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Kleptoparasitism of a Red-cockaded Woodpecker (Picoides borealis) Nest Cavity by a Red-bellied Woodpecker (Melanerpes carolinus)
David K. Delaney and Lawrence D. Carlile

Southeastern Naturalist, Volume 9, Issue 3 (2010): 624–628

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Kleptoparasitism of a Red-cockaded Woodpecker (Picoides borealis) Nest Cavity by a Red-bellied Woodpecker (Melanerpes carolinus) David K. Delaney1,* and Lawrence D. Carlile 2 Abstract - We report the kleptoparasitism of a Picoides borealis (Red-cockaded Woodpecker) cavity by a Melanerpes carolinus (Red-bellied Woodpecker). We believe this is the first video documenting kleptoparasitism of a Red-cockaded Woodpecker nest by a male Red-bellied Woodpecker in which both a nestling and an attending adult Red-cockaded Woodpecker adult were forcibly ejected. The Red-bellied Woodpecker was resolute in its attempt to usurp the nest, reaching into the cavity over 2000 times in an attempt to evict the cavity occupants. The male Red-bellied Woodpecker and his mate took over the cavity soon after. Adult Red-cockaded Woodpeckers continued to visit their nest cavity during diurnal hours for 2 days post-kleptoparasitism. It is important that resource managers incorporate proactive management techniques to lessen the impact of interspecific competition for cavities, especially in smaller or fragmented Red-cockaded Woodpecker populations. Many animals are known to use cavities excavated by Picoides borealis Vieillot (Red-cockaded Woodpecker, hereafter RCWO) (Dennis 1971, Everhart 1986, Jackson 1978). Availability of nest cavities within southern pine forests is a limiting factor for RCWOs and can lead to intense competition (USFWS 2003). Melanerpes carolinus L. (Red-bellied Woodpecker, hereafter RBWO) is considered one of the primary competitors for RCWO cavities (Jackson 1978). Usurpation of roost and nest cavities is known to negatively impact RCWOs (Jackson 1978, Kappes 1997, Kappes and Harris 1995, Ligon 1970). Kappes (1997) suggested that because usurpation is beneficial for one species over another, it is more aptly described as cavity kleptoparasitism. We use this term to describe an instance when a RBWO usurps a cavity from a RCWO by force. Kleptoparasitism (Kappes 1997) of RCWO cavities is known to occur, but has rarely been documented directly by visual observation or video surveillance. Increased activity at nest cavities (such as provisioning of nestlings) may provide visual and aural cues for cavity kleptoparasites (Conner et al. 1999). Only Ligon (1971) and Jackson (1978) have previously documented RBWOs forcibly extracting RCWOs from roost or nest cavities. Ligon (1971) observed an adult RBWO usurp a roost cavity from an adult male RCWO, while Jackson (1978) reported seeing a male RBWO remove a young RCWO from its nest. Study site. We conducted this study from 1998–2000 at Fort Stewart (31.88ºN, 81.57ºW) in southeastern Georgia. Video cameras and time-lapse recorders were used to record over 15,000 hrs of nesting behavior in 38 color-banded RCWO groups (Delaney et al., in press). The size of the video image at cluster 267, where we documented the cavity kleptoparasitism was approximately 120 × 150 cm (height versus length), and covered about 90 × 45 cm (height versus length) of the tree bole, and was centered on the nest cavity. The nest was videotaped between 04:55–20:00 (all times in EST) each day. Pre-kleptoparasitism woodpecker behavior. On 28 June 2000, at 08:41:03, we videotaped an adult male RBWO land on the lip of the nest cavity in RCWO Notes of the Southeastern Nat u ral ist, Issue 9/3, 2010 624 1US Army Construction Engineering Research Laboratory, PO Box 9005, Champaign, IL 61826. 2Directorate of Public Works, Environmental Division, Fish and Wildlife Branch, 1177 Frank Cochran Drive, Fort Stewart, GA 31314. *Corresponding author - David.Delaney@erdc. 2010 Southeastern Naturalist Notes 625 cluster 267, which contained a 24-day-old RCWO nestling. The RBWO attempted to extract the nestling by repeatedly inserting its head into the cavity. This activity lasted until 8:42:05, when an adult RCWO made 2 flights close to the RBWO’s position, possibly causing the potential kleptoparasite to leave. Immediately afterward, an adult RCWO entered the nest cavity at 8:42:09, apparently to defend the nest. Presumably the same adult male RBWO landed on the lip of the nest cavity at 8:42:11 and resumed attempting to remove the occupants of the cavity. A second adult RCWO landed twice on the tree near the nest and flew closely past the nest 3 additional times while the first RCWO was in the nest cavity, but it never directly confronted the RBWO. The RBWO inserted its head and body into the cavity every 4.1 seconds, on average, for a total of 945 times over 1.04 hrs, in an apparent attempt to kleptoparasitize the nest. The RBWO was unsuccessful in his attempt to evict the RCWOs