Behaviour and breeding biology of the cooperatively breeding Grey-backed Fiscal Shrike Lanius excubitorius in Kenya

Grey-backed Fiscal Shrikes Lanius excubitorius were studied over a 2j year period near Lake Naivasha, Kenya. Grey-backs are cooperative breeders, with group sizes ranging from two to II. Only one pair breeds per group, with all other group members aiding in the rearing of young. The study population ranged from 64 to 79 individuals that occurred in from 13 to 16 groups. Non-breeding helpers made up to 66% of the population, with male helpers being more numerous overall than females. The annual survival rate was 65%, with no differences detected between the survival of males and females, or of breeders and helpers. Only male helpers were observed to acquire breeding status within the natal territory. Some female helpers acquired breeding positions in territories adjacent to their natal territories. Group territorial displays occurred throughout the year but were most pronounced prior to breeding during rainy periods. Reproductive success was very low, with only 14.5% of the recorded breeding attempts leading to fledged young. Large groups (four or more birds) had greater reproductive success than small groups, but many factors other than, or in addition to, group size may have influenced this pattern. The breeding male contributed the most food to the incubating female and to the nestlings. Male helpers and the breeding female contributed more to nestlings than did female helpers. Observations on the post-fledging period indicate that socialization and establishment of dominance may be of importance in group-living species living in a restricted ecological and social setting.     

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.     

The Cooperative Breeding System of the Red-cockaded Woodpecker

The breeding system of the red-cockaded woodpecker is described based on data collected over six years from a population of 500 marked individuals in the Sandhills of North Carolina. Male-female pairs were the most common social unit (59%), but 30% of social units contained one or more adult helpers, and 11% consisted of solitary males. Helpers were almost exclusively male: 27% of males remained in their natal group as helpers for at least one year, whereas only four (1%) females did. Most breeding females remained as breeders in the same group from one year to the next (56%), but a surprising number (12%) moved to another group. Many movements were related to incest avoidance or mate death, but 39% involved deserting a mate, usually following successful reproduction. We suggest that females sometimes are forced from groups by immigrants or other group members. The median distance of movements by adult females was only 1.3 km. In contrast to females, no breeding males switched groups. Survival of both breeding (76%) and helper (80%) males was higher than that of breeding females (69%). Males exhibited two distinct life-history strategies. Some remained as helpers on their natal territory for one or more years, and became breeders by inheriting breeding status in the natal group (17% per year) or by replacing a deceased breeder in a nearby group (13% per year, median distance moved 1.0 km). Other males dispersed from their natal group permanently during their first year. Some of these males were floaters at age one year, others were solitary, and a few became helpers in a non-natal group, but many were breeders. In contrast to males that first functioned as helpers, those that dispersed after fledging moved long distances (median dispersal distance 4.5 km), longer even than dispersing female fledglings moved (median distance 3.2 km). The habitat saturation model of the evolution of cooperative breeding is based on selection between the two life-history strategies exhibited by male red-cockaded woodpeckers. The model therefore may be tested directly with this species. Another indication that this model is appropriate for this species is the existence of a resource (cavity trees) that might provide an ecological basis for habitat saturation.     

Paying to stay or paying to breed? Field evidence for direct benefits of helping behavior in a cooperatively breeding fish

Several hypotheses aim to explain the evolution of helping behavior,but conclusive experimental support for evaluating the relativeimportance of individual hypotheses is still lacking. We reporton two field experiments conducted to test the "territory inheritance"and "pay-to-stay" hypotheses in the cooperatively breeding cichlidfish Neolamprologus pulcher The territory inheritance hypothesiswas tested by removing one parent, which created breeding vacancies.In 39% of cases, same-sex helpers took over the breeding spot;in 44% of cases helpers continued helping new breeders, and17% were evicted by new breeders. Helpers that were closelysize matched to the removed breeder had a better chance of gainingthe breeding spot Male helpers tended to continue helping aftera takeover more often than females.The pay-to-stay hypothesiswas tested-by temporarily removing helpers. Whereas breedersdid not respond aggressively to removals, other group membersattacked the removed helpers on their return, and 29% were eventuallyevicted. The returning helpers assisted more by increasing theirrate of territory maintenance and defense and visiting the broodchamber more frequently Size and sex of removed helpers didnot explain the observed aggressive reactions of other groupmembers. Thus, our results support both hypotheses: N. pulcherneeds to pay with help to be allowed to remain protected inthe family group, and there they may inherit the natal territory.N. pulcher helpers gain direct benefits from helping behavior.     

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.     

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.     

