Population estimates of the breeding birds of the Tinhosas islands (Gulf of Guinea), the only major seabird colony of the eastern tropical Atlantic

The Tinhosas islands, in São Tomé e Príncipe, host the most important seabird breeding colony in the Gulf of Guinea, but information on its conservation status was hitherto unpublished or anecdotal, the last assessment having been performed in 1997. A two-day expedition to the Tinhosas islands was undertaken to estimate the status of breeding seabirds in 2013. Four of the five seabird species known to breed in São Tomé e Príncipe, namely Brown Booby Sula leucogaster, Sooty Tern Onychoprion fuscatus, Brown Noddy Anous stolidus and Black Noddy Anous minutus, occur on the Tinhosas. A decrease of 80% in Brown Booby numbers, possibly due to occasional exploitation, and a 30% increase in Sooty Tern and Black Noddy numbers, were found compared to 1997 data although survey methods differed. Breeding of Brown Noddy and Madeiran Storm-petrel Hydrobates castro remains unconfirmed. Our estimates confirmed that BirdLife International Important Bird and Biodiversity Area criteria are met for at least one species, the Sooty Tern. The islands are not legally protected, nonetheless, apart from moderate levels of disturbance by fishermen who land on Tinhosa Grande, no alien species were seen, and no immediate threats to the Tinhosas colony were detected. Multiple visits within and between years are recommended, to census breeders, monitor threats and establish breeding phenologies.     

The impacts of introduced House Mice on the breeding success of nesting seabirds on Gough Island

Invasive species are the main threat to island biodiversity; seabirds are particularly vulnerable and are one of the most threatened groups of birds. Gough Island, a UNESCO World Heritage Site in the South Atlantic Ocean, is an Important Bird and Biodiversity Area, and one of the most important seabird colonies globally. Invasive House Mice Mus musculus depredate eggs and chicks of most seabird species on the island, but the extent of their impact has not been quantified. We used field data and bootstrapped normal distributions to estimate breeding success and the number of surviving chicks for 10 seabird species on Gough Island, and compared estimates with those of analogous species from predator‐free islands. We examined the effects of season and nest‐site location on the breeding success of populations on Gough Island, predicting that the breeding success of Gough birds would be lower than that of analogues, particularly among small burrow‐nesting species. We also predicted that winter‐breeding species would exhibit lower breeding success than summer‐breeding species, because mice have fewer alternative food sources in winter; and below‐ground nesters would have lower breeding success than surface nesters, as below‐ground species are smaller so their chicks are easier prey for mice. We did indeed find that seabirds on Gough Island had low breeding success compared with analogues, losing an estimated 1 739 000 (1 467 000–2 116 000) eggs/chicks annually. Seven of the 10 focal species on Gough Island had particularly high chick mortality and may have been subject to intense mouse predation. Below‐ground and winter breeders had lower breeding success than surface‐ and summer‐breeders. MacGillivray's Prion Pachyptila macgillivrayi, Atlantic Petrel Pterodroma incerta and Tristan Albatross Diomedea dabbenena are endemic or near‐endemic to Gough Island and are likely to be driven to extinction if invasive mice are not removed.     

Cooperative Breeding and Group Structure in the Lake Tanganyika Cichlid Neolamprologus savoryi

As yet, cooperative breeding has been described only for some fish species. However, evidence is accumulating that it is widespread among Lake Tanganyika cichlids. We studied the cooperative breeding system of the substrate breeding cichlid Neolamprologus savoryi. Breeding groups typically consisted of a large breeding male with one to four breeding females and three to 33 helpers (mean group size: 14.3 members). Group size was significantly related to breeding male and female body sizes, and larger males had more breeding females and larger sized male helpers. The size of the largest female in the group was positively related to the number and sizes of secondary breeding females and female helpers. In case of multiple breeding females, these females usually divided the group's territory into sub‐territories, each with its own helpers (subgroups). Interspersed between groups, independent fish were detected defending an individual shelter (4.4% of all fish). In 9% of the groups no breeding female was present. All group members participated in territory defence and maintenance, and showed submissive behaviours to larger group members. As expected, the level of between‐subgroup conflicts was high compared with the level of within‐subgroup conflicts. We compare these results with data available from other cooperatively breeding fishes.     

