DOI: 10.5061/dryad.3bk3j9kqs
title: Long-term demographic data on the red-cockaded woodpecker

description: In cooperative breeding systems, inclusive fitness theory predicts that
 non-breeding helpers more closely related to the breeders should be more
 willing to provide costly alloparental care, and thus have more impact on
 breeder fitness. In the red-cockaded woodpecker (Dryobates borealis), most
 helpers are the breeders’ earlier offspring, but helpers do vary within
 groups in both relatedness to the breeders (some even being unrelated) and
 sex, and it can be difficult to parse their separate impacts on breeder
 fitness. Moreover, most support for inclusive fitness theory has
 been positive associations between relatedness and behavior, rather than
 actual fitness consequences. We used functional linear models to evaluate
 the per capita effects of helpers of different relatedness on eight
 breeder fitness components measured for up to 41 years at three sites. In
 support of inclusive fitness theory, helpers more related to the breeding
 pair made greater contributions to six fitness components. However, male
 helpers made equal contributions to increasing pre-fledging survival
 regardless of relatedness. These findings suggest that both inclusive
 fitness benefits and other, direct benefits may underlie helping behaviors
 in the red-cockaded woodpecker. Our results also demonstrate the
 application of an underused statistical approach to disentangle a complex
 ecological phenomenon.
description: We used long-term demographic monitoring data collected over 28
 to 41 consecutive years at three sites: the Sandhills region in
 south-central North Carolina (1980–2020), Marine Corps Base Camp Lejeune
 on the central coast of North Carolina (1986–2020), and Eglin Air Force
 Base in the western panhandle of Florida (1993–2020). Monitoring methods
 are described in detail by Walters et al. (1988) (see also Appendix A for
 more details on monitoring). See Walters and Garcia (2016) for how
 individuals are assigned breeder and helper status.
description: You will need both R and RStudio to use the dataset (and
 corresponding code). 
description: *Manuscript citation*: Kerr, William, and Walters (2023) Inclusive fitness
 may explain some but not all benefits derived from social behavior in a
 cooperatively breeding bird. American Naturalist. *Archive citation*:
 Kerr, Natalie; Morris, William; Walters, Jeffrey (Forthcoming 2023).
 Demographic data on the red-cockaded woodpecker [Dataset]. Dryad.
 https://doi.org/10.5061/dryad.3bk3j9kqs *Affiliated authors:* Natalie Z.
 Kerr, William F. Morris, and Jeffrey R. Walters *Corresponding author
 details:* * Email: natalie.kerr@duke.edu, * Location: Department of
 Biology, Duke University, Durham, NC 27705 To run the code file
 ("Kerr-et-al\_FLMs.rmd"), you will need to install R and
 RStudio, as well as install the packages (using install.packages()) listed
 below and at the beginning of the RMarkdown file. *List of packages and
 their versions* * mgcv (Version 1.8.42) * bbmle (Version 1.0.25) Note that
 we used these versions of these two packages for Kerr et al. 2023. The
 RMarkdown file and datafiles need to be contained within the same folder
 for the code to run. To run our FLM analyses, we archived two datasets:
 reproductive data (for the five reproductive fitness components) and the
 survival data (for the three survival fitness components). Both are part
 of the long term demographic monitoring. For the reproductive data
 (rep\_dat), each row is a group in a particular year. For the survival
 data (surv\_dat), each row is a particular individual (that is either a
 nestling N and breeder B). List of variables present in both datasets: *
 Pop - Three populations or "sites" (i.e., Bragg, Camp Lejeune,
 and Eglin) * Territory - Unique number ID for each territory *
 TerritoryByManag - Unique number ID for territory x management period *
 Year - Year data was collected * mother.age - Age of the mother (in years)
 for that territory in that particular year * father.age - Age of the
 father (in years) for that territory in that particular year * MR - For
 the rds file, MR is a matrix where each column represents a summed
 relatedness value for male helpers, rows represent unit of observation
 which is group for rep\_dat and individual birds for surv\_dat. Note that
 the columns of the MR matrix are populated into individual columns of the
 data frame for the csv file. * MN - For the rds file, MN is a
 corresponding matrix for MR that represents the number of male helpers of
 that particular summer relatedness value present in the group. Note that
 the columns of the MN matrix are populated into individual columns for the
 csv file. * FR - Similar to MR, the rds file contains a FR matrix where
 each column represents a summed relatedness value for female helpers, rows
 represent unit of observation which is group for rep\_dat and individual
 birds for surv\_dat. Note that the columns of the MR matrix are populated
 into individual columns of the data frame for the csv file. * FN - Similar
 to MN, the rds file contains a corresponding FN matrix for FR that
 represents the number of female helpers of that particular summer
 relatedness value present in the group. Note that the columns of the FN
 matrix are populated into individual columns for the csv file. List of
 variables present in the reproductive dataset: * firstnestattempt - Binary
 variable representing whether the breeding pair attempted a first nest
 (i.e., 0 or 1) * firsteggattempt - Number of eggs in the first clutch *
 eggsuccess - Binary variable representing whether one or more eggs
 successfully hatched in the first nest (i.e., 0 or 1) * nestlingsuccess -
 Binary variable representing whether one or more nestlings successfully
 fledged in the first nest (i.e., 0 or 1) * firstfledgling - Number of
 fledglings in the first nest List of variables in the survival dataset: *
 sex - Sex of the individual (i.e., F for female and M for male) * age -
 Age of the individual (in years) * status - Status of the individual bird
 (i.e., J for juvenile and B for breeder) * surtonext - Binary variable
 representing whether the bird survived to the next year ("1") or
 did not ("0") Note that columns with cells containing
 "NA" values are groups or individuals without data for that
 associated column. Please reach out to Natalie Z. Kerr, for any questions
 regarding these data and code.