Cooperative breeding by the Bald Eagle (Haliaeetus leucocephalus) on Vancouver Island, British Columbia

Cooperative breeding Bald Eagles - Dawe 35 Cooperative breeding by the Bald Eagle (Haliaeetus leucocephalus) on Vancouver Island, British Columbia Neil K. Dawe 609 Willow Street, Parksville, BC V9P 1A5; e-mail: nkdawe@shaw.ca. Abstract: An instance of cooperative breeding by the Bald Eagle (Haliaeetus leucocephalus) on Vancouver Island, British Columbia, is discussed including summary observations of the trio through two nesting seasons. Two other possible instances of cooperative breeding from British Columbia are also discussed. Key words: Bald Eagle, cooperative breeding, Haliaeetus leucocephalus, Vancouver Island Dawe, N.K. 201 6. Cooperative breeding by the Bald Eagle (Haliaeetus leucocephalus) on Vancouver Island, British Columbia. British Columbia Birds 26:35 40. First published online December 201 5. Introduction Cooperative breeding is a reproductive system in which more than a pair of individuals show parent-like behaviour towards young of a single nest or brood (Hatchwell and Komdeur 2000). A prerequisite for cooperative breeding is the evolution of delayed dispersal (Kimball et al. 2003), which may occur as the result of either life history traits (e. g. low adult mortality; Arnold and Owens 1 998) or ecological constraints (e. g. habitat saturation; Emlen 1 982). The latter is the widely accepted explanation for delayed dispersal, although Hatchwell and Komdeur (2000) argue that the two life history traits and ecological constraints likely work in concert. For detailed reviews of cooperative breeding in birds see Hatchwell and Komdeur (2000) and Kimball et al. (2003). Benefits from cooperative breeding include indirect fitness to a helper of close genetic relatives, through the passing of their closely-related genes, or direct fitness, such as the acquired skills attained by helping another breeding pair, skills that could result in future increased reproductive success. The pair benefits from increased survivorship through additional help in defending the territory and increased prey delivery to the nest (Kimball et al. 2003). Cooperative breeding in diurnal raptors is fairly widespread, occurring in 22 of 76 genera and 42 of 304 species; in the Accipitidrae it occurs in 26% of genera and 36% of species. Group composition can consist of reproductive pairs with immature helpers only, immature or adult helpers, or solely adult helpers, with the sex of the helper being male in some taxa, female in others, or both. The trios may be stable over many years (Kimball et al. 2003). Cooperative breeding in the Bald Eagle (Haliaeetus leucocephalus), has been reported only rarely. In their review, Kimball et al. (2003) note four sources of cooperative breeding in Bald Eagles. I found four additional reports, for a total of eight instances of cooperative breeding in this species, none, other than this study, from Canada (Table 1 ). In the Bald Eagle, both polyandry and polygyny have been exhibited, although the latter appears to be the more common. This paper reports an instance of cooperative breeding by the Bald Eagle on Vancouver Island, British Columbia, and includes summary observations of the trio through two nesting seasons. Two other possible instances of cooperative breeding are discussed briefly. Methods On 201 4 April 2, I began observations, as time allowed, at a Bald Eagle nest (BAEA-1 05-1 87; Wildlife Tree Stewardship Program [WiTS] 201 5) where incubation had already begun. The nest was built in 2008 and was very likely occupied by the same eagles that formerly used nest BAEA-1 05-1 71 a nest that used to be in the primary perch of this study. That nest blew down during an autumn 2006 storm; it was sketchily rebuilt in 2007 but a nesting attempt failed (Wildlife Tree Stewardship Program 201 5). My observations continued through 201 5 to include the fledging of the young, the August departure of the eagles from the area, to the first two weeks after their return in October. Volume 26, 201 6 British Columbia Birds