Impact of the introduced yellow crazy ant Anoplolepis gracilipes on Bird Island, Seychelles

The introduced yellow crazy ant or long-legged ant Anoplolepis gracilipes was first reported in Seychelles in 1969 and now occurs on at least nine islands in the Central Seychelles. We describe the yellow crazy ant's effects on vegetation and invertebrate communities on one of these, Bird Island; in 2000, Anoplolepis (first reported in 1991) occurred there at densities at least 80 times higher than on other islands in the Central Seychelles. They were associated with high densities of coccid scale insects on foliage, especially of the native tree Pisonia grandis, in some instances causing tree death. Yellow crazy ants on Bird Island also significantly affected invertebrate communities on foliage and on the ground, both in terms of taxonomic composition and the density of specific taxa, apparently causing the local exclusion of some invertebrates.     

Helper contributions in the cooperatively breeding laughing kookaburra: feeding young is no laughing matter

I studied the contributions of individuals to incubation and nestling feeding in a population of cooperatively breeding laughing kookaburras, Dacelo novaeguineae. In most cooperatively breeding birds where nest success is limited by nestling starvation, related helpers increase the overall level of provisioning to the nest, thus boosting the production of nondescendent kin. However, although partial brood loss is the largest cause of lost productivity in kookaburra nests, additional helpers failed to increase overall provisioning. Instead, all group members, but especially helpers, reduced their feeding contributions as group size increased. Breeders and helpers reduced the size of prey delivered, and helpers also reduced the number of feeding visits. An important benefit of helping in kookaburras may be to allow all group members to reduce their effort. Within groups, contributions to care depended on status, sex, group size and the brood size. Breeding males delivered the most food. Breeding females provisioned less than their partner, but their effort was comparable to that of male helpers. Female helpers contributed the least food. Incubation effort followed similar patterns. The relatedness of helpers to the brood had no impact on their provisioning. Across all group sizes, helpers generally brought larger items to the nest than breeders. Copyright 2000 The Association for the Study of Animal Behaviour.     

The Effects of Social Context on the Food-Caching Behavior of Florida Scrub-Jays (Aphelocoma coerulescens)

Florida scrub‐jays are cooperative breeders that live in family groups consisting of a breeding pair, often with several non‐breeding helpers. Florida scrub‐jays cache food by scatter‐hoarding items for later consumption. Within family groups, members have the opportunity to observe and pilfer the caches of other members. We observed jays harvesting experimentally provisioned peanuts alone and in the presence of other family members, to determine whether jays modify their food‐handling behavior relative to social context. Non‐breeding helpers were less likely to cache in the presence of the dominant male breeder than when alone and all jays tended to cache out of sight when observed by another jay. These changes in caching behavior are consistent with cache protection strategies employed by other species. However, the adaptive value of such cache protection within a sedentary cooperatively breeding family group on a year‐round territory is unclear.     

Hybridization and Back-Crossing in Giant Petrels (Macronectes giganteus and M. halli) at Bird Island,South Georgia,and a Summary of Hybridization in Seabirds

Hybridization in natural populations provides an opportunity to study the evolutionary processes that shape divergence and genetic isolation of species. The emergence of pre-mating barriers is often the precursor to complete reproductive isolation. However, in recently diverged species, pre-mating barriers may be incomplete, leading to hybridization between seemingly distinct taxa. Here we report results of a long-term study at Bird Island, South Georgia, of the extent of hybridization, mate fidelity, timing of breeding and breeding success in mixed and conspecific pairs of the sibling species, Macronectes halli (northern giant petrel) and M. giganteus (southern giant petrel). The proportion of mixed-species pairs varied annually from 0.4–2.4% (mean of 1.5%), and showed no linear trend with time. Mean laying date in mixed-species pairs tended to be later than in northern giant petrel, and always earlier than in southern giant petrel pairs, and their breeding success (15.6%) was lower than that of conspecific pairs. By comparison, mixed-species pairs at both Marion and Macquarie islands always failed before hatching. Histories of birds in mixed-species pairs at Bird Island were variable; some bred previously or subsequently with a conspecific partner, others subsequently with a different allospecific partner, and some mixed-species pairs remained together for multiple seasons. We also report the first verified back-crossing of a hybrid giant petrel with a female northern giant petrel. We discuss the potential causes and evolutionary consequences of hybridization and back-crossing in giant petrels and summarize the incidence of back-crossing in other seabird species.     

Female‐Biased Helping in a Cooperatively Breeding Bird: Female Benefits or Male Costs?