and left the RCWO’s nest cavity at 9:46:13. Kleptoparasitism event. At 08:52:42 on 29 June, we videotaped presumably the same adult male RBWO from the previous day landing on the lip of the nest at cluster 267. The RBWO attempted to extract the cavity occupants again by repeatedly inserting its head and body into the cavity every 6.4 seconds, on average, for a total of 712 times over 45.3 minutes until it successfully grasped the bill of the nestling and extracted it from the nest at 09:37:58 and released it. The RBWO then made 384 additional attempts (1 attempt every 4.8 seconds) to grasp the adult RCWO over the next 18.2 minutes and extracted the adult from the nest at 09:56:20, though this time both adult woodpeckers fell towards the ground and out of camera view. Post-kleptoparasitism woodpecker behavior. We recorded 43.5 hours of postkleptoparasitism behavior by RBWOs and RCWOs at cluster 267 from 9:56:21 on 29 June until 5:26:00 on 1 July. A male RBWO returned to the kleptoparasitized cavity 8 seconds after ejecting the adult RCWO. This was the first of 54 documented visits by RBWOs to the cavity (46 visits when only 1 RBWO was present and 8 visits when the pair was present) after the event. The majority of the 46 individual visits to the cavity were to enlarge the cavity entrance (72%), and primarily were made by the female (64%). Only the male RBWO entered the cavity to make modifications and used it as a roost for 2 nights. Video surveillance ended before we were able to determine if the RBWO pair attempted to nest. Discussion. RCWOs were observed visiting the nest cavity 7 times to maintain resin wells following the kleptoparasitism event. We did not observe any direct confrontations between woodpecker species post-kleptoparasitism, and the cavity was still occupied by RBWOs when video surveillance ended. The adult and nestling RCWOs ejected by the male RBWO were observed in good condition 2 days after the cavity kleptoparasitism occurred. The nestling was found roosting near its sibling that fledged at 6:22:02 on 29 June 2000. Cavity kleptoparasitism can impact RCWOs by: 1) preventing a RCWO group from breeding by usurping the roost cavity (Jackson 1978); 2) reducing reproductive success through partial (Ligon 1970) or whole brood loss (Jackson 1978); 3) reducing the number of woodpecker groups through impacts on reproduction (Kappes 2008); 4) restricting cavity availability (Carrie et al. 1998); 5) enlarging cavities, thereby making them unsuitable or increasing chances of predation (Saenz et al. 1998); 6) increasing intragroup competition (Kappes 2004); 7) limiting retention of breeding females (Daniels and Walters 2000); 8) increasing interspecific competition for cavities (Kappes 1997); 9) increasing predation risk when RCWOs are forced to roost in the open (Rudolph et al. 1990); 10) increasing the potential to injure or kill adult woodpeckers while usurping the cavity (Ligon 1970, Neal et al. 1992), and 11) reducing the population (Baker 1983). 626 Southeastern Naturalist Notes Vol. 9, No. 3 Few researchers have described how RCWOs respond to cavity kleptoparasitism, though it appears that species like RBWOs regularly usurp active RCWO cavities throughout the year (Jackson 1978, Kappes 2004, Kappes and Harris 1995, Ligon 1970), especially in cavity-limited habitats (Kappes 2004). Data are limited to determine if this behavior is more prominent in the breeding or non-breeding season. Most of the available literature describes how RCWOs respond to intraspecific territorial encounters or interspecific interactions (Jackson 1978, Ligon 1971, Wood 1983), but none have fully characterized cavity usurpation by describing woodpecker response during and after such an event. It has been suggested that interspecific competition for cavities destroys more nests than predation (LaBranche and Walters 1994). We only observed the forced premature fledging of an RCWO nestling. The RBWO did not actually destroy the nest as part of this interspecific competition for the cavity. RBWOs have been documented as aggressive cavity kleptoparasites (Baker and Payne 1993, Jackson 1978, Ligon 1971). The male RBWO we videotaped spent 2.06 hrs during 3 different periods over 2 days attempting to kleptoparasitize a RCWO nest before succeeding. The stationary behavior at the cavity lip for periods of 18–60 minutes and the large number of reaching motions per visit inside the cavity entrance to eject the RCWOs, clearly shows persistence by the RBWO to usurp the cavity. Ligon (1971) observed a RBWO eject an adult male RCWO from his roost, while Jackson (1978) observed a RCWO nestling removed from its nest. This is the only study to observe a RBWO eject both a nestling and attending adult RCWO from an active nest. RCWOs are known to defend their cavities from a variety of avian species, such as Dendrocopos villosus L. (Hairy Woodpecker), Sialia sialis L. (Eastern Bluebird), and RBWO (Ligon 1970). Nest defense from