36 Cooperative breeding Bald Eagles - Dawe Table 1. Reports of cooperative breeding by the Bald Eagle. Observations through Bausch and Lomb 8 x 42 binoculars and a Bausch and Lomb Elite telescope with 20 60 power eyepiece were recorded on a digital calendar app on a Nexus 7 tablet. Distances were approximated from Google Earth (201 2). Digital photographs of the eagles at the nest and on proximate perch trees were also taken. Sex of the eagles was noted, where possible, determined by the sexual size dimorphism between the females and male and by their behaviour (e. g. during copulation); however, I found it nearly impossible to determine the sex of birds in the nest or to distinguish between the females. Location The nest was located near the mouth of Parksville Bay (1 0 U 403 21 9 m E, 5465576 m N), about 1 3 0 m from the observation site. It was situated in a Class 4 (Fenger et al. 2006) Grand Fir (Abies grandis) that stood near the edge of a bluff about 20 m above the Salish Sea shoreline. A row of nearby Douglas-fir trees (Pseudotsuga menziesii; primarily Class 1 and 2) adj acent to the bluff, served as perches for the eagles. The tallest, their primary perch tree (Class 2 Douglas-fir; BAEA-1 05-1 71 ), was approximately 1 05 m southeast of the nest. A secondary perch tree (Class 4 Douglas-fir; BAEA-1 05-1 3 6), about 25 m east of the nest, was also frequently used. In addition, branches on the water side of all the taller Douglas-fir trees along the bluff edge were often used as perch sites. Results 2014 nesting season On April 1 2, I was observing the pair of eagles at the nest, the female in incubating posture, when a third adult, subsequently determined to be a female, landed in the primary perch tree. The pair in the nest ignored this third bird, which I thought was unusual behaviour. Nesting Bald Eagles are normally monogamous and territorial during the breeding season, usually defending the nesting area vigorously (Retfalvi 1 965, Mahaffy and Frenzel 1 987, Buehler 2000). Over the next two weeks I invariably saw one bird in the nest in incubating posture, either alone or sometimes with another adult present at the nest or with a male and a female nearby on the primary perch. There never were more than three adults within 1 50 m of the nest. On April 28, the activity and behaviour of the adults suggested that an egg or eggs had hatched. This would indicate that incubation began on or about March 24, based on an incubation period of 35 d (Buehler 2000). On April 30, I observed a female on the nest feeding a nestling and, for the first time since I began observations, the male and second female were on a branch adjacent to the nest. On May 7, I saw two eaglets in the nest. Subsequently, all three adults were fairly often seen at the nest tree concurrently, either in or beside the nest. Throughout the nestling period, all three adults defended the nest and territory, which included passive perching on the primary and secondary perches, threat vocalizing, and active territorial chases of intruding conspecifics. When an British Columbia Birds Volume 26, 201 6

Cooperative breeding Bald Eagles - Dawe 37 However, on November 1 5, three adults and an immature were perched in the general vicinity of the nest when another immature bird flew over the nest. Two of the adults immediately chased the young intruder, vocalizing all the while. The observations of the immature perching near the adults suggest the bird was likely related to the trio, although Beebe (1 974) is reasonably certain that most family units had broken up by this time. Nest maintenance activities began on October 1 9 when I saw an adult carrying a large branch to the nest. This activity continued regularly through November and December into 201 5 January. All three birds were involved in carrying materials (sticks, branches, green conifer boughs and sprigs) to the nest or placing and arranging materials in the nest. Over this period, the trio could often be seen together on the primary perch (Figure 2). Figure 1. Bald Eagle trio at nest vocalizing to warn off a sub-adult intruder. The nest held two young, 201 4 May 1 4. immature eagle or adult flew over, all three adults in the trio would vocalize until the intruder left the territory (Figure 1 ). Often, one or two individuals of the trio would quickly and purposefully leave the nest or perch tree to rout the intruder. The bird or birds that remained on or near the nest would vocalize loudly until the intruder left the area. Sub-adult intrusions appeared to be tolerated more than adult intrusions with sub-adults often getting