There is often a sex bias in helping effort in cooperatively breeding species with both male and female helpers, and yet this phenomenon is still poorly understood. Although sex‐biased helping is often assumed to be correlated with sex‐specific benefits, sex‐specific costs could also be responsible for sex‐biased helping. Cooperatively breeding brown jays (Cyanocorax morio) in Monteverde, Costa Rica have helpers of both sexes and dispersal is male‐biased, a rare reversal of the female‐biased dispersal pattern often seen in birds. We quantified helper contributions to nestling care and analyzed whether there was sex‐biased helping and if so, whether it was correlated with known benefits derived via helping. Brown jay helpers provided over 70% of all nestling feedings, but they did not appear to decrease the workload of breeders across the range of observed group sizes. Female helpers fed nestlings and engaged in vigilance at significantly higher levels than male helpers. Nonetheless, female helpers did not appear to gain direct benefits, either through current reproduction or group augmentation, or indirect fitness benefits from helping during the nestling stage. While it is possible that females could be accruing subtle future direct benefits such as breeding experience or alliance formation from helping, future studies should focus on whether the observed sex bias in helping is because males decrease their care relative to females in order to pursue extra‐territorial forays. Explanations for sex‐biased helping in cooperative breeders are proving to be as varied as those proposed for helping behavior in general, suggesting that it will often be necessary to quantify a wide range of benefits and costs when seeking explanations for sex‐biased helping.     

Egg investment in response to helper presence in cooperatively breeding Tibetan ground tits

Life‐history theory predicts a trade‐off between current and future reproduction to maximize lifetime fitness. In cooperatively breeding species, where offspring care is shared between breeders and helpers, helper presence may influence the female breeders’ egg investment, and consequently, survival and future reproductive success. For example, female breeders may reduce egg investment in response to helper presence if this reduction is compensated by helpers during provisioning. Alternatively, female breeders may increase egg investment in response to helper presence if helpers allow the breeders to raise more or higher quality offspring successfully. In the facultatively cooperative‐breeding Tibetan ground tit Pseudopodoces humilis, previous studies found that helpers improve total nestling provisioning rates and fledgling recruitment, but have no apparent effects on the number and body mass of fledglings produced, while breeders with helpers show reduced provisioning rates and higher survival. Here, we investigated whether some of these effects may be explained by female breeders reducing their investment in eggs in response to helper presence. In addition, we investigated whether egg investment is associated with the female breeder's future fitness. Our results showed that helper presence had no effect on the female breeders’ egg investment, and that egg investment was not associated with breeder survival and reproductive success. Our findings suggest that the responses of breeders to helping should be investigated throughout the breeding cycle, because the conclusions regarding the breeders’ adjustment of reproductive investment in response to being helped may depend on which stage of the breeding cycle is considered.     

Geographical variation in patterns of parentage and relatedness in the co‐operatively breeding Ground Tit Parus humilis

The Ground Tit is a large parid species endemic to the Tibetan Plateau. Here we describe its genetic breeding system based on 2 years of fieldwork on a population from Damshung, Tibet. Genetic relatedness and parentage were analysed using 16 microsatellite markers and sex was determined with a marker on the Z and W chromosomes. We established that 16 of 75 families (21%) were assisted by one or occasionally two male helpers, which in most cases were young from a previous brood of one or both of the breeding adults. The helpers typically stayed with their families throughout the breeding season. Helpers never obtained any paternity in their own families, but one helper was identified as the sire in the only instance of extra‐group (and extra‐pair) parentage detected. Thus, the level of extra‐group/extra‐pair parentage appears to be very low in this Ground Tit population, and sharing of reproduction within family groups is apparently absent. Our results contrast with the findings from another Ground Tit population in Gansu, further northeast in China, with respect to both the number and sex of helpers and the division of parentage within and among family groups. In Gansu, helpers regularly produced offspring and both extra‐pair and extra‐group paternity as well as maternity (egg dumping) was common. Differences in family structure, philopatry, territoriality and potential inclusive fitness benefits can probably explain this contrast, and are likely to reflect the relative costs and benefits of co‐operative breeding.     

