianus colchicus) during early stages of succession. But, the success of hens in early successional CRP, relative to late successional CRP and other grassland habitats, has yet to be evaluated. The reproductive period is especially critical for populations of pheasants, and CRP's benefits to hens and chicks may decrease as fields age because of loss of vegetative diversity, decrease in vegetation density, and accumulation of residual litter. During 2005–2006, we evaluated spatial and temporal variation in nest and brood survival for radio-marked hen pheasants in areas of northeastern Nebraska where portions of CRP fields had been recently disced and interseeded (DICRP) with legumes. Nests in DICRP tended to have a higher daily survival rate (0.984; 95% CI: 0.957–0.994) than nests in grasslands (including CRP) that were unmanaged (0.951; 95% CI: 0.941–0.972). The probability of 23-day nest success was 0.696 (95% CI: 0.631–0.762) for DICRP and 0.314 (95% CI: 0.240–0.389) for unmanaged grasslands. Daily brood survival rates varied by habitat type, brood age, and date of hatch. The probability of a brood surviving to day 21 was 0.710 (95% CI: 0.610–0.856). Brood survival rates increased with time spent in DICRP and as the brood aged. Survival decreased as broods spent more time in cropland and peaked seasonally with broods that hatched on 15 June. Brood survival probability, to 21 days, would be reduced to 0.36 (95% CI: 0.100–0.701) if broods in our sample had not used DICRP. We combined nest and brood survival in a productivity model that suggested 2,000 hens, in a landscape with no DICRP, would produce 1,826 chicks, whereas the same hens in a landscape of 100% DICRP would produce 5,398 chicks. Production of first-year roosters more than doubled when hens nested in DICRP. Without DICRP, population growth rates of pheasant populations usually declined; with DICRP, populations stabilized with at annual survival rates of 0.3 or greater. The positive response of nest and brood survival to discing and interseeding CRP provides further evidence that CRP fields must be managed to optimize wildlife benefits. © 2012 The Wildlife Society.     

Nest‐site selection by golden plover: why do shorebirds avoid nesting on slopes?

It is widely known that many upland-breeding shorebirds tend to nest on plateaus but to date no studies have put forward explanations for this phenomenon. We examined the effect of slope and habitat on the distribution of ground-nesting golden plover Pluvialis apricaria at two study sites in County Durham, U.K. Golden plovers showed strong selection for nesting on flat ground. Habitat significantly affected nest-site location on one study site (heather burnt within the past 2 years was favoured and older stands of heather were avoided) but not on the other. Fifty-nine per cent of all nests failed. We attributed 95% of all losses to predation. Seventy-five per cent of nests, in which the predator was identified, were taken by ground predators, mainly stoats Mustela erminea . Nests on flat ground had significantly higher rates of survival than those on slopes. Nest survival did not vary significantly with habitat type nor with vegetation height or density around the nest. Neither habitat type, vegetation height nor vegetation density around nests differed between nests on slopes and on flat ground. In addition, individuals that nested on flat ground tended to have less black on their underparts (a suggested indicator of dominance). We suggest that birds nesting on slopes are less efficient at avoiding nest predation than those nesting on the flat. The nature or degree of a bird's response to a predator may be related to the efficiency of individual anti-predator responses and/or to visibility from the nest. This study cannot differentiate between these two explanations. We encourage further work to investigate differences in behaviour between individuals nesting on slopes and those on flat ground.     

Improving conservation strategies of raptors through landscape ecology analysis: The case of the endemic Cuban Black Hawk

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Landscape-Level Assessment of Brood Rearing Habitat for Greater Sage-Grouse in Nevada