away with simply a vocalized warning rather than being routed by a chase, as was more often the case with adult intruders. All three adults brought prey to the nest to feed the young, primarily fish often still alive but also birds. Where I could determine the sex of the adults at the nest, most of the feeding of the young appeared to be carried out by the females. An eaglet was first noticed mantling prey on July 4. One young fledged on July 1 3 and the second young fledged on July 20; the nestling period ranged from 76 83 d, suggesting the first nestling to fledge was a male and the second a female (Bortolotti 1 986). The adults and young left the nesting territory in early August (last observations: adults - August 6; young - August 1 2). I had no further eagle observations until October 5, when I saw an adult in the primary perch tree; the pair was on territory by October 1 5. On October 1 8, the adult helper returned. An immature bird was in the territory on October 27 and little concern was expressed by the three adults. 2015 nesting season Nest maintenance by all three adults continued through much of January. It wasn't until mid-february that I saw what I took to be a pair copulating, although it was difficult to determine positively through the early morning fog and drizzle. Near the end of February all three birds were spending noticeably more time beside or on the nest. My first certain observation of copulation came on March 1 2, just before incubation began. I also observed birds copulating on March 1 8, 1 9, 21, 22, during the incubation period, and once on April 30, during the nestling period. All copulations took place on the primary perch; in each case, the second female was on the nest. However, a photograph I took on March 1 8 suggests a copulation attempt may have taken place on the nest: the male appears to be on the back of the incubating female. Figure 2. Bald Eagle trio atop the primary perch, 201 4 October 31. Volume 26, 201 6 British Columbia Birds

38 Cooperative breeding Bald Eagles - Dawe Figure 3. Bald Eagle trio at nest shortly after a fish was delivered by one of the adults perched adjacent to the nest. The female in the nest is arranging nest material and had just finished feeding the young and herself, 201 5 May 6. of July 27, when it purposefully left the nest and flew to a group of firs about 80 m away. Excluding the premature fledging, the nestling period ranged from 98 1 00 d, which suggests the young was a female (Bortolotti 1 986). All the eagles had left the nesting area by the afternoon of August 1 4 and I had no further observations until October 3, when I saw the adult male on the primary perch shortly after noon; the adult female of the pair returned a few hours later. On October 5 both birds were at the nest, the female re-arranging branches while the male perched beside the nest. On October 7, the female was again in the nest re-arranging branches and placing new branches in position that the male brought to the nest. On October 1 0, the main trunk that was supporting the eagle nest broke in strong winds and the nest fell to the ground. On the afternoon of October 1 4, the helper returned and joined the pair on the primary perch and by October 1 5, all 3 adults were taking branches to what appeared to be their new nest location in the primary perch some 1 0 1 5 m below the top. Incubation began sometime between March 1 4 and 1 6 and a young had hatched between April 1 8 and 20. This year the trio produced only one young that I could see (Figure 3). The eagle's nesting behaviour through the incubation and nestling stages was similar to my observations in 201 4 with all three birds participating in territorial defence. Prey delivery appeared similar to that in 201 4, although this year I could not determine for certain that all three birds were participating. Most feeding of the young again appeared to be done by both females. In 201 5, the female helper was noticeably absent from the area later in the nestling period; I had no observations of her for over a month, between June 5 and July 29 and only once after that. The eaglet fledged prematurely on July 20, after just having fed on a fish that had been delivered to the nest. While flapping its wings during a strong wind, the young bird was lifted off the nest. As it rose, it grasped a branch