Corticosterone Levels Correlate With Alloparental Care in a Sex‐Dependent Manner in African Striped Mice,Rhabdomys pumilio

Alloparental care of non‐breeders is the main characteristic of cooperatively breeding species. While many studies have contributed to the understanding of the evolutionary reasons why individuals provide care to young that are not their own offspring, the variables influencing and causing alloparental care are less understood. We tested in African striped mice (Rhabdomys pumilio) whether age, sex, testosterone and corticosterone were correlated with alloparental care of non‐breeding helpers. We studied 11 family groups under controlled conditions in the laboratory, each with two juvenile and two adult helpers, one being male and one being female in each age category. We predicted male helpers to show more alloparental care than female helpers, as males are the dispersing sex and might thus have to pay for staying. We also expected adult helpers to show more alloparental care than juvenile helpers and both corticosterone and testosterone to correlate negatively with alloparental care. We found high levels of alloparental care in non‐breeding striped mice, which spent a significant amount of time in the nest, huddling and licking pups. There was neither a difference between the sexes nor between age categories (although both factors were significant in interaction terms), indicating either low costs and/or high benefits of alloparental care. Mothers showed significantly more care than helpers, and fathers showed similar levels of parental care as mothers but not significantly more than helpers. Although testosterone levels differed significantly between helpers of different age and sex, with adult male helpers showing the highest levels, we did not find any relationships between testosterone and the amount of alloparental care. Corticosterone levels were negatively correlated with alloparental care, and these effects were modulated by the sex and the age of helpers. In females, less alloparental care was shown with increasing corticosterone levels, while in males, the relationship was positive. Also, younger individuals with lower corticosterone levels showed more alloparental care than older individuals with low corticosterone levels. In sum, alloparental care is well developed in male and female non‐breeding helpers of striped mice, both in adult and juvenile helpers, but independently of testosterone, with corticosterone showing an age‐ and sex‐specific relationship with alloparental care.     

No Evidence for Enforced Alloparental Care in a Cooperatively Breeding Parrot

In cooperatively breeding species, in which non‐breeding helpers assist in rearing the offspring of breeding individuals, conflicts of interest commonly occur between breeders and helpers over their respective contributions to offspring care. During such conflicts, breeders might use aggressive behavior to enforce contributions of helpers to offspring care, especially if helpers are not related to the breeders and their offspring and thus do not stand to gain indirect fitness benefits by helping. Using a combination of behavioral and genetic data, we investigated in the cooperatively breeding El Oro parakeet Pyrrhura orcesi (i) whether breeders are commonly dominant over helpers, (ii) whether they use aggressive behavior toward helpers to enforce offspring provisioning, and (iii) whether the relatedness of helpers to the nestlings affects the frequency of—or the reaction of helpers to—such aggressions. Even though breeders were generally dominant over helpers, we found no evidence for the enforcement of alloparental care. This finding was independent of the relatedness between helpers and nestlings, even though distantly related helpers overall contributed little to offspring care. We suggest that the inability of breeders to properly assess the work rates of their helpers at least partly explains the absence of enforcement. More generally, our results add to a body of evidence suggesting that enforcement might be an exceptional rather than a general mechanism underlying the expression of alloparental care in cooperatively breeding species.     

Does it always pay to defend one's nest? A case study in African penguin

Throughout the animal kingdom, individual variation in reproductive success is commonly observed, even under similar environmental conditions. However, the mechanisms behind such differences remain unclear. The notion of behavioural consistency in animals has developed rapidly since the early 21st century partly as an approach to understand among‐individual differences. In this context, a number of studies have highlighted the influence of pair assortment in personality on breeding success. In this study, we related breeding success to individual behaviour, specifically a risk‐taking behaviour, and pair assortment per behaviour in African penguins (Spheniscus demersus) over two breeding seasons of contrasting food availability. On Bird Island, Algoa Bay, South Africa, we used indices of boldness and overall mobility in penguins’ nest defence behaviour as a response to a standard pedestrian approach during chick‐rearing. These behaviours were consistent over the trials and indicated these traits may be related to personality in African penguins. Individuals were categorized as risk‐prone (“bold,” “mobile”) or risk‐averse (“shy,” “non‐mobile”). We then assessed their breeding success through chick growth and survival over 4 weeks in 2015 and 2016. There was weak positive assortment of pairs in relation to nest defence behaviour. However, pair assortment did not significantly influence birds’ breeding success. Shy penguins were generally the most successful (had the highest chick growth rates), which was especially apparent during a food shortage in 2016, possibly reflecting a higher energy investment when foraging. In contrast, chicks from bold parents grew significantly slower, especially in 2016. Bold parents may defend their nest successfully against predation or intra‐specific aggression when food is abundant, but when predation risks are limited and food availability is low, this strategy may not be beneficial. In the context of climate change, where food shortage events may become more frequent, risk‐averse individuals may be favoured and genetic diversity may be reduced in African penguins.     

