Is helping a beneficial learning experience for red-cockaded woodpecker (Picoides borealis) helpers?

Several hypotheses have been suggested to explain the evolution and maintenance of helping behavior in cooperatively breeding birds, one of which we investigated in the red-cockaded woodpecker (Picoides borealis). Helping may provide a learning experience that improves reproductive success once the helper becomes a breeder. We used data collected from a population of red-cockaded woodpeckers in the Sandhills of North Carolina to compare the reproductive success of 2-year-old breeders that were helpers at age 1 (helping experience) to those that were floaters or solitary males at age 1 (no helping experience). Reproductive success of the two groups was similar, indicating that helping provides no experience useful in reproduction. We reject the learning experience hypothesis as an explanation for the maintenance of helping behavior in red-cockaded woodpeckers. Received: 29 July 1996 / Accepted after revision: 27 April 1997     

Genetic evidence for monogamy in the cooperatively breeding red-cockaded woodpecker

We examined the genetic relationship among putative parents, offspring, and helpers in 224 red-cockaded woodpeckers (Picoides borealis) from the Sandhills of North Carolina. Comparison of DNA similarity with a pedigree constructed from 3,823 individually-marked birds observed from 1979 to 1992 provided verification of observed relatedness in the sampled population (R ² = 0.87,df = 14,P<0.001). In this population, breeding pairs may or may not have helpers, most of which are males that remain on their natal territory. Our sample included helpers assisting their parents, helpers assisting a related male and an unrelated female, and helpers assisting an unrelated pair. Band by band comparison and examination of similarity among DNA profiles indicated that all offspring from non-helped nests were sired by their putative parents (n = 28 families). Similarly, all but one offspring in helped nests were also sired by their putative parents (n = 16 families). In the exceptional case, the offspring evidently was sired by a male external to the group. Analysis of similarity values supported the conclusion that matings by helpers or extra-group males are rare or non-existent. Our results indicate that in this species advantages gained by individuals remaining on their natal territories as helpers do not generally include siring offspring. Correspondence to: S. Haig     

Optimal Allocation of Conservation Resources to Species That May be Extinct

Abstract: Statements of extinction will always be uncertain because of imperfect detection of species in the wild. Two errors can be made when declaring a species extinct. Extinction can be declared prematurely, with a resulting loss of protection and management intervention. Alternatively, limited conservation resources can be wasted attempting to protect a species that no longer exists. Rather than setting an arbitrary level of certainty at which to declare extinction, we argue that the decision must trade off the expected costs of both errors. Optimal decisions depend on the cost of continued intervention, the probability the species is extant, and the estimated value of management (the benefit of management times the value of the species). We illustrated our approach with three examples: the Dodo (Raphus cucullatus), the Ivory‐billed Woodpecker (U.S. subspecies Campephilus principalis principalis), and the mountain pygmy‐possum (Burramys parvus). The dodo was extremely unlikely to be extant, so managing and monitoring for it today would not be cost‐effective unless the value of management was extremely high. The probability the Ivory‐billed woodpecker is extant depended on whether recent controversial sightings were accepted. Without the recent controversial sightings, it was optimal to declare extinction of the species in 1965 at the latest. Accepting the recent controversial sightings, it was optimal to continue monitoring and managing until 2032 at the latest. The mountain pygmy‐possum is currently extant, with a rapidly declining sighting rate. It was optimal to conduct as many as 66 surveys without sighting before declaring the species extinct. The probability of persistence remained high even after many surveys without sighting because it was difficult to determine whether the species was extinct or undetected. If the value of management is high enough, continued intervention can be cost‐effective even if the species is likely to be extinct.     

Do stress hormones suppress helper reproduction in the cooperatively breeding red-cockaded woodpecker (<Emphasis Type="Italic">Picoides borealis</Emphasis>)?

In the cooperatively breeding red-cockaded woodpecker (Picoides borealis), male helpers are subordinate to male breeders and do not mate with females, even when unrelated to the breeding female within their group or through extra-group matings, yet exhibit reproductive hormone profiles similar to those of breeders. We investigated whether reproduction might be suppressed in helper males via high levels of the stress hormone corticosterone. We also examined effects of group size and season on corticosterone levels by comparing baseline and maximal plasma levels of corticosterone between helper males and breeding males, and among helper males and breeders of both sexes living in groups of different sizes throughout the reproductive cycle. We also measured plasma levels of estrogen, progesterone, and testosterone to examine other potential hormonal differences between helpers and breeders. Male status did not explain variation in any hormones; therefore, our data do not support the hypothesis that helper males are reproductively suppressed via corticosterone or the other hormones investigated. However, the presence of two or more helper males in a group tended to reduce baseline corticosterone in breeding and helper males, but not breeding females, suggesting that helper males reduce parental effort of other male group members. Seasonally, maximal corticosterone peaked during the nestling provisioning phase for breeding and helper males, but not breeder females, suggesting that males show an increased response to stressors posing a potential threat to survival of offspring.     