Abstract: Loss of quality brood rearing habitat, resulting in reduced chick growth and poor recruitment, is one mechanism associated with decline of greater sage-grouse (Centrocercus urophasianus) populations. Low chick survival rates are typically attributed to poor-quality brood rearing habitat. Models that delineate suitability of sage-grouse nesting or brood rearing habitat at the landscape scale can provide key insights into the relationship between sage-grouse and the environment, allowing managers to identify and prioritize habitats for protection or restoration. We used Southwest Regional Gap landcover types to identify early and late greater sage-grouse brood rearing in east-central Nevada. We conducted an Ecological Niche Factor Analysis to 1) examine the effect these landcover types and other ecogeographical variables have on sage-grouse selection of brood rearing habitat, and 2) generate landscape-scale suitability maps. We also evaluated if incorporating a fitness component (brood survival) in landscape spatial analyses of habitat quality influenced our assessment of habitat suitability. Because 36% of our 6,500-km² study area was identified as early brood rearing habitat, we believe this habitat may not be limiting greater sage-grouse populations in east-central Nevada, USA, at least in wet years. We found strong selection for particular landcover types (e.g., higher elevation, moist sites with riparian shrubs or montane sagebrush) during late brood rearing. Late brood rearing habitat on which broods were successfully reared represented only 2.8% of the study area and had a restricted distribution, suggesting the potential that such habitat could limit sage-grouse populations in east-central Nevada.     

Trade-offs in nest site selection in coastal populations of Lapwings Vanellus vanellus

In coastal populations of Lapwings Vanellus vanellus in southwestern Sweden, arable fields predominated as foraging habitat before laying. Females caught more large prey items on arable fields and shores than on pastures. Close to egg laying, females foraged mainly near their future nest sites. Arable land and pastures were used to a similar extent for nesting. We found no difference in nest predation between habitats. Egg volume varied among females and was correlated with wing-length, body mass and condition. Mean egg volume also was positively correlated with feeding time on arable land before laying. Pairs nesting on arable fields therefore generally produced larger eggs than those on pastures. The distances between nests and chick foraging areas, however, were significantly longer for birds nesting on arable land than for those on pastures. Moreover, in 2 of 3 years, the proportion of hatched chicks that survived until fledging was negatively correlated with this distance. There was no difference in chick survival between broods hatched on arable fields and pastures. We suggest that nest site selection and offspring production involve a trade-off between the benefits of nesting close to rich feeding grounds for adults and the costs of moving long distances between nest sites and chick-rearing areas.     

Multi-scale habitat models for reintroduced bird populations: a case study of South Island saddlebacks on Motuara Island

Understanding resource selection by animals is important when considering habitat suitability at proposed release sites within threatened species recovery programmes. Multi-scale investigatory approaches are increasingly encouraged, as the patchy distribution of suitable habitats in fragmented landscapes often determines species presence and survival. Habitat models applied to a threatened New Zealand forest passerine, the South Island saddleback (Philesturnus carunculatus carunculatus), reintroduced to Ulva Island (Stewart Island) found that at landscape scale breeding pairs? preferences for sites near the coast were driven by micro-scale vegetation structure. We tested these results by examining models of breeding site selection by a reintroduced saddleback population on Motuara Island (Marlborough Sounds) at two scales: (1) micro-scale, for habitat characteristics that may drive breeding site selection, and (2) landscape scale, for variations in micro-scale habitat characteristics that may influence site colonisation in breeding pairs. Results indicated that birds on Motuara Island responded similarly to those on Ulva Island, i.e. birds primarily settled at the margins of coastal scrub and forest and later cohorts moved into larger stands of coastal forest where they established breeding territories. Plant species composition was also important in providing breeding saddleback pairs with adequate food supply and nesting support. However, Motuara Island birds differed in their partitioning of habitat use: preferred habitats were used for nesting while birds were foraging outside territorial boundaries or in shared sites. These differences may be explained because Motuara has a more homogeneous distribution of microscale habitats throughout the landscape and a highly bird-populated environment. These results show that resource distribution and abundance across the landscape needs to be accounted for in the modelling of density?bird?habitat relationships. In the search for future release sites, food (invertebrates and fruiting tree species) should be abundant close to available nesting sites, or evenly spread and available throughout the landscape.     