Social organization and communal breeding in the chestnut-bellied starling (Spreo pulcher)

Chestnut-bellied starlings (Spreo pulcher) live in social groups of 10–30 individuals and during their breeding seasons maintain group ranges which show little overlap with neighbouring groups. A social group may contain two to six breeding pairs, non-breeding adults of both sexes, and immatures. Each breeding female has her own nest and she alone incubates. The parents and up to 12 other starlings feed the nestlings. Individual helpers may successively or simultaneously attend the nests of different breeders. The percentage of nests attended within a group differs for helpers of different sex, age and breeding status: immatures of 1–2 years and non-breeding adult males help most and adult females least. Nests with more helpers have higher fledging success than those with fewer helpers. These results are discussed with reference to the tentative benefits of helping behaviour and kinship relationships within the social group.     

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.     

Helpers increase long-term but not short-term productivity in cooperatively breeding long-tailed tits

Helpers at the nest in the cooperative breeding system of long-tailedtits Aegithalos caudatus exhibit kin preference in their helpingbehavior. The aim of this study was to use multivariate analysesto investigate whether helpers accrue indirect fitness benefitsthrough their cooperation by increasing the productivity ofrelatives. All birds started each season breeding independentlyin pairs, but birds that failed in their own breeding attemptoften redirected their care to help another pair provision theiroffspring. About half of all broods had one or more helpers,86% of which were male. Provisioning rates increased and therewas a corresponding increase in the mass of nestlings withinbroods as the number of helpers increased. Helpers had no significantshort-term effect on productivity because nest predation, nestlingsurvival, and brood size were unaffected by the presence ofhelpers. However, in the long term helpers had a highly significanteffect on the recruitment of fledglings, the positive effectof helpers being linear within the range of helper numbers thatwe observed. We found no evidence to suggest that these resultswere confounded by the effects of individual or habitat quality.We conclude that long-tailed tits accrue indirect fitness benefitsby helping kin. Nevertheless, the inclusive fitness benefitfrom helping is substantially lower than that of independentbreeding, showing that helpers are making the best of a badjob.     

Cichlids do not adjust reproductive skew to the availability of independent breeding options

Helpers in cooperatively breeding species forego all or partof their reproduction when remaining at home and assisting breedersto raise offspring. Different models of reproductive skew generatealternative predictions about the share of reproduction unrelatedsubordinates will get depending on the degree of ecologicalconstraints. Concession models predict a larger share when independentbreeding options are good, whereas restraint and tug-of-warmodels predict no effects on reproductive skew. We tested thesepredictions by determining the share of reproduction by unrelatedmale and female helpers in the Lake Tanganyika cichlid Neolamprologuspulcher depending on experimentally manipulated possibilitiesfor helper dispersal and independent breeding and dependingon helper size and sex. We created 32 breeding groups in thelaboratory, consisting of two breeders and two helpers each,where only the helpers had access to a nearby dispersal compartmentwith (treatment) or without (control) breeding substrate, usinga repeated measures design. We determined the paternity andmaternity of 1185 offspring from 47 broods using five to nineDNA microsatellite loci and found that: (1) helpers participatedin reproduction equally across the treatments, (2) large malehelpers were significantly more likely to reproduce than smallhelpers, and (3) male helpers engaged in significantly morereproduction than female helpers. Interestingly, in four broods,extragroup helper males had fertilized part of the brood. Nohelper evictions from the group after helper reproduction wereobserved. Our results suggest that tug-of-war models based oncompetition over reproduction within groups describe best thereproductive skew observed in our study system. Female breedersproduced larger clutches in the treatment compared to the controlsituation when the large helpers were males. This suggests thatmale breeder-male helper reproductive conflicts may be alleviatedby females producing larger clutches with helpers around.     

Social organization,demography and genetic mating system of a Tibetan cooperative breeder

Knowledge of cooperative breeding in birds from longitudinal studies is available only for a small proportion of species. This paper reports data from a 12‐year study on the Tibetan Ground Tit Pseudopodoces humilis. On average, 27.2% (range: 13.0–36.1%) of monogamous pairs in each year contained one (85.4%) or more (14.6%) male helpers, 83.7% of which were yearlings staying on natal territories. Most helpers (89.6%) helped once and then bred independently. Adults had male‐biased sex ratios, low annual survival rates (averaging 0.50) and shorter longevities (averaging 1.8 years) compared with low‐altitude avian cooperative breeders, suggesting that mate shortage promotes helping behaviour in this species. Incest occurred rarely (2.1% of pairs), probably because kin recognition occurs through year‐around living in family groups. There was a low level (3.1% of broods) of extra‐pair parentage, which could facilitate the maintenance of cooperative breeding.     