Plural Breeding in the Splendid Fairy-wren,Malurus splendens (Aves: Maluridae), a Cooperative Breeder

Groups of the cooperatively breeding splendid fairy-wren Malurus splendens may include more than one female. Previously this species has been described as singular breeding (only one female breeds). This paper describes the occurrence of plural breeding (PB) groups in 10% of group years, in which two females had separate nests. In all cases, the secondary female (Y) was related to the primary breeding female (X) and was generally a 2-year old female which had helped in the group during the previous breeding season. Plural breeding was correlated with an increase in population density and in the number of female helpers; PB groups were larger than singular-breeding groups. In most cases, the X female was occupied with her own nest or offspring when the Y female began to nest, and there was no aggression between them. Which birds helped the Y female to feed at her nest depended on the time between the hatching of the two nests. If the interval was small, some group members helped at each nest; with longer intervals, the group members began to feed at the earlier nest, and the other female was left to raise her brood alone. Female helpers were very active in feeding at single nestings, and the cost to an X female of a Y female breeding was mainly a loss of this assistance. The success of individual X nests was not affected. Effects on productivity were slight, but fewer X females in PB groups raised second broods than did experienced singular breeding females. Y females were less productive than X females, but no less productive than singular breeding novice females without helpers. It is not known whether Y females copulated with primary or secondary males within their group, or with males from outside the group. Certainly, they did not form an observable pairing with any male in the group. Plural breeding occurred in a minority of group years in response to extrinsic conditions and the current demographic situation, and shows the extreme plasticity of the mating system in M. splendens.     

Testis size depends on social status and the presence of male helpers in the cooperatively breeding cichlid Julidochromis ornatus

Cooperatively breeding animals, in which helpers may participatein reproduction with dominant breeders, are ideal species forexamining intraspecific variation in testis size because theyoften exhibit both monogamous breeding (low risk of sperm competition)and polyandrous breeding (high risk) within a population. However,little is known about testis investment as a result of spermcompetition in these animals. The substrate-brooding cichlidfish Julidochromis ornatus has a cooperatively breeding system,in which some males mate monogamously and other males reproduceas dominant breeders or helpers within cooperatively breedinggroups, in which male helpers frequently sire young. We examinedthe relationship between testis investment and male social statusin relation to the risk of sperm competition. As predicted fromsperm competition models, in groups with male helpers, boththe male breeders and the male helpers invested more in testesmass, compared to breeding males without male helpers. We alsofound a positive relationship between the testes mass of malebreeders and their male helpers, suggesting that males increasetheir investment in reproductive capability under the risk ofsperm competition. Sperm competition models also predict thatlarger testes are associated with increased siring success.Our paternity analysis supported this prediction; we found apositive relationship between testis investment by male helpersand the number of offspring they sired.     

Cooperative breeding shapes post‐fledging survival in an Afrotropical forest bird

For avian group living to be evolutionary stable, multiple fitness benefits are expected. Yet, the difficulty of tracking fledglings, and thus estimating their survival rates, limits our knowledge on how such benefits may manifest postfledging. We radio‐tagged breeding females of the Afrotropical cooperatively breeding Placid greenbul (Phyllastrephus placidus) during nesting. Tracking these females after fledging permitted us to locate juvenile birds, their parents, and any helpers present and to build individual fledgling resighting datasets without incurring mortality costs or causing premature fledging due to handling or transmitter effects. A Bayesian framework was used to infer age‐specific mortality rates in relation to group size, fledging date, maternal condition, and nestling condition. Postfledging survival was positively related to group size, with fledglings raised in groups with four helpers showing nearly 30% higher survival until independence compared with pair‐only offspring, independent of fledging date, maternal condition or nestling condition. Our results demonstrate the importance of studying the early dependency period just after fledging when assessing presumed benefits of cooperative breeding. While studying small, mobile organisms after they leave the nest remains highly challenging, we argue that the telemetric approach proposed here may be a broadly applicable method to obtain unbiased estimates of postfledging survival.     

COOPERATIVE BREEDING OF THE NORTHWESTERN CROW CORVUS CAURINUS IN BRITISH COLUMBIA

The behaviour of helpers at nests of Northwestern Crows was studied on Mandarte Island and Mitlenatch Island, British Columbia. Not all nests had a helper and there was only one helper per nest. Helpers participated in varying degrees in the defence of the territory and nest, feeding of the nestlings and fledglings and they cached food on the territory. Adult males fed helpers, and helpers obtained most of their food on the adults' territory. Adults with helpers laid larger clutches and produced more fledglings per nest than adults without helpers. It is suggested that cooperative breeding in the Northwestern Crow is of recent origin.     