Use of Restored Riparian Habitat by the Endangered Least Bell's Vireo (Vireo bellii pusillus)

A primary objective of riparian restoration in California is the creation of habitat for endangered species. Four restoration sites in San Diego County were monitored between 1989 and 1993 and evaluated for their suitability as nesting habitat for Vireo bellii pusillus (Least Bell's Vireo), a state and federally endangered obligate riparian breeder. Vegetation structure at each site was quantified annually and compared to a model of canopy architecture derived from Least Bell's Vireo territories in natural habitat. Vireo use of restored habitat was documented through systematic surveys and nest monitoring. By 1993, only one site in its entirety met the habitat suitability criteria of the model, but portions of each site during all years did so. Differences between sites in the time required to develop suitable habitat—well-developed layered vegetation from the ground to under 8m in height)—were attributable largely to variation in annual rainfall. Vireos visited restoration sites to forage as early as the first growing season, but they did not establish territories or nest there until at least part of the site supported suitable habitat as determined from the model. Placement of territories and nests coincided with patches of dense vegetation characteristic of natural nesting areas. Occupation of restored sites was accelerated by the presence of adjacent mature riparian habitat, which afforded birds nest sites and/or foraging habitat lacking in the planted vegetation. Vireos nesting in restored habitat achieved success comparable to that of vireos nesting in surrounding natural habitat, and there was no evidence that productivity was reduced in created areas. These findings indicate that creating nesting habitat for this target species is feasible and suggest that the critical components of vireo nesting habitat have been captured in both the design and quantitative assessment of restoration sites.     

Developmental changes in factors limiting colony survival and growth of the leaf‐cutter ant Atta laevigata

Many species of leaf‐cutter ants (Atta and Acromyrmex) increase in abundance following natural or anthropogenic disturbances in the vegetation. However, the mechanisms responsible for such an increase are still poorly understood. We evaluated the effects of nesting site and the availability of palatable forage on survival and growth of Atta laevigata colonies at different developmental stages. Foundress queens transplanted into man‐made clearings (dirt roads) had a much higher survival than those transplanted into the adjacent undisturbed savannah vegetation. Similarly, incipient colonies (≥3‐months old) had significantly greater survival and growth rates in dirt roads. In contrast, nesting site did not influence performance of young colonies (≥15‐months old). Both incipient and young colonies responded strongly and positively to experimental supplementation of palatable forage, and this effect was independent of the nesting habitat. Colonies that received extra food grew faster and had a significantly greater survival rate than control colonies. These results suggest that performance of A. laevigata is affected by the generally greater availability of suitable nesting sites and palatable vegetation in disturbed habitats. This may explain how these ants maintain high densities in these habitats, and since the relative importance of these factors changed with colony ontogeny, our findings highlight the importance of evaluating potential limiting factors throughout the full range of an organism's developmental stages.     

The function of habitat change during brood-rearing in the precocial Kentish plover <Emphasis Type="Italic">Charadrius alexandrinus</Emphasis>

Food availability is often variable during the breeding season. Parents with nonmobile, altricial young have no choice but to accept changes in local food availability, whereas in precocial animals, the parents may lead their young away from poor sites to areas that have rich resources and/or are safe from predators. We investigated the latter hypotheses in the Kentish plover Charadrius alexandrinus, a precocial shorebird that raises its young in two habitats: on lakeshore and in saltmarsh. Parents move with their broods from saltmarsh to lakeshore, especially late in the breeding season, and we hypothesized that lakeshores provide more food than the saltmarsh. Consistent with our hypotheses, plover chicks grew faster on the shore, and the difference in growth rates between the two habitats was amplified later in the breeding season. In addition, brood survival was higher on lakeshore than in saltmarsh and decreased with hatching date. Taken together, our results suggest that Kentish plover parents increase their reproductive success by switching brood-rearing habitats strategically.     

Human disturbance and stage‐specific habitat requirements influence snowy plover site occupancy during the breeding season