Southern Lapwing Vanellus chilensis cooperative helpers at nests are older siblings

Cooperative breeding is rare in shorebirds, and when found it is thought to be due to polygamous mating (cooperative polygamy). Here we describe the social structure of cooperatively breeding groups in Southern Lapwing Vanellus chilensis and test the prediction that offspring sex ratio is skewed towards the sex that helps. The social groups consisted of a breeding pair with one or two young (mostly males, 68%) from the previous breeding season, and offspring sex ratio was not skewed. Cooperative breeding in the Southern Lapwing is not the consequence of cooperative polygamy, but rather groups were formed by a mated pair and some of their offspring from previous nests as helpers.     

Dwarf Mongoose Helpers at the Den1

This field study investigated helper behaviour in the care of the young in a small social carnivore, the dwarf mongoose (Helogale parvula). The frequency with which individually marked mongooses, living in six packs, guarded young at the den (babysitting) and brought food to the young was recorded in all-day and morning watches. The principal helpers were the subordinate adult ♀♀ and the yearlings. In a pack of known kinship relationships, an immigrant ♀ did more babysitting than any of the other pack members, all of whom were related to the young. Possible advantages of the aid-giving behaviour to the helpers are discussed.     

Infant Carrying Behavior in Callitrichid Primates: Callithrix and Leontopithecus

Care of offspring by individuals other than the mother is ubiquitous in callitrichid primates. In spite of its widespread occurrence, however, there is considerable taxonomic variation in the timing and intensity of parental effort by breeding males and nonbreeding juvenile group members. These differences may be attributable to generic and specifies differences in the costs of reproduction for females or in ecological constraints on travel and foraging. We present data on patterns of infant carrying in social groups of two taxa of callitrichid primates (Callithrix and Leontopithecus) throughout the first 3 months of infant life. We evaluated patterns of care in small groups (two or fewer juvenile or subadult helpers) and in large groups (three or more helpers in addition to the breeding adult male and female). Group size had little effect on levels of maternal carrying effort in either marmosets or lion tamarins, and mothers ceased carrying infants by 3 months of age. Carrying efforts by fathers were significantly reduced in groups with many helpers relative to small groups. Helpers carried at consistent rates during the second and third months of infant life in Leontopithecus, while in Callithrix, carrying by helpers peaked during the second month. These results suggest that if helpers reduce energetic demands on lactating females, the mechanism by which helpers reduce these burdens is independent of maternal carrying effort.     

Flexible Helping Behaviour in the Azure-Winged Magpie

Helping to rear the offspring of others may be a way for younger birds to gain access to future reproduction especially when turnover of breeding opportunities is low. However, this explanation is not applicable to cases where adults also help, or when roles shift between helpers and breeders. Over a period of 6-yr, we studied a marked population of azure-winged Magpie (Cyanopica cyanus) breeding in a non-territorial, colonial system. Magpies bred in a highly flexible cooperative system, in which individuals helped at different stages of the breeding cycle, including nest building, feeding the incubating female and feeding the young and removing the faecal sacs. On average, 50% of hatched nests were assisted by helpers-at-the-nest, and nest success appeared to be positively related to the presence of helpers. Helpers were predominantly males. Although juveniles were more likely to help, both juvenile and adult birds helped. Individual birds behaved as helpers either as a first-option or after having attempted their own breeding (second-option helpers). An individual helper may assist more than one nest during the same breeding season and in different breeding seasons. Reversals between breeder and helper roles were common in both directions, within a breeding season and between years. Helping behaviour is an option for almost any member of the colony. Therefore, hypotheses related to the enhancement of future breeding opportunities for juveniles can be discarded as general explanation of helping in this species. Although the decision to help appeared to be influenced by proximal environmental conditions hindering successful breeding, the associated benefits of helping as opposed to simply recovering for future reproduction, especially for former breeders, deserve further study.     

Group structure,within-group conflict and reproductive tactics in cooperatively breeding Galápagos mockingbirds,Nesomimus parvulus

Reproductive conflict within groups can be an important feature of cooperative breeding systems, especially when more than one individual of a sex breeds within a social group. Relationships between group structure, dominance, within-group conflict and reproductive tactics of cooperatively breeding Galápagos mockingbirds were examined on Isla Genovesa. Territorial groups of 2–24 adults included up to three breeding females, with 42% of the groups containing more than one (plural groups); females in most plural groups nested separately. Territory size increased with group size, but the area available per pair in plural groups was smaller than in singular groups (groups with only one breeding pair). Most pairings were monogamous, and males usually outnumbered females; high-ranking males obtained mates more frequently than subordinate males. In 3 relatively dry years, but not in a wet El Niño year, subordinate pairs in plural groups fledged fewer young than dominant pairs or pairs breeding in singular groups. Interference by dominant breeders, often leading to abandonment of nests by subordinate pairs, appears to account for these differences: through nest disruption in drier years, dominant individuals may reduce the cost of sharing their territories and increase the chances of recruiting helpers. Dominant males in plural groups may also father young through extra-pair copulations with subordinate females. Despite costs imposed by within-group conflict, subordinate breeders have higher long-term reproductive success than birds that defer breeding. Plural group structure is maintained because unpredictable climatic variation favours opportunistic breeding by subordinates.     