Extinction and endemism in the New Zealand avifauna

Aim Species belonging to higher taxa endemic to islands are more likely to go extinct following human arrival. This selectivity may occur because more highly endemic island species possess features that make them uniquely vulnerable to impacts associated with human arrival, specifically: (1) restricted distribution (2) reduced predator escape response, including loss of flight, and (3) life history traits, such as large body mass, associated with greater susceptibility to hunting or habitat loss. This study aims to identify which of these features can explain the selective extinction of more highly endemic bird species in New Zealand. Location North and South Island, New Zealand. Methods Bird species breeding in New Zealand prior to human arrival were classified according to whether they became extinct or not during two periods of human settlement, prehistoric (post‐Maori but pre‐European arrival) and historic (post‐European arrival). We modelled the relationships between extinction probability, level of endemism and life history traits in both periods. Results The prehistoric extinction–endemism relationship can be explained entirely by the selective extinction of large‐bodied species, whereas the historic extinction–endemism relationship appears due to increased susceptibility to introduced predators resulting from the loss of predator escape responses, including loss of flight. Conclusions These features may explain extinction–endemism relationships more generally, given that human hunting and predator introductions are major impacts associated with human arrival on islands.     

Do Parents and Helpers Discriminate between Related and Unrelated Nestlings in the Cooperative Breeding Silver‐Throated Tit?

When benefits exceed costs, natural selection may favor adults that develop the ability to recognize and preferentially direct care toward their own offspring to maximize their fitness. Investigations into the ability of adults to recognize offspring in offspring's early development period may help to understand when the ability of kin recognition starts to develop. In birds, studies of offspring recognition have mainly been conducted on bi‐parental breeding species, but relatively seldom on cooperative breeding species, despite that kin recognition may be of particular importance for cooperative breeders. The silver‐throated tit Aegithalos glaucogularis is a small passerine in which some nests have helpers during breeding. We tested whether silver‐throated tit parents and helpers were able to distinguish between their own and alien nestlings 2–5 d before fledging when recognition mechanisms were likely to have been developed. Through two forced choice experiments, of which one was conducted right beside the experimental nests (<0.8 m) and the other far away from the experimental nests (~6 m), we found that neither parents nor helpers discriminatively fed their own and alien nestlings, which suggested that at least during the experimental nestling age, and within the 6‐m‐radius area around the nests, they might not have the ability to recognize offspring. The possibility that silver‐throated tits use a larger area (>6 m radius) around their nests as a location‐based cue for offspring recognition, or would develop an offspring recognition ability at an older nestling age and/or even after fledging, warrants future studies.     

Effects of Supplementary Feeding on the Breeding Ecology of the Buff-Throated Partridge in a Tibetan Sacred Site,China

Our goal was to document effects of year-round supplemental feeding on breeding ecology of the Buff-throated Partridge, Tetraophasis szechenyii, within a Tibetan sacred site. We evaluated effects of supplemental feeding used as religious/cultural practices which could potentially aid conservation of endangered phasianids. We compared fed breeding groups to neighboring nonfed groups. Fed groups initiated first clutches significantly earlier than nonfed groups. Earlier laying groups within fed and nonfed groups showed significantly lower hatching rates than later groups; however, fed groups showed significantly higher hatching rates than nonfed groups laying in the same period. Earlier laying increased opportunities to renest. All six fed groups with clutch failures renested compared to only one of five nonfed groups with clutch failures. Fed female breeders showed significantly greater investment in their young with larger clutches and larger eggs, which likely increased survivability of early hatchlings. We observed no predation on birds at feeding sites and recorded only four cases of predation on incubating females, which showed no detectable difference between fed and nonfed groups. Ground-nesting birds typically face high risks of predation. Ten of the 48 groups nested in trees, which occurs in few phasianid species. Tree nests showed significantly higher hatching rates compared to ground nests; however, we found no significant difference in tree nesting between fed and nonfed groups. This partridge is one of four gallinaceous species with cooperative breeding. Breeding groups with helpers had significantly greater reproductive success than single pairs, and fed female breeders with helpers laid bigger eggs than single pairs. Comparing annual reproductive output per group, fed groups not only produced significantly more independent young (≥150 days post-hatching), their young hatched significantly earlier, which likely have greater reproductive value over later hatched young of nonfed groups. Supplemental feeding year-round is likely what enabled the successes of the fed partridges.