Habitat use has important consequences for avian reproductive success and survival. In coastal areas with recreational activity, human disturbance may limit use of otherwise suitable habitat. Snowy plovers Charadrius nivosus have a patchy breeding distribution along the coastal areas on the Florida Panhandle, USA. Our goal was to determine the relative effects of seasonal human disturbance and habitat requirements on snowy plover habitat use. We surveyed 303 sites for snowy plovers, human disturbance, and habitat features between January and July 2009 and 2010. We made multiple visits during three different sampling periods that corresponded to snowy plover breeding: pre‐breeding, incubation, and brood‐rearing and used multi‐season occupancy models to examine whether human disturbance, habitat features, or both influenced site occupancy, colonization (probability of transition from an unoccupied site to an occupied site), and extinction (probability of transition from an occupied site to an unoccupied site). Snowy plover site occupancy and colonization was negatively associated with human disturbance and site extinction was positively associated with human disturbance. Interdune vegetation had a negative effect on occupancy and colonization, indicating that plovers were less likely to use areas with uniform, dense vegetation among dunes. Also, dune shape, beach debris, and access to low‐energy foraging areas influenced site occupancy, colonization, and extinction. Plovers used habitat based on beach characteristics that provided stage‐specific resource needs; however, human disturbance was the strongest predictor of site occupancy. In addition, vegetation plantings used to enhance dune rehabilitation may negatively impact plover site occupancy. Management actions that decrease human disturbance, such as symbolic fencing and signage, may increase the amount of breeding habitat available to snowy plovers on the Florida Panhandle and in other areas with high human activity. The specific areas that require this protection may vary across snowy plover life history stages.     

Habitat preferences of lesser rheas in Argentine Patagonia

This work reports the first results of a 3-year study (1998–2001) on habitat use and preferences by wild lesser rheas (Rhea pennata pennata) in the ecotone Monte–Patagonian steppe of Argentina. Ponds and four different habitat alternatives for lesser rheas were determined using satellite images and different structural vegetation characteristics: steppe, shrubland, shrub steppe, and mallines. Lesser rheas (adults, juveniles and chicks) used all habitats available to feed, although they showed preference for mallines, open areas that offer good visibility and abundant food resource. Rheas used shrub steppe and steppe for nesting, but they did not show preference for either habitat type. Nest site had a higher percentage of vegetation cover than control sites. Lesser rheas apparently preferred concealed sites for nesting since they offer protection from severe climate conditions and from predators. Our results suggest that habitat preference by lesser rheas counterbalances profitability of feeding to the corresponding cost of predation.     

Effects of fire management on northern bobwhite brood ecology

Northern Bobwhite Colinus virginianus chicks require ample invertebrates for growth and feather development. Early successional or resprouting vegetation provides invertebrates for chicks but may not provide other resources such as roosting and loafing cover that is typically provided by later successional stages. Thus, management for bobwhites provides multiple seral stages in close proximity but the effects of landscape interspersion have not been tested for bobwhite broods. During a 2-year study, we explored the effects of landscape complementation and food availability on growth and survival of bobwhite chicks. We found growth of chicks to be negatively related to home range size which was negatively correlated to the amount of area burned. We also found survival of chicks to be positively related to the amount of burned area (i.e., foraging area) within brood home ranges. To maximize the growth and survival of bobwhite chicks, it would be necessary to increase access to foraging areas while decreasing the size of brood home ranges. Access to foraging areas can be created through frequent prescribed fire at small spatial scales.     

Breeding ecology of Yellow Wagtails Motacilla flava in an arable landscape dominated by autumn-sown crops

Capsule Yellow Wagtails successfully raised two consecutive broods in landscapes dominated by autumn-sown crops and did not require spring crops or fallow plots for later nesting.

Aims To assess whether arable landscapes dominated by winter cropping provide habitats that allow Yellow Wagtails to raise two successful broods. To assess the utility of spring cultivated agri-environment fallow plots as a nesting and foraging habitat for Yellow Wagtails.

Methods Nesting success and foraging behaviour of Yellow Wagtails was monitored on lowland arable farmland dominated by winter cropping during two successive breeding seasons.

Results Yellow Wagtails successfully raised first and second broods mainly in winter cereals, with later nests being more successful. Some nests were initiated in bean fields where egg-stage failure rates were high, probably as a consequence of depredation. Fallow plots were not used for nesting but, along with other areas of sparse vegetation, were regularly used for foraging.

Conclusions Yellow Wagtails breeding at relatively low densities achieved high rates of reproductive success in a landscape dominated by winter cereals and with few spring-sown crops. This study suggests that spring-sown crops may not be necessary for Yellow Wagtails to rear two successful broods each summer within arable landscapes.     

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.