competitors like RBWOs appears to be part of the daily activities of RCWOs (Jackson 1978) and can be detrimental (Ligon 1971, Neal et al. 1992). We documented minimal outside cavity defense (i.e., close flights) against the RBWOs. An adult RCWO made 2 close passes by a RBWO, which allowed an adult RCWO to enter and defend the cavity from the inside. Most of the defensive behavior we videotaped occurred inside the cavity by adult RCWOs directed at the RBWO. We did not detect any close diving passes at the RBWO once an adult was on the nest to defend it from the inside. Interestingly, most of the RCWO activity near the usurped nest cavity occurred when adult RCWOs were observed conducting resin well maintenance. Aggressive defense of nest cavities previously has been shown by RCWOs towards RBWOs during attempts to usurp their nest (Ligon 1970). We only documented one instance of cavity kleptoparasitism of a RCWO nest by RBWOs among 38 clusters (over 15,000 hrs of video surveillance) that were intensively monitored as part of a larger project studying woodpecker response to military training operations (Delaney et al., in press). It appears that habitat quality, cavity availability, and population size strongly influence the level of interspecific competition for cavities and the impact of kleptoparasites within RCWO habitat (Neal et al. 1992). For large, healthy populations of RCWOs, such as Fort Stewart’s, cavity kleptoparasitism may not be a limiting factor in the short term when proactive management techniques are employed, such as periodic prescribed fires and provisioning of artificial cavities and drilled starts. Smaller or more fragmented RCWO populations may be impacted more by kleptoparasites. It is important to note that artificial cavities may only be a temporary fix to the problem of cavity availability; more long-term solutions for maintaining open mature forests through periodic prescribed fires and maturation of current pine forests are the ultimate goals (Neal et al. 1992, 2010 Southeastern Naturalist Notes 627 USFWS 2003). It is important that resource managers understand the impact that kleptoparasites can have on RCWO populations and attempt to incorporate proactive management techniques (e.g., frequent periodic prescribed fires, provisioning of artificial cavities and drilled starts) to lessen the impact of cavity kleptoparasitism, especially in smaller or fragmented RCWO populations. Acknowledgments. This study was supported by US Army Forces Command and Fort Stewart Army Installation with funding from the Strategic Environmental Research and Development Program, under Conservation Project No. CS-1083 and the Construction Engineering Research Laboratory, which is part of the Engineer Research and Development Center for the US Army Corps of Engineers. The Fish and Wildlife Branch at Fort Stewart provided logistical support and conducted most RCWO nest surveys. We thank T. Brewton, H. Erickson, M. Fay, T. Hasty, M. Huffman, M. Klich, S. Kovac, B. Platt, A. Rinker, and A. Walde for assisting with data collection. We thank T. Brewton, T. Grubb, and A. Walde for assisting in placing video cameras and A. Cone, B. MacAllister, L. Nguyen, C. Smith, and A. Walde for assisting in reviewing videotapes. We thank T. Hayden for his comments and suggestions on an earlier draft of this manuscript. We also thank two anonymous reviewers for their helpful comments. Literature Cited Baker, J.L. 1983. Decline and extirpation of a population of Red-cockaded Woodpeckers in northwest Florida. Pp. 41–43, In D. A. Wood (Ed.). Red-cockaded Woodpecker Symposium II. Florida Game and Freshwater Fish Commission, Tallahassee, fl. Baker, J.L., and R.L. Payne. 1993. Nest usurpation of a Starling nest by a pair of Red-bellied Woodpeckers. Florida Field Naturalist 21:33–34. Carrie, N.R., K.R. Moore, S.A. Stephens, and E.L. Keith. 1998. Influences of cavity availability on Red-cockaded Woodpecker group size. Wilson Bulletin 110:93–99. Conner, R.N., D.C. Rudolph, R.R. Schaefer, D. 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Sequential occupation of cavities by Redcockaded Woodpeckers and Red-bellied Woodpeckers in the Ouachita National Forest. Proceedings of the Arkansas Academy of Science 46:106–108. Rudolph, D.C., H. Kyle, and R.N. Conner. 1990. Red-cockaded Woodpecker vs. Rat Snakes: The effectiveness of the resin barrier. Wilson Bulletin 102:14–22. Saenz, D., R.N. Conner, C.E. Shackelford, and D.C. Rudolph. 1998. Pileated Woodpecker damage to Red-cockaded Woodpecker cavity trees in eastern Texas. Wilson Bulletin 110:362–367. Wood, D.A. 1983. Observations on the behavior and breeding biology of the Red-cockaded Woodpecker in Oklahoma. Pp. 92–93, In D.A. Wood (Ed.). Red-cockaded Woodpecker Symposium II. Florida Game and Freshwater Fish Commission. Tallahassee, fl. US Fish and Wildlife Service (USFWS). 2003. Recovery plan for the Red-cockaded Woodpecker (Picoides borealis): Second revision. Atlanta, GA. 296 pp.