An analysis of reciprocal exchange of helping behavior in the red-cockaded woodpecker (Picoides borealis)

We examine the frequency of reciprocal exchanges of helping behavior in three red-cockaded woodpecker populations to determine if such exchanges might constitute a significant fitness benefit of helping. Specifically, we determine how often helpers, once they become breeders, are assisted by young they previously helped raise. The estimated frequencies of reciprocal exchange of helping based on survival and status transition probabilities are low (2%). Observed frequencies of reciprocity are significantly higher than estimated frequencies in two of the three populations studied, suggesting that male fledglings more often remain on their natal territory as helpers if another helper is already present. High rates of retention of young males as helpers on high-quality territories or preferential helping of kin, as well as preferential helping of former care-givers, might explain this result. When the analysis is restricted to helpers unrelated to the young they help raise, which controls for preferential helping of kin and largely eliminates effects of territory quality, empirical estimates of the frequency of reciprocity do not differ from estimates calculated from population demography. We conclude that young males do not preferentially help former care-givers and that reciprocal exchange of help is not an important factor in the evolution of helping behavior in the red-cockaded woodpecker. In this species, all known benefits of helping behavior are indirect (i.e., derived through kin selection ) rather than direct. We discuss the occurrence of unrelated helpers in this context. Received: 10 August 1999 / Received in revised form: 13 February 2000 / Accepted: 19 February 2000     

Genetic evidence for monogamy in the cooperatively breeding red-cockaded woodpecker

We examined the genetic relationship among putative parents, offspring, and helpers in 224 red-cock-aded woodpeckers (Picoides borealis) from the Sandhills of North Carolina. Comparison of DNA similarity with a pedigree constructed from 3,823 individually-marked birds observed from 1979 to 1992 provided verification of observed relatedness in the sampled population (R ² = 0.87, df = 14, P < 0.001). In this population, breeding pairs may or may not have helpers, most of which are males that remain on their natal territory. Our sample included helpers assisting their parents, helpers assisting a related male and an unrelated female, and helpers assisting an unrelated pair. Band by band comparison and examination of similarity among DNA profiles indicated that all offspring from non-helped nests were sired by their putative parents (n = 28 families). Similarly, all but one offspring in helped nests were also sired by their putative parents (n = 16 families). In the exceptional case, the offspring evidently was sired by a male external to the group. Analysis of similarity values supported the conclusion that matings by helpers or extra-group males are rare or non-existent. Our results indicate that in this species advantages gained by individuals remaining on their natal territories as helpers do not generally include siring offspring. Correspondence to: S. Haig     

Effects of helpers on breeder survival in the red-cockaded woodpecker (Picoides borealis)

Helpers can gain future indirect fitness benefits by increasing the survival of breeders that produce offspring related to the helper. Helping augments group size through the helper's presence and, in some cases, by increasing fledging success. Breeders may then experience enhanced survivorship because of the benefits of living in large groups. Helping may also reduce the workload of the breeder, which in turn may increase the likelihood that the breeder will survive to breed again. We used Cox's proportional hazards model to examine whether breeders' survival in two populations of the red-cockaded woodpecker (Picoides borealis) was enhanced when group size was increased in the presence of (1) the helper itself, or (2) "extra" fledglings (fledglings produced by the breeder because of helping behavior). We found that in the presence of helpers, the risk of a breeder dying declined by 21–42% for males and 0–14% for females. Our results suggest reduced breeder workload as one mechanism to explain reduced breeder mortality in the presence of helpers: breeders spent less time incubating and provisioning nestlings when assisted by helpers. The risk of a breeder dying declined by 16–42% in males and 26–43% in females in the presence of "extra" fledglings. We speculate on possible mechanisms by which fledglings might affect breeder survival. Our results support the hypothesis that helpers gain future indirect benefits by reducing breeder mortality. Electronic Publication