on the nest edge but the branch gave way and the eaglet and branch were lifted up and over the nest edge where the eaglet then lost lift and fell through adjacent fir branches finally landing on a branch of the nest tree some 1 0 m below. I happened to be filming the young at the time and caught much of this on video. Immediately, both young and adults began vocalizing. The young stayed on the same branch for the next three days. In the early morning of July 23, for the first time in three days, I saw an adult perched beside the young bird, feeding it a fish. On July 24, the young left the branch it had been on for four days and moved to another, closer to the edge of the bank. Sometime later that day I saw the young back in the nest. It stayed in or near the nest until the early afternoon Discussion Circumstantial evidence suggests that this trio may have formed in 2004. A record from the WiTS database (Wildlife Tree Stewardship Program 201 5) indicates that on both June 7 and July 5 of that year, the observer reported an adult perched with an immature ( immature possibly 2003 ) in the secondary perch (BAEA-1 05-1 36), about 82 m from the original nest (BAEA-1 05-1 71 ). Retfalvi (1 965) discusses a situation where the adults tolerated intrusion by a juvenile into their territory; however, in two other territories he was studying, the adults chased subadults. Turrin and Watts (201 4) found that immature Bald Eagle intrusions were allowed closer to the nest than adult intrusions and elicited a response from the breeders only 47% of the time compared with 78% of the time with adult intruders. They suggest that the breeders' perceived threat may be different for adult versus immature intruders and their incentives for intrusion, which may allow them to tolerate immature intruders, especially when they are previous year's offspring. This could result in helping behaviour from the immature that arises as a result of ecological constraints such as habitat saturation and limited availability of quality territories in stable environments, or a lack of suitable mates. By 2009, evidence suggests the trio was using the current nest (BAEA-105-187): on April 14, the WiTS observer notes one adult incubating, calling a few times to other neighbouring eagle pair at their nest tree ~150m East. At about the same time for nest BAEA-105-171, the observer reported 2 adults together at top ofnest tree. Only a small pile ofsticks visible, no organized nest structure in this tree. Neighbour says 'no or very little nest building activity seen this year, this very active eagle pair still British Columbia Birds Volume 26, 201 6

Cooperative breeding Bald Eagles - Dawe 39 using nest tree and many other favoured perches daily.' This is precisely the behaviour I had often observed over the past two years during the incubation period: an incubating bird in the nest and the other two birds of the trio perched atop the old nest tree, now their primary perch. Considering the proximity of the two nests, it seems most likely they belonged to the same pair. Bald Eagles have high levels of fidelity to their nest sites (Buehler 2000), making it likely that this pair had been using the territory since at least 2002. Buehler (2000) notes that in the case of trios reported at nests, extra-pair copulations have not been observed but adds that trios are usually not reported until during or after incubation, thus extra-pair copulations would not likely be observed. However, even if they were observed, unless the researcher could distinguish between the helper and the female of the pair, it would be impossible to determine extrapair copulations. In 201 4, I witnessed one copulation after the young had hatched. In 201 5 I saw six actual and one possible copulations, one before incubation began, five during incubation, and one after the young had hatched. In none of those cases could I distinguish the female helper from the female of the pair. All three adults were observed defending the nesting territory, maintaining the nest, and delivering prey to the nest, which has also been documented in earlier studies (e. g. Hopkins et al. 1 993, 1 995; Nye 1 994; Garcelon et al. 1 995; Sharpe and Dooley 2001; Ortego et al. 2009) In other studies, all three birds in the trio also played a role in incubation, and brooding and feeding the young (Hopkins et al. 1993, 1995; Garcelon et al. 1995; Ortego et al. 2009) and that was likely the case with this trio, as well, but was not confirmed. Nest departure for the eaglets in 201 4 fell within the limits of the 8 1 4 weeks noted by Buehler (2000); in 201 5 departure was at the high end of the period and maybe slightly exceeding it. This extended nestling period could have been influenced by the young's premature departure from the nest. The young eagles stayed in the territory from 1 8 23 d after fledging and then left abruptly along with the adults. The birds were likely taking advantage of salmon-spawning runs along the coast as far north as Alaska or those in the Interior of the province (Beebe 1 974, Servheen and English 1 979, Hunt et al. 1 992). The adults were away from the territory a little over 50 days, returning in early October in both years. In 201 4, nest maintenance activities began two weeks after the pair were back on territory; in 201 5, nest maintenance began four days after their autumn arrival. This is about five months prior to egg-laying, longer than the generally... 1 3 months noted in Buehler (2000). There have been at least two other instances of what appear to be cooperative breeding by Bald Eagles in British Columbia; however, details are sketchy: Guy Monty (Pers. comm., Nanoose Bay, B.C.) found a trio tending a nest on the Nanaimo River estuary. Monty notes, The nest on Oak Island at the Nanaimo River Estuary had three adults tending it for at least two years, about 1 0 years ago. Unfortunately, additional details have been lost and there is no mention of the trio in the WiTS database (BAEA-1 05-031 ). A second report, suggestive of cooperative breeding by Bald Eagles comes from Gabriola Island through the British Columbia Wildlife Tree Stewardship Atlas (Wildlife Tree Stewardship Program 201 5). In 2006, a monitor for nest BAEA-1 01-002 reported the nest was occupied by three adults that fledged at least one chick. In 2007, for the same nest, the following was reported: "Neighbour assured me of fledged chicks. It appeared to her that three adults were occupying nest. I contacted the neighbour, Arlene Catling, in 201 4 for more details and she responded that For 3 or 4 years there hasn t been any activity. Before that, for four or five years, there were three eagles that tended the nest two mature adults and one immature. The immature took his/her turn sitting on the eggs but if memory serves me correct he/she did not sit on the chicks but sat perched near the nest when he/she was there the two mature eagles sat all of the time (Pers. comm., Arlene Catling, Gabriola Island, B.C. ). Because of the discrepancy between the ages of the third eagle reported in the atlas and Catling's response, I again contacted Catling to verify which was correct. She replied, I'm absolutely positive it was two mature and one immature [Bald Eagles] (Pers. comm., Arlene Catling). Further details were unavailable. Cooperative breeding by Bald Eagles may be more common than has been previously reported. This particular territory was likely occupied by the trio since 2004 and had been surveyed most years, through to 201 3; however, the trio tending this nest had not been reported. This is not surprising. A high density of nesting eagles in the area could make three or more birds in proximity not an unusual event causing the true nature to be overlooked; e.g. there are eight Bald Eagle nests that have been active within a radius of 3 km at one time or another over the period this nest has been under observation (Wildlife Tree Stewardship Program 201 5). Since the Bald Eagle is monogamous, observers seeing additional birds within a territory could view them simply as transients (Kimball et al. 2003). In addition, the amount of time all three birds can be seen on or at the nest together is small. Such a relatively rare occurrence coinciding with the random arrival of an occasional observer to the nest site further decreases the odds of recording such an event. Acknowledgements A number of people helped me in the preparation of this paper. Jess Thompson, Cottonwood Photography, and Dale Schmidt and Brent Ortego, Texas Parks & Wildlife Volume 26, 201 6 British Columbia Birds