Helpers in a cooperatively breeding cichlid stay and pay or disperse and breed, depending on ecological constraints

The theory of family-group dynamics predicts that group structure, helping behaviour and social interactions among group members should vary with the opportunities of subordinates to breed independently. We investigated experimentally whether unrelated mature helpers in the cooperatively breeding cichlid Neolamprologus pulcher reduce costly social and cooperative behaviour and choose to disperse and breed independently when offered vacant breeding sites. As predicted by the ecological constraints hypothesis,when breeding substrate was available, (i) helpers spent more time in dispersal areas and it was mainly large helpers that left the group to breed independently; (ii) all helpers invested less in costly submissive behaviours towards other group members and large helpers reduced help, supporting the 'pay-to-stay' hypothesis; and (iii) large helpers, particularly those that dispersed and bred, increased more in body mass in the treatment than those without breeding options, suggesting status-dependent strategic growth of helpers.We conclude that helpers of N. pulcher decide whether to stay and pay or disperse and breed in response to constraints on independent breeding.     

Reproductive roles in the cooperatively breeding acorn woodpecker: incest avoidance versus reproductive competition

Incest is rare in the cooperatively breeding acorn woodpecker (Melanerpes formicivorus) despite a polygynandrous mating system in which nearly all group members are close relatives. Here we test the relative importance of avoiding matings between close relatives (incest avoidance) and within-sex competition for breeding opportunities (reproductive competition) in determining the mating system of acorn woodpeckers by examining how reproductive roles change following breeding vacancies. In 83% of cases in which helpers of the same sex were present in the group, reproductive vacancies were resolved when new unrelated immigrants filled the vacancy to the exclusion of resident same-sex helpers, who generally emigrated or did not breed while they remained in the group. Helpers of the opposite sex, especially when male, were significantly more likely to remain in their natal group and in about half the cases inherited and bred following reproductive vacancies. This result was not explainable by reproductive competition, since the number of immigrants was often less than or equal to the number of same-sex helpers in the group. Apparent incest resulted in 5% of cases. The time required to resolve reproductive vacancies was significantly longer for groups with helpers of the same sex as the vacancy. These results confirm that both incest avoidance and reproductive competition are important factors determining reproductive roles within groups of this highly social species.     

Kin and social relationships in splendid fairy-wrens: recognition by song in a cooperative bird

Cooperatively breeding splendid fairy-wren,s Malurus splendens, were tested with songs recorded from individuals of known social and kin relationships. Both males and females sang and responded aggressively to songs of wrens from other social groups. Wrens responded similarly to songs of non-kin and songs of close kin in the absence of social familiarity with them. Breeding females responded more intensely to songs of helpers from other groups than to songs of helpers in their own group. The songs of male and female helpers elicited similar responses by breeding females. The response to other females may be associated with competition for breeding status and helpers. Two females sometimes breed in social groups with two older females; no interference was observed. Song may allow individuals to recognize other wrens in their group and to direct their behaviour towards non-dispersing relatives by location and social familiarity rather than by kinship identifiers.     

Experimental evidence for helper effects in a cooperatively breeding cichlid

Neolamprologus pulcher is a cooperatively breeding cichlid fish,in which helpers stay in their natal territory and help withbrood care, territory defense, and maintenance. In this studywe investigated helper effects by an experimental group sizereduction in the field. After this manipulation, focal helpersin reduced groups tended to feed less, and small helpers visitedthe breeding shelter significantly more often than same-sizedhelpers in control groups. No evidence was found that remaininghelpers compensated for the removed helpers by increasing territorydefense and maintenance behavior. Breeders, however, did showa lower defense rate, possibly caused by an increase in broodcare effort. Survival of fry was significantly lower in removalthan control groups, which provides the first experimental proofin a natural population of fish that brood care helpers do effectivelyhelp. The data suggest that in small, generally younger, helpers,kin selection may be an important evolutionary cause of cooperation.Large helpers, however, who are generally older and less relatedto the breeders than small helpers are suggested to pay to beallowed to stay in the territory by helping. All group membersbenefit from group augmentation.