40 Cooperative breeding Bald Eagles - Dawe Department, provided information about the Texas nesting trio. Jeff Williams, Alaska Maritime National Wildlife Refuge, Aleutian Islands Unit, provided a reference to the Amchitka Island nesting trios. Peter Sharpe, Institute for Wildlife Studies in California, provided information on the Santa Catalina nesting trio. Renate Sutherland provided some of the observation data for the Parksville nesting trio. Gavin Hanke, Royal British Columbia Museum, provided identifications for some of the fish prey items captured in my photographs. The following people were associated with the British Columbia Wildlife Tree Stewardship Atlas: Ian Moul provided me with initial information about the Gabriola Island nest and Darlene Mace directed me to the nest monitor, Arlene Catling, who provided additional details of that nesting trio; Sandra Gray, who has recorded nest success for this pair since 2002, provided past history details about the Parksville nests and answered a number of my questions. I thank them all. Literature cited Arnold, K.E., and I.P.F. Owens. 1998. Cooperative breeding in birds: a comparative test of the life history hypothesis. Proceedings ofthe Royal Society oflondon B 265:739 745. Beebe, F.L. 1 974. Field studies of the Falconiformes of British Columbia: Vultures, eagles, hawks and falcons. Occasional Paper No. 1 7, B.C. Provincial Museum, Victoria. Bortolotti, G.R. 1 986. Influence of sibling competition on nestling sex ratios of sexually dimorphic birds. The American Naturalist 1 27:495 507. Buehler, D.A. 2000. Bald Eagle (Haliaeetus leucocephalus). The Birds of North America Online. (A. Poole, ed.). Cornell Lab of Ornithology, Ithaca, N.Y. <http://bna. birds. cornell. edu/bna/species/506> [201 4 November 25]. Emlen, S.T. 1 982. The evolution of helping. I. An ecological constraints model. The American Naturalist 11 9:29 39. Fenger, M., T. Manning, J. Cooper, S. Guy, and P. Bradford. 2006. Wildlife & trees in British Columbia. Lone Pine, Edmonton. Fraser, J.D., L.D. Frenzel, J.E. Mathisen, and M.E. Shough. 1 983. Three adult Bald Eagles at an active nest. Journal ofraptor Research 1 7:29 30. Garcelon, D.K., G.L. Slater, C.D. Danilson, and R.C. Helm. 1 995. Cooperative nesting by a trio of Bald Eagles. Journal ofraptor Research 29:21 0 21 3. Google Earth. 2012. Google Earth. <http://www.google.- com/earth/index.html>. [2012 February 6]. Hatchwell, B.J., and J. Komdeur. 2000. Ecological constraints, life history traits and the evolution of cooperative breeding. 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Occurrence and evolution of cooeprative breeding among the diurnal raptors (Accipitridae and Falconidae). The Auk 1 20:71 7. Mahaffy, M.S., and L.D. Frenzel. 1 987. Elicited territorial responses of northern Bald Eagles near active nests. Journal ofwildlife Management 51 :551 554. Nye, P.E. 1 994. Polygamy in Bald Eagles. p. 63 in Abstracts of presentations made at the annual meeting of the Raptor Research Foundation, Inc., held at Charlotte, North Carolina, on 3 7 November 1 993. Journal ofraptor Research 28:45 71. Ortego, B., C. Gregory, D. Mabie, M. Mitchell, and D. Schmidt. 2009. Texas Bald Eagles. Bulletin of the Texas Ornithological Society 42:1 1 7. Retfalvi, L. 1 965. Breeding behavior and feeding habits of the Bald Eagle (Haliaeetus leucocephalus L.) on San Juan Island, Washington. MF Thesis, University of British Columbia, Vancouver, B.C. Servheen, C., and W. English. 1 979. Movements of rehabilitated Bald Eagles and proposed seasonal movement patterns of Bald Eagles in the Pacific Northwest. Journal ofraptor Research 1 3:79 88. Sharpe, P.B., and J. Dooley. 2001. Restoration and Management of Bald Eagles on Santa Catalina Island, California, 2001. U.S. Fish and Wildlife Service, Sacramento, Calif. <http://www.globalraptors.org/grin/researchers/uploads/464/catalina_bald_eagle_report_2001 %5B1%5D.pdf>. [201 4 April 30]. Sherrod, S.K., C.M. White, and F.S.L. Williamson. 1 976. Biology of the Bald Eagle on Amchitka Island, Alaska. Living Bird 1 5:1 43 1 82. Thompson, J. 201 4. Cottonwood Photography. <http://cottonwoodphoto.photoshelter.com/>. [201 4 November 7]. Turrin, C., and B.D. Watts. 201 4. Intraspecific Intrusion at Bald Eagle Nests. Ardea 1 02:71 78. Wildlife Tree Stewardship Program. 201 5. Nest Tree Report. The Community Mapping Network. <http://www.cmnmaps.ca/wits/>. [201 5 March 1 8]. British Columbia Birds Volume 26, 201 6